Zootaxa 5468 (2): 361–378 ISSN 1175-5326 (print edition)

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Copyright © 2024 Magnolia Press
Article ZOOTAXA
ISSN 1175-5334 (online edition)
https://doi.org/10.11646/zootaxa.5468.2.7
http://zoobank.org/urn:lsid:zoobank.org:pub:D794AFA5-EDE1-436B-8A19-E1841FCBA942

Description of a new sandstone-dwelling species of genus Cnemaspis (Sauria:

Gekkonidae) from Gunung Santubong National Park, southwestern Sarawak,
Malaysia
TAKAKI KURITA1*, KANTO NISHIKAWA2,6, MOHAMAD YAZID HOSSMAN3, TAKAFUMI MIZUNO2,7,
HIROTOSHI SATO2,8 & MELVIN GUMAL4,5
1
Natural History Museum and Institute Chiba, 955-2 Aoba-cho, Chuo-ku, Chiba 260-8682, Japan
� [email protected]; https://orcid.org/0000-0002-2857-3834
2
Graduate School of Global Environmental Studies, Kyoto University, Yoshida Honmachi, Sakyo-ku, Kyoto 606-8501, Japan
3
Research & Development Division, Forest Department Sarawak, Km10 Jalan Penrissen, 93250 Kuching, Sarawak, Malaysia
� [email protected]; https://orcid.org/0000-0002-1230-9131
4
Sarawak Forestry Corporation, Jalan Sungai Tapang, Kota Sentosa, Kuching 93250, Sarawak, Malaysia
5
School of Engineering and Science, Swinburne University of Technology, Sarawak Campus. Jalan Simpang Tiga, 93350 Kuching,
Sarawak, Malaysia
� [email protected]; https://orcid.org/0000-0001-7872-5907
6�
[email protected]; https://orcid.org/0000-0002-6274-4959
7�
[email protected]; https://orcid.org/0000-0001-7535-3052
8�
[email protected]; https://orcid.org/0000-0003-4489-6569
*Corresponding author

Abstract

A newly discovered sandstone-dwelling species of the rock gecko, genus Cnemaspis, is described from Santubong National
Park, 25 km north of Kuching, southwestern Sarawak, Malaysian Borneo. Molecular phylogenetic analysis confirmed that
Cnemaspis puterisantubongae sp. nov. is closely related to other Cnemaspis species in southwestern Sarawak, except for
C. kendallii, although these relationships remain unclear. The new species differs from closely related species in having
a moderate snout–vent length of up to 66.7 mm; 10–14 precloacal pores in males; 9–14 obviously-convex precloacal
pore-bearing scales in females; enlarged, smooth, flat median subcaudal scales; sharp-edged, broad black markings on
the trunk dorsum; and a black–gray banding pattern on the posterior part of the original tail, with white median subcaudal
scales in males. This discovery of this new species from Santubong National Park, which is highly accessible due to its
proximity to Kuching, highlights the need for extensive inventory surveys throughout the national parks of Sarawak.

Key words: Borneo, Cnemaspis nigridia group, saxicolous, Puteri Santubong, PUBS, SATREPS

Introduction

The spatial distributions of animal species can be regulated by their substrate environments. Geckos, which include
a large number of saxicolous and arboreal species, frequently occur exclusively on specific substrate types. For
example, genus Cnemaspis, a polyphyletic group distributed in South Asian to Southeast Asian tropical regions
(Pyron et al. 2013; Grismer et al. 2014; see Malonza & Bauer 2022 for a genus-level revision), comprises more than
200 species (Uetz et al. 2023). The habitat types of the Southeast Asian members of Cnemaspis were compiled by
Grismer et al. (2014), who reported that 38 of 47 listed species were distributed in specific habitats (e.g., limestone,
granite, vegetation, and terrestrial), of which 32 are limestone and/or granite specialists. Indeed, karst areas are
presumed to generate and preserve the diversity of gecko species (e.g., Luu et al. 2016; Grismer et al. 2020).
In Borneo, Cnemaspis species have been recorded in both Sarawak, Malaysia and West Kalimantan, Indonesia
(Grismer et al. 2014; Maulidi et al. 2019; Maulidi et al. 2019; Nashriq et al. 2022), both of which are located on
the western part of the island. Recent studies by Nashriq & Das (2021) and Nashriq et al. (2022) revealed the
underestimated species diversity of genus Cnemaspis in karst environments of Sarawak; four of eight Bornean

Accepted by A.M. Bauer: 20 May 2024; published: 12 Jun. 2024 361

Cnemaspis species (Cnemaspis paripari Grismer & Chan, 2009; Cnemaspis matahari Nashriq, Davis, Bauer &
Das, 2022; Cnemaspis sirehensis Nashriq, Davis, Bauer & Das, 2022; and Cnemaspis lagang Nashriq, Davis, Bauer
& Das, 2022) are limited to karst habitats. However, the diversity of this genus on other substrates in this region
remains poorly understood. The granite-related species Cnemaspis nigridia (Smith, 1925) and the habitat generalist
Cnemaspis kendallii (Gray, 1845) have been found in southwestern Sarawak, and Cnemaspis dringi Das & Bauer,
1998 has been found in central Sarawak. Additionally, Cnemaspis leucura Kurita, Nishikawa, Matsui & Hikida,
2017 was recently found in a sandstone area of southwestern Sarawak (Das 2017; Kurita et al. 2017).
In 2022, a Cnemaspis gecko was photographed in Gunung Santubong National Park, Kuching, southwestern
Sarawak (Fig. 1), during a field survey under the Project on Development of Management Systems for Multiple
Utilization of Biodiversity in the Tropical Rainforests at the Protected Areas in Sarawak (PUBS). Gunung Santubong
mainly comprises several types of sandstone formations (Hazebroek 2019). Based on one specimen, the gecko species
C. kendallii, which has a pale gray, pale yellow, or brown background colour with lighter and darker markings on
the dorsum, was recorded in Gunung Santubong (Das et al. 2019; Nashriq & Das 2021). However, this individual
had gray dorsal colouration in the middorsal region and obvious dark yellow in the paravertebral and flank regions;
although it superficially resembled C. leucura this species has been recorded only from other mountain regions
(Kurita et al. 2017; Nashriq et al. 2022). Therefore, we conducted a further investigation in the national park and
attempted to verify the taxonomy of this population. Morphological and genetic analyses allowed us to confirm that
the population was a previously unrecognised evolutionary branch of Bornean Cnemaspis. Therefore, we describe
it as a new species.

Materials and methods

Genetic analyses—The phylogenetic positions of the taxonomically unassigned Cnemaspis geckos collected from
Santubong National Park were inferred based on the NADH dehydrogenase subunit 2 (ND2) gene from mitochondrial
DNA (mtDNA) and recombination activating gene 1 (RAG1) from nuclear DNA. Total DNA was extracted from
ethanol-preserved liver tissues of unidentified and other putatively closely related Cnemaspis species using the
DNeasy Blood & Tissue Kit (Qiagen). The ND2 and RAG1 genes were amplified through standard PCR using the
primer pairs M112F–M1123R (Doughty et al. 2012) and R13–RAG1-r-Squamata (Groth & Barrowclough 1999;
Kurita et al. 2020), respectively.
The ND2 and RAG1 sequences of the putatively undescribed species were determined using an Illumina
sequencer in the Research & Development Division, Forest Department Sarawak because Sanger sequencers could
not be accessed due to the lack of a necessary export permit for materials. PCR products were processed to create
Illumina sequencing libraries using DNA Prep (Illumina), with one-fourth of the amount of reagent recommended by
the manufacturer. In this process, PCR amplicons were randomly fragmented to approximately 300-bp molecules and
adapters including sample-specific indices and flow cell-binding sites were ligated onto each end of the fragments.
The library was sequenced using 75 bp paired-end reads via MiSeq (Illumina), producing a randomly selected 2 ×
75-bp region of the target gene as a read. For all other samples, amplicons were processed using a standard cycle
sequencing reaction with BigDye terminator v3.1 (Thermo Fisher Scientific) and sequenced using a ABI PRISM
3130xl Genetic Analyzer (Thermo Fisher Scientific).
Raw reads produced by the Illumina sequencer were processed for quality control using Fastp (Chen et al. 2018).
Bases with low quality scores (4-base sliding window average of Q < 20) were cut from the 3´-end of each read,
and reads with ≥ 30% low-quality bases, five or more undetermined bases, and/or length < 50 bp were discarded;
these steps were followed by broken mate pair removal. The processed reads were mapped to a ND2 sequence of C.
nigridia in the DDBJ/EMBL/GenBank database (accession no. LC158342) and a RAG1 sequence of Gekko gecko
(AY662625) using the “snap-aligner paired” function in SNAP (Zaharia et al. 2011). Read pairs uniquely mapped to
the reference were selected and visualised using the Integrative Genomic Viewer (Robinson et al. 2011). Majority
consensus sequences of each individual were obtained from the alignments. The obtained sequences were combined
with ND2 and RAG1 sequences of putatively closely related species from the DDBJ/EMBL/GenBank database,
then aligned using MAFFT (Katoh & Standley 2013) and the “tranalign” function of EMBOSS (Rice et al. 2000).
Newly obtained sequences from this study were deposited in the database (accession nos. LC782916–782937), and
sequences analysed below are listed in Supplementary Data 1.

362 · Zootaxa 5468 (2) © 2024 Magnolia Press KURITA ET AL.

 Maximum likelihood trees were estimated for ND2, RAG1, and a concatenated assembly of these genes using
IQ-TREE (Nguyen et al. 2015). Nucleotide substitution models for the first, second, and third codons of ND2 and
RAG1 were separately estimated using ModelFinder (Kalyaanamoorthy et al. 2017). Support values for the internal
branches were evaluated using 1000 ultrafast bootstrap replicates and 1000 SH-aLRT replicates (Guindon et al.
2010; Hoang et al. 2018).
Morphological examination and analyses—We examined morphologies of Bornean Cnemaspis specimens
deposited in the following collections: Sarawak Forest Corporation (SFC), Sarawak Research Collection, Sarawak
Forest Department (SRC), Graduate School of Human and Environmental Studies, Kyoto University (KUHE),
Zoological Collection of Kyoto University (KUZ), and Lee Kong Chian Natural History Museum (ZRC). We did not
examine specimens of C. dringi and C. sirehensis; quantitative and qualitative characteristics of these species were
obtained from the work of Nashriq et al. (2022). The following specimens were examined in this study: Cnemaspis
sp. (Santubong), SFC-GRB-00515–00519, and 00528–00531; C. nigridia, KUZ-R 27082, 27091, 27093, 27095,
27101–27104, 27127–27129, 27133, 27134, 27137, 27144–27148, and 27164; C. paripari, ZRC 2.6813, 2.6814,
KUHE 57121, 57122, 57124, 57268, 57285–57287, 57289–57291, 1808-S109, and 1808-S123–1808-S126; C.
matahari, KUHE 57447–57451; C. leucura, KUHE 57421, 57422, 57423, 57424, 1808-S210, 1808-S211, 1808-
S212, SRC 00021 (KUZ-R 70773), 00022 (KUZ-R 70774), and 00329 (KUHE 55469); C. lagang, KUHE 56015,
59162–59164, 60320, 1808-S15, 1808-S16, 1808-S83, and 1808-S84; C. kendallii, KUZ-R 12161, 12162, 12164,
12165, 12332, 12334, 12336, 12425, 12426, 12885, 12886, 12888, 27059, 27060, 27065, 27085–27087, 27100,
27110, 27111, 27141, 27149, 27157, 27158, 27286, 48609, and 56042.
Our examination of morphometric and meristic characters mainly followed the methods of Grismer et al. (2014)
and Nashriq & Das (2022). Morphometric characters were measured using dial calipers to the nearest 0.1 mm;
meristic and several qualitative characteristics were observed with a microscope. Bilateral morphometric characters
were measured from the left side of body unless damaged, and bilateral meristic characters were counted on each
side.
Our morphometric measurements included snout-vent length (SVL), head length (tip of snout to occipital
protrusion), head width, head depth (deepest part of head; usually immediately posterior to the eyes), head depth at
the level of the centre of the eye, snout length (snout tip to anterior edge of orbit), beak length (tip of snout to most
convex point of snout), beak depth (most convex point of snout to ventral surface of head), jaw length (snout tip to
externally visible end of mouth slit; usually immediately posterior to the centre of the eye), jaw width (between the
ends of the mouth slit), lower jaw length (snout tip to lower jaw angle), eye diameter (diameter of the visible surface
of the eyeball), interorbital distance (narrowest part of the parietal bone), anterior interorbital distance (breadth the
of bony part immediately anterior to the edges of the orbits), nape width, axilla–groin length, tail length, brachial
length, antebrachial length, femur length, crus length, pes length (interdigital slit between first and second toes
to the posterior edge of the pes), first and fourth finger lengths, and first and fourth toe lengths. The ratio of each
measurement to SVL was calculated, except for the ratios of the nape width, due to unstable measurement points
defined by soft parts; tail length, because the tail was broken or regenerated in a significant number of specimens;
and brachial length, which was difficult to measure in some specimens due to the fixation style. Principle component
analysis for these ratios was only conducted for individuals with SVL ≥ 80% that of the largest individual of each
species, using R v4.2.0 (R Core Team 2022).
The meristic characters examined included supralabial, infralabial, postnasal (granular scales bordered with
nasal), internasal (between supranasals and bordered with rostral), postmental scales; paravertebral tubercles
between the forelimb and hind limb insertions; precloacal pores or pore-bearing scales; poreless scales separating
the precloacal pore series; postcloacal tubercles; and subdigital lamellae from the first to fifth fingers and toes.
Scale conditions, such as the distribution of tubercles and other enlarged scales, keeling, and arrangement, were
also observed.

Results

Based on maximum likelihood phylogenetic inference performed using concatenated 1041-bp ND2 and 1038-bp
RAG1 sequences, the unidentified Cnemaspis individuals sampled from Santubong National Park were confirmed
to belong to the C. nigridia species group sensu Grismer et al. (2014) (hereinafter referred to as the nigridia group).

New Cnemaspis from Santubong Zootaxa 5468 (2) © 2024 Magnolia Press · 363
The Santubong population was phylogenetically distinct from any known Bornean species analysed in this study,
with significant genetic divergence (15.71–17.20% and 0.40–0.67% uncorrected p distances for ND2 and RAG1,
respectively; Table 1 and Fig. 1). The phylogenetic position of the population was clearly distinct from other
populations in the maximum likelihood trees based on each gene (Supplementary Data 2). The nigridia group
consisted of C. nigridia, C. paripari, C. matahari, C. sirehensis, C. leucura, and the Santubong population (ultrafast
bootstrap/SH-aLRT = 100/100); the other examined Bornean species, C. kendallii and C. lagang, were identified
as sister groups of non-Bornean species. Phylogenetic relationships among species in the nigridia group were
poorly resolved. Cnemaspis nigridia, C. paripari, the Santubong population, and a well-supported clade including
C. leucura, C. matahari, and C. sirehensis (100/100) formed a polytomy. Neither of the limestone-specific nor the
sandstone-specific species in the nigridia group were monophyletic.

Table 1. Uncorrected p distances between the Cnemaspis nigridia-group species including the Santubong population.
The genetic distances based on ND2 and RAG1 genes are shown below and above diagonal, respectively. Numbers in
parentheses indicate standard error.
Santubong nigridia paripari leucura matahari sirehensis
Santubong 0.0067 (0.0024) 0.0057 (0.0022) 0.0040 (0.0019) 0.0050 (0.0022) -
nigridia 0.1594 (0.0114) 0.0067 (0.0024) 0.0050 (0.0021) 0.0060 (0.0024) -
paripari 0.1622 (0.0115) 0.1685 (0.0115) 0.0035 (0.0017) 0.0045 (0.0021) -
leucura 0.1571 (0.0113) 0.1420 (0.0113) 0.1416 (0.0102) 0.0010 (0.0010) -
matahari 0.1637 (0.0119) 0.1502 (0.0113) 0.1451 (0.0104) 0.0904 (0.0083) -
sirehensis 0.1720 (0.0122) 0.1610 (0.0119) 0.1512 (0.0110) 0.1110 (0.0099) 0.1016 (0.0099)

FigUre 1. The type localities of the nigridia group of the Bornean Cnemaspis (thus not shown for C. lagang and C. kendallii)
and phylogenetic relationship of its members. The map is tilted 45° counterclockwise. Igneous and sedimentary bases within
Sarawak were shown in light and dark gray, respectively. Support of the internal branches of the maximum likelihood tree was
evaluated by ultrafast bootstrap/SH-aLRT values, and asterisks indicate those higher than 95/95. The black, brown, and gray
bands under the tips of the tree indicate granite, sandstone, and limestone dwelling species, respectively.

364 · Zootaxa 5468 (2) © 2024 Magnolia Press KURITA ET AL.

 The results of our principle component analysis based on body proportion (Fig. 2) demonstrated some overlap
between C. nigridia and the Santubong species; they clearly differed from C. paripari, C. matahari, and C. leucura
in having a larger head (head length/SVL = 26.4% in C. nigridia and the Santubong population versus 25.3% in
the remaining species on average; head width/SVL = 18.0% versus 17.2%), a longer beak (beak length/SVL =
8.0% versus 7.4%), and shorter antebrachium/crus and digits (antebrachial length/SVL = 17.7% versus 18.9%;
crus length/SVL = 21.0% versus 22.2%; fourth finger/SVL = 15.4% versus 13.9%; fourth toe/SVL = 15.3%
versus 17.3%). With respect to meristic and qualitative characteristics, the Santubong population differed from the
remaining species in SVL, precloacal pores/pore-bearing scales, postmental scales, dorsal tuberculation condition,
subcaudal scale morphology, and colouration.
The genetic and morphological features of the Santubong population were clearly distinct from those of any
known Cnemaspis species, indicating the evolutionary independence of the population. Therefore, the Santubong
population should be treated as a separated species.

FIGURE 2. The result of principle component analysis based on the ratios of morphometric characters to snout-vent length with
80% confidence ellipses. The closed squares, crosses, open squares, open rounds, and closed rounds indicate Cnemaspis sp., C.
nigridia, C. paripari, C. matahari, and C. leucura, respectively.

Systematics

Cnemaspis puterisantubongae sp. nov. Kurita & Nishikawa

English: Santubong rock gecko
Malay: Cicak puteri Santubong

New Cnemaspis from Santubong Zootaxa 5468 (2) © 2024 Magnolia Press · 365
Holotype. Adult male, SFC-GRB-00517 (field no.: KUHE Swk23-186) collected on September 24, 2023, by
T. Kurita and K. Nishikawa from Gunung (= Mt.) Santubong, Kuching, Kuching District, Sarawak, Malaysia
(1.75791°N, 110.32685°E, ca. 250 m a.s.l.).
Paratype. Five adult males and three adult or young females (SFC-GRB-00515, 00516, 00518, 00519, SFC-
GRB-00528–00531; field nos. KUHE Swk23-184, 185, 187, 188, 197–200, respectively) collected on September
24–25, 2023, by T. Kurita and K. Nishikawa from the type locality.
Diagnosis. Cnemaspis puterisantubongae differs from all other species in the unique combination of the
following characters: SVL up to 66.7 mm; 9–12 supralabial scales; 9–12 infralabials; 0–1 internasal; 4–7 postmentals;
head, ventral, forearm, tibial, subcaudal, and dorsal tubercles keeled; 29–34 paravertebral tubercles between limb
insertions, not linearly arranged; few or no tubercles on ventrolateral surface of trunk; chevron arranged 10–14
precloacal pore scales in adult males and 9–15 pore-bearing scales in adult females, separated to right and left at
midline by 1–3 poreless scales; no femoral pores; 1–3 postcloacal tubercles on each side; enlarged submetatarsal
scales beneath first toe; 27–33 subdigital lamellae on fourth toe; tubercles present within lateral caudal furrow,
at least anterior part; ventrolateral caudal tubercles present anteriorly; single enlarged, smooth median row of
subcaudals; yellow ocher head, trunk, and limb dorsum in both sexes; distinct paravertebral black blotches; faint
yellow dotted transverse markings on flanks; black and white banding pattern on original tail.
Description of holotype (Fig. 3). Adult male; SVL 60.9 mm; head moderate in size (head length/SVL = 25.9%),
oblong in dorsal profile (head width/head length = 66.7%), flat (head depth/head length = 42.2%), distinct from neck
(nape width/head width = 62.9%); snout short (snout length/head length = 43.0%), beak concave at middle between
snout tip and anterior edges of eyes in lateral profile (beak depth/head depth = 73.7%), raised at middorsal region,
dented posterior to nostril; head slightly concave middorsally and convex at orbital edges; canthus rostralis smoothly
rounded; nostrils round, open posterodorsally; scales of rostrum juxtaposed, conical, weakly keeled; occipital area
covered with conical scales smaller than those of rostrum, intermixed with larger conical scales posterior to the level
of anterior edge of ear opening; eye moderate in size (eye diameter/head length = 16.5%); eyeball located about
three granular scales inward from lateral margin of snout and not visible from the ventral side; pupil round; strongly
prominent, pointed extrabrillar fringe scales from anterior to dorsal edge of eye, largest anterior; ear opening oval,
oriented dorsoventrally; rostral rectangular, wider than high in dorsal and lateral views, high enough to cover
entire snout tip, bordered by first supralabials, two large supranasals, and nostrils; nostrils bordered by rostral,
first supralabials, supranasals, and six/seven postnasals; medial longitudinal groove of rostral reaches two thirds of
height on dorsal edge; 11/12 supralabials and 10/10 infralabials, gradually decreasing in size posteriorly; mental
large, fan-shaped with U-shaped posterior edge extending to the level of middle of third infralabials, bordered
posteriorly by six postmentals; postmentals bordered by first infralabials elongate anteroposteriorly, larger than the
others; gular scales juxtaposed, slightly keeled, raised, oval, gradually decreasing in size posteriorly; throat scales
larger than gular scales, slightly imbricate, keeled, weakly raised, oval.
Body slender, elongate (axilla–groin length/SVL = 47.3%); small, smooth or weakly keeled granular scales
throughout trunk dorsum, intermixed with keeled conical tubercles more or less randomly arranged, infilled with
a few small amorphous scales between granular scales and tubercles; one long median keel on anterior part of
dorsal tubercle, with an additional pair of dorsolateral short keels on larger tubercles; tubercles dense dorsally,
decreasing in number laterally, absent on ventrum; 31/33 paravertebral tubercles; abdominal scales square, flat,
keeled, slightly imbricate anteriorly, stretching longitudinally, slightly raised, strongly keeled, imbricate posteriorly;
precloacal scales oval, raised, keeled, juxtaposed; six/seven precloacal pores or pore-bearing scales forming chevron
arrangement, medially separated by two keeled, poreless scales; precloacal depression absent; no femoral pores;
postcloacal scales oval, raised, keeled, slightly imbricate; five/five enlarged, a pair of postcloacal tubercles on
lateral surfaces of hemipenal swellings at base of tail.
Brachium and antebrachium long (brachial length/SVL = 13.6%, antebrachial length/SVL = 17.0%); forearms
covered with conical, keeled, slightly imbricate dorsal scales and low-conical, keeled, juxtaposed ventral scales;
palmar scales conical, weakly keeled, slightly enlarged on wrist; fingers long with an inflected joint, increasing in
length from first to fourth, with fifth slightly shorter than fourth; interdigital webbing weak; subdigital lamellae
smooth, almost entirely wide and saddle-shaped, unnotched; lamellae on joint between second and third distal
phalanges (base of digit in first finger) distinctively enlarged; subdigital lamellae from first to fifth fingers 15, 20,
26, 27, 20/16, 20, 26, 27, 20; claws recurved, bordered by a smooth dorsal scale and distalmost subdigital lamella;
hind limbs longer (femur length/SVL = 21.6%, crus length/SVL = 20.0%), more robust than forelimbs; hind limbs

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covered with conical, keeled, juxtaposed dorsal scales; ventral scales on hind limbs weakly raised, keeled, slightly
imbricate proximally, gradually increasing in size, degree of keel distally; plantar scales round, raised, smooth,
slightly imbricate; slightly-enlarged flattened smooth metatarsal scales beneath first metatarsal; toes long with an
inflected joint, increasing in length from first to fourth with fifth slightly shorter than fourth; subdigital lamellae
similar to fingers; toe lamellae from first to fifth digit 13, 19, 26, 29, 24/12, 20, 25, 27, 24; claws recurved, bordered
by a smooth dorsal scale and distalmost lamella.

FIGURE 3. The holotype (SFC-GRB-00517; temporal field number KUHE Swk23-186) of Cnemaspis puterisantubongae sp.
nov. The lateral (upper), dorsal (lower left), and ventral views (lower right) were provided before fixation, with close-up pictures
of some discriminant characters after fixation. The label KUHE Swk23-186 is a temporal field number.

Tail original, long (tail length/SVL = 121.7%), segmented; shallow contiguous middorsal caudal furrow from
first to ninth segments; lateral caudal furrow continues from first to 14th segments; caudal scales arranged in
segmented whorls; dorsocaudal scales round, weakly raised, smooth or keeled, slightly imbricate; ventrocaudal
scales oval, strongly keeled, slightly imbricate, larger and more flattened than dorsocaudals; single median row of
enlarged, smooth, imbricate, subcaudal scales, with 4–5 scales per caudal segment in anterior part of tail; conical

New Cnemaspis from Santubong Zootaxa 5468 (2) © 2024 Magnolia Press · 367
enlarged caudal tubercles with strong keel arranged on paravertebral, dorsolateral, lateral (in lateral furrow), and
ventrolateral rows, gradually decreasing in size posteriorly; 1–2 ventrolateral tubercles per caudal segment on basal,
one per segment on distal; lateral tubercles in lateral caudal furrows on basal segments; paravertebral tubercles
largest, decreasing in size towards venter; tubercles not encircling tail.
In life, head yellow ocher, with scattered faint yellow and black spots posterior to the level of anterior eye
edge; black longitudinal stripes extending from nostrils to middorsal forehead; eight black lines radially extending
from eye, of which two on anterior eye edge reach supralabials, three on dorsal edge short, other three on posterior
edge stretching to nape, anterior part of lateral trunk, and anterior edge of forelimb insertion; supralabials yellowish
gray with ocher blob; infralabials yellowish gray; iris brownish yellow and black; nape yellow ocher dorsally, gray
laterally with faint yellow spots; trunk dorsum yellow ocher, with yellowish gray region around forelimb insertions;
vertebral region of trunk faint yellowish gray; eight faint yellow blotches along vertebrae from nape to base of tail;
eight clear black dorsal blotches from nape to base of tail, of which first is on middorsal and others are separated to
right and left along faint vertebral region; two black dorsal blotches on shoulder region separated paravertebrally
and dorsolaterally; dorsal surface of forelimbs, hind limbs, and digits yellow ocher with black and faint blown
transversally elongated blotches; four faint yellow dotted lines on lateral sides of trunk; abdominal surface of head,
trunk, and limbs gray, with slightly darker stain on head ventrum; yellow spots on marginal regions of head, trunk,
and limbs in abdominal view; seven poorly-demarcated black and seven maculate white bands encircling or forming
saddles on tail; tail with black tip; black bands approximately twice as long as adjacent white bands; enlarged
median subcaudal scales same colouration as caudal bands on 10 proximal caudal segments, immaculate white on
subsequent segments.
Variation. Male paratypes are highly similar to the holotype in colouration and pattern. Compared with males,
female paratypes have pore-bearing precloacal scales, obviously convex at centre (versus precloacal pores), clear
black and white bands encircling original tail (versus white midventral and adjacent scales on posterior part of tail),
and darker body colouration (Fig. 4). Tails regenerated from tip are dorsally black and ventrally white in males,
uniformly dark gray in females; those regenerated from the middle are dorsally black and ventrally yellow in males,
unknown in females. Scales on regenerated tail are small, raised, keeled, slightly imbricate, and equal in size.
Scalation and other meristic characters are described in Table 2.
Comparisons (Table 3). Cnemaspis puterisantubongae sp. nov. is differentiated from all other Southeast Asian
congeners except C. leucura, C. matahari, and probably C. sirehensis in having an ocher yellow dorsal surface of the
head, trunk, and limbs and in black and white banding/saddling of the posterior part of original tail; the regenerated
tail has a yellow subcaudal portion in males. Cnemaspis puterisantubongae is distinct from known Bornean species
in the following characters: moderate size, with SVL up to 66.7 mm (versus 76.9 mm in C. nigridia and < 60 mm in
C. paripari, C. matahari, C. sirehensis, C. dringi, C. lagang, and C. kendallii); 4–7 postmental scales (versus 10 in
C. dringi and 8–10 in C. lagang); 10–14 precloacal pores in males (versus 2–5 in C. paripari, 7–8 in C. matahari,
4–9 in C. leucura, 5–6 in C. dringi, 4–9 in C. lagang, and absent in C. kendallii); 9–14 putatively precloacal pore-
bearing convex scales in females (versus no obvious convex precloacal scales in C. paripari, C. matahari, C.
leucura, C. lagang, and C. kendallii); 29–34 paravertebral tubercles (versus indistinct dorsal tubercles in C. lagang
and 12–18 in C. kendallii); at least a few caudal tubercles present in a lateral furrow (versus absent in C. sirehensis
and C. dringi); single median row of smooth, flat subcaudal scales (versus smooth but folded scales in C. paripari,
C. matahari, and possibly C. sirehensis, and keeled scales in C. dringi, C. lagang, and C. kendallii); 1–3 postcloacal
tubercles (versus absent in C. dringi); sharp tetragonal dorsal black marks (versus diffuse thin marks in C. paripari,
C. matahari, C. sirehensis, and C. leucura; sharp thin marks in C. lagang; and sharp round marks in C. kendallii);
and black–gray dorsal banding and white median subcaudal of original tail (versus black–dark yellow banding in
C. nigridia, immaculate white in C. paripari and C. lagang, and black-blobbed dorsal and immaculate ventral in C.
leucura).
Etymology. The generic name is feminine in gender. The specific epithet puterisantubongae is derived from
Puteri (= princess) Santubong, who is one of two princesses appearing in a local myth. These princesses are said
to have descended from the celestial realm to a human village; their remarkable beauty and weaving skills are
represented in the species by yellow ocher body colouration, distinct bold black marks, and pale/faint yellow spots
in both sexes. Santubong is also the name of the promontory for which the Santubong National Park is named.
Distribution. Cnemaspis puterisantubongae is known from Santubong National Park, Kuching District,
Sarawak, East Malaysia (Fig. 1).

368 · Zootaxa 5468 (2) © 2024 Magnolia Press KURITA ET AL.

 Table 2. Quantitative and qualitative character status of type series of Cnemaspis puterisantubongae sp. nov. Bilateral meristic characters are shown as left/right. Unit of
New Cnemaspis from Santubong

measurements is mm.
SFC-GRB-
SFC-GRB- SFC-GRB- SFC-GRB- SFC-GRB- SFC-GRB- SFC-GRB- SFC-GRB- SFC-GRB-
Voucher number 00517
00529 00515 00528 00531 00518 00530 00516 00519
(holotype)
Sex male male male male male male female female juvenile
SVL 60.9 66.7 66.4 61.5 58.5 57.2 64.7 62.6 50.5
Head length 15.5 17.7 17.2 16.4 15.1 15.6 17.5 16.5 13.7
Head width 10.5 12.1 12.1 11.7 9.8 10.4 11.8 11.5 9.3
Head depth 6.7 7.8 8.0 7.6 6.1 7.1 7.7 7.6 5.8
Head depth at eye centre 5.8 7.1 7.2 6.8 5.5 6.0 6.9 6.9 5.0
Snout length 6.7 7.4 7.8 7.1 6.4 7.0 7.4 7.3 6.0
Beak length 4.9 5.9 4.6 5.0 4.9 4.9 5.6 5.5 4.2
Beak depth 3.5 3.9 4.1 3.8 3.4 3.7 4.0 4.0 2.8
Jaw length 9.5 10.2 10.1 10.5 9.0 10.2 11.2 10.6 8.3
Jaw width 9.7 11.1 10.6 9.8 8.8 9.3 10.9 10.6 8.3
Lower jaw length 15.8 16.5 17.2 16.3 14.3 15.1 16.7 16.3 13.4
Eye diameter 2.6 3.6 3.2 3.7 2.7 3.4 3.5 4.5 2.5
Interorbital distance 1.8 2.4 3.1 2.1 1.5 1.7 3.3 2.7 1.3
Anterior interorbital 4.7 5.3 4.8 5.0 4.3 4.5 4.8 4.5 3.9
Zootaxa 5468 (2) © 2024 Magnolia Press ·

distance

Internostril distance 1.8 1.9 1.4 1.6 1.7 1.6 1.9 1.9 1.4
Axilla–groin length 28.8 32.8 32.3 29.1 27.5 27.4 31.8 29.8 24.5
Brachial length 8.3 8.5 8.3 8.5 8.1 8.3 9.7 8.7 7.5
Antebrachial length 10.4 11.9 12.2 10.8 10.4 10.6 11.4 11.1 9.1
Femur length 13.2 13.8 14.4 14.1 12.5 13.4 14.8 13.8 12.0
Crus length 12.2 12.7 13.3 13.1 11.7 12.5 12.6 12.5 10.3
Pes length 5.9 5.9 6.1 5.8 5.5 6.1 5.7 5.3 4.5
First finger length 4.8 5.4 7.1 5.3 5.2 5.3 5.5 5.6 4.5
Fourth finger length 8.6 9.3 9.2 7.8 7.7 8.4 9.2 8.9 7.5
First toe length 4.2 4.2 4.3 3.5 3.2 4.3 4.4 3.8 3.3
......Continued on the next page
369
370 · Zootaxa 5468 (2) © 2024 Magnolia Press
Table 2. (Continued)
SFC-GRB-
SFC-GRB- SFC-GRB- SFC-GRB- SFC-GRB- SFC-GRB- SFC-GRB- SFC-GRB- SFC-GRB-
Voucher number 00517
00529 00515 00528 00531 00518 00530 00516 00519
(holotype)
Fourth toe length 9.3 9.3 10.1 13.0 8.4 9.4 9.6 9.4 8.3
Supralabials 11/12 10/11 11/11 11/11 11/9 10/10 10/11 12/11 12/9
Infralabials 10/10 11/11 12/11 12/10 10/8 10/9 11/10 11/11 10/10
Postnasals 6/7 7/6 6/6 7/8 7/7 5/5 8/8 6/6 5/6
Internasal 0 0 0 1 0 0 1 0 0
Postmental 6 6 6 6 5 6 6 7 4
Precloacal pores (pore- 6/7 6/8 7/5 7/7 5/7 5/5 (7)/(7) (4)/(5) (8)/(7)
bearing convex scales)

Scales separating 2 1 3 1 3 2 1 2 3
precloacal pores
Paravertebral tubercles 31/33 32/30 29/30 31/31 30/33 31/32 31/30 34/33 31/32
Tubercles on lower present present absent present present absent absent absent absent
flanks
Anterior ventrolateral present present present present present present absent present present
caudal tubercles

Postcloacal tubercles 1/1 3/1 2/3 1/1 2/2 1/2 2/2 1/1 1/1
Finger lamellae: 15,20,26,27, 17,23,31,31, 16,20,28,29,20/ 16,21,25,27,22/ 17,22,27,28,22/ 15,23,28,29,21/ 14,20,26,27,21/ 15,22,28,30,21/ 17,22,28,29,
1,2,3,4,5th digits 20/16,20,26, 24/19,25,28, 16,23,26,28,21 16,22,27,27,20 16,22,28,30,23 16,23,27,28,21 14,21,26,26,22 16,22,29,29,22 23/17,22,27,
27,20 32,22 29,23

Toe lamellae: 1,2,3,4,5th 13,19,26,29, 13,22,29,32, 12,20,26,31,22+ 13,20,26,29,23/ 14,22,28,30,26/ 13,22,26,30,25/ 13,19,25,27,24/ 14,21,26,30,27/ 14,21,27,30,
digits 24/12,20,25, 25/14,23,29, /13,20,27,30,26 13,18,25,29,23 14,23,27,31,25 14,22,26,29,26 12,19,25,29,24 14,22,28,32,27 26/14,22,27,
27,24 33,28 30,26
KURITA ET AL.
 Table 3. Morphological comparison among the Bornean Cnemaspis. For quantitative characters, average of SVL and modes of other characters were indicated, followed by
New Cnemaspis from Santubong

the range in brackets, and if the character could be observed in a part of the sample, number of individuals in parentheses. Precloacal pores and pore-bearing scales were shown
as “modes of pore counts in one side of body [ranges; ranges of total precloacal pore/pore-bearing scale numbers] (examined individuals).” For qualitative characters, status of
the character is shown followed by the number of individuals possessing the status. "ND" indicates missing data. Character values/states in bold italic indicate those clearly differ
from Cnemaspis puterisantubongae sp. nov.
Cnemaspis C. nigridia C. paripari C. matahari C. sirehensis * C. leucura C. dringi * C. lagang C. kendallii
puterisantubongae
sp. nov.
Number of examined 9 20 17 5 3 10 2 9 28
individuals
SVL 61.0 [57.2–66.7] 63.0 [38.4– 48.0 [37.1– 50.7 [49.7– [45.1–48.8] 59.5 [57.4– 52.9 [46.3– 46.3 [45.2– 54.5 [49.1–
76.9] 52.9] 56.8] 62.7] 59.5] 49.0] 59.2]
Supralabials 11 [9–12] 11 [10–13] 12 [10–13] 10 [10–13] [12–13] 12 [10–13] 11 14 [11–14] 10 [9–13]
Infralabials 10 [8–12] 11 [9–12] 10 [8–11] 11 [9–11] [10–12] 10 [8–12] [9–13] 11 [11–13] 10 [7–11]
Postnasals 6 [5–8] 6 [4–8] (19) 6 [4–7] 7 [5–8] ND 6 [5–7] ND 7 [3–7] 5 [3–6] (25)
Internasals 0 [0–1] 0 [0–1] 0 [0–0] 1 [0–1] 1 ** 1 [0–2] ND 1 [1–1] 1 [0–1] (27)
Postmentals 6 [4–7] 6 [5–8] 7 [5–10] 5 [5–9] [6–7] 5 [4–7] 10 10 [8–10] 3 [2–3] (27)
Ventral scales keeled keeled keeled keeled keeled keeled keeled keeled keeled
Precloacal pores in male **** 7 [5–8; 10–14] (6) 7 [6–8; 13– 2 [1–3; 2–5] 4 [3–4; 7–8] ND 3 [1–5; 4–9] [ND ; 5–6] 3 [2–5; 4–9] absent (24)
14] (6) (12) (2) (5) (6)
Pore-bearing convex scales in 7 [4–7; 9–14] (2) 6 [3–7; 8–13] absent (3) absent (2) ND absent (5) ND absent (3) absent (3)
Zootaxa 5468 (2) © 2024 Magnolia Press ·

female **** (7)

Scales separating precloacal 2 [1–3] (8) 3 [1–3] (13) 4 [3–6] (12) [3–4] (2) ND 4 [1–4] (5) ND 3 [2–4] (6) -
pores ****
Shape of precloacal pores **** round (8) round (13) round (12) round (2) ND round (5) round round (6) -
Paravertebral tubercles 31 [29–34] 32 [25–38] 28 [23–29] 31 [23–36] ND 28 [24–31] ND indistinct 15 [12–18]
tubercles
Dorsal tubercle arrangement random random random random ND random ND - random
(25)/slightly
arranged (3)
Tubercles on lower flanks abs. (5)/pres. (4) abs. (14)/ abs. (16)/ abs. (4)/pres. ND abs. (2)/pres. ND absent abs. (21)/
pres. (6) pres. (1) (1) (8) pres. (7)
Lateral caudal furrows present present (19) present (15) present present present present present present (20)
......Continued on the next page
371
372 · Zootaxa 5468 (2) © 2024 Magnolia Press
Table 3. (Continued)
Cnemaspis C. nigridia C. paripari C. matahari C. sirehensis * C. leucura C. dringi * C. lagang C. kendallii
puterisantubongae
sp. nov.
Caudal tubercles in lateral present abs. (18)/ abs. (2)/pres. present absent abs. (2)/pres. absent abs. (5)/pres. abs. (19)/
furrow pres. (1) (13) (8) (4) pres. (1)
Anterior ventrolateral caudal abs. (1)/pres. (8) present (19) absent (15) absent absent present ND absent present (20)
tubercles
Lateral caudal tubercle row present present (19) present (15) present present present ND present present (20)
Caudal tubercles restricted to a no no (19) no (15) no no no no no no (20)
single paravertebral rows

Subcaudals keeled keeled (19) keeled (15) keeled keeled keeled keeled keeled keeled (20)
Single median row of smooth, flat smooth, flat smooth, smooth, keeled *** smooth, flat keeled keeled keeled (20)
subcaudals (19) folded (15) folded
Caudal tubercles encircle tail no no (19) no (15) no no no no no yes (20)
Enlarged median subcaudal present present (19) present (15) present present present present present (7) present (20)
scale row
Postcloacal tubercles 1 [1–3] 3 [0–6] 2 [1–3] 3 [2–4] [1–2] 5 [3–8] absent 1 [1–2] 2 [1–4]
Enlarged femoral scales absent absent absent absent absent absent absent absent absent
Shield-like subtibial scales absent absent absent absent absent absent absent absent absent
Subtibial scales keeled keeled keeled keeled keeled keeled keeled keeled keeled
Enlarged scales on base of 1st present present present present present present present present present
metatarsus
Dorsal black marks sharp, broad sharp/ diffuse, thin diffuse, thin diffuse, thin diffuse, thin ND sharp, thin sharp,
diffused, round
broad
Posterior original tail colour black-gray banding, black-dark immaculate immaculate ND black-blobbed ND immaculate gray-black
pattern in male white median yellow white, white with dorsal, white banding
subcaudal scales banding occasionally small black immaculate dorsal,
with dorsal blobs on ventral immaculate
black blobs dorsum ventral
Subdigital lamellae beneath 1st 16 [14–19] 15 [14–17] 15 [13–17] 15 [13–17] [12–13] 15 [14–19] [14–16] (1) 16 [14–18] 15 [12–18]
KURITA ET AL.

fingers
—2nd fingers 22 [20–25] 20 [18–23] 20 [18–22] 21 [20–23] [19–23] 22 [21–26] [22–24] (1) 22 [19–24] 21 [19–24]
......Continued on the next page
 New Cnemaspis from Santubong

Table 3. (Continued)
Cnemaspis C. nigridia C. paripari C. matahari C. sirehensis * C. leucura C. dringi * C. lagang C. kendallii
puterisantubongae
sp. nov.
—3rd fingers 28 [25–31] 25 [22–29] 26 [23–30] 27 [25–30] [21–28] 27 [26–32] 27 (1) 26 [24–29] 28 [24–32]
(19)
—4th fingers 29 [26–32] 29 [26–32] 27 [24–31] 28 [26–32] [24–31] 30 [27–36] 29 (1) 27 [27–32] 31 [27–33]
—5th fingers 21 [20–24] 21 [18–24] 20 [17–21] 20 [18–24] [18–21] 22 [20–27] [19–21] (1) 21 [19–24] 22 [19–25]
Subditital lamellae beneath 1st 14 [12–14] 12 [11–15] 12 [11–14] 12 [11–13] [10–13] 12 [11–14] 12 (1) 12 [11–14] 12 [10–13]
toes
—2nd toes 22 [18–23] 19 [16–21] 19 [17–21] 19 [19–21] [17–24] 21 [18–23] [20–21] (1) 20 [18–23] 20 [18–23]
—3rd toes 26 [25–29] 26 [22–28] 24 [22–26] 24 [23–28] [23–27] 26 [24–31] [25–26] (1) 25 [22–28] 28 [23–31]
—4th toes 30 [27–33] 31 [26–32] 28 [25–32] 29 [27–33] [26–30] 31 [27–34] [29–30] (1) 31 [28–33] 31 [28–35]
—5th toes 26 [23–28] 24 [22–27] 23 [21–26] 23 [22–27] [21–25] 26 [23–28] 26 (1) 26 [24–31] 26 [23–31]
* transcribed from Nashriq et al. (2022).
** supranasals are excluded.
*** probably the same as “smooth, folded.”
**** counted for individuals with SVL 80% or more of the largest individual.
Zootaxa 5468 (2) © 2024 Magnolia Press ·
373
FIGURE 4. The dorsal (upper) and ventral views (middle) of an adult female paratype (SFC-GRB-00516; temporal field
number KUHE Swk23-185) of Cnemaspis puterisantubongae sp. nov. The magnified view of precloacal pore-bearing scales on
the left side of body (bottom) are indicated by black arrows.

Natural History. The species has been found exclusively on the surfaces of sandstone boulders or outcrops
containing pebble conglomerate, sometimes along small streams in lowland mixed dipterocarp forest (Fig. 5).
Individuals have been observed and/or collected at about 01:00, 09:00, 16:00, and 20:00. From morning to dusk,
geckos positioned themselves on the edges and inside of large rock crevices. Geckos tended to avoid humans and
scattered when approached, particularly at distances of several metres. However, at around midnight, they strayed
farther from crevices and remained exposed even when approached by humans, suggesting that this species is
diurnal.
Cnemaspis puterisantubongae shares the same habitat with the sympatric species C. kendallii, Gekko monarchus
(Schlegel, 1836 in Duméril & Bibron, 1836), and Tropidophorus brookei (Gray, 1845). The two sympatric Cnemaspis
species appear to use different microhabitats. Compared with C. kendallii, which uses trees, logs, and the upper and
central parts of large boulders above human height, C. puterisantubongae has never been observed on such open

374 · Zootaxa 5468 (2) © 2024 Magnolia Press KURITA ET AL.

surfaces but remains within closed or complex interspaces (Fig. 5). Typically, one Cnemaspis individual is found
on a single boulder; on a larger boulder two or three individuals can be observed, sometimes including both C.
puterisantubongae and C. kendallii, while at other times only one of the two species is present. This behaviour may
indicate a competitive relationship among these individuals. However, although both species have been observed
in partially open spaces around midnight, they were slow to react, and interspecific interactions were not observed
during surveys. Whether microhabitat selection by either species is affected by the occurrence of the other remains
unknown because the range of C. puterisantubongae is within that of C. kendallii.
One female C. puterisantubongae individual had two large eggs and others did not (Fig. 4), suggesting that the
reproductive season of these species includes September. We observed more than 20 individuals, but none exhibited
tail waving when they were threatened or startled as opposed to C. kendallii, which frequently waves or wags its
tail when threatened.

FIGURE 5. The habitat (sandstone boulders) of Cnemaspis puterisantubongae sp. nov. in Santubong National Park (upper).
The microhabitat preferences of C. puterisantubongae (lower left; inner surface of boulders) and co-occurring C. kendallii
(lower right; both of inner and outer surfaces) are shown, with arrows pointing the positions of geckos.

New Cnemaspis from Santubong Zootaxa 5468 (2) © 2024 Magnolia Press · 375
Discussion

Neither the sandstone dwelling species (C. leucura and C. puterisantubongae) nor the limestone dwelling species,
(C. paripari, C. matahari, and C. sirehensis) were monophyletic in the maximum likelihood tree (Fig. 1), which
suggests that the substrate preference of these species has changed during their evolutionary history. This finding
is consistent with the evolutionary flexibility inferred from the pattern of substrate preference across Southeast
Asian Cnemaspis species, as noted by Grismer et al. (2014). However, as indicated by the close distributional
ranges of C. matahari and C. sirehensis, and the divergences among four southern Vietnamese Cnemaspis species
within narrow ranges (Nguyen et al. 2018), Cnemaspis geckos have diverged within relatively small regions. Thus,
accessible substrates are geographically and physically limited for saxicolous Cnemaspis species. Future studies
should explore whether the substrate specialisation of the nigridia group is the result of actual habitat preferences
among species or simply a reflection of their small ranges in the context of Cnemaspis gecko evolution.
Gunung Santubong National Park is only 25 km from Kuching, the capital city of Sarawak. This park is easily
accessible for both local and foreign visitors, including researchers. Indeed, various herpetofauna surveys, including
public participation events, have been conducted in the park within the last few decades, although these surveys
were not necessarily focused on geckos (e.g., Denzer 1996; van Rooijen 2009; Das et al. 2014, 2019). The discovery
of the new species indicates that inventory studies may be insufficient to provide a comprehensive understanding
of gecko species diversity, even when such studies are conducted in popular and easily accessible national parks.
Cnemaspis puterisantubongae may have remained hidden for a long duration due to its narrow habitat selection.
This new species was not found in coastal areas of the mountain, where C. kendallii was common, despite the
presence of large, exposed sandstones. Our surveys indicated that this species occurs only sporadically further
inland. In this study, we were unable to characterise microenvironment difference between coastal and inland areas,
which could limit the distribution of C. puterisantubongae within the national park. The cryptic nature of this gecko,
particularly in daytime, also may have caused it to remain overlooked. To reveal further biodiversity, extensive
surveys should be conducted at various times and target broad ranges of national parks, including both trails and
backcountry areas.
Southwestern Sarawak lies on complex geological basements, with mountains and hilly areas mainly composed
of scattered sandstone and granite. The distributional ranges of the nigridia group within this region are poorly
understood. No members of the nigridia group have been recorded in areas that are geographically similar to
Santubong National Park, such as Kubah National Park, Bako National Park, or several non-protected regions
(Hamanaka et al. 2019; Nashriq & Das 2021; Nashriq et al. 2022). Many such regions are located within the
Kuching area; it would be useful to reassess these areas, which may have greater species diversity than previously
thought, to further elucidate the evolutionary history of Bornean Cnemaspis, particularly focusing on the nigridia
group.

Acknowledgements

This research was conducted in accordance with the Memoranda of Understanding signed between the Sarawak
Forestry Corporation (SFC) and the Japan Research Consortium for Tropical Forests in Sarawak (JRCTS) and
with that between Forest Department Sarawak (FDS) and JRCTS. The Sarawak Government kindly permitted
us to conduct the surveys (SFC.730-1/1/3/TPA/Jilid.3(218), NPW.907.4.4.(Jld.14)-129, NCCD.907.4.4(Jld.14)-61,
(134)JHS/NCCD/600-7/2/107, Park Permit No.66/2017, NO.WL72/2018). Research and Development Division
(RDD), FDS, provided all facilities for conducting research. We are grateful to Zolkipli Mohd Aton for approving
our work in Sarawak. We thank Paschal Anak Dagang and Azroie Bin Denel, SFC, for curation of the type series.
The naming of the species was done in consultation with SFC. We thank Runi Ak Sylevester Pungga, Paulus Ak
Meleng, and other staff of the RDD and Santubong National Park for supporting our field surveys. We thank Sena
Fujii, Asami Kimoto, and Ibuki Fukuyama for critical information of the distribution of C. puterisantubongae, and
thank Tomoyuki Komai for advice on etymology. We thank Textcheck (http://www.textcheck.com/) for English
language editing. This survey was conducted as a part of the PUBS (directed by Takao Itioka and Runi Sylvester
Pungga) supported by Science and Technology Research Partnership for Sustainable Development (SATREPS;
JPMJSA1902) in collaboration between Japan Science and Technology Agency (JST) and Japan International
Cooperation Agency (JICA).

376 · Zootaxa 5468 (2) © 2024 Magnolia Press KURITA ET AL.

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Supplementary materials

Supplementary materials are available at figshare: https://doi.org/10.6084/m9.figshare.25358146

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