Herpetologica, 64(2), 2008, 224234

E 2008 by The Herpetologists League, Inc.

A NEW SPECIES OF BENT-TOE GECKO (GEKKONIDAE:

CYRTODACTYLUS) FROM SULAWESI ISLAND, EASTERN INDONESIA
CHARLES W. LINKEM1,6, JIMMY A. MCGUIRE2, CHRISTOPHER J. HAYDEN3, MOHAMMED IQBAL
SETIADI4, DAVID P. BICKFORD5, AND RAFE M. BROWN1
1

Department of Ecology and Evolutionary Biology and Natural History Museum and Biodiversity Institute, University of
Kansas, Dyche Hall, 1345 Jayhawk Blvd, Lawrence, KS 66045, USA
2
Museum of Vertebrate Zoology, University of California Berkeley, Berkeley, CA 94720, USA
3
Museum of Natural Science, 119 Foster Hall, Louisiana State University, Baton Rouge, LA, 70803, USA
4
McMaster University, 1280 Main St., West Hamilton, Ontario, L8S 4K1, Canada
5
Conservation Ecology Laboratory, National University of Singapore, Block S2 14 Science Drive 4, Singapore 117543
ABSTRACT: A new species of Cyrtodactylus is described from Lore-Lindu National Park, Sulawesi Island,
Indonesia. It is distinguished from all other Cyrtodactylus by a unique suite of scalation characters and a
distinctive color pattern. The new species is the fourth Cyrtodactylus known from the island of Sulawesi and
one of two new species found in 2004. These recent discoveries suggest that the diversity of the herpetofauna
in Wallacea, a poorly studied biological hotspot, may be far richer than previously thought.
Key words: Cyrtodactylus; Gekkonidae; Indonesia; Lore-Lindu National Park; New species; Southeast
Asia; Squamata; Sulawesi

THE GENUS Cyrtodactylus contains 95 described species distributed throughout the

Indo-Australian Archipelago westward to India (Bauer and Henle, 1994). Although many
species recently have been reassigned to other
genera such as Tenuidactylus, Cyrtopodion,
Nactus, and Geckoella (Golubev and Szczerbak, 1985; Kluge, 1983, 1991, 1993, 2001;
Macey et al., 2000; Szczerbak and Golubev,
1984, 1986), the number of species currently
or formerly in this genus continues to grow.
New species have been recently described
from Myanmar (Bauer, 2002, 2003), Sri Lanka
(Batuwita and Bahir, 2005), Malaysia (Grismer, 2005; Grismer and Leong, 2005; Youmans and Grismer, 2006), Thailand (Bauer et
al., 2002, 2003; Pauwels et al., 2004), Vietnam
(Heidrich et al., 2007; Orlov et al., 2007;
Quang et al., 2007; Ziegler et al., 2002), and
southern Laos (David et al., 2004).
For the island of Sulawesi, Boulenger
(1897) and de Rooij (1915) listed three species
of Cyrtodactylus: C. fumosus, C. jellesmae,
and C. marmoratus. Based on overlap in pore
characters, Brongersma (1934) synonomized
C. fumosus with C. marmoratus thereby
reducing the number of species on the island
to two. Hayden et al. (2008) recently described a third species of Cyrtodactylus from
6

CORRESPONDENCE: e-mail, cwlinkem@[Link]

the southwestern peninsula of Sulawesi.

Herein we describe a fourth species of
Cyrtodactylus from Sulawesi that differs
dramatically from all known congeners.
MATERIALS AND METHODS
A herpetological biotic survey was conducted on Sulawesi between September and
December 2004. Specimens were tissued,
preserved in 10% buffered formalin and
transferred to 70% ethanol approximately
two months later. The following measurements (after Bauer, 2002) were made on
preserved specimens with dial calipers to the
nearest 0.1 mm: snoutvent length (SVL);
trunk length (TrunkL); crus length (CrusL);
tail length (TailL); tail width (TailW); head
length (HL); head width (HW); head height
(HH); ear length (EarL); forearm length
(ForeaL); orbit diameter (OrbD); nares to
eye distance (NarEye); eye to ear distance
(EyeEar); internarial distance (Internar); interorbital distance (InterOrb). Bauers (2002)
snout to eye distance (SnEye) is referred to as
rostrum length (RostL) to reflect the preferred definition of rostrum as the portion of
the head anterior to the orbit. In contrast,
snout length (SnL) is defined as the portion of
the head anterior to the nares.
Sex was determined by gonadal inspection
and scoring of prominent secondary sexual
224

June 2008]

HERPETOLOGICA

225

FIG. [Link] of holotype, MZB 7024, showing pattern and tail curling behavior.

characteristics. We scored the following scale

counts following Grismer (2005): postmentals
(and their degree of medial contact); supralabials; infralabials; number of longitudinal
tubercle rows; number of paravertebral tubercles; number of ventral scales; number and
type of subdigital lamellae on fourth toe.
Additional scale counts in this manuscript are:
number of spines on ventrolateral fold, count
of spines between fore- and hind limb along
the ventrolateral fold; number of caudal
annuli, count of annuli down length of tail.
For the recognition of the new species, we
adopted the General Lineage Species Concept of de Queiroz (1998, 1999) as the natural
extension of the Evolutionary Species Concept (Wiley, 1978). Application of lineagebased species concepts to island endemics is
straightforward because of the known history
of isolation of island populations (Brown and
Guttman, 2002; Brown and Diesmos, 2002).
We consider as new species morphologically
diagnosable forms for which the hypothesis of
conspecificity can be rejected.
Images of specimens were taken using a
copy stand and a manual-focus digital camera.
Four images were taken at different focal
lengths and then combined using the program
CombineZE to create one image with full
depth of field.

RESULTS
Cyrtodactylus spinosus sp. nov.
[Link] 7024 (BSI-FS 1694)
(Fig. 1, 2, 3), adult male, collected 8 November 2004 at 19:35 h, from Indonesia: Sulawesi
Island: Sulawesi Tenggah Province: Kabupaten Donggala: Kecematan Kulawi: Desa Mataue: Lore-Lindu National Park (01.44883 S,
119.99483 E) at 696 m. Collected by CJH,
RMB, JAM and CWL.
[Link] 70259 (BSI-FS
140811) collected 3 November 2004: 7025
adult male, 70268 adult females, 7028 with
two eggs. Specimen MZB 7029 collected 6
November 2004: adult male. All other data are
the same as for the holotype.
[Link] spinosus is distinguished from all other Cyrtodactylus species by the following characters: a row of
spines along ventrolateral body fold; six lateral
rows of small, unkeeled body tubercles, with
most ventral row intermixed with spines; two
spines on temporal region of head; 31 spineadorned annuli encircling original tail; tubercles on fore- and hind limbs; spines on
postantefemoral portion of hind limb. Additional characters distinguishing this species
include: proximal subdigital lamellae transversely expanded; 1921 subdigital lamellae

226

HERPETOLOGICA

[Vol. 64, No. 2

FIG. [Link] of holotype features. (A) Dorsal view of head showing temporal spines, (B) lateral view of head,
(C) ventral view of head showing complete separation of post-mentals, variable within the species. (D) Palmar surface of
right pes showing differential size between proximal and distal subdigital lamellae.

on toe IV; 3844 mid body ventral scales; most

scales in femoral region small, granular; 712
enlarged femoral series scales lacking pores;
presence of pre-cloacal groove in males
(absent in females); presence of pre-cloacal
pores (1213) in a chevron-shaped groove;
subcaudals not transversely expanded; three
chevron-shaped dark bands on a grayishbrown background.
Description of the [Link] male
SVL 70.9 mm. Head moderately long (HL/
SVL 0.3), wide (HW/HL 0.6), somewhat
depressed (HH/HL 0.3), distinct from neck,
and spade-shaped in dorsal profile; lores
weakly raised, prefrontal region concave,
canthus rostralis rounded and granular; rostrum short (RostL/HL 0.4) and narrow in
dorsal profile. Eye large (OrbD/HL 0.3); pupil
vertical with crenulated margin. Ear opening

tear-shaped, small (EarL/HL 0.05); eye-to-ear

distance slightly greater than diameter of eye.
Rostral scale rectangular, width twice height,
partially divided dorsally, bordered posteriorly
by large left and right supranasals and two
small internasals; external nares bordered
anteriorly by rostral, dorsally by large supernasal, posteriorly by three small postnasals,
and ventrally by the first supralabial. Supralabials square, 11/12 extending to center of
eye, first supralabial larger than remainder.
Infralabials 11/11 extending to posterior of
orbit; first five scales of series largest. Scales of
rostrum, lores, top of head, and occiput small
and granular; scales of occiput and top of head
with infrequent, large tubercles; spines present on temporals and gular regions. Dorsal
and ventral supraciliaries circular; mental
triangular, bordered laterally by infralabials

June 2008]

HERPETOLOGICA

FIG. [Link]-cloacal region of the holotype and

paratopotype (MZB 7026) showing the sexual dimorphism
in pre-cloacal pores. The male holotype (A) has a precloacal groove and pore-bearing scales. The female (B)
lacks a pre-cloacal groove and pores.

and posteriorly by left and right square

postmentals; postmentals not in contact,
separated by large gular scale. One slightly
enlarged and elongate row of sublabials
extending posteriorly to the third infralabial;
gular scales small and granular, grading
posteriorly into slightly larger, flatter, imbricate pectoral and ventral scales (Fig. 2).
Body relatively elongate (TL/SVL 0.4) with
moderate ventrolateral folds that contain
large, semiregular, spines; dorsal scales small
and granular interspersed with moderatesized, triangular, semiregularly arranged
keeled tubercles; tubercles extending from
occiput to anterior portion of tail; tubercles on
occiput irregularly spaced, those on nape and
anterior of body largest; approximately six
longitudinal rows of tubercles; 40 flat, imbricate ventral scales between ventrolateral body
folds, ventral scales larger than dorsals; two
rows of enlarged pre-cloacal scales bordering
pre-cloacal groove; 12 pre-cloacal pores occurring within groove.

227

Forelimbs slender, relatively short (ForeL/

SVL 0.2); granular scales of forearm slightly
larger than those of body; tubercles present;
scales on palmar surface slightly elevated;
digits well developed, inflected at basal
interphalangeal joints; subdigital lamellae
transversely expanded proximal to joint inflections, digits narrow distal to joints; claws well
developed, sheathed by dorsal and ventral
scales; relative lengths of fingers: IV . III .
II . V . I.
Hind limbs more robust than forelimbs,
moderately long (CrusL/SVL 0.2), covered
dorsally with flat, granular scales interspersed
with larger, raised tubercles; ventral scales of
femora oval and larger than dorsals; ventral
tibial scales granular, imbricate; nine enlarged
scales in pore-bearing femoral series, lacking
pits; scales on plantar surface oval, imbricate,
elevated; toes well developed, inflected at
basal interphalangeal joints; subdigital lamellae transversely expanded proximal to inflected joints, digits narrow distal to joints; nine
expanded subdigital lamellae, 12 unexpanded
subdigital lamellae on right toe IV; claws well
developed, sheathed by dorsal and ventral
scales. Relative lengths of toes: IV . V 5 III
. II . I (Fig. 2D).
Tail 90 mm long, complete, 4.3 mm in
width at base, tapering to a point; dorsals
and caudals granular; spines oriented with
recurved points facing posteriorly, numbering
four per caudal annulus, situated at the
posterior edge of each annulus; two small,
oval, smooth, postanal scales on either side of
tail base.
Coloration in [Link] ground color of
head, neck, trunk, limbs and tail brown with
dark mottling. Three chevron-shaped interleaved black and tan bands on the dorsal
surface between limbs (Fig. 1); pattern continues down tail, transitioning to stripes; trunk
lacking a pattern, but having numerous white
spines along ventrolateral body fold; nuchal
region covered with a black triangle, bordered
distally by a tan nuchal stripe extending from
eye to first tan chevron on nape; two tan-white
spines along tan nuchal stripe at apex of
postocular region; a postocular black patch
extends to insertion of forelimb ventral to
nuchal stripe; labial region mottled anteriorly,
transitioning posteriorly to two white subocu-

228

HERPETOLOGICA

[Vol. 64, No. 2

TABLE [Link] characteristics of the holotype and five paratopotypes. Abbreviations of measurements are the same
as in methods. Indications of character states in the table are presence of a character state (1), and absence of a character
state (0).
MZB

Sex
SVL
Postmentals
Degree of postmental contact
Supralabials
Infralabials
Longitudinal rows of tubercles
Ventrolateral spines
Paravertebral tubercles
Ventral scales
Expanded lamellae on 4th toe
Narrow lamellae on 4th toe
Enlarged pre-cloacal scale patch
Pre-cloacal groove
Pre-cloacal pores
Number of pre-cloacal pores
Femoral pore-like scales
TrunkL
CrusL
TailL
TailW
HL
HW
HH
EarL
ForeaL
OrbD
NarEye
RostL
EyeEar
Internar
Interorb

7024

7025

7026

7027

7028

7029

Mean

Standard
deviation

Male
70.9
2
0
11
11
8
9
26
40
9
12
1
1
1
12
9
30.7
14.0
90.1
4.4
22.1
14.3
8.6
1.1
10.4
6.0
7.0
8.5
5.7
2.4
7.7

Male
70.0
2
25%
9
8
8
12
25
44
7
12
1
1
1
13
12
31.1
11.2
58.7
5.1
20.8
13.4
8.2
0.9
9.5
4.8
6.7
9.0
5.5
2.5
7.6

Female
79.2
2
25%
10
9
8
12
30
40
8
12
1
0
0
13
12
34.9
13.2
95.8
5.3
23.2
15.4
9.1
1.0
11.5
6.4
7.5
9.5
7.2
2.9
9.1

Female
78.3
2
0
8
8
8
11
25
38
8
13
1
0
0
12
7
34.4
13.9
79.6
4.7
23.1
16.4
9.8
1.4
11.4
5.9
8.2
9.7
7.1
2.7
8.9

Female
83.2
2
25%
9
8
8
11
25
41
8
13
1
0
0
12
9
40.9
14.6
NA
4.3
23.6
16.1
9.6
1.5
11.7
5.9
7.7
10.1
7.3
2.9
9.2

Male
58.4
2
25%
9
7
8
11
26
42
7
12
1
1
1
13
9
27.5
10.2
NA
NA
18.2
11.5
6.8
1.4
8.6
4.3
5.8
7.5
4.8
2.2
6.2

73.3

33.3
12.9
54.0
4.0
21.8
14.5
8.7
1.2
10.5
5.6
7.2
9.1
6.3
2.6
8.1

8.9

5.0
1.7
8.2
0.5
2.2
2.0
1.2
0.2
1.3
0.8
0.9
1.1
1.1
0.3
1.3

lar bands separated by a black band; canthus

rostralis dominated by black scales; with
mottled brown/yellow scales on both sides of
canthal ridge; postnasal region yellow; supracilliary scales banded yellow and dark brown;
eye golden-brown with dark brown venation.
Limbs uniform brown with darker mottling;
digits banded with dark and light; most body
spines tan-to-white, becoming lightest at their
apices. Caudal spines match overlying banding pattern coloration (i.e., dark bands with
black spines); ventral ground color gray with
dark mottling; scales flecked with black spots
throughout ventral surface.
[Link] paratopotypes closely resemble the holotype. All specimens except
MZB 7029 are adults. There is sexual
dimorphism in pre-cloacal pore structure.
Males posses large, pore-bearing scales in a

chevron shape with a large groove; females

posses differentiated scales in the porebearing series (arranged in a chevron shape)
but without a groove and lacking pores
(Fig. 3). The numbers of differentiated scales
in the pore-bearing series do not differ
between the sexes. Specimen MZB 7028 has
two large eggs. Coloration is similar among all
specimens, with each possessing three chevrons across the back. Chevrons on MZB 7029
are less complete than others. Meristic
variation in the type series is presented in
Table 1.
Natural [Link] specimens were
found in secondary-growth forest (neighboring selectively logged first growth forest) of
Lore Lindu National Park. The first specimens were found at night on shrubs (#1.5 m
above the ground), although we searched in

June 2008]

HERPETOLOGICA

FIG. [Link] of Sulawesi Island showing the type

locality of Cyrtodactylus spinosus.

the same habitat for several more nights and

did not encounter any other lizards. Four
nights later, one individual was found on the
trunk of a tree at a height of 45 m. Although
the new species appears to be primarily
arboreal (possibly a canopy specialist that
was driven to lower forest strata by heavy
rains prior to collection), limited sampling
precludes confident inference of the species
preferred microhabitat.
[Link] species is only known
to occur at the type locality of Lore Lindu
National Park, Kecematan Kulawi, Kabupaten
Donggala, Sulawesi Tenggah Province, Sulawesi Island, Indonesia (Fig. 4). It is possible
that this species occurs outside of this area,
but not likely given the low abundance of
quality forest in Sulawesi. This species is likely
an island endemic and should be considered
threatened due to its limited range.
[Link] name is derived from the
Latin word spina which means spiny or
thorny in reference to the unique spines
possessed by this species.
Comparison to coastal Sunda Shelf, Philippine, Indonesian and Papuan [Link] spinosus can be distinguished
from all other congeners by the following
suite of characteristics: the presence of spines
along the ventrolateral fold, temporal region,
tail and postantefemoral portion of hind limbs

229

distinguish it from all other species; a

maximum SVL of 83 mm distinguishes it from
the larger species: C. aurensis, C. consobrinus, C. darmandvillei, C. derongo, C. irianjayensis, C. lateralis, C. louisiadensis, C.
mimikianus, C. novaeguineae, C. pequensis,
C. pulchellus, and C. tiomanensis; a minimum
SVL of 70 mm distinguishes C. spinosus from
the smaller species: C. annulatus, C. elok, C.
laevigatus, C. papuensis and C. semenanjungensis; the presence of pre-cloacal pores
distinguishes C. spinosus from C. darmandvillei, C. jellesmae, C. laevigatus, C. semenanjungensis, C. sermowaiensis and C. thirakhupti; the presence of a pre-cloacal groove
distinguishes it from all species except C.
annulatus, C. aurensis, C. cavernicolous, C.
marmoratus, C. papuensis, C. philippinicus,
C. pubisulcus, C. pulchellus, C. redimiculus,
C. semenanjungensis and C. tiomanensis;
absence of a continuous series of pre-cloacal
and femoral pores distinguishes it from: C.
angularis, C. chanhomeae, C. jarunjini, C.
loriae, C. louisiadensis, C. novaeguinea, C.
pulchellus, C. seribuatensis, C. thiakhupti;
absence of enlarged subcaudal scales distinguishes it from: C. angularis, C. aurensis, C.
baluensis, C. chanhomeae, C. consobrinus, C.
darmandvillei, C. derongo, C. ingeri, C.
intermedius, C. irianjayensis, C. jarunjini, C.
louisiadensis, C. malayanus, C. mimikianus,
C. peguensis, C. pulchellus, C. redimiculus, C.
sumonthai and C. thirakhupti; the presence of
broad subdigital lamellae distinguish it from:
C. tigroides and C. yoshii; the presence of
more than 17 subdigital lamellae distinguishes
it from: C. angularis; the presence of fewer
than 22 subdigital lamellae distinguishes it
from: C. cavernicolous, C. consobrinus, C.
derongo, C. ingeri, C. irianjayensis, C. louisiadensis, C. marmoratus, C. matsuii, C.
mimikianus, C. noveaguinea, C. philippinicus,
C. sermowaiensis, and C. yoshii; and the
presence of enlarged scales in the porebearing femoral series distinguishes it from:
C. angularis, C. annulatus, C. aurensis, C.
cavernicolous, C. elok, C. ingeri, C. intermedius, C. irianjayensis, C. jellesmae, C. laevigatus, C. lateralis, C. malayanus, C. matsuii,
C. papuensis, C. peguensis, C. philippinicus,
C. pubisulcus, C. quadrivirgatus, C. semenanjungensis, C. sermowaiensis, C. sumonthai, C.

spinosus
aaroni
agusanensis
angularis
annulatus
aurensis
baluensis
brevipalmatus
chanhomeae
cavernicolous
consobrinus
darmandvillei
derongo
elok
ingeri
intermedius
irianjayensis
irregularis
jarujini
jellesmae
laevigatus
lateralis
loriae
louisiadensis
malayanus
marmoratus
matsuii
mimikianus
novaeguineae
papuensis
pequensis
philippinicus
pubisulcus

Taxon

SVL

7083
7086.5
70103
?
4570
9295
7286
6473
79
6481
97121
85
105112
5668
6576
5781
147163
5671
70
6375
43
85
?
110122
7073
76
105
95
115129
6165
85
7492
5974

1
1
1
?
1
0
1
1
1
1
1
1
1
0
1
1
1
1
1
1
0
1
1
1
1
1
1
1
1
1
1
1
1

1
1
1
?
?
0
1
1
1
0
1
0
1
1
1
1
1
1
0
1
?
1
1
1
1
1
1
1
1
1
1
1
0

1
1
1
?
?
0
1
1
1
1
1
1
1
1
1
1
1
1
1
1
?
1
1
1
1
1
1
1
1
1
1
1
1

1
1
1
?
?
0
1
0
?
1
1
1
1
0
1
1
1
1
1
1
0
1
1
1
1
1
1
1
1
1
1
1
1

1
1
1
0
1
0
1
1
1
1
1

1
1
0
1
0
0
1
0

1
1
1
?
?
0
0
1
1
1
1
1

4043
3440
4651
4042
5060
4551
4045
3545
3638
5158
5865
3640
4648
44
4043
4050
3641
4146
28
3258
30
6064
4449
3038
5862
4050
51
3448
3546
?
2938
3642
4355

0
0
0
1
0
1
1
0
1
0
1
1
1
0
1
1
1
0
1
0
0
0
0
1
1
0
0
1
0
?
1
0
0

1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1

1821
2128
2328
15, 16
1823
1823
2123
1619
2123
2226
2328
17
2426
18, 19
2327
21
3135
19, 20
22
1925
14
21, 22
2024
2331
2123
23, 24
22
2228
2833
?
1618
2224
1722

0
?
0
?
?
0
1
1
?
0
0
1
1
0
0
1
1
0
1
0
0
?
1
1
0
1
1
?
1
1
?
0
0

10

0
1
1
1
0
0
1
1
0
0
1
?
1
0
0
1
1
1
1
0
0
?
1
1
1
1
0
1
1
1
0
0
0

11

12

47
1120
314
0
0
0
69
67
3234
0
16
18, 19
0
0
0
0
0
78
19
0
0
0
5370
3664
0
310
0
1012
3842
0
0
0
0

1
0
0
0
1
1
0
0
0
1
0
0
?
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
1
0
1
1

13

1
1
1
1
1
1
1
1
?
1
1
?
1
1
1
1
1
1
0
0
0
1
1
1
1
1
0
1
1
1
1
1
1

14

15

812
58
711
3
47
7
910
910
3234
4
910
0
0
8
8
8
716
8
19
0
0
13
5370
3664
810
1216
7
714
3842
8, 9
79
810
79

0
0
0
1
0
0
0
0
1
0
0
0
?
0
0
0
0
0
1
0
0
0
1
1
0
0
0
0
1
0
0
0
0

16

0
0
0
0
1
1
1
0
0
1
1
0
0
0
1
0
0
0
0
0
0
0
0
0
1
0
1
0
0
0
1
0
0

17

1
1
0
0
1
1
0
0
1
1
1
0
0
0
0
0
1
1
1
0
0
0
0
1
1
0
0
1
1
1
1
1
0

18

0
0
1
1
1
0
1
1
0
0
0
1
1
1
1
1
0
0
0
1
1
1
1
0
0
1
1
0
0
1
0
0
1

19

0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0

20

HERPETOLOGICA

56
K
L
8H
6
C
E
C
H
J
J
C

5
A
9B
C
4C
D
1D
E
F
1C
1
1G
1H
I
C
C
J
C

TABLE [Link] table of characters used to distinguish Cyrtodactylus spinosus from other Cyrtodactylus species. Indications of character states in the table are presence of
a character state (1), absence of a character state (0), and data missing from the literature (?). Characters are notated as 1tuberculation moderate to strong, 2tubercles on
forelimbs, 3tubercles on hind limbs, 4tubercles on head and/or occiput, 5tubercles on at least 1/3 of tail, 6number of ventral scales, 7enlarged median subcaudals,
8proximal subdigital lamellae broad, 9number of subdigital lamellae on toe IV, 10contact of posterior thigh scales abrupt, 11enlarged femoral scales, 12number of
femoral pores, 13pre-cloacal groove, 14enlarged pre-cloacal scales, 15number of pre-cloacal pores, 16pre-cloacal and femoral pores/scales continuous, 17reticulate
pattern on head, 18body banded, 19body blotched, 20body striped. Where derived from the literature, references are abbreviated by letters as follows: AGunther and
Rosler (2002), BBrown and Alcala (1978), CYoumans and Grismer (2006), DGrismer (2005), ETaylor (1963), FBauer et al. (2003), Gde Rooij (1915), HBrown
and Parker (1973), IDring (1979), JBrongersma (1934), KDarevsky (1964), LWerner (1896), MBauer et al. (2002), NSmith (1925), ODas and Lim (2000).

230
[Vol. 64, No. 2

0
1
?1
0
0
0
0
0
0
0
0
0
0
0
?
1
1
1
0
1
0
0
0
1
0
1
0
1
F
O
C

M
N

C
C

pulchellus
quadrivirgatus
redimiculus
semenanjungensis
seribuatensis
sermowaiensis
sumonthai
sworderi
thirakhupti
tigroides
tiomanensis
yoshii

C
C

115
5171
78
5969
75
6893.5
71
6377
7280
83
84
7596

1
1
?
1
1
1
1
1
1
1
1
1

1
1
?
1
1
1
1
1
1
1
1
1

1
1
?
1
1
1
1
1
1
1
1
1

1
1
?
1
1
1
1
1
1
?
1
1

1
1
?
?
1
0
0

3335
3442
?
4853
2839
3038
3336
4248
3740
34
3640
5058

1
0
1
0
0
0
1
0
1
0
0
0

1
1
?
1
1
1
1
1
1
0
1
0

19, 20
19, 20
2024
1721
1922
22
18
20
20, 21
1923
2022
2530

1
0
?
1
1
0
?
1
1
?
1
0

1
1
?
0
1
0
0
1
1
0
1
0

1418
0
89
0
4245
0
0
0
0
57
0
0

1
0
1
1
0
0
0
0
0
0
1
0

1
1
1
1
1
0
1
1
1
1
1
0

68
04
58
0
4245
0
2
5, 6
0
89
35
812

1
0
?
0
1
0
0
0
1
0
0
0

0
0
?
0
0
0
0
0
1
0
0
0

1
0
?
1
0
0
1
0
1
1
1
0

20
18
17
16
15
14
13
12
11
10
9
8
7
6
5
4
3
2
1
SVL
n
Taxon

TABLE [Link].

HERPETOLOGICA

19

June 2008]

231

sworderi, C. thirakhupti, C. tiomanensis and

C. yoshii (Table 2).
DISCUSSION
We consider herpetological diversity on the
island of Sulawesi to be underestimated owing
to a lack of attention by taxonomists and
comprehensive survey work (Brown et al.,
2000; Evans et al., 2003a,b; Iskandar and Tjan,
1996; McGuire et al., 2007). Recent biotic
survey and inventor efforts have found new
species on Sulawesi and adjacent islands
including two new species of flying lizard,
genus Draco (McGuire et al., 2007) and
additional species that are in need of description. Genetic analyses of some groups: Limnonectes (Evans et al., 2003a), Bufo celebensis
(Evans et al., 2003b) and Lamprolepis (unpublished data) have revealed cryptic lineages. Exploration of the northern, central and
southwestern regions of the island have
resulted in the discovery of two new species
of Cyrtodactylusa new species described by
Hayden et al. (2008) and C. spinosus. We
suspect that additional species of Cyrtodactylus await discovery and several taxa currently
masquerade under the widespread species C.
jellesmae.
Among SE Asian members of the genus,
Cyrtodactylus spinosus is morphologically
distinct, owing to the presence of spines,
unique scalation, and distinct color pattern; it
is consequently unlikely to be confused with
any other species. Indeed, this species is so
unique we cannot speculate on its closest
relatives within the genus.
Discovery of this new species within LoreLindu National Park further emphasizes the
need for conservation efforts targeting this
major center of SE Asian biodiversity. Despite
the parks having been designated as a
protected area, farmers in the area continue
to encroach the parks boundary, using slashand-burn methods to clear land for agriculture. Both small-scale timber poaching and
large-scale logging operations continue to
degrade the park, especially along its unguarded southern boundary (Bickford et al.,
2007). Improved security of the park boundary is needed to protect this stronghold of
biodiversity that likely houses a large diversity
of endemic species yet to be discovered.

232

HERPETOLOGICA

[Link] thank N. Andayani (Univ. of

Indonesia; Wildlife Conservation Society), J. Supriatna
(Univ. of Indonesia; Conservation International), Mumpuni (Museum Zoologicum Bogoriense, LIPI), and A.
Riyanto (Museum Zoologicum Bogoriense, LIPI) for their
assistance in obtaining permits to work in Indonesia, as
well as with other logistical issues. We thank the following
government officials who provided research and export
permits, as well as specimen loans: A. Budiman (formerly
of LIPI), S. Prijono (former Director of Museum
Zoologicum Bogoriense). For assistance and companionship in the field, we are grateful to F. Chain, D. Halliwel,
A. Rosyid, Shobi, and Ted Townsend (plus others). For
discussion and/or comments on the manuscript, we thank
L. Trueb. This work was supported by the National
Science Foundation (DEB-BSI 0328700).

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APPENDIX
Specimens Examined
Codes for institutions are CAS, California Academy of
Sciences; KU, University of Kansas Natural History
Museum; LSUMZ, Louisiana State University Museum
of Zoology; MVZ, Museum of Vertebrate Zoology,
University of California, Berkeley; MZB, Museum Zoologicum Bogoriense, Bogor Indonesia; NMPNG, National
Museum of Papua New Guinea; UPNG, University of
Papua New Guinea; CCA, Christopher C. Austin; RMB,
Rafe M. Brown; JAM, Jimmy A. McGuire. Field collection
numbers are in parentheses.
Cyrtodactylus agusanensis: PHILIPPINES: MINDANAO:
Province of Davao del Norte: 70 km S Bislig: LSUMZ
41601, 41602 (males), LSUMZ 41603 (female), LSUMZ
41604, 41605 (males); DINAGAT: Province of Surigao del
Norte: Municipality of Loreto: KU 3055645; SAMAR:
Province of Eastern Samar: Municipality of Taft: Barangay
San Rafael: KU 305569. Cyrtodactylus annulatus: PHILIPPINES: MINDANAO: Province of Davao del Norte:
70 km S Bislig: LSUMZ 41606 (female), 41608, 41609
(males). Cyrtodactylus baluensis: EAST MALAYSIA:
SABAH: Kina Balu Peak: CAS 25626 (male). Cyrtodactylus
cavernicolus: EAST MALAYSIA: SARAWAK: Niah Cave:
CAS 23726 (male). Cyrtodactylus consobrinus: EAST
MALAYSIA: SARAWAK: Bintulu District: Sungei Seran:
CAS 105993 (male); Kapit District: Sungai Mengiong:
MVZ 11782 (male); BRUNEI: Temburang District: Sungai
Belalang: LSUMZ 55833 (male); INDONESIA: PROPINSI
SUMATERA UTARA: Bukit Lawang: Bahorak: MZB 4355
(male), 4356 (juvenile). Cyrtodactylus darmandvillei:
INDONESIA: PROPINSI NUSA TENGGARA BARAT: Pulau
[Link] 81732 (JAM 3176) (male). Cyrtodactylus derongo: PAPUA NEW GUINEA: WESTERN PROVINCE:
Derongo: NMPNG R23452 (female). Cyrtodactylus

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HERPETOLOGICA

jellesmae: INDONESIA: PROPINSI SULAWESI SELATAN:

Anabanua: JAM 5628, 5631 (male); Harapan: JAM 5643
(female); Takalasi: JAM 5670, 5671, 5678 (male), 5677
(female); Maroangin: JAM 5680, 5683, 5684, 5686, 5688,
5704 (males), 5681, 5682, 5685, 5687 (females); Enrekang:
JAM 5705, 5747, 5749, 5768, 5769, 5771 (males), 5770,
5772, 5773 (females); Tapung: JAM 5783, 5784 (females);
Pecinong: JAM 5850, 5851 (females), 5852 (juvenile);
Mariorilau: JAM 5892, 5895, 5897, 5899 (males), 5893,
5896, 5898 (females), 5900 (juvenile); PROPINSI SULAWESI
BARAT: Kabiraan: JAM 63416343, 6346 (males), 6339,
6340, 6344, 6345, 6347 (females), 6338 (juvenile);
PROPINSI SULAWESI TENGGAH: Donggala Kabupaten:
LSUMZ 8403 (male), 8400 (female), 8401 (juvenile); Poso
Kabupaten: LSUMZ 8402 (male); Luwuk Kabupaten:
LSUMZ 8405 (male), 8404, 8406 (females). Cyrtodactylus
loriae: PAPUA NEW GUINEA: CHIMBU PROVINCE:
Karimui: CAS 117964 (male), CAS 118018 (female),
NMPNG R23625, R8347, R8348; PAPUA NEW GUINEA: UPNG 5687 (male); WESTERN HIGHLANDS PROVINCE:
Kaironk: UPNG 0976 (female); MOROBE PROVINCE: Wau.:
NMPNG 24730. Cyrtodactylus louisiadensis: PAPUA
NEW GUINEA: BOUGAINVILLE: Kunua: KU 98481; MILNE
BAY PROVINCE: Halowina: CCA 4096 (female), 4426, 4427,
4582, (males), 4428 (juvenile). Cyrtodactylus malayanus:
INDONESIA: PROPINSI KALIMANTAN TIMUR: Maruwai:
MZB 29272929 (males); PROPINSI KALIMANTAN BARAT,

[Vol. 64, No. 2

Taman Nasional Bentuang Karimun: MZB 3920 (female).

Cyrtodactylus marmoratus: INDONESIA: JAVA: Djurung
Sriti: Mount Merapi: KU 153796; PROPINSI JAVA BARAT:
Sukabumi: LSUMZ 81868 (male), LSUMZ 8186981870
(females), LSUMZ 8187181872 (males), LSUMZ 81873
(female), LSUMZ 81874 (subadult male), LSUMZ 81875
(female), LSUMZ 81876 (male). Cyrtodactylus novaeguineae: PAPUA NEW GUINEA: SANDAUN PROVINCE: Utai:
CCA 3142 (female), 3415 (juvenile); Bewani Station: CCA
3674 (male). Cyrtodactylus quadrivirgatus cf.: INDONESIA: SUMATRA: Propinsi Bengkulu: 46 km East of
Bengkulu:.MVZ 239338 (male), 239578 (female). Cyrtodactylus philippinicus: PHILIPPINES: LUZON: Camarines del Sur Province: Municipality of Naga: KU 305571;
NEGROS: Negros Oriental Province: Municipality of
Dumagete, Baragay Valencia: KU 305572; LUBANG:
Occidental Mindoro Province: Municipality of Lubang,
Barangay Vigo: KU 303846, 303844. Cyrtodactylus
sermowaiensis: PAPUA NEW GUINEA: SANDAUN PROVINCE: Utai: CCA 3108, 3143, 3407, 3411, 34483451, 3506,
3507, 3517 (males), 2895, 2911, 3034, 3144, 3188, 3224,
3233, 3358, 3359, 3490, 3530 (females); Bewani Station:
CCA 3603 (male), 3604 (juvenile); MADANG PROVINCE:
Sapi Creek: NMPNG R22712; South Naru: NMPNG
R2480324806; EAST SEPIK PROVINCE: Maprik: NMPNG
R22796; SANDAUN PROVINCE: Idam River: UPNG 3914
3916.