a new gekko from the babuyan islands, northern philippines

Herpetologica, 64(3), 2008, 305–320
E 2008 by The Herpetologists’ League, Inc.
A NEW GEKKO FROM THE BABUYAN ISLANDS,
NORTHERN PHILIPPINES
RAFE M. BROWN1,4, CARL H. OLIVEROS1,2, CAMERON D. SILER1,
1
AND
ARVIN C. DIESMOS3
Natural History Museum and Biodiversity Research Center, Department of Ecology and Evolutionary Biology,
The University of Kansas, Lawrence, KS 66045-7561, USA
2
ISLA Biodiversity Conservation; 9 Bougainvillea St, Manuela Subdivision, Las Pinas City 1741 Philippines
3
National Museum of the Philippines, Rizal Park, Padre Burgos Ave. Ermita 1000, Manila, Philippines
ABSTRACT: We describe a new species of gekkonid lizard on the basis of 21 recently acquired specimens
from Babuyan Claro Island, Babuyan Islands group, northern Philippines. The new species differs from other
Philippine Gekko by characteristics of external morphology, color pattern, and body size. The new species has
been found low on trunks and buttresses of mature closed-canopy climax forest trees at low elevation near the
island’s coast. It is known from only Babuyan Claro Island and is likely endemic to this single small, isolated
landmass. The remaining gekkonid fauna of the Babuyans and Batanes island groups is understudied and in
need of comprehensive review.
Key words: Babuyan Claro; Babuyan Islands; Gekkonidae; New species; Philippines
PHILIPPINE lizards of the family Gekkonidae
are represented by 10 genera and 37 species:
Cyrtodactylus (4), Gekko (9), Gehyra (1),
Hemidactylus (5, including H. platyurus, a
species formerly assigned to Cosymbotus),
Hemiphylodactylus (1), Lepidodactylus (6),
Luperosaurus (6), Pseudogekko (4), and Ptychozoon (1) (Brown and Alcala, 1978; Brown
et al., 2007; Gaulke et al., 2007; Taylor,
1922a,b). Although the lack of definitive
characters distinguishing the genera Luperosaurus and Gekko have clouded generic
boundaries (Brown and Alcala, 1978; Brown
et al., 2000a; Brown et al., 2007; Russell,
1979), most authors have applied the name
Gekko to Philippine species that possess (1)
moderate to large body size and longer, more
slender limbs; (2) near or complete absence of
interdigital webbing; (3) a lack of cutaneous
expansions, save for ventrolateral adipose
folds and very slight expansions bordering
the posterior margins of the hind limbs; (4)
convex to conical enlarged dorsal tubercles
arranged in longitudinal rows; (5) enlarged,
plate-like subcaudals; (6) differentiated, slender, and elongate postmentals; and (7) minute
and non-imbricate rows of dorsal scales
between tubercle rows (minute dorsals differentiated from the enlarged, imbricate scales of
the venter (Brown and Alcala, 1978; Brown et
al., 2000a; Brown et al., 2007).
4
CORRESPONDENCE: e-mail, rafe@ku.edu
305
Philippine members of the genus Gekko
include seven endemic species (G. athymus,
G. ernstkelleri, G. gigante, G. mindorensis, G.
palawanensis, G. porosus, and G. romblon;
Brown and Alcala, 1978; Brown et al., 2007;
Roesler et al., 2006; Taylor, 1922a), two
species shared with neighboring countries
(G. gecko and G. monarchus; Manthey and
Grossman, 1997; Wermuth, 1965), and one
species (G. hokouensis) that is probably
included in the country’s gekkonid fauna in
error (Brown and Alcala, 1978; Ota et al.,
1989).
Several additional Philippine gekkonid taxa
presently await description, including multiple
unnamed species from the northern Philippines. During recent biological inventory
work in the Babuyan archipelago, we discovered a distinctive species of Gekko on
Babuyan Claro Island, now represented in
U.S. and Philippine collections by 21 specimens. In this paper we describe the new
species and discuss morphological diversity,
patterns of gekkonid endemism, and probable
processes of evolutionary diversification in the
Philippine island radiation of Gekko.
MATERIALS AND METHODS
We (RMB and CDS) collected data from
fluid-preserved specimens deposited in U.S.
and Philippine collections (see Acknowledgments). Sex was determined by inspection of
gonads or by scoring of prominent secondary
306
HERPETOLOGICA
sexual characteristics (Brown, 1999; Brown et
al., 1997, 2000a;) when dissection was not
possible. Measurements (to the nearest
0.1 mm) were taken with Fowler digital
calipers following character definitions by
Ota and Crombie (1989), Brown et al.
(1997), Brown (1999), and Brown et al.
(2007). Character abbreviations are: snout–
vent length (SVL); tail length (TL); head
length, (HL); head width (HW); head depth
(HD); snout length (SNL); eye diameter,
(ED); eye–narial distance (END); auricular
opening diameter (AO); inter-narial distance
(IND); inter-orbital distance (IOD); axilla–
groin distance (AGD); femur length (FL);
tibia length, (TBL); Toe I length (TiL); Toe IV
length (TivL); tail width (TW); and tail depth
(TD); number of supralabials (SUL) and
infralabials (IFL) to the center of the eye
and posteriorly to the point at which point
labials are no longer differentiated; enlarged
circumorbitals dorsoanterior to orbit (CO);
modified spiny circumorbitals (cilaria;) dorsoposterior to orbit (SC); differentiated preanal
pore-bearing scales (PS); femoral pore-bearing scales (FS); differentiated subdigital scansors on fingers 1–5 (FS I–V); subdigital
scansors on toes 1–5 (TS I–V); midbody
ventral transverse scale rows counted between
lateral body folds (MBVS); midbody dorsal
transverse scale rows counted across dorsum
between lateral body folds (MBDS); midbody
transverse tubercle rows counted across dorsum between dorsolateral body folds (MBTR);
paravertebrals between midpoints of limb
insertions (PVS); ventrals within AGD (counted midventrally between limb insertion; VS)
paravertebral tubercles within AGD (counted
along right side of vertebral axis between limb
insertions; PVT); tail annuli (in complete tails;
TA); and subcaudals (beneath complete tails;
SC). For the recognition of the new species,
we adopted the General Lineage Species
Concept (GLC) of de Queiroz (1998, 1999)
as the natural extension of the Evolutionary
Species Concept (Wiley, 1978). Application of
lineage-based species concepts to island
endemics is straightforward because of the
known history of isolation of island populations (Brown and Diesmos, 2002; Brown and
Guttman, 2002). We consider as new species
morphologically diagnosable, isolated popula-
[Vol. 64, No. 3
tions where the hypothesis of a distinct
evolutionary lineage cannot be rejected.
SYSTEMATICS
Gekko crombota sp. nov.
Figs. 2–5
Holotype.—PNM 9280 (Field no. RMB
5954; formerly KU 304833), an adult male
collected at night (2125 h on 13 March 2005)
on the buttress of a large dipterocarp tree
by RMB at an area known locally as
‘‘Asked,’’ Barangay Babuyan Claro, Municipality of Calayan, Cagayan Province, W
coast of Babuyan Claro Island, Philippines
(19.5041u N, 121.9120u E; 20 m above sea
level).
Paratopotypes.—KU 304825 adult male;
KU 304808, 304814, 304821, 304848, PNM
9281–82 (formerly KU 304829, 304832), two
male juveniles and four juveniles of undetermined sex; KU 304807, subadult female; KU
304809, 304826, 3043830, 304836, 304845,
PNM 9283–84 (formerly KU 304847 and
304849), adult females, collected by RMB,
CO, Jason Fernandez and Boying Fernandez;
same date, locality, and circumstances of
capture (19:45–22:45 h) as holotype.
Other paratypes.—PNM 9095–96 (Field
Nos. MGDP 110 and 113), adult females;
and PNM 9097 (Field No. MGDP 111), adult
male, collected by CO and Marisol Pedigrosa,
23 April 2004 at Ayumit, Municipality of
Calayan, Cagayan Province, Babuyan Claro
Island (19.545u N, 121.958u E, 360 m); PNM
9098, subadult male, collected by CO and
Marisol Pedigrosa, 27 April 2004 at Corog,
Municipality of Calayan, Cagayan Province,
Babuyan Claro Island 19.489u N, 121.948u E,
between sea level and 100 m; PNM 9090
(Field No. MGDP 119), adult male, collected
by CO and Marisol Pedigrosa, 27 April 2004 at
Rakwaranom, Municipality of Calayan, Cagayan Province, Babuyan Claro Island 19.561u
N, 121.953u E, between sea level and 100 m.
Diagnosis.—Gekko crombota differs from
all other species of Philippine Gekko (i.e., G.
athymus, G. ernstkelleri, G. gecko, G. gigante,
G. mindorensis, G. monarchus, G. palawanensis, G. porosus, and G. romblon) in the
following combination of features (1) large
body size (SVL 85.5–117.9 for adult males;
September 2008]
HERPETOLOGICA
307
FIG. 1.—Map of the northern Philippines (inset) showing the type locality (1: Asked) of Gekko crombota on the SE
coast of Babuyan Claro Island (A). Additional collection localities include (2) Corog, (3) Rakwaranom, and (4) Ayumit.
85.1–106.9 mm for females); (2) dorsum
medium brown to gray, marked with cream
colored, transverse, tri-lobed bars with a black
anterior border; (3) high numbers of dorsal
body scales (107–132 transverse midbody
scales; 192–226 paravertebrals); (4) high
numbers of sharply conical dorsal body
tubercle rows (17–21 midbody; 27–33 paravertebrally); (5) preanofemorals arranged in a
non-continuous series (preanals and femoral
pore-bearing scale series separated by one or
two non-pored scales in 75% of specimens) of
58–74 differentiated, slightly enlarged scales.
Comparison with similar species.—Gekko
crombota differs from its phenotypically most
similar Philippine congener, Gekko porosus
(Batan and Itbayat islands, of the Batanes
Islands group, north of the Babuyans), by its
attainment of a larger maximum male body
size (males: SVL 85.5–117.9 vs. 91.0–96.7 in
G. porosus; Fig. 5); swollen (presumptively
hypertrophied) adductor and temporal musculature of the head, leading to a wider,
vaguely triangular head shape (Fig. 3B,C;
versus adductor and temporal musculature
distinctly less enlarged in G. porosus, head
more slender); brown to gray dorsal coloration, interrupted by distinct, tri-lobed, creamcolored bars (Fig. 2; versus indistinct dark
transverse bands and/or circular vertebral
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HERPETOLOGICA
[Vol. 64, No. 3
FIG. 2.—Photographs of northern Philippine Gekko in life. (A, B) Adult female Gekko crombota (KU 304849; SVL 5
111.8 mm); (C, D) juvenile G. crombota of undetermined sex (KU 304829; SVL 5 58.5 mm); (E) Adult presumably
male Gekko porosus and (F) juvenile G. mindorensis of undetermined sex; both photographed (specimens not collected)
in Crystal Cave, Batan Island, 2006, by RMB.
blotches in G. porosus); a greater number
(107–132) of transverse midbody dorsal scales
(vs. 88–103 in G. porosus); a greater number
(192–226) of paravertebral scales (vs. 173–191
in G. porosus); the presence of sharply
protuberant (vs. merely convex or slightly
raised) dorsal tubercles; a greater number
(17–21) of midbody dorsal tubercle rows (vs.
15–17 in G. porosus); and a greater number
(27–33) of paravertebral tubercle rows (vs.
17–24); separation (1 or 2 undifferentiated
scales) between the preanal and femoral series
of pore-bearing scales (in 75% of specimens;
vs. absent in G. porosus); the presence of
fewer (58–74) preanofemoral pore-bearing
scales (vs. 74–82 in G. porosus); and the
absence of a modified distal femoral porebearing patch (vs. present in G. porosus,
composed of a short series of 2 or 3 rows of
pore-bearing scales (Fig. 4C).
September 2008]
HERPETOLOGICA
309
FIG. 3.—Lateral (A), dorsal (B), and ventral (C) scalation of head of holotype of Gekko crombota (male PNM 9280).
Scale bar 5 5 mm.
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HERPETOLOGICA
[Vol. 64, No. 3
FIG. 4.—(A) Ventral view of left hand of holotype of Gekko crombota (male PNM 9280; Scale bar 5 5 mm); (B) left
side of preano-femoral pore-bearing scale series of holotype of Gekko crombota (male PNM 9280) and (C) pore-bearing
series of an adult male of Gekko porosus (male USNM 266517; Scale bar 5 5 mm). Note single undifferentiated, nonpore-bearing scale interrupting pre-anal and femoral series in G. crombota (arrow).
September 2008]
HERPETOLOGICA
311
FIG. 5.—Photographs of left side of heads of preserved specimens of (upper) Gekko porosus (juvenile female holotype
CAS 60526), (middle) Gekko porosus adult male (USNM 266517), and (lower) Gekko crombota (adult male holotype,
PNM 9280); scale bar 5 5 mm.
The new species differs from Gekko monarchus, G. mindorensis, G. romblon, G.
gigante, G. ernstkelleri and G. palawanensis
in having a larger maximum male body size
(85.5–117.9 mm SVL; Table 1); the only
Philippine species approaching the body size
of G. crombota are G. gecko (SVL 120.0–
153.8 mm) and G. athymus (99.2–119.9). The
number of preanofemorals distinguishes G.
crombota from G. monarchus, G. gigante, G.
ernstkelleri, G. athymus, and G. gecko (Table 1). The presence of separated preanal and
femoral pore-bearing scales is shared at
variable frequencies (Table 1) by at least G.
crombota, G. romblon, G. gigante, and G.
palawanensis and distinguishes these species
from G. porosus, G. monarchus, G. mindorensis, G. ernstkelleri, G. athymus, and G.
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HERPETOLOGICA
[Vol. 64, No. 3
FIG. 6.—Photographs of the coastal forest habitat of G. crombota on Babuyan Claro Island. View of uninhabited east
coast of the island (A) and view of the type locality (Asked) from the southeast side of the island (B). In both images,
thick volcanic igneous layers are in the foreground, intact coastal forest can be seen above the uninhabited beach, and
the largely forested Mt. Pangasun is visible in the background. Photos courtesy of G. Broad.
4m; 9f;
crombota
1f (juv)
porosus holotype
3m
porosus adults
4m; 12f
monarchus
3
2
1
4m; 6f
romblon
2
12–13
convex
1–2
5
37–42
102–108
12–15
18–24
175–195
63–66
2
12–14
protuberant
1
5
40–47
102–125
16–20
17–26
180–195
58–63
55.0–88.2
62.7–89.2
68.2–70.9
58.6–72.5
dark thin
light + dark
transverse
vertebral
bands
blotches
11–13
11–14
52–66
71–84
2
+ (33%)
22m; 13f
mindorensis
4m; 8f
ernstkelleri
3m; 5f
palawanensis
3m; 2f
athymus
gecko
9m; 12f
65–74
175–207
19–28
41–50
123–135
12–18
5
1
16–19
protuberant
2
58–62
178–200
17–25
42–48
112–127
10–16
5–6
17–19
protuberant
to conical
1
2
54–58
155–170
23–27
38–43
114–121
10–20
5
1
16–19
conical
2
66–72
158–179
—
30–36
92–104
—
5
1
18–22
absent
2
60–64
91–102
18–22
30–35
94–106
10–12
5–6
1–3
17–20
conical
2
89.7–104.7
82.0–92.1
57.2–65.7
99.2–119.9 120.1–166.1
79.7–87.9
78.0–88.0
44.5–61.8
88.2–117.1 119.2–144.1
dark paired white circular dark paired light + dark rust colored
blotches
spots
spots
inverted
spots
V-shaped
11–13
15–16
12–14
11–13
12–14
52–66
36–42
64–70
20–24
12–20
+ (28%)
2
+ (39%)
2
2
5m; 4f
gigante
Defined as all enlarged/differentiated supralabials, counted posteriorly to the point at which scales were no longer differentiated.
Taylor (1922) reported 80 preanofemorals; Brown and Alcala (1974) reported 70–72; we count 82 enlarged scales in the pore-bearing series
This character cannot be confidently assessed in the immature female holotype (CAS 60526) due to the absence of pores in the preanofemoral series of females.
111.8–117.9
91.0–96.7
56.2–80.7
85.5–106.9
49.3
91.0–96.7
40.6–69.7
light tri- light circular indistinct
dark transverse
lobed
blotches
transverse spot rows
bars
dark bands
13–15
12
12–13
11–13
Supralabials1
74–80
31–40
Preanofemorals
58–74
822
3
2
2
Preanal &
+ (75%)
?
femoral series
interrupted
+
2
Distal femoral
2
?3
pore-bearing
patch present
Toe IV scansors
15–18
16
14–16
13–15
Dorsal
protuberant
convex
convex
protuberant
Tubercles
to conical
Internasals
1
1
1
1–2
contacting
rostral
Scales contacting
5
5
5
5
nostril
Midbody ventrals 38–42
40
35–40
38–44
Midbody Dorsals 107–132
100
88–103
96–112
Midbody
18–22
15
15–17
16–20
tubercle rows
Vertebral
29–33
18
17–24
18–23
tubercles In
AGD
Paravertebrals in 192–226
175
173–191
171–203
AGD
Ventrals in AGD
67–85
64
64–74
5761
male SV:L
female SVL
Vertebral
coloration
n5
TABLE 1.—Distribution of selected diagnostic characters in Gekko crombota and other Philippine species of Gekko. For simplicity, bilaterally symmetrical characters are
presented for the left side only. Measurements are presented in mm and all specimens (with the exception of the G. porosus holotype) are considered adults. Data from seven
juvenile paratypes of G. crombota were exclude.
September 2008]
HERPETOLOGICA
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HERPETOLOGICA
[Vol. 64, No. 3
TABLE 2.—Continuous morphometric variation in the type series of G. crombota. Data are presented for adults only
(subadults and juveniles excluded). See Materials and Methods for character definitions. Entries are means 6 1 SD
(range in parentheses).
Character
SVL
TL1
HL
HW
HD
SNL
ED
END
AO
IND
IOD
AGD
FL
TBL
TIL
TIVL
TW
TD
1
Males (n 5 4)
101.1
101.7
27.7
21.2
11.6
12.7
5.9
9.9
2.5
3.3
4.7
48.1
22.8
17.4
6.0
11.6
9.8
7.3
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
16.2 (85.5–117.9)
18.7 (80.5–115.7)
3.6 (24.5–31.1)
3.3 (18.3–24.6)
2.0 (9.6–13.5)
1.7 (11.2–14.6)
0.6 (5.3–6.7)
1.5 (8.6–11.5)
0.7 (1.8–3.2)
0.9 (2.5–4.2)
0.3 (4.4–5.2)
8.8 (40.0–58.1)
3.8 (19.4–26.1)
3.8 (14.9–19.9)
1.0 (4.8–7.2)
1.8 (9.9–13.2)
1.0 (8.4–10.8)
1.0 (6.4–8.2)
Females (n 5 9)
98.7
83.0
26.7
20.2
11.2
12.7
5.1
9.3
2.0
3.5
5.0
47.2
21.8
17.0
5.5
10.6
9.0
7.3
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
8.4
8.6
1.9
1.7
1.2
0.7
0.6
0.6
0.3
0.3
0.4
3.7
2.4
1.4
0.7
0.7
1.2
0.8
(85.1–106.9)
(71.9–94.0)
(23.5–28.1)
(17.7–21.8)
(10.0–12.7)
(11.3–13.1)
(4.0–5.7)
(8.4–10.2)
(1.7–2.4)
(3.1–3.9)
(4.5–5.6)
(42.3–52.0)
(17.8–23.6)
(14.8–18.2)
(4.3–6.3)
(9.7–11.9)
(7.5–10.6)
(6.3–8.2)
Includes data on regenerated tails (4m; 7f).
gecko. The new species is further distinguished from G. athymus by the presence of
dorsal body tubercles (absent from G. athymus)
and from G. porosus and G. romblon by the
presence of sharply conical dorsal tubercles (vs.
tubercles only slightly convex in these species).
Finally, the presence of distinct transverse light
cream-colored tri-lobed bars across the dorsum
appears to distinguish G. crombota from the
highly variable suite of color patterns exhibited
by other Philippine species of Gekko. These and
other differences among Philippine Gekko
species are summarized in Table 1.
Description of holotype.—Adult male in
excellent condition (Figs. 3–4), with a small
incision in the sternal region (portion of liver
removed for tissue specimen), and hemipenes
partially everted. SVL 118.0 mm; habitus
robust, 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 wide, characterized by extremely
hypertrophied temporal and adductor and
temporal musculature; noticeably broader
(1.2 times) body at widest point; snout
subtriangular, rounded at tip in dorsal and
lateral aspect (Fig. 3A,B,C); HW 79.1% of HL
and 20.8% of SVL; SNL 59.3% of HW and
46.9% of HL; dorsal surfaces of head relatively homogeneous, with only slightly pronounced concave postnasal, prefrontal, interoribital, and parietal depressions; auricular
opening large, round, angles slightly lateroposteriorly from beneath temporal swellings
on either side of head; tympanum very deeply
sunken; orbit large, bordered by only slightly
distinct supraorbital crest; eye large, pupil
vertical, margin wavy (Fig. 3A); AO 49.1% of
ED; limbs relatively long and slender; femoral
segments of hind limbs thick, bulky; TBL
16.9% of SVL, 76.2% of FL.
Rostral large, subrectangular, not quite
twice as broad as high, with two dorsomedial
depressions between raised posterodorsal
projections that form the anterolaterally-projecting edge of the nares and suture anteriorly
with the supranasals; nostril surrounded by
rostral, the first labial, a single lower postnasal,
an enlarged upper postnasal, and an enlarged,
round, convex supranasal; supranasals separated on either side by a single large
internasal; supranasals and internasal followed
posteriorly by a pair of slightly enlarged
posterosupranasals on each side; supranasals
and posterosupranasals each separated by a
single minute median scale; scales immediately posterior to posterosupranasals only
slightly enlarged.
September 2008]
HERPETOLOGICA
Total number of differentiated supralabials
12/12 (L/R; 9/8–9 to center of eye), bordered
dorsally by one row of slightly differentiated
snout scales; total number of differentiated
infralabials 11/11 (8–9/9 to center of eye),
bordered ventrally by one row of enlarged
scales and 3 rows of only slightly differentiated
chin scales; mental triangular; mental and first
three infralabials on left (first four on right)
greatly enlarged and wrapping onto ventral
surfaces of chin, nearly twice the size of
individual infralabials 4–11; mental followed
by a pair of slender, elongate postmentals;
postmentals bordered posterolaterally by a
secondary pair, approximately one half the
length of first pair, and a tertiary pair of lateral
postmentals, one quarter the length of primary postmentals; postmental scale series bordered posteriorly by a single series of slightly
enlarged scales; followed immediately by a
sharp transition to nondifferentiated chin and
gular scales; postrictal scales slightly enlarged,
2 or 3 times the size of gular scales; remainder
of undifferentiated gular scales very small,
round, nonimbricate, juxtaposed (Fig. 3C).
Dorsal cephalic scales highly heterogeneous
and varied in shape, disposition, and distribution; scales of rostrum enlarged, round, oval to
subrectangular, and convex to conical; postnasal, prefrontal, and interorbital depressions
possess noticeably 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, flat to irregularly convex, reducing in size
posteriorly, interspersed with numerous
slightly enlarged conical tubercules, and
making a smooth transition through nuchal
region to small, juxtaposed, flat trunk scales
interspersed with enlarged, sharply conical
body tubercles; throat and chin scales small,
juxtaposed, nonimbricate; gular and pectoral
regions with enlarged cycloid, imbricate
scales, continuing to increase in size through
ventral abdomen, becoming very enlarged and
strongly imbricate.
Ornamental cephalic scalation includes
numerous conical tubercles on posterolateral
portions of head (temporal, supratympanic,
and postrictal regions) and a short curved
series of 4–5 enlarged, sharply conical (with
315
blade-like ridges) preorbital scales (Fig. 3A,
B); 38/42 circumorbitals in total, differentiated into the following distinct regions: (1)
minute precircumorbitals, (2) enlarged, flat,
squarish circumoribtals dorsoanterior to orbit
(7 on left, 8 on right), (3) transverse elongation
and modification into fringe-like points (spiny
ciliaria, 11 on left, 9 on right) across
dorsoposterior margin of orbit, gradually
reducing to (4) minute postcircumorbitals; a
total of 35 interorbital scales (straight line
distance from center of each eye, across both
eyelids).
Axilla–groin distance 49.2% of SVL; undifferentiated dorsal body scales round to
irregularly octagonal, nonimbricate, relatively
homogeneous in size, convex; dorsals sharply
transition to imbricate ventrals along the
ventrolateral adipose fold; dorsals lack surrounding interstitial granules but are interspersed with 18 irregularly transverse rows (33
paravertebral rows) of highly enlarged and
protuberant, to strongly conical dorsal body
tubercles; each dorsal tubercle with a raised,
thorn- to keel-like posteriorly-projecting
point; each tubercle surrounded by a circle
of very slightly enlarged adjacent dorsals; 109
transverse midbody dorsals; 202 paravertebrals between midpoints of limb insertions; 42
transversely arranged ventrals; scales on
dorsal surfaces of limbs larger than dorsals,
with interspersed enlarged tubercles extending down limbs and terminating 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 dimpled pore-bearing scales
(Fig. 4B) in a near-continuous preanofemoral
series (18 preanals on left, 19 on right;
separated from femorals by a single undifferentiated scale; 19 femorals on left; 18 on right)
each punctured by pore bearing dark orange,
exudate, arranged in a wavy, widely obtuse,
inverted ‘‘V’’ formation and continuing to just
before the patellar region; preanal pores 2–3
316
HERPETOLOGICA
times the diameter of femoral pores; preanals
situated atop a substantial preanal bulge that
folds over into precloacal region in preserved
specimen but was erect and protuberant in
life; preanals preceded by five similarly
enlarged but non-dimpled scale rows; preanals followed by five enlarged scales rows,
roughly forming a triangular patch of scales
before vent; femoral series lacks preceding or
following enlarged scale rows; scales lateroposterior to preanofemoral 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 lack interdigital webbing; subdigital scansors of
manus: 11/12, 10/11, 14/15, 14/16, and 13/12
on left/right digits I–V respectively; pes: 13/12,
11/12, 16/15, 15/15, and 15/14 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; remaining terminal claw-bearing phalanges compressed, with large recurved claws, not rising
free at distal end until they extend beyond
dilated hyperextensible portion of digit.
Tail base bordered by a single, greatly
enlarged conical postcloacal spur on each side
of vent; postcloacal swellings pronounced;
hemipenes incompletely everted, their distal
structures not evident; tail long, 98.0% of
SVL; tail not depressed, subcylindrical, divided into distinct fracture planes/autotomy
grooves (5 whorls or annulations); distal tail
portions not original; tail with clear autotomy
scar and distally regenerated portion; 11
caudal annulations before autotomy scar
(22.2 mm), 12–13 annulations estimated in
autotomized portion based on length
(47.1 mm), for an estimated total possible
annulation count of 25–26; TD (not including
basal postcloacal swelling) 75.9% of TW;
dorsal tail (following description based on
original portions of tail) as heavily adorned as
dorsum with tubercles; caudal tubercles
concentrated along posterior edge of caudal
annulation; caudals similar in size to dorsals;
[Vol. 64, No. 3
subcaudals enlarged, plate-like, 3–4 rows per
annulations; subcausals widely expanded to
cover majority of ventral surface of tail, or split
into a pair of subcaudals along posterior
margin of each annulation.
Variation.—Ranges of selected diagnostic
meristic characters are presented in Table 1.
Morphometric variation in mensural characters of adult paratypes is presented in Table 2.
The type series contains two large mature
males, two small but presumably mature
males (with swollen preanal buldges and
hemipenal tail base swellings), nine mature
females (all gravid, with large white oblong
eggs visible through skin of posteroventral
body wall) and eight juveniles/subadults, four
of which are males, with hemipenes partially
everted.
Coloration of holotype in ethanol.—Dorsal
ground coloration of head, body, tail, and
dorsal surfaces of limbs medium gray with
scattered indistinct light gray blotches and
black flecks; light tri-lobed bars traverse the
body in nuchal region, above limb insertions,
and across torso to base of tail; anterior
margin of tri-lobed light gray dorsal bars
bordered by faintly darker pigmentation;
posterior edge of light gray transverse bars
colored as dorsal trunk ground coloration.
Dorsal and lateral surfaces of head similar
to dorsal ground coloration, but with distinct
light gray spots; a light cream bar extends
posteriorly from the orbit; palpebra dark gray;
rostral and supralabials medium gray; infralabials very light gray.
Limbs colored as torso, lacking transverse
banding; dorsal surfaces of hands and feet
light gray; digits light gray with dark gray
surfaces of expanded distal portions; tail
medium gray with dark gray bands corresponding to two caudal annuli; regenerated
portion of tail flat gray.
Ventral head, neck, and torso light cream
ventral surfaces of limbs slightly darker;
ventral surfaces of digits (scansors) dark gray;
preanofemoal region white with orange pore
exudate; ventral surfaces of tail medium gray
(both original and regenerated portions).
Coloration of holotype in life.—(from photographs of holotype before preservation;
Fig. 2A–D) Dorsal ground coloration dark
purplish-gray to yellowish-brown, with indis-
September 2008]
HERPETOLOGICA
tinct dark gray to black blotches; dorsum with
six light cream tri-lobed bars traversing the
axilla-groin region, each bordered anteriorly
by an accompanying thin transverse wavy
black band (the darkest above insertion of
hind limbs) and posteriorly by normal trunk
ground coloration.
Dorsal nuchal region and posterior portions
of head very similar to trunk coloration but
with distinct, round, cream-colored spots;
similar cream bars radiate out from the orbit;
postrictal region flat gray; labial scales purplish-gray with cream spots on every third
labial scale; darker black blotches and flecks
congregate on snout, interorbital region, and
parietal region; infralabial region and chin
gray to light gray; snout scale purplish gray;
gular region light gray to brownish-tan.
Dorsal surfaces of limbs light gray with
numerous dark brown flecks; dorsal surfaces
of digits dark gray with slightly lighter claws;
dorsal and lateral portions of tail banded
alternating dark gray and cream (corresponding to tail annuli); distal autotomy regrowth
dark brown.
Ventral body and limbs yellow with scattered gray and dark brown flecks; preanofemoral region bright yellow with dark orange
pores; palmar and plantar surfaces of manus
and pes yellowish with light gray subdigital
scansors; ventral tail cream with brown
transverse bars, and solid brown ventral
coloration distal to autotomy scar and subsequent regrowth.
Color variation.—Our sample of four adult
males, a subadult male, two juvenile males,
nine adult females, and five juveniles of
undetermined sex exhibits moderate color
variation. The adult male paratype (KU
304825) and two female paratypes (KU
304809, 304836) have darker dorsal coloration
than that of the holotype, with very dark gray
anterior margins of the light transverse bars.
Two females (KU 304826, 3004849) have
much darker dorsal ground coloration than
do any of the other specimens. In these
specimens, the tri-lobed light gray transverse
bars take on a more striking, contrasting
appearance. Three adult female paratypes
(KU 304845, 304847, 304851) have a light to
medium gray, nearly patterenless dorsum,
with only faint vestiges of transverse banding.
317
Juveniles are patterned more brightly than
adults, with more intensely contrasting light
and dark dorsal and tail coloration. In some
juveniles (KU 304821) the tail banding
contrast is nearly extreme black and white.
Juveniles (KU 304814, 304821, 304829
304832) also possess cream spots on the
dorsal surfaces of articulations between adjacent phalanges. This gives the fingers and toes
a light and dark banded appearance. This
pattern is faint in the subadult paratype (KU
304807) and absent in all adults.
Ventral coloration is nearly invariant, with
the exception that some specimens have very
dark gray to black ventral tail surfaces (e.g.,
KU 304809, 304826, 304849) while others
have very light gray ventral tail surfaces that
are only slightly darker than ventral body
coloration (e.g., KU 304845, 304847). Only in
two juveniles is the black and white transverse
banded caudal pigmentation strong enough to
wrap around on to the ventral surface of the
tail (KU 304814, 304821).
Distribution and natural history.—The new
species is known only from Babuyan Claro
Island where it was collected low (,3 m) on
trunks and buttresses in primary dipterocarp
forest at low elevations, close to the island’s
coast. Given the island’s volcanic origin and
geological history of isolation (Marini et al.,
2005; McDermott et al., 1993) we do not
expect G. crombota to be distributed on other
islands in the Babuyan Island group, the
Batanes Island group, or mainland Luzon.
Our impression is that the new species is very
common at the type locality. No vocalizations
by this species were heard during our brief
visits to the type locality. The only other
gekkonids encountered during our three night
stay on Babuyan Claro were Hemidactylus
frenatus, Cyrtodactylus philippinicus, and
Luperosaurus macgregori (see Discussion in
Brown et al., 2007).
Etymology.—The specific epithet crombota
is a group of letters derived from the names
Crombie and Ota, and treated as a noun in
apposition. We employ the amalgam crombota
to jointly honor Ronald Crombie and Hidetoshi Ota in recognition of their collaborative
survey efforts in the Babuyan Islands (Ota and
Crombie, 1989; Ota and Ross, 1994) and their
continued work on the gekkonid fauna of this
318
HERPETOLOGICA
archipelago (H. Ota and R. Crombie, unpublished data). Suggested common name: Babuyan Claro Gecko.
DISCUSSION
The description of Gekko crombota brings
the total number of endemic Philippine Gekko
species to eight (ten total, when G. hokouensis
is excluded and the nonendemic G. monarchus and G. gecko are included). We are
certain that this number represents an underestimate of species diversity and we enumerate the following series of unresolved taxonomic issues that need to be addressed before
a full realization of Philippine gekkonid
diversity can be achieved. First, the Babuyans
and Batanes island groups require additional
survey work before we can be reasonably
certain that the total gekkonid fauna is known.
We are aware of at least four additional,
undescribed Gekko species in the Babuyans
alone and we suspect that several additional
undescribed species in the Babuyans and
Batanes await discovery on small isolated
islands surrounded by deep water. Second,
although morphologically distinct species are
still being described from other parts of the
archipelago (Roesler et al., 2006), the majority
of the Philippines’ more subtle or possibly
cryptic species diversity undoubtedly masquerades under the widespread species Gekko
mindorensis and Gekko monarchus. Preliminary molecular sequence data (C. Siler, A.
Diesmos, and R. Brown, unpublished data)
suggest that these taxa may be comprised of
numerous cryptic, unrecognized evolutionary
lineages, and potentially are worthy of specific
rank. At a minimum, the major Pleistocene
Aggregate Island Complexes of the Philippines (Brown and Diesmos, 2002; Brown et
al., 2007; Gaulke et al., 2007) might each be
expected to harbor endemic species diversity
in both of these species complexes. Third,
numerous deep water islands (e.g., not
connected to adjacent islands during the last
glaciations; Brown and Diesmos, 2002) have
not been exhaustively surveyed for herpetofauna and are thus ripe for the potential
discovery of additional Gekko species. Islands
is this category are are Lubang, Camiguin
Norte, Camiguin Sur, Calayan, Dalupiri,
Fuga, Masbate, Siquijor, Dinagat, Siargao,
[Vol. 64, No. 3
Sarangani, Maranduque, Coron, Busuanga,
Burias, Ticao, Semira, Semirara, Maestre de
Campo, Cuyo, Basilan, Jolo, Tawi-Tawi, and
many other similarly small, isolated landmasses. Fourth, it is clear that comprehensive
efforts to survey gekkonid fauna of the
Philippines would do well to target isolated
limestone karst areas and natural caves. Such
habitats are not only patchily distributed (and
thus can be expected to have promoted
evolutionary divergence via isolation) but are
also heavily imperiled and increasingly under
threat from overexploitation by humans for a
variety of natural resources (Clements et al.,
2006). The discovery of G. ernstkelleri in an
isolated limestone outcop on Panay Island (C.
D. Siler, personal observations; Roesler et al.,
2006) demonstrates that hidden gekkonid
diversity awaits field herpetologists willing to
target long-overlookecd limestone habitats.
Similarly, G. gigante (endemic to the limestone landbridge Gigante island group; W.
Brown and Alcala, 1978; R. Brown and Alcala,
2000) is proof that geological isolation (e.g.,
landmasses separated by deep water) may not
be necessary to promote gekkonid diversification if limestone habitats have been isolated
over geological timescales.
Finally, numerous isolated mountain ranges
on larger islands (Luzon, Palawan, SamarLeyte, Mindanao) can be expected to support
as yet undocumented gekkonid diversity.
Recent discoveries from the Sierra Madre
mountain range of Luzon (Brown et al.,
2000b, 2007; A. Diesmos, unpublished data)
suggest that these remaining forested regions
all warrant extensive biodiversity surveys.
Whatever the circumstances, we are certain
that the diversity of Philippine Gekko species
is substantially underestimated. Given the
undeniable fact that destructive exploitation
of Southeast Asian forests is partly related to
an ignorance of their biodiversity, it is crucial
that future faunal inventories throughout the
Philippines pay careful attention to habitats
known to harbor endemic gekkonids. We
would not be surprised if the eventual number
of known endemic Philippine Gekko species
were to double in the very near future.
Acknowledgments.—For the loans of specimens or
assistance while visiting museum collections, we thank
the following individuals and their respective institutions
September 2008]
HERPETOLOGICA
(museum abbreviations follow Leviton et al., 1985): A.
Resetar, M. Kearney, and H. Voris (FMNH); J. Vindum,
R. Drewes, and A. Leviton (CAS); J. Simmons and L.
Trueb (KU), A. Wynn, R. Wilson, R. Heyer, and K. de
Queiroz (USNM) and R. Sison (PNM). Support for
fieldwork was provided by the University of Kansas, and
the National Science Foundation. The Stearns Fellowship
of the California Academy of Sciences provided support
that allowed RMB and ACD to undertake multiple visits
to CAS. We thank the Department of the Environment
and Natural Resources, and the Protected Areas and
Wildlife Bureau (especially C. Custodio, T. M. Lim, and
A. Tagtag), for facilitating research and export permits for
this and related studies, KU IACUC for approving
research protocols, and M. Pedregosa, and M. A. Reyes,
N. Antoque, B. Fernandez, and J. Fernandez for untiring
assistance in the field. We also thank Municipal DENR
authorities of Calayan, Cagayan Province for logistical
support. Thanks are due to M. Garfield for scientific
illustration and J. Weghorst, A. Bauer, and an anonymous
reviewer for comments on previous versions of the
manuscript.
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.Accepted: 3 June 2008
.Associate Editor: Christopher Raxworthy
APPENDIX I
Comparative Material Examined
All specimens examined are from the Philippines.
Numbers in parentheses indicate the number of specimens examined for each species and museum abbreviations follow Leviton et al. (1985).
Gekko athymus.—(7) PALAWAN ISLAND, 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 Samariñana; Mt. Mantalingahan:
KU 309331–34.
Gekko ernstkelleri.—(10) PANAY ISLAND, ANTIQUE
PROVINCE, Municipality of Pandan, Barangay Duyong,
Duyong Hillside (5 ‘‘Mt. Lihidian’’), 300 m.a.sl.: PNM
9152–54; KU 300196–202.
Gekko gecko.—(13) LUBANG ISLAND, OCCIDENTAL
MINDORO PROVINCE, Municipality of Lubang, Barangay
Paraiso: KU 303960–72.
Gekko gigante.—(8) SOUTH GIGANTE ISLAND,
ILOILO PROVINCE, Municipality of Carles, Tantangan: CAS
124315–17 (paratypes); NORTH GIGANTE ISLAND,
[Vol. 64, No. 3
ILOILO PROVINCE, Municipality of Carles: CAS 124866–67
(Paratypes); Barangay Asloman: KU 305138–40.
Gekko hokouensis.—(1) ‘‘Philippines’’ FMNH 17812
(Luperosaurus amissus holotype).
Gekko mindorensis.—(56) NEGROS ISLAND, NEGROS
ORIENTAL PROVINCE, Himangpangon Cave, Manjayod:
CAS-SU 28656–60; GUIMARAS ISLAND, GUIMARAS
PROVINCE, Municipality of Buenavista, Barangay Old
Poblacion: KU 302721, 302725; NEGROS ISLAND,
NEGROS OCCIDENTAL PROVINCE, Municipality of Cauayan,
Barangay Camalandaan: 302722–24; MASBATE ISLAND, MASBATE PROVINCE, Municipality of Mandaon,
Barangay Poblacion: 302726–28; PANAY ISLAND, CAPIZ
PROVINCE, Municipality of Pilar, Barangay Natividad:
302729–32; LUBANG ISLAND: OCCIDENTAL MINDORO
PROVINCE, Municipality of Lubang, Barangay Vigo: KU
303913–16, 303917–951.
Gekko monarchus.—(3) PALAWAN ISLAND, PALAPROVINCE, ca. 1.5 km. W.S.W. of Iwahig: CAS-SU
28416; ca. 5 km SSE of Iwahig: CAS-SU 28496; ca. 7 km
WNW of Iwahig: CAS-SU 28554.
WAN
Gekko palawanensis.—(25) PALAWAN ISLAND, PAPROVINCE, 7 km WNW of Iwahig: CAS 17318; 8 km
W of Iwahig: CAS 17319; ca. 9 km W of Iwahig: CAS
17320–22; KU 30948, 309171, 309279–95, 309468.
LAWAN
Gekko porosus.—(4) BATANE ISLAND, BATANES
PROVINCE, 3 km ENE of Basco Town: USNM 266519,
291387; Mahatao: USNM 266517; ITBAYAT ISLAND:
CAS 60526 (holotype).
Gekko romblon.—(12) SIBUYAN ISLAND, ROMBLON
PROVINCE, Taclobo Barrio: CAS 139180–82 (paratypes);
ROMBLON ISLAND, ROMBLON PROVINCE, Municipality
of Romblon, Barangay Li-O: KU 302736–42, 303977–78.