Society for the Study of Amphibians and Reptiles

Phylogeny of Gekko from the Northern Philippines, and Description of a New Species from
Calayan Island
Author(s): Rafe M. Brown, Carl Oliveros, Cameron D. Siler and Arvin C. Diesmos
Source: Journal of Herpetology, Vol. 43, No. 4 (Dec., 2009), pp. 620-635
Published by: Society for the Study of Amphibians and Reptiles
Stable URL: http://www.jstor.org/stable/25599264
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JournalofHerpetology,Vol. 43, No. 4, pp. 620-635, 2009
Copyright 2009 Society for the Study of Amphibians and Reptiles

Phylogeny of Gekko from the Northern Philippines, and Description of

a New Species from Calayan Island

Rafe M. Brown,1,2 Carl Oliveros,1,3 Cameron D. Siler,1,4 and Arvin C. Diesmos5

Natural History Museum and Biodiversity Institute, Department of Ecology and Evolutionary Biology, University of
Kansas, Lawrence, Kansas 66045-7561 USA
3ISLA Biodiversity Conservation; 9 Bougainvillea St, Manuela Subdivision, Las Pinas City, Philippines 1741; E-mail:
carl_oliveros@yahoo. com
5National Museum of the Philippines, Rizal Park, Padre Burgos Avenue, Ermita 1000, Manila, Philippines; E-mail:

[email protected]

Abstract.?We use mitochondrial gene sequences to estimate relationships among Gekko populations
from the northern Philippines. These data, plus morphological and biogeographical evidence, suggest that
the Babuyan and Batanes island groups (north of Luzon Island) are inhabited by a minimum of six distinct

evolutionary lineages, only two of which

(Gekko porosus Taylor from the Batanes and Gekko crombota from
Claro) have been formally as distinct species. In this paper, we provide a description of
Babuyan recognized
another new species, the endemic Gekko from Calayan Island. This isolated species is
geographically
diagnosed on the basis of a distinct color pattern, body size, scalation, and significant divergence in
mitochondrial gene sequences. The new species has been found on rocky outcrops and limestone caves in
forested areas and on trunks of mature forest trees <500 m above sea level, away from the island's coast.
Given the history of geological isolation of Calayan Island and the distinctiveness of the endemic Gekko

populations there, we are confident in diagnosing this gecko as a unique evolutionary lineage; it is unlikely
that this species will be found on neighboring land masses. The remaining major islands of the Babuyans

group (Camiguin Norte, Fuga, and Dalupiri islands) each contain similarly distinct endemic species that
await description; additional surveys throughout the Batanes and Babuyan islands will be necessary to arrive
at an estimate of total species diversity for this isolated gekkonid radiation.

The Philippine gekkonid lizard fauna consists specimens matching its description
or the
of 10 genera and 40 described species: Cyrto holotype have been collected since Taylor,
dactylus (5 species); Gekko (10 or 11); Gehyra (1); suggesting
erroneous attribution to the archi
a (Brown and Alcala, 1978; Ota et al.,
Hemidactylus (5; including platyurus, species pelago
formerly assigned to Cosymbotus); Hemiphyllo 1989).
of Gekko are consid
dactylus (1); Lepidodactylus (6); Luperosaurus (7); Eight species confidently
Pseudogekko (4); and Ptychozoon (1) (Taylor, ered endemic to the Philippines; these include
1922a,b; Brown and Alcala, 1978; Brown et al., Gekko athymus, Gekko gigante, Gekko mindorensis,
1997, 2007, 2008, in press; Brown, 1999; Gaulke Gekko palawanensis, Gekko porosus, Gekko romblon
et al., 2007; Welton et al., in
press). (Taylor, 1922a; Brown and Alcala, 1978), Gekko
Philippine species of the genus Gekko include
ernstkelleri (Roesler et al., 2006), and Gekko

three taxa shared with countries crombota (Brown et al., 2008). All endemic
neighboring
(Gekko gecko, Gekko monarchus, and G. hokouensis; Philippine Gekko share (1) moderate body size
and longer, slender limbs; (2) near complete
Taylor, 1922a,b; Brown and Alcala, 1978; Ota et or
absence of cutaneous
al., 1989). Although G. gecko and G. monarchus interdigital webbing
are common in collections, Gekko hokouensis has body expansions; (3) enlarged dorsal tubercles
been represented
in the
country's gekkonid arranged in longitudinal rows on the dorsum
fauna by only a single specimen originally (except G. athymus,dorsal tubercles absent); (4)
scales of dorsum between tubercle rows minute,
described as amissus
by Taylor Luperosaurus
nonimbricate; (5) scales of venter enlarged, flat,
(Taylor, 1922a,b), later considered to be Gekko
imbricate; (6) differentiated postmentals elon
japonicus (Brown and Alcala, 1978) and eventu and and (7) subcaudals
ally determined to be G. hokouensis (Ota et al.,
gate slender; enlarged,
(Brown and Alcala, 1978; Brown et al.,
The was collected platelike
1989). specimen apparently
2007, 2008).
from Tablas Island (Taylor, 1962), and no recent inventories of the
During biological
Babuyan Islands (Fig. 1), we obtained new
2 and for a study of
Author. E-mail: [email protected] specimens tissues, allowing
4Corresponding
E-mail: [email protected] the genetic variation exhibited by isolated

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 PHYLOGENY AND NEW SPECIES OF NORTHERN PHILIPPINE GEKKO 621

_> ,.;-1-TT^
12r larged cloacal spurs in males (both absent in
N Itbayat
/a females; Brown, 1999; Brown et al., 1997, 2007).
gff T &*?DR>go Gonadal was whenever
Jy inspection performed
possible. Measurements (to the nearest 0.1 mm)
were taken with digital calipers following
?^L A? X? Batan character definitions by Ota and Crombie
,bah08'&
Sabtang
(1989), Brown et al. (1997, 2007, 2008) and
Brown (1999). Character abbreviations include:

-lo^^ft
snout-vent length (SVL); tail length (TL); tail
& /Tp width (TW); tail depth (TD); head length (HL);
head width (HW); head depth (HD); snout
M?-""i22' i26?| Babuyan
length (SNL); eye diameter (ED); eye-narial
distance (END); auricular opening diameter
(AO); internarial distance (IND); interorbital
distance (IOD); axilla-groin distance (AGD);
aK?l ^Catayan femur length (FL); tibia length (TBL); Toe I
>1200|
C (l\Dalupiri Camiguin length (TIL); Toe IV length (TIVL); number of
supralabials (SUL); infralabials (IFL; counted
both to the center of the eye and posteriorly to
the point at which labials were no longer
Fuga
differentiated); enlarged circumorbitals dor
Palaui soanterior to orbit (CO); modified spiny circu
r\ y^-x morbitals (cilaria) dorsoposterior to orbit; dif
ferentiated precloacal pore-bearing scales (PS);
femoral pore-bearing scales (FPS); differen
/^^^^^^^^B Luzon
>>^^f) tiated subdigital scansors on Fingers I-V
(FS I-V); subdigital scansors on Toes I-V (TS
I-V); ventral transverse scale rows
midbody
Fig. 1. Map of the most northern islands of the (MBVS); midbody dorsal transverse scale rows
Philippines (inset) showing the type locality of Gekko
(MBDS); midbody transverse tubercle rows
rossi on Calayan Island circle) in relation to
(shaded (MBTR); paravertebrals in AGD (PVS; counted
other landmasses discussed in the text. Additional
northern Philippine include Gekko porosus along middorsum between limb insertions);
lineages ventrals (VS; counted midventrally between
(A), Gekko crombota (B), and three undescribed species
on
limb insertion); tail annuli (TA); and subcaudals
Dalupiri, Fuga, and Camiguin Norte islands (C-E).
(SC).
Molecular from 21
Data.?Samples specimens
representingnine species of Gekkowere included
northern Philippine Gekko populations. In this inphylogenetic analyses. These include fiveof the
paper, we present a
phylogenetic analysis of the northern Philippine species, G. monarchus (a
northern Philippine Gekko and a description of a species sharedwith Sulawesi Island of Indonesia;
Appendices 1 and 2) and outgroup species G.
new from Island.
species Calayan
gecko and Gekko smithi.For all 21 samples, we
Materials and Methods sequenced themitochondrial gene NADH Dehy
drogenase Subunit 2 (ND2) and components of
Specimens and tissues were collected by the four flanking transfer RNA genes (tRNAmet,
authors between 2004 and 2008. Specimens tRNA^, tRNA*1*, tRNAasn). All sequences were
were initiallypreserved in 10% buffered forma in GenBank 2).
deposited (Appendix
lin and then transferred to 70% ethanol 1-3 Genomic DNA was extracted from liver
months later. Tissues were
preserved
in 95% tissue following the guanidine thiocyanate
ethanol. Voucher specimens
are
deposited
in method of Esselstyn et al. (2008). The external
U.S. and Philippine museum collections (insti primers Metf6 (5'-AAGCTTTCGGGCCCA
tutional abbreviations follow Leviton et al.,
TACC-3') and COIH (5'-AGRGTGCCAATG
1985). TCTTTGTGRTT-3') (Macey et al., 1999) were
Morphological Data.?We (CO, CDS, and used to amplify the target fragment using the
RMB) collected data from fluid-preserved spec polymerase chain reaction (PCR). Cycle se
imens (Appendix 1). Sex was determined by quencing reactions were
performed with com
eversion of hemipenes
during preservation binations of nested and internal primers de
(males) and confirmed by noting prominent signed for this study to collect complete double
sexual characteristics such as the stranded We used two nested
secondary sequences.
presence of precloacofemoral pores and en
primers, GekND2.NestFl (5'-CCATACCCCGA

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 622 R. M. BROWN ET AL.

CAATGTTGGWAC-3') and GekND2.NestRl used to find appropriate models of evolution for

(5'-AGCTGTAGACTYATRTACGGRGG-3'), and our data. The best-fitmodel for each of the
two internal GekND2.IntFl (5'-TCW seven was the general time reversible
primers, partitions
TAGCYTWYTCATCAATYGC-3') and GekND2. (GTR) model, with a proportion of invariable
IntR4 (5'-GATGARWARGCTAWGATTTTTCG sites (I) and a
parameter for variation in rates
GG-3'). among sites (r). Partitioned ML were
analyses
We used the following thermal cycler profile: then run under the same model (GTR + I + T)
4 min at 94?, followed by 35 cycles of 94? for with RAxMLHPC v7.0, with 100 replicate best
30 sec, 52-53? for 30 sec, and 72? for 1 min tree inferences. Each inference was
performed
30 sec, and a final extension at 72? for with the options "-d" to start each search with
phase
7 min. were visualized on a random tree, and "-f d" to run each
Amplified products starting
analysis with the rapid hill-climbing algorithm
1.5% agarose Clean bands of the
gels. single
targetproduct were purified with 1 |iL of a 20% implemented
in the new version of RAxML.
diluted solution of ExoSAP-IT (US78201, Amer Nodal support was assessed with 1,000 boot
sham Biosciences, NJ) on the
Piscataway, strap pseudoreplicates.
following thermal cycler profile: 31 min at 37?, Taxonomic Decisions.?We adopted the Gen
followed by 15min at 80?. Cycle sequencing eral Lineage Species Concept (GLC) of de
reactions were run using ABI Prism BigDye Queiroz (1998, 1999) as the natural extension
Terminator chemistry (Vers. 3.1; Applied Bio of the Evolutionary Species Concept (Simpson,
systems, Foster City, CA), and purified with 1961; Wiley, 1978; Frost and Hillis, 1990).
Sephadex Medium (NC9406038, Amersham Application of lineage-based species concepts
Biosciences, NJ) in 96 to recognition of island taxa is straightforward
Piscataway, Centri-Sep
spin plates (CS-961, Princeton Separations, because of the high probability of a history of
Princeton, NJ). Purified was
product analyzed isolation (Brown et al., 2000, 2008; Brown and
with an ABI Prism 3130x1 Genetic Analyzer
Diesmos, 2002; Brown and Guttman, 2002) of
(Applied Biosystems). Gene sequence contigs these distinct lineages (McDermott et al., 1993;
were assembled and initially edited using et
Yang al., 1996; Marini et al., 2005). We
4.8 (Gene Codes Ann Arbor, as new
Sequencher Corp., consider and
species morphologically
MI).
genetically diagnosable allopatric populations
Alignment and PhylogeneticAnalysis.?An ini forwhich the hypothesis of a distinct evolu
tial alignment was produced inMuscle (Edgar, cannot be
tionary lineage rejected.
2004), and manual were made in
adjustments
MacClade 4.08 (D. R. Maddison and W. P.
Maddison, MacClade: Analysis of Phylogeny
Results
and Character Evolution, Sinauer, Sunderland,
were con Phylogeny.?The aligned dataset contained
MA, 2005). Phylogenetic analyses
ducted using both parsimony and likelihood 1,282 total characters, of which 659 were

criteria. were variable, and 503 were parsimony-informative.

optimality Parsimony analyses
conducted in PAUP* 4.0 L. The maximum recovered a
(D. Swofford, parsimony analysis
tree =
PAUP*. Phylogenetic Analysis Using Parsimo single most parsimonious (length 1,114).
The 100 inferences from the maximum likeli
ny [*and Other Methods], Sinauer, Sunderland,
2002) with gaps treated as missing data hood showed an likelihood
MA, analysis average
and all characters Most score of -In L 6077.327465, with a tree
weighted equally. single
parsimonious
trees were estimated using heu having the highest likelihood score of -In L
ristic searches with 1,000 random addition 6077.326765 (inferred by RAxML). The trees
and tree bisection and recovered from the two methods had a
sequence replicates nearly
reconnection (TBR) branch swapping. To assess identical topology (Fig. 2), differing only by the
nodal support, nonparametric bootstrapping position of short terminal branches within
was conducted 1,000 bootstrap Most Batanes and
using replicates, species. Babuyans popula
each with 100 random tions are substantial
addition-sequence repli separated by genetic
cates and TBR branch divergences (9.3-14.6% uncorrected pairwise
swapping.
Partitioned maximum likelihood (ML) analy genetic sequence divergence; only 2.6% differ
ses were conducted in RAxMLHPC v7.0 (Sta ence between Dalupiri and Camiguin Norte
matakis, 2006). The data setwas partitioned by lineages), and no instances of multiple sympat
codon for the protein-coding of ric lineages island were detected (5-10
position region per
ND2, and each of the four flanking tRNAs individuals per island were
sequenced,
but

(tRNAmet, tRNA^, tRNAala, tRNAasn) were only 2-3 are included in the presented analysis
as The Akaike for computational following confir
analyzed separate partitions. simplicity,
Information Criterion (AIC) as implemented in mation that thesewere genetically identical). All
Modeltest v3.7 (Posada and Crandall, 1998) was relationships were highly supported with the

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 PHYLOGENY AND NEW SPECIES OF NORTHERN PHILIPPINE GEKKO 623

-Gekko smithi
-Gekko gecko

100/1001"" Gekko monarchus

_ L Sulawesi Island
100/100

' Gekko mindorensis

t00/100
U MindoroIsland
100/100]

100/98 wk*v sd
100/100 i- Gekko ?p. Ar\
-***-
Dalupiri Island

100/100

U Gekko sp. B
99/1
100| Camiguin Norte Island
_100/100

loo/loo Gekko crombota

I Babuyan Claro Island

91/63 P
loo/loo Gekko porosus
I Batanes Isl.Group
100/100

o.i lop/loo Gekko rossi, n. sp.

subst/site Island
Calayan

Fig. 2. The preferred topology, inferred in both partitioned likelihood and equally weighted parsimony
analyses of ND2 sequence data (1282 nucleotide positions). Numbers to nodes are likelihood/
adjacent
parsimony bootstrap support values.

exception ofmoderate support (91ML/63 MP) Taxonomy

inferred for the between G. crom
relationship Gekko rossi sp. nov.
bota (Babuyan Claro Island) and the sister
Figures 3-5
couplet consisting of Gekko porosus
species
Gekkomonarchus Part (Calayan Island) Stejne
(Batan Island) and Gekko rossi (Calayan Island).
The southern Babuyans lineages (Dalupiri and ger, L. 1907. Proc. U.S. Nat. Mus. 33:545-546

Norte Islands) form a Holotype.?PNM 9543 (Field number RMB

Camiguin strongly sup
ported clade, to the exclusion of the lineages 5998; formerly KU 304877), an adult male
from the Northern Batanes and collected at 2245 h 2 m from the ground on the
Babuyans
trunkof a large (1.2m dbh) treeby RMB at an area
(Calayan and Babuyan Claro). Other Gekko
species Gekkomindorensis (Mindoro Island) and
known as "Macarra,"
locally Barangay Magsidel,
G. monarchus here with Municipality of Province,
(represented samples Calayan, Cagayan
from Sulawesi Island) cluster
together
and are Calayan Island, Philippines (19.294?N, 121.409?E;
sister to the northern Philippine lineages to the 245 m above sea level), on 15March 2006.
exclusion of outgroups G.
gecko
and G. smithi Paratopotypes.?KU 304919, 304927, 304931,
(Fig. 2). In summary,
genetic data support the 304885, and PNM 9538, 9540 (formerly KU
recognition of theCalayan Island population as 304935, 304938) adult males; KU 304916
a distinct and endemic lineage that is 304918, 304923-304924, PNM 9542, 9539, 9537
highly
divergent from other described Babuyan and (formerlyKU 304934, 304936, 304937), gravid
Batanes island
populations (G. porosus and adult females; PNM 9541 (formerlyKU 304939)
Gekko crombota). and KU 304876, juveniles of undetermined sex,

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 624 R. M. BROWN ET AL.

Fig. 3. (A-C) Adult male Gekko rossi holotype in life (PNM 9543; SVL = 98.4 mm); (D) Gekko crombota from
Claro Island (KU 304849); and (E) Gekko porosus from Batan Island (specimen not collected).
Babuyan

collected by RMB, C. Oliveros, and J.Fernandez; for females); (2) dorsum brown with six diffuse
15-17 March 2006, same locality,microhabitat, transverse black bars adjacent
to six transverse
and circumstances of capture (tree trunks, < series of two or three cream spots; (3) high
3 m; 1830-2330 h) as holotype. numbers of dorsal body scales (125-170 trans
Other Paratype.?PNM 9091 (Field No. MGDP verse midbody scales; 251-281 paravertebrals);
129), adult male; collected by C. Oliveros and M. (4) high number of sharply conical dorsal body
Pedregosa on large limestone boulders in tubercle rows (16-18 midbody; 31-37 paraver
second growth forest, 12 May 2004 at Sitio tebrally); (5) 77-88 enlarged precloacal-femorals
Longog, Municipality of Calayan, Cagayan arranged
in a continuous, uninterrupted series

Province, Calayan Island. (pore bearing in males; lacking pores in

Diagnosis.?Gekko rossi differs from all other females).
species of Philippine Gekko (i.e., G. athymus,G. Comparison with Similar Species.?Gekko rossi
crombota, G. ernstkelleri, G. gecko, G. gigante, G. differs from its phenotypically most similar
mindorensis, G. monarchus, G. palawanensis, G. Philippine congener, Gekko porosus (Batan and
porosus, and G. romblon) by the following Itbayat islands, of the Batanes Island group,
combination of characters (1) larger body size Fig. 1) by having dorsum brown with six
(SVL 95.5-108.2 mm for adult males; 86.8-100.0 diffuse transverse black bars adjacent to six

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 PHYLOGENY AND NEW SPECIES OF NORTHERN PHILIPPINE GEKKO 625

Fig. 5. (A) Ventral view of left hand and (B) left

side of precloacal-femoral pore-bearing scale series of
holotype of Gekko rossi holotype, hemipenes excluded
for simplicity (male PNM 9543). Scale bars = 5 mm.

three rows of cream

longitudinal paravertebral
spots adjacent to diffuse transverse black bars
(vs. presence of trilobed cream bars) on the
body trunk; a tendency toward higher midbody
dorsal scale counts (125-170 vs. 107-132); a
Fig. 4. Lateral (A), dorsal (B), and ventral (C) greater number (251-281) of paravertebral
scalation of head of holotype of Gekko rossi (male PNM scales (vs. 192-226); fewer (16-18) midbody
= 5 mm.
9543). Scale bar dorsal tubercle rows (vs. 18-22); precloacal
femorals in a continuous series (vs. 1
arranged
transverse series of three cream vs. or 2 scale between and
spots (Fig. 3; separation precloacals
dark transverse bands and circular vertebral femorals; Brown et al., 2008:fig 4B); and a
blotches in G. porosus, light spots absent), a greater number (77-88) of precloacal-femoral
number (125-170) of transverse mid scales(vs. 58-74).
greater pore-bearing
dorsal scales (vs. 88-103), a The new differs from G. monarchus, G.
body greater species
number (251-281) of paravertebral scales (vs. mindorensis, G. romblon, G. ernstkelleri, and G.
173-191), the presence of sharply protuberant paiawanensis by larger male body size (95.5
(vs. merely convex or
slightly raised) dorsal 108.2mm SVL; Table 1); the only remaining
tubercles, a
greater number (31-37) of paraver
Philippine species overlapping the body size of
tebral tubercle rows (vs. 17-24), and by the G. rossi are G. gecko (SVL 120.0-153.8 mm), G,
absence of a modified distal femoral pore gigante (89.7-104.7), and G. athymus (99.2-119.9).
bearing patch (vs. present, composed of a short The number of precloacals distinguished G. rossi
series of 2 or 3 rows of pore-bearing scales from G. monarchus, G. mindorensis, G. romblon, G.
[Brown et al., 2008:fig. 4C]). Finally, although gigante,G. ernstkelleri, G. paiawanensis, G. athy

body
sizes of these two
species overlap, adult mus, and G. gecko (Table 1). The absence of
males of G. rossi (SVL 95.5-108.2 mm) tend tobe separated precloacal and femoral pore-bearing
larger than G. porosus (91.0-96.7). scales is shared (Table 1) with G. porosus, G.
The new species differs from themorpholog monarchus, G. mindorensis, G. ernstkelleri, G.

ically similar G. crombota (Brown et al., 2008) by athymus, and G. gecko. The new
species has

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 626 R. M. BROWN ET AL.

fewer Toe IV scansors than G. gigante, G. Total number of differentiated supralabials

ernstkelleri, G. palawanensis, and G. athymus. 13/14 (L/R; 10/11 to center of eye), bordered
The new species has fewer (33-41) .midbody dorsally by one row of slightly differentiated
ventrais than G. gigante (41-50) and G. ernstkel snout scales; total number of differentiated
leri (42-48), more midbody dorsals than G. infralabials 12/12 (9/9 to center of eye),
monarchus (96-112), G. mindorensis (102-125), G. bordered
ventrally by
one row of
enlarged
romblon (102-108), G. palawanensis (114-121), scales and four rows of only slightly differen
and G. (92-104). The new has tiated chin scales; mental mental and
athymus species triangular;
thehighest paravertebral count (251-281) of any first five infralabials greatly enlarged and
Philippine Gekko (Table 1) and more ventrais wrapping
onto ventral surfaces of chin, at least
than all
Philippine species except G. crombota twice the size of individual infralabials 6-13;
(Table 1). Additionally, the new species has mental followed by
a
pair of slender, elongate
more tubercle rows in the axilla bordered
paravertebral postmentals; postmentals posterolat
groin region than all Philippine species except erally by
a
secondary pair,
more than one-half
G. crombota (Table 1) and is further distin the length of firstpair, and a tertiarypair of
from G.
guished athymus by the presence (vs. nonelongate, hexagonal lateral postmentals,
not
absence) of dorsal tubercles. more than the of primary
body Finally, one-quarter length
dorsal coloration (dorsum brown with six bordered
postmentals; postmental posteriorly
diffuse transverse black bars adjacent
to six by 2 or 3 scale rows of slightly enlarged,
transverse series of two or three cream spots) irregular scales; followed immediately by a
distinguishes G. rossi from the highly variable sharp transition to nondifferentiated chin and
range of patterns exhibited by other Philippine gular scales; postrictal scales slightly enlarged, 2
Gekko. These and other differences are summa or 3 times the size of gular scales; remainder of
rized in Table 1. undifferentiated gular scales very small, round,
Description of Holotype.?Adult male (PNM juxtaposed (Fig. 4C).
9543, formerlyKU 304877; fieldNo. RMB 5998; Dorsal cephalic scales highly heterogeneous
Figs. 3-5). SVL 98.4 mm; habitus robust, limbs and varied in and
shape disposition; scales of
well developed, relatively slender (but with rostrum somewhatenlarged, round, oval to
femoral segments of hind limbs hypertrophied); subrectangular,
and convex; postnasal, prefron
TBL 16.7% SVL, 72.6% FL; tail relatively long; tal, and interorbital depressions with
noticeably
margins of limbs smooth, cutaneous smaller scales; scales
lacking palpebral heterogeneous,
flaps or dermal folds; a thin adipose line with some scales as small as
adjacent interor
ventrolateral bital and others as as
(cutaneous fold) running along region large and raised

margin of trunk. rostral scales; undifferentiated temporal region

Head large, characterized by lightly hyper scales granular, flat to
irregularly convex,
and adductor musculature, in size with
trophied temporal reducing posteriorly, interspersed
as wide as at widest snout numerous and
body point; highly enlarged, protuberant
rounded at in dorsal and conical tubercles; nuchal with small,
subtriangular, tip region
lateral aspect (Fig. 4A, B, C); HW 69.2% HL and juxtaposed, flat trunk scales interspersed with
18.3% SVL; HL, 26.4% SVL; SNL 59.4% HW and enlarged sharply conical body tubercles; throat
41.2% of HL; dorsal surfaces of head relatively and chin scales small, juxtaposed; gular and
smooth, with concave with imbri
pronounced postnasal, pectoral regions enlarged cycloid,
interoribital, and cate scales, ven
prefrontal, parietal depres increasing posteriorly through
sions; auricular oriented ter, becoming and
opening large, round, very enlarged strongly
slightly lateroposteriorly frombeneath temporal imbricate.

swellings on either side of head; tympanum Ornamental occiput scalation includes nu

very deeply sunken; orbits large, bordered by merous conical tubercles on

posterolateral por
slightly distinct supraorbital crests; eye large, tions of head (temporal, supratympanic, and

pupil vertical, itsmargin wavy (Fig. 3A); AO postrictal regions; Figs. 3, 4) and a short curved
59.6% ED, IOD 23.8% HW. series of 4-5 enlarged, bluntly conical preorbital
Rostral large, subrectangular, nearly
twice as scales (Figs. 3B, 4A, B); 45/41 total circumorbi
broad as high, with two dorsomedial depres tals, differentiated into the following distinct
sions between raised posterodorsal projections regions: (1) 17/15 minute precircumorbitals, (2)
that form the anterolateral^ projecting edge of 7/8 enlarged, flat, squarish circumoribtals dor
the nares and suture with the soanterior to orbit, (3) 9/8 transversely
anteriorly elongat
the across
supranasals; nostril surrounded by rostral, ed fringe-like spiny ciliaria dorsoposter
first labial, and an
enlarged, round, convex iormargin of orbit, gradually reducing to (4)
supranasal,
an
enlarged posterosupranasal,
and 12/10 minute postcircumorbitals; a total of 45
two small
postnasals; supranasals separated
on interorbital scales (straight line distance from
a center across
either side by single large internasal. of each eye, both eyelids).

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 ^1

conical
toto
conical 2

6.7 monarchus56.2-80.7 romblon

62.7-89.2 gigante
89.7-104.7 82.0-92.1ernstkelleri 57.2-65.7 palawanensis
99.2-119.9 athymus
ft
m;f 4m;
9f If(juv) 3m 2m;
If 13f
4m; 6f
22m; 5m;
3f 4m;
6f 3m;
5f 3m;2f C
91.0-96.7 40.6-69.7 mindorensis J
light
dark
Vertebral
indistinct
tri-
thin
dark
paired
white
+
55.0-88.2
58.6-72.5 79.7-87.9 78.0-88.0 44.5-61.8 88.2-117.1 W
dark spot rows blotches spots spots
?c
Preanaland
+(28%)
+(39%)
<+(33%)
+(75%)
- - -
68.2-70.9 bands

diagnostic
Gekko
Distribution
1.
Philippine
Table
selected
of
in
the
characters
and
(exclusive
Gekko
rossi
other
of
gekko
hokouensis
widespread
of
and
species
nt protuberant
[a
Japanese
included
Philippine
probably
in
the
species
erroneously convex
herpetofauna]).
(with
presented
Measurements
millimeters
in
of
all
the
and
specimens
exception
are coloration
transverse
lobed
circular
transverse
transverse
transverse
vertebral
blotches
circular
paired
inverted
^
convex protuberant convex protuberant protuber
Midbody
dorsals 125-170 107-132 100 88-103 102-125
102-108
Paravertebrals
251-281
192-226
175
173-191
171-203
180-195
175-195
175-207
178-200
155-170
158-179
* 123-135 112-127 114-
protuberant 96-112
Midbody
16-18
18-22
15
15-17
16-20
16-20
12-15
12-18
10-16
10-20
?^
Midbody
ventraisSupralabials3
33-41
13-16
11-13
12-14
16
13-15
13
15,
^12,
12
11-14 38-42 40
scansors 35-40
18-22
16-19
17-19
IV
Toe
13
16
15-18
10-16
12,
12-14
13-15
^14-16 40-17
? 37-42
Vertebral
23-27
29-33
31-37
17-25
17-26
18-23
17-24
19-28
18-24
18
W 41-50 42-48 38-43
74-80 inAGD 74-104 52-66 67-85 71-84
Ventrais 64
38-44 64-74 52-66
58-63
63-66 65-
31-40 57-61

(1978)
b
Taylor
Alcala
70-72;
82
(1922a,b)
80
Brown
the
preanofemorals;
reported
and
pore-bearing
count
enlarged
in
scales
series.
we

(CAS
female
holotype
because
character
impossible
60526)
in
is
the
{5
immature
This
to
the
pores
cof
absence
females.
in
assess
Gekko
the
holotype)
porosus
Data
juvenile
from
considered
adults.
two
paratypes
of
rossi
excluded.
also
G.
are
were

porosus
^d

Defined
all
supralabials
ato
the
counted
as
point
differentiated.
posteriorly
which
at
longer
scales
no
were

asals 1,2 1 1

light
bands
CJ
spots

Scales 5 5 5 5 5
^ iQ femoral
Distal
?c+
----
--
contacting
fiG
series patch
qpresent
?n gtubercle
rows
mterrupted
W 5C O
in
tubercles
femoral pore-bearing ?d
nostril
contacting
grostral O
g AGD
AGD in

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 628 R. M. BROWN ET AL.

Axilla-groin distance 46.9% SVL; undifferen but first (inner) claw greatly reduced to a
tiated dorsal body scales round to irregularly conical nail; other terminal claw-bearing pha
ho with
octagonal, convex, nonimbricate, relatively langes compressed, large recurved claws,
mogeneous; dorsals sharply transition to imbri not freeuntil beyond dilated portion of digit.
cate ventrals
along
the ventrolateral adipose Tail base bordered by a single, greatly
fold; dorsals lack interstitial granules but enlarged conical postcloacal spur on each side
with 18 transverse rows of vent;
interspersed irregularly postcloacal swellings pronounced;
(36 paravertebral rows) of highly enlarged and hemipenes completely everted; regenerated
protuberant,
to
strongly conical dorsal body portion long, 122% SVL; TD (not including
tubercles; each dorsal tubercle with a raised, basal postcloacal swelling) 93.7% TW; tail not
thornlike point; 144 transverse midbody dor depressed, nearly cylindrical, divided into
sals; 250 between of distinct fracture grooves (=
paravertebrals midpoints planes/autotomy
limb insertions; 38 transversely ven whorls or annulations); dorsal surface
arranged heavily
trals; scales on dorsal surfaces of limbs adorned with raised tuber
larger spinose, posteriorly
than dorsals, with interspersed tuber cles, concentrated along posterior of
enlarged edge
cles extending down limbs, especially concen annulations, caudals similar in size to dorsals;
trated on radioulnar segment of forelimb (and subcaudals enlarged, platelike,
3-5 rows per
covering entiretyof hind limb), and terminating annulus, widely expanded
to cover most of
at the dorsal surfaces of hands and feet; ventral surface or split into a pair of subcaudals

enlarged patches of distinct imbricate scales along posterior margin of each annulus; distal
on wrist, anterior surface tail with clear autotomy
present (prebrachial) portions regenerated;
of upper arm and
thigh,
on knee, and on distal scar and dis tally regenerated portion, 15 annuli
ventral surface of hind limb, just before attach before autotomy
scar (80 mm), 7-8 annulations
ment of foot; scales on dorsal surfaces of hands estimated in autotomized based on
portion
and feet similar to dorsal limb scales; ventral length (39mm), for an approximate total of 22
scales flat, cycloid, imbricate, or 23 annuli.
body strongly
much than lateral or dorsal body scales, Variation.?The type series contains eight
larger
at mid ventral line. adult males with hemipenes everted, two
largest large
Seventy-nine dimpled pore-bearing scales presumably juvenile females, and eightmature
(Fig. 5B)
in a continuous precloacal-femoral
females (all gravid, with a pair of large white
series (42 on L/37 on R) each punctured with subovate eggs visible through skin of poster
dark exudates; oventral wall). Ranges of selected diag
pore bearing orange precloacal body
femorals in a wavy, obtuse, inverted nostic meristic characters are in
arranged presented
"V" and continuing to the ends of the knee; Table 1.
inferred "precloacal" pores 2-3 times the Coloration of Holotype in Ethanol.?Dorsal
diameter of inferred "femoral" pores (Fig. 5B); ground coloration of head, body, tail, and dorsal
situated a substantial surfaces of limbs medium gray with scattered
precloacals atop precloa
cal bulge that folds over into precloacal region indistinct light and dark (almost black) gray
in but was erect and blotches; six diffuse black bars traverse the
preserved specimen
protuberant in life (Fig. 5B); precloacal-femorals axilla-groin distance, with two more above both
five but non the fore- and hind-limb insertions; accompany
preceded by similarly enlarged
dimpled scale rows; precloacals followed by ing each bar are three light gray spots,
one
minute scales and two scales rows vertebral and a of ones on
enlarged pair paravertebral
(absent following femorals); femoral series lacks each side of the dorsal midline, bars and spots
or scale rows; of consistent trunk.
preceding following enlarged intensity throughout
scales lateroposterior
to
precloacal-femoral
se Dorsal and lateral surfaces of head similar to
ries (i.e., along ventroposterior surfaces of hind dorsal ground color; temporal and nuchal
to minute of cream
region with
limb) reduce in size sharply scales small, distinct spots; palpe
the posterior edge of the hind limb. bra darker gray than surrounding occiput;
and covered on rostral and medium gray
Digits moderately expanded supralabial regions
surfaces unnotched, with cream on infralabials
palmar/plantar by bowed, spots supralabials;
undivided scansors (Fig. 5A); no interdigital very lightgray; infra-auricular region lightgray,
webbing; scansors ofmanus: 13/13, 12/12, 14/
with dark gray supra-auricular
contrasting
15, 13/15, and 13/14 on left/rightdigits I-V, region.
as transverse
respectively; pes: 10/13, 12/13, 15/16, 15/12, Limbs colored torso, lacking
and 15/14 on left/rightdigits I-V, respectively; banding; dorsal surfaces of hands and feet light
scansors of manus and pes bordered basally (on gray; digits medium gray with cream spots; tail
palmar and plantar surfaces) by \-A slightly medium gray with dark gray bands every two
scales that form a near-continuous or three caudal annuli; dark
enlarged regenerated portion
series with scansors; all digits clawed, gray.
enlarged

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 PHYLOGENY AND NEW SPECIES OF NORTHERN PHILIPPINE GEKKO 629

Ventral head, neck, and torso cream; broken into transverse blotches. In
light irregularly
ventral surfaces of limbs slightly darker with others (KU 304885, 304924, 304927, 304931;
black flecks;ventral surfaces of digits (scansors) PNM 9091), bands are distinctly interrupted
dark gray; palmar and plantar surfaces medium vertebrally and dorsolaterally, with paired
gray; precloacofemoral region white with black squarish spots on either side of the
ventral surfaces of tail medium vertebral line the transverse
orange pores; (accompanying
not banded. series of three cream In
gray, paravertebral spots).
Coloration ofHolotype in Life.?From photo three specimens (KU 304916, 304919, 304923),
graphs by RMB (Fig. 3A-C). Dorsal ground dark transverse
banding
is faint or absent and
coloration dark yellowish-brown (laterally) to dorsal coloration is limited to paired cream
dark brown (dorsally), with indistinct, irregu paravertebral spots
on a brown
background.
larly scattered lightgray and black blotches and Juveniles (KU 304876, PNM 9541) are pat
small spots; dorsum with six diffuse bars light terned more brightly than adults, with more
cream trilobed bars traversing the axilla-groin intensely contrasting light and dark dorsal
region, each bordered posteriorly by
an accom coloration with tail banding nearly black and
set of three cream one situated white. In the of
panying spots, place paired paravertebral
vertebrally, and flanked by a paravertebral pair cream
spots, juveniles possess trilobed cream
on either side. bars, reminiscent of adult coloration in G.
Dorsal nuchal and crombota.
region posterior portions
of head similar to trunk coloration but darker Ventral coloration nearly
invariant: ventral
and with distinct, subcircular, cream
spots and body surfaces are cream to lightgray (yellowish
black blotches; similar cream
spots span a line in life); subcaudal coloration is light to dark
between the orbit and auricular opening; post gray. Only in the two juveniles does the black
rictal region flat purplish gray; labial scales and white transverse banded caudal pigmenta
purplish gray with cream spots on every third tion wrap around to the ventral surface of the
labial scale; rostral and circumnasal scales dark tail. In life,males had a more brightly colored
purplish gray; interorbital region, and parietal yellow (nearly orange in some
specimens)
medium gray with white infra of in the
region spots; congregation pigment precloacal
labial region and chin gray to light gray; gular region, along the line of femoral scales, overly
to brownish-tan. on wrists
region light gray ing the hemipenal bulge, and and
Dorsal surfaces of limbs light gray with ankles. Eye coloration
was
consistently gold in
numerous dark brown, black, and yellowish adults, but silvery in juveniles.
cream knees and elbows flat gray; dorsal Distribution and Natural new
spots; History.?The
surfaces of digits dark gray, with cream spots; species is known only from Calayan Island
dorsal and lateral portions of tail banded where itwas collected low (<5 m) on trunks
alternating dark gray and cream (corresponding and buttresses of trees or on the faces of rocky
to every third or fourth tail annulus); distal outcrops and in limestone caves in
primary
dark brown to dark gray, forest at moderate elevations (300
autotomy regrowth dipterocarp
nonbanded. 400 m) far from the island's coast (Fig. 6). Given
Ventral and limbs yellowish-cream with the island's of isolation
body geological history
scattered et al., et al.,
gray and dark brown flecks, especially (McDermott 1993; Yang 1996;
on chin and ventral surfaces of limbs; precloa Marini et al., 2005), we consider it extremely
calfemoral with dark that G. rossi will be encountered on
region yellow orange unlikely
pores, with other islands in the island group, the
contrasting distinctly posterior Babuyan
surfaces of thighs; limbs gray, fading to yellow Batanes island group, or mainland Luzon.
at wrists and ankles; and Our is that the new is very
palmar plantar impression species
surfaces ofmanus and pes yellowish with light common in internal forested regions
at the
type
gray distal portions and darker subdigital locality,but we note thatwe did not find itnear
scansors; ventral tail
yellowish, fading
to cream the island's denuded coast. It is possible that
distally, lacking
transverse bars, regenerated only the forested areas ofCalayan Island (Fig. 6)
portion solid brown; eye gold, with radiating are optimal habitat for this species.
purple lines surrounding crenulated black Other gekkonids encountered on Calayan
pupil. Island include Gehyra mutilata, Hemidactylus
Color Variation.?Dorsal color pattern is some and sp. Calayan Island
frenatus, Lepidodactylus
what variable in the type series. In some is the type locality of the rare Luperosaurus
specimens, transverse is it has not been on
banding relatively macgregori, although reported
complete and continuous (KU 304876, PNM this island since the original description (Stej
9538, 9539, 9541) and in others (KU 304917, neger, 1907) but has recently been rediscovered
304918; PNM 9540), the diffuse black transverse on
Babuyan Claro Island (Brown and Diesmos,
banding is more or less discontinuous and 2000; Brown et al, 2007, 2008).

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 630 R. M. BROWN ET AL.

""""
B

Fig. 6. Gefcfco rossi habitat at the type locality. Forest interior at 1,000 m (A); forest edge at 300 m (B).
Photographs by M. Babon and A. Bajarias Jr.

Etymology.?We
are
pleased
to name this new realized. With regard to the Gekko fauna of the
species of Gekko for Charles Andrew Ross in northern Philippines, theBabuyans and Batanes
recognition of his numerous contributions to the island groups require additional survey work
systematics of reptiles and amphibians of the before we can be reasonably
certain that the

Philippines and his particular enthusiasm for total gekkonid fauna is known. Endemism in
the herpetofauna of the Babuyan islands. both island groups (Ota and Crombie, 1989;
Suggested
common name: Ross' Calayan Gecko. Lazell, 1992a; Ota and Ross, 1994) is expected to
be high because of their isolation (Yang et al.,
1996;Marini et al, 2005; Brown et al., 2008). The
Discussion Batanes support endemic populations of pit
The description of G. rossi brings the total vipers (Leviton, 1964), gekkonid lizards (Taylor,
number of endemic Philippine species of the 1922a; Ota and Crombie, 1989), agamid lizards
genus Gekko to nine (11 species when nonen (Lazell, 1992), and snakes of the genus Lycodon
demics G. gecko and G. monarchus are included (Ota and Ross, 1994). The Babuyan Islands are
and the single doubtful record of G. hokouensis is home to endemic lizards of the genera Luper
excluded). We are certain that this number osaurus (Stejneger, 1907) and Gekko (Brown et al.,
underestimates actual species diversity,
and we 2008) and multiple species of snakes in the
have previously noted (Brown et al., 2008) genus Lycodon (Ota and Ross, 1994).
numerous taxonomic that must be We are aware of at least three additional,
problems
addressed before a
comprehensive
understand morphologically distinct, undescribed Gekko
ing of Philippine gekkonid evolution can be species in the Babuyans (Fuga, Dalupiri, and

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 PHYLOGENY AND NEW SPECIES OF NORTHERN PHILIPPINE GEKKO 631

Camiguin Norte island populations; H. Ota and support the recognition of a minimum of six
R. Crombie,
unpubl. data), and we suspect that
evolutionary lineages (= species) that occur
several additional new
species will be discov north of Luzon Island.With the exception of the
ered ifbiologists are able to visit other Philip two undescribed species
on
Dalupiri
and
islands isolated water. In partic Norte, which are
pine by deep Camiguin separated by only
ular, northern populations (all currently 2.6% uncorrected sequence divergence (but are
considered G. porosus) require reevaluation with morphologically highly distinct species; RMB
fresh and data once the and CO, pers. obs.), all taxa in our
specimens genetic represented
Batanes Islands have been show 9.3-14.6% and all taxa
thoroughly surveyed. analysis divergence
The endemic Gekko of Lanyu Island, south of are readily distinguishable on the basis of
Taiwan (G. kikuchii;Oshima, 1912) appears tobe scalation, body size, and color pattern.
more closely related to Philippine taxa than to If island emergence is followed by coloniza
other Taiwanese and Ryukyu Archipelago tion (with some necessary lag time as habitats
G. G. hokouensis, G. become suitable to terrestrial verte
species (e.g., japonicus, support
yakuensis,
G. shibatai, G. vertebralis; Ota, 1989; brates), we might expect island age to be
Ota et al, 1989; Toda et al, 2001, 2008), but correlated with the level of genetic divergence
phylogenetic appraisal of this taxonomic expec exhibited by endemic lineages inhabiting those
tation must await the availability of genetic islands. If the processes of dispersal and
samples. Gekko kikuchiihas been phenotypically establishment of populations is
relatively
con
allied to G. mindorensis (Bauer, 1994; Ferner et stant, this correlation be
expected may strong
al., 2001; R. Crombie and H. Ota, pers. comm.); (Steppan et al., 2003). Alternatively, a lack of a
it is not morphologically similar to the new correlation would the presumption that
support
species described here. historical and chance may be
contingency
The unexpected relationships elucidated by responsible for establishment of
lineages
on
our estimate of northern oceanic islands. if a strong correlation is
phylogenetic Philip Finally,
Gekko warrant comment. The southern and individual outliers can be identi
pine present
Babuyans lineages (Dalupiri and Camiguin fied, the identificationof recent dispersal events
Norte Islands) form a strongly supported clade, may be possible.
whereas the Gekko of the northern Babuyans There is no clear
relationship between genetic
(Calayan and Babuyan Claro) and Batan Island divergence
and estimated island age in northern

appear to be
closely related. This result is
Philippine Gekko. Radiometric ages of volcanic
contrary to
expectations based on
previous flows of Babuyan and Batanes Islands (Yang et
discussions of species distributions that have al., 1996) a estimate or minimum
provide rough
emphasized longitudinal patterns of species date of the volcanic activity that forced these
endemism. Most studies to date have identified islands above sea level. Although some highly
endemic or on either such as G. porosus and G.
species species complexes divergent lineages
the eastern Babuyan-Batanes Island bank or the rossi are endemic to the old islands of
relatively
western bank (Ota and Crom Batan (9.36-2.0 and (6.50-4.37
Babuyan-Batanes mya) Calayan
bie, 1989; Lazell, 1992; Ota and Ross, 1994).We mya), other highly divergent lineages such as G.
await the results of several crombota are found on islands
enthusiastically extremely young
additional phylogenetic studies involving other such as
Babuyan Claro (1.70-0.80 mya), sug
northern to deter error in radiometric
Philippine reptile lineages gesting dating techniques,
mine whether the latitudinal (north-south di recent dispersal from an unidentified source, or
or (east-west diver a combination of these and other factors.
vergence) longitudinal
gence) pattern prevails. Finally, the close sister relationship (with a
Surprisingly, G. rossi appears to be most pairwise genetic distance of only 2.6%) between
closely related to G. porosus, rather than to the the lineages endemic to Dalupiri (Gekko sp. A)
more
geographically proximate island lineages and Camiguin Norte (Gekko sp. B) cannot be
such as those on Dalupiri or Camiguin Norte explained by the estimated island ages (Dalu
(Fig. 2). Relationships supported by phyloge piri: 22.40 mya; Camiguin Norte: 2.82-0.40 mya)
netic analysis of the ND2 locus were very well unless we assume recent from the
dispersal
supported (likelihood and parsimony boot former to the latter. Thus, in the absence of a

straps of 99 to 100 throughout) with the straightforward correlation, we feel it more

exception of the node subtending the (G. likely, at least for this relatively young group
crombota [G. rossi + G. porosus]) clade. Thus, we of islands and their endemic geckos, that
are unable to assert with
complete confidence colonization of the islands north of Luzon has
the of G. crombota from not been a constant
systematic position steady, process (see discus
Claro Island. However, we can state sion in Lazell, 1992). Rather, we assume that
Babuyan
with a high degree of confidence that the dispersal to, and successful colonization of,
combination of + data islands north of Luzon is a stochastic
genetic morphological process

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 632 R. M. BROWN ET AL.

that is dependent an of abiotic our recent studies in the Babuyans

upon array encouraging
factors (weather, wind, ocean current, and and Batanes.

chance) and biotic components of the environ

ment numbers of colonists,
(e.g., competition
from previously established populations, Literature Cited
gecko
etc.). Bauer, A. M. 1994. Liste der rezenten Amphibien und
Aside from the new here, I (Australia). Das Tierreich,
species reported Reptilien: Gekkonidae
numerous Philippine gekkonid taxa await de Vol. 108, W. de Gruyter and Co., Berlin, Germany.
scription, including several additional species of Brown, R. M. 1999. New species of parachute gecko
Gekko from the Babuyans and Batanes island (Squamata; Gekkonidae; Genus Ptychozoon) from

groups, morphologically cryptic Gekko lineages northeastern Thailand and Central Vietnam. Co
in the of "wide peia 1999:990-1001.
masquerading taxonomy
Brown, R. M., and A. C. Diesmos. 2000. The lizard
species such as G. mindoresis
spread" complexes con
taxonomy, history, and
(RMB, CDS, and ACD, unpubl. data), several genus Luperosaurus:
servation for some of the world's
prospects
undiagosed lineages of the genus Cyrtodactylus rarest lizards. Sylvatrop: Technical Journal of
(Welton et al., in and undescribed
press), Philippine Ecosystems and Natural Resources
of the genus (Brown et al., 10:107-124.
species Luperosaurus
2007; Gaulke et al., 2007; Brown et al, in press). -. 2002. Application of lineage-based species
It is very clear that Philippine gekkonid concepts to oceanic island frog populations: the

diversity is substantially underestimated (Ota effects of differing taxonomic philosophies on the

and Crombie, 1989; Brown et al., 2007, 2008; estimation of Philippine biodiversity. Silliman

Gaulke et al., We the numbers of Journal 42:133-162.

2007). expect
Brown, R. M., and S. I. Guttman. 2002. Phylogenetic
Philippine gekkonid taxa tominimally double
in the near and we an systematics of the Rana signata complex of Philip
future, hope improved and Bornean stream frogs: reconsideration of
pine
understanding of the evolutionary diversity in Huxley's modification of Wallace's Line at the
this of Philippine verte
conspicuous group Oriental-Australian faunal zone interface. Biolog
brates will contribute to conservation efforts ical Journal of the Linnean Society 76:393^161.
aimed at the forested Brown, R. M., J.W. Ferner, and A. C. Diesmos. 1997.
preserving remaining
of the country. Definition of the Philippine Parachute Gecko,
regions
Ptychozoon intermedium Taylor 1915 (Reptilia: La
loans of specimens or certilia: Gekkonidae): redescription, designation of
Acknowledgments.?For
a neotype, and comparisons with related species.
assistance while museum collections,
visiting 53:357-373.
we thank the following individuals and their Herpetologica
Brown, R. M., J.A. McGuire, and A. C. Diesmos. 2000.
institutions: A. Resetar and H. Voris
respective Status of some Philippine frogs referred to Rana
(FMNH); J.Vindum and A. Leviton (CAS); J. everetti (Anura: Ranidae), of a new
description
Hanken, J. Losos, and J. Rosado (MCZ); J. R. igorota Taylor 1922.
species, and resurrection of
McGuire (MVZ); K. de Queiroz, A. Wynn, and Herpetologica 56:81-104.
R. Heyer (USNM); and V. Palpal-latoc (PNM). Brown, R. M., A. C. Diesmos, and M. V. Duya. 2007. A
We thank the Department of the Environment new Luperosaurus (Squamata: Gekkonidae) from
and Natural Resources and the Protected Areas the Sierra Madre of Luzon Island, Philippines.
andWildlife Bureau for facilitatingresearch and Raffles Bulletin of Zoology 55:167-174.
Brown, R. M., C. Oliveros, C. D. Siler, and A. C.
export permits for this and related studies, KU Diesmos. 2008. A new Gekko from the Babuyan Islands,
IACUC for approving research and
protocols, northern Philippines. Herpetologica 64:305-320.
M. and M. A. N. Antoque, T.
Pedregosa, Reyes,
Brown, R. M., A. C. Diesmos, M. V. Duya, H. J. D.
Peterson, B. Fernandez, and J. Fernandez for
Garcia, and E. L. Rico. In press. A new forest gecko
assistance in the field.The Stearns Fellowship of Gekkonidae; Genus Luperasaurus) from
(Squamata;
the California Academy of Sciences provided Mt. Mantalingajan, Southern Palawan Island,
support that allowed RMB, ACD, and CDS to Philippines. Journal of Herpetology.
undertake multiple research visits to CAS. We Brown, W. C, and A. C. Alcala. 1978. Philippean
also thank municipal DENR authorities of lizards of the Family Gekkonidae. Silliman Uni

Calayan, Cagayan
Province for logistical sup versity Press, Dumaguete City, Philippines.
de Queiroz, K. 1998. The general lineage concept of
port. Our fieldworkwas conducted with finan and the process of
cial support from theNational Science Founda species, species criteria,
In D. J. Howard and S. H. Berlocher
tion (DEB 0743491 to RMB and DEB 0328700 to speciation.
(eds.), Endless Forms: Species and Speciation,
J. McGuire and D. Cannatella [G. monarchus Press, New York.
pp. 57-75. Oxford University
samples]) and theUniversity of Kansas. Thanks-. 1999. The general lineage concept of species
to M. Garfield for scientific illustration; C. and the defining properties of the species category.
R. Crombie, and J. Esselstyn for review
Linkem, In R. A. Wilson (ed.), Species: New Interdisciplin
of previous drafts of the manuscript; and H. Ota ary Essays, pp. 49-89. Massachusetts Institute of
and R. Crombie for sharing information and Technology Press, Cambridge.

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Appendix 1
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and C. A. Ross. 1994. Four new Examined
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Philippines. Copeia 1994:159-174. unless otherwise noted. Numbers in parentheses

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All use subject to JSTOR Terms and Conditions
 634 R. M. BROWN ET AL.

indicate the number of specimens examined for each PANAY ISLAND, Capiz Province, Municipality of
species. Pilar, Barangay Natividad: KU 302729-302732; LU
Gekko athymus.?(7) PALAWAN Palawan BANG ISLAND: Occidental Mindoro Province, Mu
ISLAND,
Province, 10 km west-southwest of nicipality of Lubang, Vigo: KU 303913
approximately Barangay
303916, 303917-303951.
Iwahig: CAS 137677; approximately 8-9 km south of
Balico: CAS-SU 23119 (holotype); approximately Gekko monarchus.?(6) PALAWAN ISLAND, Pala
20 km southwest of Iwahig: CAS-SU 23121 (paratype); wan Province, 1.5 km west-southwest
approximately
Municipality of Brooke's Point, Barangay Mainit: KU of Iwahig: CAS-SU 28416; approximately 5 km south
309335; Barangay Samarinana; Mt. Mantalingahan, southeast of Iwahig: CAS-SU 28496; approximately
900m: KU 309331-309334. 7 km west-northwest of Iwahig: CAS-SU 28554; Mu
Gekko crombota.?(21) BABUYAN CLARO ISLAND, nicipality of Brookes Point, Mt. Mantalingahan: KU
Cagayan Province, Municipality of Calayan, Barangay 309362; INDONESIA, SULAWESI ISLAND: BSI 340,
Claro: PNM 9280 (holotype); KU 304807 819 (specimens at Museum
Babuyan deposited Zoologicum
304809, 304814, 304821, 304825-304826 3043830, Bogoriense, Cibinong, Jakarta, Indonesia).
304836, 304845, 304848; PNM 9281-9284,PNM 9090, Gekko paiawanensis.?(24) PALAWAN ISLAND, Pa
9095-9098. lawan Province, 7 km west-northwest of Iwahig: CAS
Gekko ernstkelleri.?(10) PANAY ISLAND, Antique 17318; 8 km west of Iwahig: CAS 17319; approximate
Province,Municipality of Pandan, Barangay Duyong, ly 9 km west of Iwahig: CAS 17320-17322; 19
at KU
Duyong Hillside (= "Mt. Lihidian"), 300 m.a.sl.: PNM currently uncataloged specimens (RMB 7460,
9152-54; KU 300196-300202. 7531, 7561-7562, 7589-7590, 7615, 7642. 7726-7727,
Gekko gecko.?(13) LUBANG Occidental 7780, 7878, 7922-7924, 7930-7931, 7937-7938).
ISLAND,
Mindoro Province, Municipality of Lubang, Gekko porosus.?(8) BATAN ISLAND, Batanes Pro
Barangay
Paraiso: KU 303960-303972. vince, 3 km east-northeast of Basco Town: USNM
Gekkogigante.?(13) SOUTH GIGANTE ISLAND, 266519,291387;Mahatao: USNM 266517;Municipality
of Basco, of Basco near
Iloilo Province, Municipality of Carles, Tantangan:
outskirts Town, airport: PNM
CAS 124315-124317 (paratypes);NORTH GIGANTE 9532-9536; ITBAYAT ISLAND: CAS 60526 (holotype).
ISLAND, Iloilo Province, of Carles: Gekko romblon.?(15) SIBUYAN ISLAND, Romblon
Municipality
CAS 124866-124867 Asloman: Province, Taclobo Barrio: CAS 139180-139182 (para
(Paratypes); Barangay
KU 302716-302720, 305138-305140. types); ROMBLON ISLAND, Romblon Province, Mu
Gekko hokouensis.?(1) "Tablas Island" nicipality of Romblon, Barangay Li-O: KU 302733
(presumably
in error): FMNH 17812 (Luperosaurus amissus holo 302742, 303977-303978.

type). Gekko sp. A.?(35) DALUPIRI ISLAND, Cagayan

Gekko mindorensis.?(56) Province, Municipality of Calayan, Creek; KU
NEGROS ISLAND, Negros Nipa
Oriental 307022-307039, 307040-307057.
Province, Himangpangon Cave, Manjayod:
CAS-SU 28656-28660; GUIMARAS ISLAND, Guimaras Gekkosp. B.?(24) CAMIGUIN NORTE ISLAND,
Province, Municipality of Buenavista, Barangay Old Cagayan Province, Municipality of Calayan, Barangay
Poblacion: KU 302721, 302725; NEGROS ISLAND, Balatubat; KU 304583,304585,304586,304588,304605
Negros Occidental Province, Municipality of Cauayan, 304611, 304617, 304673, 304728-304733, 304738,
Barangay Camalandaan: KU 302722-302724; MAS 307990, 308043; Magas-asok: PNM; 9099; Pomoctan
BATE ISLAND, Masbate Province, Municipality of Island (small island to Camiguin Norte
adjacent
Mandaon, Barangay Poblacion: KU 302726-302728; Island): PNM 9100.

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