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ZOOTAXA
ISSN 1175-5326 (print edition)
ISSN 1175-5334 (online edition)
Accepted by M. Heinicke: 18 Jan. 2022; published: 24 Mar. 2022 1
Zootaxa 5120 (1): 001–029
https://www.mapress.com/zt/
Copyright © 2022 Magnolia Press Article
https://doi.org/10.11646/zootaxa.5120.1.1
http://zoobank.org/urn:lsid:zoobank.org:pub:53246B55-8BC9-4BB2-99EB-6B34CF48F6D0
Three New Species of Cnemaspis (Sauria: Gekkonidae) from Sarawak, East
Malaysia, Borneo
IZNEIL NASHRIQ1, 4, HAYDEN R. DAVIS2, 3, AARON M. BAUER2 & INDRANEIL DAS1
1Institute of Biodiversity and Environmental Conservation, Universiti Malaysia Sarawak, 94300 Kota Samarahan, Sarawak,
Malaysia.
�
idas@unimas.my, https://orcid.org/000-0001-9522-2228
2Department of Biology and Center for Biodiversity and Ecosystem Stewardship, Villanova University, 800 Lancaster Avenue, Villanova,
Pennsylvania, 19085, USA.
�
hrdavis1@uw.edu, https://orcid.org/0000-0003-1401-1221
�
aaron.bauer@villanova.edu, https://orcid.org/0000-0001-6839-8025
3Department of Biology, and Burke Museum of Natural History and Culture, University of Washington, Seattle, Washington, USA,
91895.
4Corresponding author:
�
izneilnshrq@gmail.com, https://orcid.org/000-0002-3849-7732
Abstract
Three new species of Cnemaspis are described from karst regions of Sarawak, Malaysia, on the island of Borneo. These
are Cnemaspis matahari sp. nov. and C. sirehensis sp. nov. from limestone hills located in the Serian Division of western
Sarawak, and C. lagang sp. nov. from Gunung Mulu, Miri Division, in northern Sarawak. All can be distinguished from
congeners using mitochondrial DNA as well as an enlarged metatarsal scales on the first toe. Individually, each species
can be diagnosed by differences in subcaudal scale morphology: Cnemaspis lagang sp. nov. lacks enlarged subcaudals;
Cnemaspis matahari sp. nov. has keeled subcaudals bearing an enlarged median row of smooth scales; and Cnemaspis
sirehensis sp. nov. has an enlarged median row of weakly keeled scales. The discovery of these species suggests that
additional unrecognized species may exist within the genus on Borneo, especially in association with karst formations.
High endemism and species diversity notwithstanding, these karst formations are under severe pressure from limestone
extraction and deforestation.
Key words: Cnemaspis, taxonomy, new species, ecology, conservation, endemic, karst, Sarawak
Introduction
The tropical island of Borneo is a known biodiversity hotspot, with many endemic vertebrates (Van Paddenburg et
al. 2012). One group which has attracted the attention of taxonomists in the recent past is the Old World gekkonid
genus Cnemaspis Strauch 1887. The genus currently comprises ~198 species that are distributed across tropical
Africa and Asia, although molecular data indicates its polyphyletic nature (Gamble et al. 2012; Grismer et al. 2014),
with three distinct and distantly related clades corresponding to African, south Asian, and south-east Asian groups.
Morphology is known to be highly conserved across Cnemaspis, with most species being small to medium-sized,
cryptically-coloured geckos, with broad, flattened heads, large, forward and upwardly directed eyes, a flattened body,
long widely splayed limbs and long, inflected digits. Members of the genus occupy habitats ranging from lowland
dipterocarp forests to primary and old-growth forests, often within karst, granite or sandstone landscapes (Das &
Bauer, 1998; Das & Sengupta, 2000; Das, 2004; Amarasinghe et al. 2016; Iskandar et al. 2017) The south-east Asian
Cnemaspis group has been reported from areas of Vietnam, Cambodia, Laos, Thailand, Peninsular Malaysia and its
offshore islands, Singapore, Sumatra and Borneo, with the highest diversity on the Thai-Malay Peninsula (Grismer
et al. 2014). Despite the south-east Asian clade being the most species rich, only five species have been described
from the island of Borneo (Koch & Schulz, 2014; Kurita et al. 2017).
The distribution of known Bornean species of Cnemaspis is restricted to the state of Sarawak, East Malaysia
and West Kalimantan, Indonesia (Maulidi et al. 2019), with no records from the state of Sabah, East Malaysia or the
NASHRIQ ET AL.
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independent nation of Brunei Darussalam. The greatest species richness occurs in western Sarawak. Phylogenetic
analyses of south-east Asian Cnemaspis (Grismer et al. 2014) reveals two divergent lineages: the southern Vietnamese
insular endemics referred to as the Ca Mau clade and a clade containing three major subclades: the Pattani clade,
restricted to the southernmost portion of peninsular Thailand and the Northern Sunda and Southern Sunda sister clades;
and the Northern Sunda clade extending from Vietnam to central Peninsular Malaysia; and the Southern Sunda clade
extending from southern Peninsular Malaysia and Singapore, eastward through the Seribuat, Anambas, and Natuna
Archipelagos to northern Borneo. The Bornean elements of the Southern Sunda clade comprise two genetically distinct
groups (the kendallii group and the nigridia group) the former with one representative Cnemaspis kendallii (Gray,
1845) and the latter with four, Cnemaspis nigridia (Smith, 1925), Cnemaspis dringi Das & Bauer, 1998, Cnemaspis
paripari Grismer & Chan, 2009 and Cnemaspis leucura Kurita, Nishikawa, Matsui & Hikida, 2017.
During recent fieldwork conducted in karst regions of Sarawak, we identified three undescribed species of
Cnemaspis. Using a combination of morphological and molecular data for two of the lineages and only morphological
data for a third, we provide evidence to recognize three new species of Cnemaspis in Borneo. These new discoveries
and an expanded genetic dataset enhance our understanding of the evolutionary history of the genus, especially
considering the relatively low diversity of Cnemaspis in Borneo.
Materials and Methods
Sampling. Specimens were collected by hand during visual encounter surveys. Coordinates and elevational data
were recorded using a Garmin GPSMap 76CS receiver, with the datum set to WGS84. Materials examined are
deposited in the Institute of Biodiversity and Environmental Conservation (IBEC) Systematics Lab in Universiti
Malaysia Sarawak, Kota Samarahan, Sarawak, Malaysia (UNIMAS). Specimens examined were from the following
collections: The Natural History Museum, London, UK (BMNH); Field Museum, Chicago, USA (FMNH);
Herpetological Collection of the Universiti Kebangsaan Malaysia, Bangi, Selangor, Malaysia (HC); La Sierra
University Herpetological Collection, Riverside, USA (LSUHC); Museum of Comparative Zoology, Cambridge,
USA (MCZ); Sarawak Biodiversity Center, Kuching, Sarawak, Malaysia (SBC); Universiti Malaysia Sarawak, Kota
Samarahan, Sarawak, Malaysia (UNIMAS); United States National Museum, Washington, DC, USA (USNM); Lee
Kong Chian Natural History Museum, Singapore (ZRC). All specimens labeled as IN field series are deposited in
the UNIMAS museum collection.
Photographs of live individuals were taken using a Nikon D600 DSLR camera and 105 mm Micro-Nikkor
f/2.8 D lens, illuminated by a Speedlight flash unit (SB800), using a Lastolyte softbox. Specimens were euthanized
by intracoelomic injection of 6 mg/mL of sodium pentobarbital solution. Sexes were determined by hemipeneal
eversion. A small incision was made on the abdominal region of the specimens and liver samples were taken
and preserved in 95% ethanol for DNA analysis. Morphometric measurements were taken using digital caliper.
Colour identification was determined using the colour swatches in ‘Naturalist’s Color Guide’ (Smithe, 1975, 1981).
Radiographic examination of vertebrae and phalanges were conducted with a VistaScan Mini Plus x-ray unit (Dürr
Dental SETM). Specimens were fixed in 10% formalin solution for 4–5 days. Once the specimen became rigid, the
formalin was washed off in tap water and the specimens were transferred to 70% ethanol solution for long term
storage. A total of 92 specimens were examined (Appendix I) representing all known species from the State of
Sarawak.
DNA Extraction and Analysis. Genomic DNA was extracted following the technique of Aljanabi and Martinez
(1997). The mitochondrial NADH dehydrogenase subunit 2 (ND2) locus was amplified using a double-stranded
Polymerase Chain Reaction (PCR) under the following conditions: 2.5 μl 5X buffer, 2.5 μl 10X buffer, 2.5 μl
dinucleotide pairs, 2.5 μl forward and reverse primer, 0.18 μl Taq polymerase, 9.82 μl ultra-pure H2O, and 2.5 μl of
DNA for a total of 25.0 μl. PCR reactions were executed on an Eppendorf Mastercycler gradient thermocycler under
the following conditions: initial denaturation at 95 ºC for 35 s, annealing at 50 ºC for 35 s, and a cycle extension at 72
ºC for 35 s, for a total of 34 cycles. All PCR products were viewed on a 1.0% agarose gel electrophoresis. Successful
PCR products were purified using a home-made magnetic bead solution (Rohland & Reich, 2012), and sequenced
using the ABI Big-Dye Terminator v3.1 Cycle Sequencing Kit on an Eppendorf Mastercycler gradient thermocycler.
Cycle sequencing reactions were purified using an in-house protocol (see Davis et al. 2019) and sequenced on an
ABI 3730xl DNA Analyzer. Primers used for amplification and sequencing are presented in Appendix II.
THREE NEW CNEMASPIS FROM BORNEO Zootaxa 5120 (1) © 2022 Magnolia Press · 3
FIGURE 1: ML tree of south-east Asian Cnemaspis with placement of the two new species C. matahari sp. nov. and C.
sirehensis sp. nov. within the nigridia group of the Southern Sunda clade.
NASHRIQ ET AL.
4 · Zootaxa 5120 (1) © 2022 Magnolia Press
Sequences were uploaded to Geneious © v11.1.2 (Kearse et al. 2012) for alignment and to check the quality of
the sequences by searching for miscalled bases. Forward and reverse sequence that formed contigs were then aligned
and edited when necessary. In total, 10 new sequences were generated for this study, which were combined with
ND2 sequences available on GenBank. Sequences were aligned using the MAFFT algorithm (Katoh & Standley
2013). The final ND2 alignment consisted of 1339 bp and 109 individuals comprising all four major south-east Asia
clades with multiple representatives from each group. Outgroup selection for the phylogenetic inference was based
on prior, more comprehensive phylogenetic trees incorporating Cnemaspis (Grismer et al. 2014, 2015; Wood et al.
2017).
To determine the phylogenetic placement for the new species, a gene tree was inferred using a Maximum
Likelihood (ML) approach. The ML tree was generated using IQ-TREE (Nguyen et al. 2015). ModelFinder was
used to determine the best-fit evolutionary model, and 1000 ultrafast bootstrap pseudoreplicates were utilized
(Hoang et al. 2017; Kalyaanamoorthy et al. 2017). Nodes having UFBoot2 (UF) values of 95 and above were
considered significantly supported (Minh et al. 2013). The 10 newly generated sequences are available on GenBank.
All sequences and locality data are shown in Appendix III.
Morphological Analyses. Morphological differences were examined by measuring 16 variables along with
33 meristic characters. Measurements of individuals were taken using a MitutoyoTM digital caliper (to nearest 0.1
mm), under an OlympusTM SZX10 dissecting microscope on the left side of the body (where appropriate): snout-
vent length (SVL), taken from the tip of snout to the vent; tail length (TL), taken from the vent to the tip of the tail;
tail width (TW), taken at the base of the tail immediately posterior to the postcloacal swelling; forearm length (FL),
taken on the dorsal surface from the posterior margin of the elbow while flexed 90º to the inflection of the extended
wrist; tibia length (TBL), taken on the ventral surface from the posterior surface of the knee while flexed 90º to the
base of the heel; axilla to groin length (AG), taken from the posterior margin of the forelimb at its insertion point
on the body to the anterior margin of the hind limb at its insertion point on the body; head length (HL), the distance
from the posterior margin of the retroarticular process of the lower jaw to the tip of the snout; head width (HW),
measured at the angle of the jaws; head depth (HD), the maximum height of head from the occiput to the underside of
the jaw; eye diameter (ED), the greatest horizontal diameter of the eyeball; eye to ear distance (EE), measured from
the anterior edge of the ear opening to the posterior edge of the eyeball; eye to snout distance (ES), measured from
anteriormost margin of the eyeball to the tip of snout; eye to nostril distance (EN), measured from the anteriormost
margin of the eyeball to the posterior margin of the external nares; inter-orbital distance (IO), the width of the frontal
bone at the level of the anterior edges of the orbit; ear length (EL), the greatest vertical distance of the ear opening;
and internarial distance (IN), measured between the medial margins of the nares across the rostrum.
The following morphological characters were noted: number of supralabial and infralabial scales counted from
below the middle of the orbit to the rostral and mental scales; number of internasal and postmental scales; the texture
of the dorsal and ventral, anterior margin of the forearm and subtibial scales; number, shape and arrangement of
precloacal pores in males; the number of postcloacal tubercles on each side of the tail base (left/right); the degree
and arrangement of tail tuberculation; the relative size and morphology of the subcaudal scales, and submetatarsal
scales beneath the first metatarsal; the number of subdigital lamellae for both left and right digits counted from
the base of the first phalanx to the base of the claw. On the original portion of the tail, longitudinal rows of caudal
tubercles and subcaudal scales vary between species and are useful in differentiating several taxa. When present,
the rows of caudal tubercles and subcaudal scales are usually restricted to the proximal portion of the tail, and
occasionally there may be a row of tubercles within the lateral caudal furrow. Bilateral scale counts were given as
left and right (L/R).
Results
The Southern Sunda clade is a well-supported group comprising of C. limi from Tioman and Tulai islands in the
Seribuat Archipelago as the sister lineage (UFBoot2 = 100), differing morphologically from other members of the
clade in having no precloacal pores; randomly arranged body tubercles; paravertebral and lateral row of caudal
tubercles present; no ventrolateral caudal tubercles; subcaudals smooth bearing a median, weakly enlarged, scale
row; large, black, round spots on nape and anterior portion of body; dorsal caudal tubercles white; at least posterior
one-half of subcaudal region white. The kendallii group, containing species from east Malaysia and Indonesia with
THREE NEW CNEMASPIS FROM BORNEO Zootaxa 5120 (1) © 2022 Magnolia Press · 5
C. kendallii as the sole representative from Borneo (UFBoot2 = 99), forms a morphologically supported clade.
This group comprises the only Cnemaspis species in the Southern Sunda clade that lack precloacal pores which is
considered derived in that precloacal pores are widespread throughout the Gekkonidae (Kluge 1987); and excluding
C. sundainsula, they are the only species of Cnemaspis in the Southern Sunda clade that have caudal tubercles
encircling the tail.
The related species C. nigridia, C. paripari and C. leucura are referred to as the nigridia group (UFBoot2 =
100), and phylogenetic analyses placed the two new species for which molecular data are available, C. matahari sp.
nov. and C. sirehensis sp. nov., as sister taxa within the nigridia group with strong support (UFBoot2 = 97), and as
the sister clade to C. leucura (Figure 1). The nigridia group contains species from north-western Sarawak and the
monophyly of this group is further supported in that these are the only species in the Southern Sunda clade to have
enlarged scales beneath the first metatarsals. This character state occurs in the two species of the Ca Mau clade,
four species of the Northern Sunda clade and does not occur in the Pattani clade, therefore, it is considered to be
derived in the nigridia group. The nigridia group is diagnosed as having weakly to strongly keeled ventral scales;
2–16 contiguous, pore-bearing precloacal scales with round pores; randomly to weakly aligned dorsal tubercles on
the body; no tubercles on flanks or in the lateral caudal furrows; caudal tubercles do not encircle tail; lateral row
of caudal tubercles present; subcaudals keeled and bearing a median row of enlarged, keeled or smooth scales;
1–4 postcloacal tubercles on either side of the base of the tail; no enlarged femoral or subtibial scales; subtibials
keeled; and submetatarsal scales of first toe enlarged. Genetic material for one of the three new species is currently
unavailable. Given that these new populations form well-supported independent lineages coupled with high genetic
distances and a unique set of morphological and colour pattern characteristics that separate them from all members
of their respective groups, we describe these three populations as new species.
SYSTEMATICS
Cnemaspis lagang sp. nov.
North Sarawak Day Gecko; Cicak Gua Mulu
Fig. 2, Fig. 3, Table 1
Holotype. Adult male, UNIMAS 9562, collected by Izneil Nashriq on 10 February 2020 at 22:45 hrs, from the
base of Gunung Api (4.13608°N, 114.891304°E; 90 m), Gunung Mulu National Park, Miri District, Sarawak, East
Malaysia (Borneo).
Paratype. Adult male, UNIMAS 9591, collected by Hayden Davis and Izneil Nashriq, on 22 July 2017 at 22:00
hrs, from Lagang Cave (4.050144°N, 114.824199°E; 100 m), Gunung Mulu National Park, Miri District, Sarawak,
East Malaysia (Borneo).
Diagnosis. SVL up to 46 mm; 13–14 supralabials; 10–11 infralabials; 3 internasals; 9–10 postmentals; adult
males with 7–8 discontinuous pore-bearing, precloacal scales with round pores, arranged in a chevron, separated
at midline by 2–4 poreless scales; paravertebral and lateral row of caudal tubercles present; ventrolateral caudal
tubercles absent; caudal tubercles not encircling tail; subcaudals keeled, not bearing row of enlarged median
subcaudal scales; 1–2/1–2 (L/R) postcloacal tubercles on each side of tail base; no enlarged femoral or subtibial
scales; submetatarsal scales of first toe enlarged; 29 subdigital fourth toe lamellae; faint Dusky Brown and Trogon
Yellow caudal bands anteriorly; tail immaculate posteriorly; and regenerated tail also immaculate.
Description of holotype. Male with original tail, immaculate posteriorly; supralabials 13/14 (L/R); infralabials
11/11 (L/R); snout-vent length 46 mm; head short (HL/SVL 0.29), narrow (HW/SVL 0.28), depressed (HD/HL 0.33),
distinct from neck; snout long (ES/HW 0.63), much longer than eye diameter (ED/ES 0.55); scales on snout and
forehead tuberculate, with posterior portion of each scale raised; scales on snout larger than those on occipital region;
eye small (ED/HL 0.23); orbits of eyes with extra-brillar fringes; pupil round; enlarged supraciliaries on top half of
orbit; tympanum deep, oval shaped, greatest diameter vertically, narrow (EL/HL 0.10); eye to ear distance greater
than diameter of eyes (EE/ED 1.28); rostral half as deep as wide, contacted posteriorly by 2 nasals and 3 internasals,
and rostrals in contact with supralabial I. Nostrils oval, situated within nasals, and oriented dorsally; nostrils not in
contact with supralabial I. 7 postnasals bound nasal; mental ovate, much deeper than wide, 9 postmentals border
mental; chin scales meet infralabials.
NASHRIQ ET AL.
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FIGURE 2: (A) Cnemaspis lagang sp. nov., holotype, UNIMAS 9562, SVL 46 mm, adult male from Gunung Api, within
Gunung Mulu National Park, Miri Division, Sarawak; (B) type series displaying dorsal side of the body (from left to right:
UNIMAS 9562 and UNIMAS 9591).
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FIGURE 3: Ventral view of Cnemaspis lagang sp. nov. type series (from left to right: UNIMAS 9562 and UNIMAS 9591);
Close up of body parts (A) Postmental, (B) Cloaca, (C) Subcaudal, (D) Manus and (E) Pes.
NASHRIQ ET AL.
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TABLE 1: Morphometric measurements of Cnemaspis lagang sp. nov. examined (nearest 0.1 mm).
Type Museum Number Field Number Sex SVL HL HW HD ED EE ES EN IO EL IN TL TW FL TBL AG
Holotype 9562 ID 9540 M 46.0 13.4 13.0 4.4 3.1 4.0 5.7 4.2 2.2 1.4 2.0 61.1 3.0 8.5 10.3 19.8
Paratype 9591 MCZ A-36586 M 46.0 13.2 8.4 5.1 2.8 4.0 6.0 4.8 1.6 1.1 1.5 24.5 4.5 9.1 10.8 19.4
TABLE 2: Morphometric measurements of Cnemaspis matahari sp. nov. examined (nearest 0.1 mm).
Type Museum Number Field Number Sex SVL HL HW HD ED EE ES EN IO EL IN TL TW FL TBL AG
Paratype 9573 IN 077 M 55.7 16.1 10.6 6.3 3.2 4.2 6.8 5.0 2.0 1.4 1.8 69.3 4.7 11.6 13.6 25.4
Paratype 9574 IN 078 F 54.2 16.5 9.9 5.8 3.5 3.5 7.1 5.3 2.7 1.3 1.6 58.1 3.9 10.1 12.2 23.5
Holotype 9602 MCZ A-36622 M 55.4 14.7 9.8 6.3 3.2 3.8 6.4 4.5 1.6 1.6 1.4 71.0 3.6 10.9 12.6 22.5
Paratype 9603 MCZ A-36623 M 52.7 14.5 9.1 5.6 3.6 3.3 6.4 5.3 2.0 1.1 1.7 60.7 3.3 11.6 13.0 23.3
Paratype 9606 MCZ A-36648 M 54.0 15.2 10.3 5.9 3.5 4.4 6.6 5.2 2.1 1.1 1.7 63.0 4.0 10.7 13.3 23.8
Paratype 9607 MCZ A-36649 M 55.5 15.8 10.0 6.2 3.6 4.4 6.7 5.1 2.2 1.7 2.0 54.8 3.9 10.9 14.3 24.5
Paratype 9608 MCZ A-36650 M 54.9 15.8 10.1 5.9 3.3 4.3 7.4 5.6 2.5 1.5 2.1 64.5 4.0 10.7 12.3 25.2
Paratype 9611 MCZ A-36659 M 48.3 13.9 8.2 5.1 3.0 3.8 6.1 5.0 1.8 0.9 1.4 51.3 3.9 9.2 10.9 20.4
Paratype 9615 MCZ A-36671 M 47.1 13.9 8.7 4.9 2.8 3.6 5.8 4.1 1.8 1.0 1.8 44.1 3.6 9.7 11.8 19.6
TABLE 3: Morphometric measurements of Cnemaspis sirehensis sp. nov. examined (nearest 0.1 mm).
Type Museum Number Field Number Sex SVL HL HW HD ED EE ES EN IO EL IN TL TW FL TBL AG
Holotype 9609 MCZ A-36654 F 47.2 14.1 8.3 5.1 3.1 3.7 5.7 4.3 1.6 1.2 1.5 57.6 3.6 9.4 11.1 20.3
Paratype 9610 MCZ A-36655 F 48.8 13.7 8.7 5.1 3.1 3.7 5.9 4.7 2.1 1.2 1.6 61.8 3.7 9.7 11.2 20.7
Paratype 9715 - F 45.1 13.0 7.8 4.8 3.0 3.5 5.8 4.9 2.0 1.0 1.0 57.0 3.3 8.3 11.6 23.2
THREE NEW CNEMASPIS FROM BORNEO Zootaxa 5120 (1) © 2022 Magnolia Press · 9
Body slender, short (AG/SVL 0.43); ventral scales keeled, increase in size from chin region to gular, pectoral
and abdominal regions. Dorsal scales increase in size from head to nape and body. Scales on dorsum at midbody
approximately equal to those of venter at same level; vertebral scales not reduced; no paravertebral rows of tubercles
on dorsum; pectoral and abdominal scales distinctly elongated, imbricate and unicarinate; discontinuous rows of 4/4
(L/R), pore-bearing, precloacal scales with round pores, arranged in a chevron, separated at midline by 4 poreless
scales, in adult males; no femoral pores; no preanal groove.
Forelimbs relatively long, slender, shorter than hindlimbs (FL/SVL 0.19, TBL/SVL 0.22). Dorsal scales on
forelimbs, raised, unicarinate, juxtaposed, reduced in size posteriorly; ventral scales of forelimbs slightly raised,
slightly unicarinate, juxtaposed, reduced in size posteriorly. Dorsal scales of hindlimbs raised, unicarinate,
juxtaposed, reduced in size posteriorly; ventral scales of hindlimbs slightly raised, slightly unicarinate, juxtaposed,
reduced in size posteriorly. Palmar and plantar scales smooth, granular, raised. Digits elongate, all bearing claws that
are slightly recurved; subdigital scansors entire, except for 1–2 fragmented at base of digits, unnotched; an enlarge
scansor towards base of digits, which is over twice width of other scansors; interdigital webbing absent. Subdigital
lamellae (manus) I (13); II (21); III (23); IV (27); V (21); (pes) I (11); II (19); III (23); IV (29); V (24).
Original tail longer than snout-vent length (TL/SVL 1.33); tail base distinctly swollen; tail arranged in segmented
whorls; caudal tubercles keeled, arranged in paravertebral and lateral rows; caudal tubercles do not encircle tail at
each whorl; 1/1 (L/R) postcloacal tubercles; tail with distinct pair of furrows laterally; subcaudal scales unicarinate
and no enlarged median subcaudals.
Variation in paratype. Male with a regenerated tail; SVL 46 mm; supralabials 14/14 (L/R); infralabials 10/10
(L/R); head short (HL/SVL 0.29), narrow (HW/SVL 0.18), depressed (HD/HL 0.38), distinct from neck; snout long
(ES/HW 0.72), longer than eye diameter (ED/ES 0.46); eye small (ED/HL 0.21); tympanum deep, oval shaped,
greatest diameter vertically, narrow (EL/HL 0.08); eye to ear distance greater than diameter of eyes (EE/ED 1.45);
mental bell-shaped, much deeper than wide, 10 postmentals border mental. Body slender, short (AG/SVL 0.42);
discontinuous rows of 3/4 (L/R), pore-bearing, precloacal scales with round pores, arranged in a chevron, separated
at midline by 4 poreless scales; no femoral pores; no precloacal groove. Forelimbs moderately long, slender shorter
than hindlimbs (FL/SVL 0.20, TBL/SVL 0.24). Subdigital lamellae (manus) I (14); II (19); III (24); IV (25); V (22);
(pes) I (13); II (19); III (23); IV (29); V (24). Regenerated tail shorter than snout-vent length (TL/SVL 0.53). 1/2
(L/R) postcloacal tubercles; Tail with distinct pair of furrows laterally. Subcaudal scales unicarinate.
Skeletal notes. Both specimens (UNIMAS 9591 and UNIMAS 9562) have 25 presacral and 2 sacral vertebrae.
The low presacral count is uncommon in geckos in general (26 is typical; Hoffstetter and Gasc, 1969). The
phalangeal formulae are plesiomorphic for geckos: 2–3–4–5–3 manus and 2–3–4–5–4 pes. Both specimens are
skeletally mature, showing fusion of the long bone epiphyses.
Colouration in life. Holotype; Raw Umber head, Venetian Blue shade interorbitally and around postorbital;
ventral surface of head Raw Umber with Pratt’s Payne’s Gray spots; body and limbs Trogon Yellow; ground colour
of nape and shoulder region Olive-Brown bearing Dusky Brown lines from orbit; a pair of medium, amorphous,
Dusky Brown spots in shoulder region; Sulfur Yellow flecks on flank, forelimbs and hind limbs; small, scattered,
Dusky Brown spots between limb insertion; rows of Dusky brown lines along vertebral column; proximal half of
tail Trogon Yellow bearing faint, Dusky Brown bands; ventral surfaces Cinnamon with Verona Brown except for tail
which is Trogon Yellow and posterior tail immaculate.
Paratype; Dorsal ground colour of head, body and limbs dark Brownish Olive; head bearing small, occipital
flecks; ground colour of nape and shoulder region Olive-Brown bearing paravertebral patches of irregularly shaped,
dark blotches; transverse, Sulfur Yellow flecks between forelimb and hind limb insertions; anterior one-half of
tail Brownish Olive bearing faint, dark bands; all ventral surfaces Dark Grey except for tail which is Buff Yellow;
regenerated tail Buff Yellow with whitish tip.
Etymology. The epithet lagang is a noun in apposition derived from the paratype locality of Lagang Cave,
within Gunung Mulu National Park.
Natural history. The species is nocturnal and was collected between 2000 and 2300 hrs, below 100 m asl
of Melinau Limestone formation to which the species is apparently restricted (Figure 4). Ambient temperature
was 26°C, and relative humidity 89.9% RH under clear night skies at the time of collection. The holotype was
encountered perching head-down on a stalactite at the base of Gunung Api, while the paratype was on the entrance
wall of Lagang Cave. They occur sympatrically with Cyrtodactylus consobrinus, C. miriensis and C. muluensis but
only syntopically with the last of these.
NASHRIQ ET AL.
10 · Zootaxa 5120 (1) © 2022 Magnolia Press
FIGURE 4: Habitat (A) and microhabitat (B) of Cnemaspis lagang sp. nov. in Gunung Mulu National Park, Miri, Sarawak; (C)
egg shells found in the habitat.
Comparisons. Cnemaspis lagang sp. nov. differs from other Bornean Cnemaspis by having Raw Umber head
with Venetian Blue shade on interorbital region; 13–14 supralabials (versus 11–13 in C. leucura and 12–13 in C.
sirehensis sp. nov.); 9–10 postmentals (versus 3 in C. kendallii, 5–7 in C. nigridia, 6–9 in C. paripari, 4–6 in C.
leucura, 5–9 in C. matahari sp. nov., and 6–7 in C. sirehensis sp. nov.); discontinuous rows of 7–8, pore-bearing,
precloacal scales (versus none in C. kendallii, 5–6 in C. dringi, and 14–15 in C. nigridia,); 1–2/1–2 (L/R) postcloacal
tubercles (versus 2–3/2–3 in C. kendallii, 2–3/2–3 in C. paripari, 3–8/3–8 in C. leucura and 2–7/2–7 in C. matahari
sp. nov.); ventrolateral caudal tubercles absent (versus present in C. kendallii, C. nigridia, C. paripari, C. leucura
and C. matahari sp. nov.); keeled subcaudals versus smooth in C. nigridia; no enlarged median subcaudal scales;
tail with Faint Dusky Brown and Trogon Yellow caudal bands anteriorly and immaculate posteriorly (versus Dusky
Brown and Buff-Yellow caudal bands in C. kendallii, Dusky Brown and Orange Yellow caudal bands in C. nigridia,
THREE NEW CNEMASPIS FROM BORNEO Zootaxa 5120 (1) © 2022 Magnolia Press · 11
Pratt’s Payne’s Gray anteriorly and immaculate posteriorly in C. paripari, faint Dusky Brown and Lavender caudal
bands anteriorly, immaculate posteriorly in C. matahari sp. nov., and Brownish Olive and Sulfur Yellow caudal
bands in C. sirehensis sp. nov.); 25–27 subdigital fourth finger lamellae (versus 29 in C. dringi and 26–31 in C.
matahari sp. nov.).
Cnemaspis matahari sp. nov.
White Rock Gecko; Cicak Batu Putih
Fig. 5, Fig. 6, Table 2
Holotype. Adult male, UNIMAS 9602, collected by Hayden Davis and Izneil Nashriq on 5 August 2017, from the
Serian-Tebedu limestone hills (1.131015°N, 110.443988°E; 50 m), Serian, Sarawak, East Malaysia (Borneo).
Paratypes. Serian-Tebedu limestone hills (1.131015°N, 110.443988°E; 50 m), Sarawak, East Malaysia (Borneo)
UNIMAS 9603 collected by Hayden Davis and Izneil Nashriq on 5 August 2017; UNIMAS 9606, UNIMAS
9607, UNIMAS 9608, collected by Hayden Davis and Izneil Nashriq on 28 May 2018; UNIMAS 9573, UNIMAS
9574, collected by Izneil Nashriq and Indraneil Das on 22 June 2019; Kampung Mambong, Siburan (1.355571°N,
110.350767°E; 50 m) UNIMAS 9611, collected by Hayden Davis and Izneil Nashriq on 4 June 2018; Jambusan-
Semadang limestone hills (1.318342°N, 110.254339°E; 50 m), Sarawak, East Malaysia (Borneo) UNIMAS 9615,
collected by Hayden Davis and Izneil Nashriq on 6 June 2018.
Diagnosis. SVL up to 56 mm; 11–14 supralabials; 9–13 infralabials; 2–3 internasals; 5–9 postmentals; ventral
scales keeled; adult males with a discontinuous row of 6–12 pore-bearing, precloacal scales with round pores
arranged in a chevron, separated at midline by 2–4 poreless scales; paravertebral and lateral row of caudal tubercles
present; ventrolateral caudal tubercles absent; caudal tubercles not encircling tail; subcaudals keeled, bearing an
enlarged median row of keeled subcaudal scales; 2–7/2–7 (L/R) postcloacal tubercles on each side of tail base; no
enlarged femoral or subtibial scales; submetatarsal scales of first toe enlarged; 24–31 fourth toe subdigital lamellae;
faint Dusky Brown and Lavender caudal bands anteriorly, immaculate posteriorly; and regenerated tail Spectrum
Yellow.
Description of holotype. Male with an original tail; snout-vent length 55 mm; supralabials 14/13 (L/R);
infralabials 12/12 (L/R); head short (HL/SVL 0.26), narrow (HW/SVL 0.18), depressed (HD/HL 0.43), distinct
from neck; snout long (ES/HW 0.65), much longer than eye diameter (ED/ES 0.50); scales on snout and forehead
weakly keeled, with posterior portion of each scale raised; scales on snout larger than those on occipital region;
eye small (ED/HL 0.22); orbits of eyes with extra-brillar fringes; pupil round; enlarged supraciliaries on top half of
orbit; tympanum deep, oval shaped, greatest diameter vertically, narrow (EL/HL 0.11); eye to ear distance greater
than diameter of eyes (EE/ED 1.19); rostral half as deep as wide, contacted posteriorly by 2 nasals and 2 internasals,
rostrals is in contact with supralabial I. Nostrils oval, situated within nasals, and oriented dorsally; nostrils are in
narrow contact with supralabial I; 6 postnasals bound nasal; mental large, subtriangular, much deeper than wide,
five postmentals border mental; chin scales meet infralabials.
Body slender, short (AG/SVL 0.41); ventral scales equal in size from chin region to gular; increase in size
from gular to pectoral and abdominal regions, weakly keeled. Dorsal scales increase in size from head to nape
and subequal throughout trunk. Scales on dorsum at midbody approximately equal to those of venter at same
level; vertebral scales not reduced; paravertebral rows of tubercles on dorsum present; dorsal tubercles extend from
occiput to base of tail; tubercles dense dorsally and absent on lower flanks; pectoral and abdominal scales distinctly
elongated, imbricate and unicarinate; precloacal scales oval, unicarinate; discontinuous row of 6/6 (L/R), pore
bearing, precloacal scales with round pores arranged in a chevron, separated by 3 keeled, poreless scales; no femoral
pores; no preanal groove.
Forelimbs moderately long, slender, shorter than hindlimbs (FL/SVL 0.20, TBL/SVL 0.23). Dorsal scales
on forelimbs, raised, unicarinate, juxtaposed, reduced in size posteriorly. ventral scales of forelimbs slightly
raised, slightly unicarinate, juxtaposed, reduced in size posteriorly. Dorsal scales of hindlimbs raised, unicarinate,
juxtaposed, reduced in size posteriorly; ventral scales of hindlimbs slightly raised, slightly unicarinate, juxtaposed,
reduced in size posteriorly. Palmar and plantar scales smooth, granular, raised. Digits elongate, all bearing claws that
are slightly recurved; subdigital scansors entire, except for 1–2 fragmented at base of digits, unnotched; an enlarge
scansor towards the base of digits, which is more than twice width of other scansors; interdigital webbing absent.
Subdigital lamellae (manus) I (15); II (20); III (26); IV (29); V (20); (pes) I (12); II (20); III (24); IV (29); V (23).
NASHRIQ ET AL.
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FIGURE 5: (A) Cnemaspis matahari sp. nov., paratype UNIMAS 9573, SVL 56 mm, adult male from Serian–Tebedu limestone
hills, Serian District, First Division, Sarawak. (B) type series displaying dorsal side of the body (from left to right: above,
UNIMAS 9608, UNIMAS 9607, UNIMAS 9606, UNIMAS 9603, UNIMAS 9602; below, UNIMAS 9574, UNIMAS 9573,
UNIMAS 9615 and UNIMAS 9611).
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FIGURE 6: Ventral view of Cnemaspis matahari sp. nov. type series (from left to right: above, UNIMAS 9608, UNIMAS
9607, UNIMAS 9606, UNIMAS 9603, UNIMAS 9602; below, UNIMAS 9574, UNIMAS 9573, UNIMAS 9615 and UNIMAS
9611); Close up of body parts (A) Postmental, (B) Cloaca, (C) Subcaudal, (D) Manus and (E) Pes.
NASHRIQ ET AL.
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FIGURE 7: Habitat and microhabitat of Cnemaspis matahari sp. nov. in Serian limestone region, Serian District, Sarawak,
East Malaysia (Borneo)
Original tail, longer than snout-vent length (TL/SVL 1.28); tail base distinctly swollen; tail arranged in segmented
whorls; caudal tubercles keeled, arranged on paravertebral and lateral rows; caudal tubercles do not encircle tail at
each whorl; 2/2 (L/R) postcloacal tubercles; tail with distinct pairs of furrows laterally; subcaudal scales unicarinate;
a single median row of enlarged, keeled, imbricate, subcaudal scales with 3–4 scales per segment.
Variation in paratype. The seven males (UNIMAS 9573, 9603, 9606-9608, 9611, 9615) and one female
paratype (UNIMAS 9574) closely resemble the holotype in colour and pattern, but differ in the shade of colouration.
Spectrum Yellow on upper flanks extends to paravertebral region or further to axillary region. 10–14 supralabials;
9–13 infralabials; 2–3 internasals; 5–9 postmentals; discontinuous row of 6–12 precloacal scales with round pores
separated at midline by 2–4 poreless scales; 2–7/2–7 (L/R) postcloacal tubercles on side of tail base; tail arranged
in segmented whorls; caudal tubercles do not encircle tail at each whorl; subcaudals with a single median row of
imbricated, enlarged, keeled, raised, subtriangular scales, with 3–4 scales per segment. All show regenerated tails
with differing proportions of original tail present. Tails regenerated from base or middle are yellow. Scales from
dorsal to ventrolateral surfaces of regenerated tail are small, keeled, slightly imbricate, and subequal in size.
Skeletal notes. Holotype UNIMAS 9602 and paratype UNIMAS 9603 have 26 presacral and 2 sacral vertebrae.
The phalangeal formulae are 2–3–4–5–3 manus and 2–3–4–5–4 pes. Both specimens are skeletally mature, showing
fusion of epiphyses.
Colouration in life. Dorsal ground colour of body and limbs Trogon Yellow; head Spectrum Yellow; ground
colour of nape and shoulder region Pratt’s Payne’s Gray bearing a pair of medium, round, Dusky Brown spots
in shoulder axillary; white flecks on nape to shoulder region; faint Sulfur Yellow banding on body; digits Pratt’s
Payne’s Gray with faint dark banding; anterior one-half of tail Lavender bearing faint, Dusky Brown bands; posterior
one-half of original tail immaculate; all ventral surfaces Pratt’s Payne’s Gray except for posterior half of tail, which
is white; and regenerated tail Spectrum Yellow.
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Etymology. The epithet matahari is Malay for sun, in reference to the yellowish tint on the species. It is used
as a noun in apposition.
Natural history. Found on limestone escarpments and outcrops and is exclusive to the Siburan and Serian
Districts. Individuals were collected between 2000–2200 hrs. Surveys suggest the geckos are mostly active (emerging
from rock crevices) when moisture and humidity are high. During dry conditions, when the temperature is relatively
high, geckos are not observed in the open. Water droplets can be seen dripping from the limestone formations at
the sites where this gecko occurs. Small streams are present with their sources within the karst. Limestone hills are
surrounded by vegetation (dipterocarp forest and herbaceous plants, Figure 7). Cnemaspis matahari sp. nov. occurs
sympatrically with the Cyrtodactylus geckos C. limajalur, C. consobrinus and C. pubisulcus, with Cnemaspis found
on the lower part of the limestone hill close to the ground, whereas Cyrtodactylus spp. occur from ground level up
to > 10 m.
Comparisons. Cnemaspis matahari sp. nov. differs from other Bornean Cnemaspis by having a bright yellow
head, body and limbs; 10–14 supralabials; 9–13 infralabials (versus 10–11 in C. lagang sp. nov.); 5–9 postmentals
(versus 3 in C. kendallii, 10 in C. dringi, 10–11 in C. lagang sp. nov.); discontinuous rows of 6–12, pore bearing,
precloacal scales (versus none in C. kendallii, 5–6 in C. dringi, 14–15 in nigridia); 2–7/2–7 (L/R) postcloacal
tubercles (versus 1–2/1–2 in C. lagang sp. nov. and 1–2/1–2 in C. sirehensis sp. nov.); 26–31 subdigital fourth
toe lamellae (versus 29 in C. dringi); having weekly keeled enlarged median subcaudal scales (versus smooth
enlarged median subcaudal in C. nigridia, C. leucura, and C. paripari); faint Dusky Brown and Lavender caudal
bands anteriorly, immaculate posteriorly and regenerated tail Spectrum Yellow (versus Dusky Brown and Buff-
Yellow caudal bands in C. kendallii, Dusky Brown and Orange Yellow caudal bands in C. nigridia, Pratt’s Payne’s
Gray anteriorly and immaculate posteriorly in C. paripari, Faint Dusky Brown and Trogon Yellow caudal bands
anteriorly and immaculate posteriorly in C. lagang sp. nov., and Brownish Olive and Sulfur Yellow caudal bands
in C. sirehensis sp. nov.).
Cnemaspis sirehensis sp. nov.
Blue Day Gecko; Cicak Gua Sireh
Fig. 8, Fig. 9, Table 3
Holotype. Adult female, UNIMAS 9609, collected by Hayden Davis and Izneil Nashriq, on 2 June 2018, from
Gua Sireh (1.180407°N, 110.463391°E; 50 m), Gunung Nambi, Kampung Bantang, Serian District, Sarawak, East
Malaysia (Borneo).
Paratype. Adult female, UNIMAS 9610, collected by Hayden Davis and Izneil Nashriq, on 2 June 2018,
and UNIMAS 9715, collected by Izneil Nashriq, Wong Jye Wen and Indraneil Das, from Gua Sireh (1.180407°N,
110.463391°E, 50 m), Gunung Nambi, Kampung Bantang, Serian District, Sarawak, East Malaysia (Borneo).
Diagnosis. SVL up to 49 mm; 12–13 supralabials; 10–12 infralabials; 3 internasals; 6–7 postmentals; subtibial
scales keeled; ventrolateral caudal tubercles absent; paravertebral and lateral row of caudal tubercles present; caudal
tubercles not encircling tail; subcaudals keeled, bearing an enlarged median row of weakly keeled scales; 1–2/1–2
(L/R) postcloacal tubercles on each side of tail base; no enlarged femoral or subtibial scales; submetatarsal scales of
first toe enlarged; 26–30 fourth toe subdigital lamellae.
Description of holotype. Adult female with original tail, dark banded. Snout-vent length 47 mm; supralabials
12/12 (L/R); infralabials 10/10 (L/R); head oblong, short (HL/SVL 0.30), narrow (HW/SVL 0.18), depressed
(HD/HL 0.36), distinct from neck; snout moderately long (ES/HW 0.69), longer than eye diameter (ED/ES 0.54);
scales on snout and forehead weakly keeled, with posterior portion of each scale raise; scales on snout larger than
those on occipital region; eye small (ED/HL 0.22); orbits of eyes with extra-brillar fringes; pupil round; enlarged
supraciliaries on top half of orbit; tympanum deep, oval shaped, greatest diameter vertically, narrow (EL/HL 0.09);
eye to ear distance greater than diameter of eyes (EE/ED 1.19); rostral half as deep as wide, contacted posteriorly by
2 nasals and 3 internasals, rostrals is in contact with supralabial I. Nostrils oval, situated within nasals, and oriented
dorsally; nostrils are in narrow contact with supralabial I. 5 postnasals bound nasal; mental large, subtriangular,
much deeper than wide, 7 postmentals border mental; chin scales meet infralabials.
Body slender, short (AG/SVL 0.43); ventral scales equal in size from chin region to gular; increase in size
from gular to pectoral and abdominal regions, weakly keeled. Dorsal scales increase in size from head to nape
and subequal throughout trunk. Scales on dorsum at midbody approximately equal to those of venter at same
NASHRIQ ET AL.
16 · Zootaxa 5120 (1) © 2022 Magnolia Press
FIGURE 8: Cnemaspis sirehensis sp. nov., paratype, UNIMAS 9715, SVL 45 mm, adult female from Gua Sireh, Gunung
Nambi, Serian District, First Division, Sarawak. (B) type series displaying dorsal side of the body (from left to right: UNIMAS
9715, UNIMAS 9609 and UNIMAS 9610).
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level; vertebral scales not reduced; paravertebral rows of tubercles on dorsum present; dorsal tubercles extend from
occiput to base of tail; tubercles dense dorsally and absent on lower flanks; pectoral and abdominal scales distinctly
elongated, imbricate and unicarinate; no femoral pores and preanal groove.
FIGURE 9: Ventral view of Cnemaspis sirehensis sp. nov. type series (from left to right: UNIMAS 9715, UNIMAS 9609 and
UNIMAS 9610); close up of body parts (A) Postmental, (B) Cloaca, (C) Subcaudal, (D) Manus and (E) Pes.
NASHRIQ ET AL.
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Forelimbs moderately long, slender, shorter than hindlimbs (FL/SVL 0.20, TBL/SVL 0.23). Dorsal scales
on forelimbs, raised, unicarinate, juxtaposed, reduced in size posteriorly; ventral scales of forelimbs slightly
raised, slightly unicarinate, juxtaposed, reduced in size posteriorly. Dorsal scales of hindlimbs raised, unicarinate,
juxtaposed, reduced in size posteriorly; ventral scales of hindlimbs slightly raised, slightly unicarinate, juxtaposed,
reduced in size posteriorly. Palmar and plantar scales smooth, granular, raised. Digits elongate, all bearing claws that
are slightly recurved; subdigital scansors entire, except for 1–2 fragmented at base of digits, unnotched; an enlarge
scansor towards base of digits, which is over twice width of other scansors; interdigital webbing absent. Subdigital
lamellae (manus) I (13); II (19); III (21); IV (24); V (18); (pes) I (10); II (17); III (23); IV (26); V (23).
Original tail, longer than snout-vent length (TL/SVL 1.22); tail base distinctly swollen; tail arranged in segmented
whorls; caudal tubercles subspinous, raised, weakly keeled, arranged on paravertebral and lateral row; tubercles do
not encircle tail; 2/1 (L/R) postcloacal tubercles; tail with a distinct pair of furrows laterally; a single median row of
enlarged, weakly keeled, imbricate, subcaudal scales with 3 or 4 scales per segment.
Variation in paratypes. A total of two adult females (UNIMAS 9610 and 9715) indistinguishable from holotype
apart from aspects of scale morphology and colouration; 12–13 supralabials; 11–12 infralabials; 6–7 postmentals.
They have yellow regenerated tail. Scales from dorsal to ventral surface of regenerated tail are small, keeled,
slightly imbricate, and equal in size.
Colouration in life. Dorsal ground colour of head, body and limbs Raw Umber; head Trogon Yellow, bearing
small, occipital flecks; nape and shoulder region bearing Dusky Brown patches of irregularly shaped, dark blotches;
Pratt's Payne's Gray patches on paravertebral; small, scattered, Spectrum Yellow flecks between limb insertion;
irregular Citrine bands on forelimbs and hind limbs; Tail with Brownish Olive and Sulfur Yellow banding from tail
base to tip; and Trogon Yellow regenerated tail.
Natural History. The species is nocturnal, with individuals being collected between 2000–2200 hrs from
limestone escarpments surfaces, within narrow rock crevices, outside the entrance of Gua Sireh, in Serian. No
individuals were found inside the cave. Small streams originating from the limestone formation are present. The
limestone hills are edged by dipterocarp forests as well as secondary forests, as well as plantations at the foot of the
hills (Figure 10). Cnemaspis sirehensis sp. nov. occurs sympatrically with Cyrtodactylus geckos (C. consobrinus
and C. pubisulcus) and Hemidactylus sp.
TABLE 4: Min–Max measurement of Bornean Cnemaspis examined (nearest 0.1 mm).
C.
kendallii
C.
nigridia
C. dringi C. paripari C. leucura C. matahari
sp. nov.
C. lagang
sp. nov.
C. sirehensis
sp. nov.
N = 52 N = 5 N = 2 N = 13 N = 6 N = 9 N = 2 N = 3
SVL 33.1–60.4 57.0–68.9 43.3–45.5 40.3–54.4 49.7–63.3 47.1–55.7 46.0 45.1–48.8
HL 13.1–16.7 16.1–20.5 8.4–8.6 11.4–16.5 14.6–17.6 13.9–16.5 13.2 –13.4 13.0–14.1
HW 6.8–10.3 10.4–12.6 6.9–7.0 7.4–10.3 9.1–11.4 8.2–10.6 8.4–13.0 7.8–8.7
HD 4.0–6.7 6.3–7.7 4.5–5.0 4.0–6.3 5.9–7.1 4.9–6.3 4.4–5.1 4.8–5.1
ED 2.0–4.2 3.3–4.3 2.6–3.0 2.4–3.7 4.2–3.0 2.8–3.6 2.8–3.1 3.0–3.1
EE 2.1–4.8 4.0–5.3 3.0–3.2 2.6–4.3 3.1–4.4 3.5–4.4 4.0 3.5–3.7
ES 5.7–7.7 7.1–9.0 5.5–5.7 4.8–7.4 6.0–7.9 5.8–7.4 5.7–6.0 5.7–5.9
EN 3.5–6.2 5.4–6.7 4.3–4.4 3.7–6.0 4.4–6.3 4.1–5.6 4.2–4.8 4.3–4.9
IO 1.4–3.5 1.7–2.8 2.9 1.5–2.5 1.5–2.8 1.6–2.7 1.6–2.2 1.6–2.1
EL 0.9–2.3 1.3–2.1 0.9–1.2 0.9–1.7 1.4–2.1 0.9–1.7 1.1–1.4 1.0–1.2
IN 1.0–2.4 1.6–2.1 1.6–1.7 1.1–1.7 1.5–2.2 1.4–2.1 1.5–2.0 1.0–1.6
TL 0–78.2 22.7–72.3 31.4–62.7 2.4–70.0 35.6–76.3 44.1–71.0 24.5–61.1 57.0–61.8
TW 2.1–6.1 4.5–6.5 4.7–4.9 2.4–5.7 3.6–5.6 3.3–4.7 3.0–4.5 3.3–3.7
FL 7.4–12.2 10.1–13.2 7.6–8.5 8.1–11.0 9.8–12.0 9.2–11.6 8.5–9.1 8.3–9.7
TBL 7.8–14.5 12.7–15.8 10.0–10.9 9.5–13.1 12.5–14.5 10.9–14.3 10.3–10.8 11.1–11.6
AG 18.3–32.1 19.5–28.5 19.4–20.9 19.5–24.5 23.4–30.8 19.6–25.4 19.4–19.8 20.3–23.2
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TABLE 5: Summary of variation in morphology of Bornean species of Cnemaspis. Left/Right counts are given as (L/R). ? indicates missing data.
kendallii nigridia dringi paripari leucura matahari sp.
nov. lagang sp. nov. sirehensis sp.
nov.
Supralabials 9–13/10–14 12–15/12–14 11/11 10–13 12–13/11–13 11–14/10–14 13–14/14 12–13/12–13
Infralabials 9–12/9–12 10–11/10–12 9–13 9–12 10–13/10–12 9–13/10–12 10–11/10–11 10–12/10–12
Internasals 2–3 2 ? 2–3 2–3 2–3 3 3
Postmentals 3 5–7 10 6–9 4–6 5–9 9–10 6–7
Ventral scales Keeled Keeled Keeled Keeled Keeled Keeled Keeled Keeled
Forearm scales Keeled Keeled Smooth Keeled Keeled Keeled Keeled Keeled
Subtibial scales Keeled Keeled Keeled Keeled Keeled Keeled Keeled Keeled
Number of precloacal pores in
males Absent 14–15 5–6 4–8 8–9 6–12 7–8 ?
Precloacal pores continuous or
separated Absent Separated Separated Separated Separated Separated Separated ?
Precloacal pores elongate or round Absent Round Round Round Round Round Round ?
Number of postcloacal tubercles 2–3/2–3 1–3/1–3 Absent 2–3/2–3 3–8/3–8 2–7/2–7 1–2/1–2 1–2/1–2
Lateral caudal furrows Present Present Present Present Present Present Present Present
Caudal tubercles in lateral furrows Absent Absent Absent Absent Absent Absent Absent Absent
Caudal tubercles restricted to a
single paravertebral row on each
side
Absent Absent Absent Absent Absent Absent Absent Absent
Paravertebral caudal tubercles Present Present Present Present Present Present Present Present
Lateral caudal tubercles Present Present ? Present Present Present Present Present
Ventrolateral caudal tubercles Present Present ? Present Present Present Absent Absent
Caudal tubercles encircle tail Present Absent Absent Absent Absent Absent Absent Absent
Anterior ventrolateral caudal
tubercles Present Present ? Present Present Present Absent Absent
Dorsal caudal scales Keeled Keeled Keeled Keeled Keeled Keeled Keeled Keeled
Subcaudal scales Keeled Smooth Keeled Keeled Keeled Keeled Keeled Keeled
Median row of subcaudals Keeled Smooth Keeled Smooth Smooth Keeled Keeled Keeled
Enlarged median subcaudal scale
row Present Present Present Present Present Present Absent Present
Enlarged femoral scales Absent Absent Absent Absent Absent Absent Absent Absent
......Continued on the next page
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TABLE 5. (Continued)
kendallii nigridia dringi paripari leucura matahari sp.
nov. lagang sp. nov. sirehensis sp.
nov.
Shield–like subtibial scales Absent Absent Absent Absent Absent Absent Absent Absent
Enlarged submetatarsal scales on
1st toe Absent Present Present Present Present Present Present Present
Finger lamellae on 1st digit 12–15 14–15 14/16 12–15 14–15 12–15 13–14 12–13
… on 2nd digit 17–23 19–21 24/22 18–24 19–23 19–25 19–21 19–23
…on 3rd digit 23–30 23–24 27/27 21–28 23–29 25–29 23–24 21–28
… on 4th digit 26–33 25–29 29/29 24–31 23–30 27–31 25–27 24–31
… on 5th digit 16–25 18–21 19/21 18–21 18–22 19–24 21–22 18–21
Toe lamellae on 1st digit 11–15 12–13 12/12 10–15 13–15 12–14 11–13 10–13
… on 2nd digit 18–25 20–20 20/21 17–24 17–22 18–22 19 17–24
… on 3rd digit 24–29 23–24 26/25 20–27 24–29 24–28 23 23–27
…on 4th digit 27–32 27–28 30/29 26–31 29–31 26–31 29 26–30
…on 5th digit 20–26 22–24 26/26 22–25 24–26 22–25 24 21–25
THREE NEW CNEMASPIS FROM BORNEO Zootaxa 5120 (1) © 2022 Magnolia Press · 21
FIGURE 10: Habitat (above) and microhabitat (below) of Cnemaspis sirehenesis sp. nov. in Gua Sireh, Gunung Nambi,
Kampung Bantang, Serian District, Sarawak, East Malaysia (Borneo).
NASHRIQ ET AL.
22 · Zootaxa 5120 (1) © 2022 Magnolia Press
Etymology. The epithet sirehensis comes from the type locality, Gua Sireh (Sireh Cave), Kampung Bantang,
Serian and translates to betel in Malay. Betel is a type of plant of the family Piperaceae, consumed as betel quid or
in ‘paan’. Gua Sireh is also known as an important archaeological site in south-western Sarawak.
Comparison. Cnemapsis sirehensis sp. nov. differs from other Bornean Cnemaspis by having a reddish-brown
head, body and limbs; 12–13 supralabials (versus 13–14 in C. lagang sp. nov.); 10–12 infralabials; 6–7 postmentals
(versus 3 in C. kendallii, 10 in C. dringi, and 10–11 in C. lagang sp. nov.); a pair of medium, subtriangular, dusky
brown patches in shoulder region; 1–2/1–2 (L/R) postcloacal tubercles on each side of tail base (versus 2–3/2–3 in C.
kendallii, 2–3/2–3 in C. paripari, 3–8/3–8 in C. leucura and 2–7/2–7 in C. matahari sp. nov.); ventrolateral row of
caudal tubercles absent (except C. lagang sp. nov., C. matahari sp. nov.); keeled median subcaudals (versus smooth
in C. leucura, C. nigridia, C. paripari); tail with Brownish-Olive and Bright Yellow banding from tail base to tail
tip (versus Dusky Brown and Buff-Yellow caudal bands in C. kendallii, Dusky Brown and Orange Yellow caudal
bands in C. nigridia, Pratt’s Payne’s Gray anteriorly and immaculate posteriorly in C. paripari, Faint Dusky Brown
and Trogon Yellow caudal bands anteriorly and immaculate posteriorly in C. lagang sp. nov.); and regenerated tail
Trogon Yellow (versus Spectrum Yellow in C. matahari sp. nov. and Buff Yellow in C. lagang sp. nov.).
A summary of Bornean Cnemaspis morphology is presented in Tables 4 and 5. Compared to nominal south-
east Asian Cnemaspis the Bornean Cnemaspis differ in multiple morphological aspects. The Cau Mau clade (C.
boulengerii and C. psychedelica) have fewer supralabials (7–10 versus 9–15 in Bornean species) and infralabials (5–8
versus 9–13 in Bornean species); caudal tubercles restricted to single paravertebral rows; no lateral caudal furrows;
and plate-like femoral and subtibial scales. The Pattani clade (C. monachorum, C. biocellata, C. niyomwanae,
and C. kumpoli) have smooth ventral scales; and in C. niyomwanae and C. kumpoli, no lateral and ventrolateral
row of caudal tubercles. The Northern Sunda clade, comprising the chantaburiensis group, are the only species of
Cnemaspis that have a dark, mid-gular line, thought to be synapomorphic; the siamensis group exhibit the states
of light-coloured prescapular crescent, a yellow belly and ventral surface of hindlimbs and caudal tubercles not
restricted to a single paravertebral row and encircling tail; the argus group have no lateral or ventrolateral caudal
tubercles; and the affinis group have a lateral row of caudal tubercles, no median row of enlarged subcaudal scales
and submetatarsal scales on first toe.
Discussion
The discovery of the three new Cnemaspis species (C. lagang sp. nov., C. matahari sp. nov. and C. sirehensis
sp. nov.) suggest the possibility of additional cryptic species on Borneo. The current distributions of Bornean
Cnemaspis species are limited to particular geological formations and intact forests within such formations, with six
species (C. kendallii, C. nigridia, C. paripari, C. leucura, C. matahari sp. nov. and C. sirehensis sp. nov.) recorded
in west Sarawak and two recorded species (C. dringi and C. lagang sp. nov.) in the north of Sarawak. The newly
described species occupy isolated formations of west Sarawak, the Serian and Padawan limestone formations (C.
matahari sp. nov. and C. sirehensis sp. nov.) and in north Sarawak, the Mulu limestone formation (C. lagang sp.
nov.). The distribution of the three newly described species is reflected in Figure 11.
Sarawak consists of three distinct geological provinces, coinciding with geographic demarcations -West
Sarawak and Central-North Sarawak- with the Lupar Line bounding the Sibu Zone and Kuching Zone, and the
Bukit-Mersing Line bounding the Miri Zone and Sibu Zone (James 1984; Hutchison 1989). The 15 km wide east-
southeast trending suture zone has been associated with faunal distributional disjunctions, supported in a variety
of taxonomy groups, for instance, lineages of the dipterocarps (Ashton 1972), distributional pattern mirrored by
several lineages (described in detail by Bornbusch & Lundberg 1989; Davis et al. 2021). The distributional limits
to the south on the island of Borneo appear to be coincident with the hill ranges of Penrissen, and the west-flowing
Kapuas, which periodically connected to the east coast of Sumatra (another large island inhabited by Cnemaspis,
of the kendallii group; Amarasinghe et al. 2015), most recently during the Pleistocene. Weber (1921) summarised
geological and biological evidence for such a connection, that is currently referred to as the Riau Pocket (sensu
Corner 1958; 1960; see also Kiew and Saw 2019).
The northern-most representative, C. lagang sp. nov., is found in the Baram Basin, at the headwaters of the
Sungei Baram. The southern-most species, C. leucura, has been recorded from Gunung Penrissen as has C. kendallii,
with which it is occasionally sympatric (but non-syntopic, occupying vegetation such as tree trunks, rather than rock
THREE NEW CNEMASPIS FROM BORNEO Zootaxa 5120 (1) © 2022 Magnolia Press · 23
surfaces). The known distribution of Cnemaspis in Sarawak is fragmented by human intervention (mining activities,
development and land conversion to agriculture). Shifting agriculture and mining activities are intense in Sarawak
and if not mitigated or done sustainably, will not only affect the assemblages of this genus, but in a wider context
will result in loss of diversity in other taxa (Nashriq & Das 2021). This study was focused on western Sarawak. The
formations in western Sarawak were easily accessible compared to those of central and northern Sarawak. Future
efforts should be directed in finding species of Cnemaspis in central and northern Sarawak, especially along the
karst limestone regions of the State.
FIGURE 11: Distribution of the three newly described Cnemaspis species in Sarawak, East Malaysia, Borneo.
Acknowledgements
We are grateful to the staff of the Institute of Biodiversity and Environmental Conservation, Gabriel Tonga Noweg,
Andrew Alek Tuan, Mohd-Azlan Jayasilan, Felicia Anthony Reyap, Pasey Lisus, Rahah Mohamad Yakup, Hasri
Al-Hafiz Haba, Mohsin Zainalabidin, Ketty Daun and Sauyah Su’aut for support and supplies. We are grateful to
the following curators, collection managers and institutions for access to material under their care: BMNH, London
(E.N. Arnold and C.J. McCarthy); FMNH, Chicago (H.K. Voris, R.F. Inger and A. Resetar); MCZ, Cambridge, MA
(J.E. Cadle, J. Hanken and J. Rosado); SBC, Semenggoh (C. Yeo and M. Naming); SM, Kuching (P. Kedit and
C. Leh); USNM, Washington, D.C. (W.R. Heyer, R.I. Crombie and G.R. Zug); and ZRC, Singapore (P.K.L. Ng
and K.K.P. Lim). We thank the Sarawak Forest Department for research permits (147) JHS/NCCD/600-7/2/107/
Jld.2 and Park Permit N0.74/2019. We thank Abang Abdul Rahman Zohari Abang Openg, the Honourable Chief
Minister of Sarawak for approval of the new names of the species on 2 December 2021, and to the staff of the
Sarawak Forestry Corporation, Haji Zolkipli Mohamad Aton and Oswald Braken Tisen for their assistance. The
Sarawak Forestry Corporation, allowed us entrance to the National Parks and other protected areas. At Gunung
Mulu National Park, we thank the authorities, in particular Ellen McArthur for facilitating our research. Funding
came from a grant from the Niche Research Grant Scheme of the Ministry of Higher Education, Government of
Malaysia: NRGS/1087/2013(01) to Indraneil Das. We also thank the Herpetofaunal Biology team, Pui Yong Min,
NASHRIQ ET AL.
24 · Zootaxa 5120 (1) © 2022 Magnolia Press
Anthony K. Pine, Siti Nor Baizurah, Wong Jye Wen, Veronica Martin, Veronica Leah anak Buma, Awang Khairul
Ikhwan and Chien C. Lee for field assistance and company.
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APPENDIX I. List of Comparative Material Examined
Cnemaspis dringi Das & Bauer, 1998. “Labang Camp (03º20’N; 113º29’E), Bintulu District, Fourth Division,
Sarawak, East Malaysia, Borneo” (FMNH 148588, ex-FMNH FS 19914; holotype); “Sungai Segaham (02º44’N;
113º53’E), Belaga District, Seventh Division, Sarawak, East Malaysia” (FMNH 221478, ex-FMNH FS 34321;
paratype).
Cnemaspis kendallii (Gray, 1845). “Borneo” (BMNH XXII.92a, lectotype); Bako National Park, Kuching
Division (FMNH 223201; MCZ 157158–59; SM uncatalogued, ZRC 2.4662 [three specimens]; UNIMAS 9556,
9588); Bau, Kuching Division (BMNH 1911.1.30.7–9); Bidi, Kuching Division (BMNH 1902.12.12.12); Bukit
Kawa, Bau, Kuching Division (SBC R.103); Bukit Meraja, Bau, Kuching Division (SBC R.60); Bukit Poing,
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Bau, Kuching Division (SBC R.104); Bukit Tongga, Bau, Kuching Division (SBC R.27); Gua Pari Pari, Bau,
Kuching Division (UNIMAS 9545, 9548–51, 9587); Gua Silabur, Serian, Samarahan Division (UNIMAS 9504–
05); Gumbang (UNIMAS 9577–82); Gunung Matang, Kuching Division (BMNH RR1962.182; BMNH 99.12.8.1);
Gunung Gading, Lundu, Kuching Division (USNM 76633; UNIMAS 9569, 9593, 9595–96); Gunung Penrissen,
Kuching Division (SM c.c.1.2.2.B; in addition to a second uncatalogued SM specimen; UNIMAS 9623–25);
Gunung Santubong, Kuching Division (SM c.c.1.2.1; ZRC uncatalogued; UNIMAS 9557–58, 9566, 9597–9601);
Gunung Serambu (UNIMAS 9626–28); Gunung Singai (UNIMAS 9542–43); Kampung Haji Baki (UNIMAS 9552);
Kampung Mambong (UNIMAS 9612–14); Kampung Temurang (UNIMAS 9618); Kubah National Park, Kuching
Division (UNIMAS 9554–55, 9617); Kuching, Kuching Division (SM c.c.2.1.D: three specimens); Matang Wildlife
Center, Kuching Division (UNIMAS 9559–60); Ranchan Pool Forest, Serian, Samarahan Division (UNIMAS 9553;
ZRC 2.4938–39); Samunsam Wildlife Sanctuary (UNIMAS 9575–76); Semenggoh Wildlife Sanctuary, Kuching
Division (ZRC 2.4958); Tanjung Datu National Park (UNIMAS 9630–31).
Cnemaspis leucura Kurita, Nishikawa, Matsui & Hikida, 2017. Borneo Highlands Resort, Gunung Penrissen
(UNIMAS 9563–65, 9585).
Cnemaspis nigridia (Smith, 1925) “Mt. Gadin” (BMNH 1946.8.22.90, ex-BMNH 1925.9.1.8; holotype);
BMNH 1974.3790; UNIMAS 9561; 9567–9568; 9592; 9594; SM c.c.1.2.3; individually numbered SM 7659; 7667;
8055; 8060–65; in addition to eight uncatalogued SM specimens; UNIMAS 7980; ZRC 2.1114–115; 2.5324–25;
also, PNM 7948; Lundu, Kuching Division (MCZ 15250); Gunung Pueh, Kuching Division (BMNH 1925.9.1.9–
10); Gunung Beremput, Kuching Division (UNIMAS 7559–7562).
Cnemaspis paripari Grismer & Chan, 2009 “Gua Pari-pari, Bau District, Sarawak, Malaysia (01º22.867
N, 110º07.164 E)” (ZRC 2.6812, holotype; UNIMAS 9544–47; 9584–86); “Gua Angin, Bau District, Sarawak,
Malaysia (01º24.882 N, 110º08.255 E)” (ZRC 2.6813–14, paratypes; UNIMAS 9570–72; 9589–90).
APPENDIX II. Primers used for amplification and sequencing in the current study.
Primer Name Primer Reference Primer Sequence
ND2-METF1 Macey et al. (1997) 5’ AAGCTTTCGGGCCCATACC 3’
ND2-COIRI Macey et al. (1997) 5’ AGRGTGCCAATGTCTTTGTGRTT 3’
APPENDIX III. GenBank accession numbers and locality for the taxa included in the phylogenetic analyses. The
accession numbers represent all ND2 sequences of gekkonid lizards of the genus Cnemaspis and outgroups used to
generate phylogenetic relationships.
Species Accession No. Museum No. Locality
Ailuronyx seychellensis KY038014.1 PL17 NA
Cnemaspis affinis KM024684.1 LSUHC:6757 Pulau Pinang, Penang, Malaysia
Cnemaspis affinis KM024685.1 LSUHC:6758 Pulau Pinang, Penang, Malaysia
Cnemaspis argus KM024687.1 LSUHC:8304 Gunung Lawit, Terengganu, Malaysia
Cnemaspis argus KM024691.1 LSUHC:10859 Gunung Tebu, Terengganu, Malaysia
Cnemaspis aurantiacopes KM024692.1 LSUHC:8610 Hon Dat Hill, Vietnam
Cnemaspis baueri KM024696.1 LSUHC:7302 Pulau Aur, Johor, Malaysia
Cnemaspis baueri KM024697.1 LSUHC:7303 Pulau Aur, Johor, Malaysia
Cnemaspis bidongensis KM024704.1 LSUHC:11445 Pulau Bidong, Terengganu
Cnemaspis bidongensis KM024706.1 LSUHC:11447 Pulau Bidong, Terengganu
Cnemaspis biocellata KM024707.1 LSUHC:8789 Gua Kelam, Perlis, Malaysia
Cnemaspis biocellata KM024709.1 LSUHC:8818 Kuala Perlis, Perlis, Malaysia
Cnemaspis boulengerii KM024710.1 LSUHC:9278 Con Dao Archipelago, Vietnam
Cnemaspis caudanivea KM024712.1 LSUHC:8577 Hon Tre Island, Vietnam
Cnemaspis caudanivea KM024713.1 LSUHC:8578 Hon Tre Island, Vietnam
,,,,,Continued on the next page
THREE NEW CNEMASPIS FROM BORNEO Zootaxa 5120 (1) © 2022 Magnolia Press · 27
APPENDIX III (Continued)
Species Accession No. Museum No. Locality
Cnemaspis chanardi KM024715.1 LSUHC:9567 Thum Thong Panra, Thailand
Cnemaspis chanthaburiensis KM024716.1 LSUHC:9338 Phnom Dalai, Cambodia
Cnemaspis flavigaster KM024718.1 LSUHC:8835 Kepong, Selangor, Malaysia
Cnemaspis flavigaster KM024720.1 LSUHC:10380 Ulu Gombak, Selangor, Malaysia
Cnemaspis flavolineata KM024721.1 LSUHC:8079 Fraser’s Hill, Pahang, Malaysia
Cnemaspis grismeri KM024722.1 LSUHC:9969 Lenggong, Perak, Malaysia
Cnemaspis grismeri KM024723.1 LSUHC:9970 Lenggong, Perak, Malaysia
Cnemaspis hangus KM024727.1 LSUHC:9358a Bukit Hangus, Pahang, Malaysia
Cnemaspis hangus KM024729.1 HC225 Bukit Hangus, Pahang, Malaysia
Cnemaspis harimau KM024730.1 LSUHC:9665 Gunung Jerai, Kedah, Malaysia
Cnemaspis harimau KM024732.1 LSUHC:9668 Gunung Jerai, Kedah, Malaysia
Cnemaspis huaseesom KM024734.1 LSUHC:9457 Sai Yok National Park, Thailand
Cnemaspis huaseesom KM024735.1 LSUHC:9458 Sai Yok National Park, Thailand
Cnemaspis karsticola KM024736.1 LSUHC:9054 Gunung Reng, Kelantan, Malaysia
Cnemaspis karsticola KM024737.1 LSUHC:9055 Gunung Reng, Kelantan, Malaysia
Cnemaspis kendallii KM024740.1 LSUHC:9172 Gunung Gading, Lundu, Sarawak,
Malaysia
Cnemaspis kendallii KM024743.1 LSUHC:9178 Gunung Santubong, Sarawak, Malaysia
Cnemaspis kendallii OL770080 UNIMAS 9604 Lobang Batu, Serian, Sarawak, Malaysia
Cnemaspis kendallii OL770081 UNIMAS 9605 Lobang Batu, Serian, Sarawak, Malaysia
Cnemaspis kendallii OL770082 UNIMAS 9618 Padawan, Sarawak, Malaysia
Cnemaspis kumpoli KM024746.1 LSUHC:8848 Perlis State Park, Perlis, Malaysia
Cnemaspis leucura LC205721.1 KUHE:57421 Kampung Sadir, Padawan, Sarawak,
Malaysia
Cnemaspis leucura LC205722.1 KUHE:57424 Kampung Sadir, Padawan, Sarawak,
Malaysia
Cnemaspis leucura LC205723.1 KUHE:57423 Kampung Sadir, Padawan, Sarawak,
Malaysia
Cnemaspis leucura LC205724.1 KUHE:57422 Kampung Sadir, Padawan, Sarawak,
Malaysia
Cnemaspis limi KM024747.1 LSUHC:3902 Pulau Tioman, Pahang, Malaysia
Cnemaspis limi KM024750.1 LSUHC:3888 Pulau Tioman, Pahang, Malaysia
Cnemaspis lineogularis KY091232.1 BYU:62536 Wat Khao Daen, Thailand
Cnemaspis lineogularis KY091233.1 ZMKU: R00728 Wat Khao Daen, Thailand
Cnemaspis mahsuriae KT250633.1 LSUHC:11828 Gunung Raya, Kedah, Malaysia
Cnemaspis mahsuriae KT250634.1 LSUHC:11829 Gunung Raya, Kedah, Malaysia
Cnemaspis matahari sp. nov. OL770084 UNIMAS 9602 Serian-Tebedu, Sarawak, Malaysia
Cnemaspis matahari sp. nov. OL770085 UNIMAS 9603 Serian-Tebedu, Sarawak, Malaysia
Cnemaspis matahari sp. nov. OL770086 UNIMAS 9607 Serian-Tebedu, Sarawak, Malaysia
Cnemaspis matahari sp. nov. OL770087 UNIMAS 9608 Serian-Tebedu, Sarawak, Malaysia
Cnemaspis mcguirei KM024752.1 LSUHC:8854 Bukit Larut, Perak, Malaysia
Cnemaspis mcguirei KM024753.1 LSUHC:8855 Bukit Larut, Perak, Malaysia
Cnemaspis monachorum KM024754.1 LSUHC:9114 Pulau Langgun, Kedah, Malaysia
Cnemaspis monachorum KM024756.1 LSUHC:10808 Pulau Langgun, Kedah, Malaysia
Cnemaspis mumpuniae KM024760.1 MZBLace10166 Sekunyam Forest Reserve, Riau
,,,,,Continued on the next page
NASHRIQ ET AL.
28 · Zootaxa 5120 (1) © 2022 Magnolia Press
APPENDIX III (Continued)
Species Accession No. Museum No. Locality
Cnemaspis mumpuniae KM024761.1 MZBLace10167 Sekunyam Forest Reserve, Riau
Cnemaspis narathiwatensis KM024762.1 USMHC1347 Sungai Enam, Perak, Malaysia
Cnemaspis narathiwatensis KM024765.1 USMHC1350 Sungai Enam, Perak, Malaysia
Cnemaspis neangthyi KM024767.1 LSUHC:8515 O’Lakmeas, Cambodia
Cnemaspis neangthyi KM024769.1 LSUHC:8517 O’Lakmeas, Cambodia
Cnemaspis nigridia KM024770.1 LSUHC:9168 Gunung Gading, Lundu, Sarawak,
Malaysia
Cnemaspis nigridia KM024771.1 LSUHC:9169 Gunung Gading, Lundu, Sarawak,
Malaysia
Cnemaspis nigridia KM024772.1 LSUHC:9170 Gunung Gading, Lundu, Sarawak,
Malaysia
Cnemaspis nigridia LC158342.1 KUZR27145 Gunung Gading, Lundu, Sarawak,
Malaysia
Cnemaspis niyomwanae KM024773.1 LSUHC:9568 Thum Khao Ting, Thailand
Cnemaspis niyomwanae KM024774.1 LSUHC:9571 Thum Khao Ting, Thailand
Cnemaspis nuicamensis KM024776.1 LSUHC:8647 Nui Cam Hill, Vietnam
Cnemaspis nuicamensis KM024778.1 LSUHC:8649 Nui Cam Hill, Vietnam
Cnemaspis omari KM024779.1 LSUHC:9978 Perlis State Park, Perlis, Malaysia
Cnemaspis paripari OL770083 UNIMAS 9585 Gua Pari Pari, Bau, Sarawak, Malaysia
Cnemaspis paripari LC205727.1 KUHE:57286 Gua Pari Pari, Bau, Sarawak, Malaysia
Cnemaspis paripari LC205728.1 KUHE:57289 Gua Pari Pari, Bau, Sarawak, Malaysia
Cnemaspis paripari LC205731.1 KUHE:57122 Gua Angin, Bau, Sarawak, Malaysia
Cnemaspis paripari LC205732.1 KUHE:57268 Gua Angin, Bau, Sarawak, Malaysia
Cnemaspis paripari KM024783.1 LSUHC:9186 Gua Angin, Bau, Sarawak, Malaysia
Cnemaspis pemanggilensis KM024785.1 LSUHC:8011 Pulau Pemanggil, Johor, Malaysia
Cnemaspis pemanggilensis KM024788.1 LSUHC:8014 Pulau Pemanggil, Johor, Malaysia
Cnemaspis peninsularis KM024805.1 LSUHC:8122 Selai, Johor, Malaysia
Cnemaspis peninsularis KM024810.1 LSUHC:9376 Pulau Tenggol, Terengganu, Malaysia
Cnemaspis perhentianensis KM024820.1 LSUHC:8699 Pulau Perhentian Besar, Terengganu,
Malaysia
Cnemaspis perhentianensis KM024821.1 LSUHC:8700 Pulau Perhentian Besar, Terengganu,
Malaysia
Cnemaspis phangngaensis KY091234.1 BYU:62537 Phang Nga, Thailand
Cnemaspis phangngaensis KY091235.1 BYU:62538 Phang Nga, Thailand
Cnemaspis pseudomcguirei KM024824.1 LSUHC:9145 Bukit Larut, Perak, Malaysia
Cnemaspis pseudomcguirei KM024825.1 LSUHC:9146 Bukit Larut, Perak, Malaysia
Cnemaspis psychedelica KM024827.1 LSUHC:9243 Hon Khoai Island, Vietnam
Cnemaspis psychedelica KM024828.1 LSUHC:9244 Hon Khoai Island, Vietnam
Cnemaspis punctatonuchalis KY091236.1 BYU:62539 Thap Sakae, Thailand
Cnemaspis punctatonuchalis KY091237.1 BYU:62540 Thap Sakae, Thailand
Cnemaspis roticanai KM024829.1 LSUHC:9430 Gunung Raya, Pulau Langkawi,
Malaysia
Cnemaspis roticanai KM024830.1 LSUHC:9431 Gunung Raya, Pulau Langkawi,
Malaysia
Cnemaspis
selamatkanmerapoh
KM024833.1 LSUHC:11016 Gua Gunting, Pahang, Malaysia
,,,,,Continued on the next page
THREE NEW CNEMASPIS FROM BORNEO Zootaxa 5120 (1) © 2022 Magnolia Press · 29
APPENDIX III (Continued)
Species Accession No. Museum No. Locality
Cnemaspis shahruli KM024835.1 LSUHC:9163 Pulau Jerejak, Penang, Malaysia
Cnemaspis shahruli KM024836.1 LSUHC:9586 Sungai Sadim, Kedah, Malaysia
Cnemaspis siamensis KM024838.1 LSUHC:9474 Pathio, Thailand
Cnemaspis siamensis KM024839.1 LSUHC:9485 Pathio, Thailand
Cnemaspis sirehensis sp.
nov.
OL770088 UNIMAS 9609 Gua Sireh, Serian, Sarawak, Malaysia
Cnemaspis sirehensis sp.
nov.
OL770089 UNIMAS 9610 Gua Sireh, Serian, Sarawak, Malaysia
Cnemaspis stongensis KM024841.1 LSUHC:11090 Gunung Stong, Kelantan, Malaysia
Cnemaspis stongensis KM024842.1 LSUHC:11091 Gunung Stong, Kelantan, Malaysia
Cnemaspis sundainsula KM024845.1 MZBLace10156 Pulau Natuna Besar
Cnemaspis sundainsula KM024846.1 MZBLace10157 Pulau Natuna Besar
Cnemaspis temiah KM024850.1 LSUHC:9739 Cameron Highland, Pahang, Malaysia
Cnemaspis temiah KM024851.1 LSUHC:9816 Cameron Highland, Pahang, Malaysia
Cnemaspis thachanaensis KY091243.1 BYU:62543 Tha Chana, Thailand
Cnemaspis thachanaensis KY091244.1 BYU:62544 Tha Chana, Thailand
Cnemaspis tucdupensis KM024852.1 LSUHC:8631 Tuc Dup Hill, Vietnam
Cnemaspis tucdupensis KM024853.1 LSUHC:8632 Tuc Dup Hill, Vietrnam
Urocotyledon inexpectata KY038015.1 2MA29 NA
