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Accepted by C. Siler: 16 Feb. 2017; published: 3 May 2017
ZOOTAXA
ISSN 1175-5326 (print edition)
ISSN
1175-5334
(online edition)
Copyright © 2017 Magnolia Press
Zootaxa 4258 (6): 539
–
550
http://www.mapress.com/j/zt/
Article
539
https://doi.org/10.11646/zootaxa.4258.6.3
http://zoobank.org/urn:lsid:zoobank.org:pub:D8121CF9-5F38-45B1-AB19-CAD03A95CC0B
A new species of Tropidophorus Duméril & Bibron, 1839 (Squamata: Sauria:
Scincidae) from Sarawak, East Malaysia (Borneo)
YONG MIN PUI
1
, BENJAMIN R. KARIN
2,3
, AARON M. BAUER
2
& INDRANEIL DAS
1
1
Institute of Biodiversity and Environmental Conservation, Universiti Malaysia Sarawak, 94300, Kota Samarahan, Sarawak, Malaysia.
E-mail: pui8783ibec@gmail.com; idas@unimas.my
2
Department of Biology, Villanova University, 800 Lancaster Avenue, PA 19085, USA. E-mail: aaron.bauer@villanova.edu
3
Museum of Vertebrate Zoology and Department of Integrative Biology, 3101 Valley Life Sciences Building, University of California
Berkeley, Berkeley, CA 94720, USA. E-mail: benkarin@berkeley.edu
Abstract
A new species of the genus Tropidophorus is described from Putai, upper Baleh, Kapit districts, Sarawak, East Malaysia
(Borneo). Tropidophorus sebi sp. nov. is diagnosable from congeners from Borneo by the combination of the following
characters: head shields present, dorsal and lateral scales smooth; parietal scales in two pairs; supraciliaries eight; supraoc-
ulars four; supralabials seven; infralabials four; postmental undivided; longitudinal scale rows 58; ventrals 53; transverse
scale rows at midbody 34; subcaudals 98; preanals enlarged, single; and subdigital lamellae of Toe IV 19. In addition, we
determine the phylogenetic position of this species within the Tropidophorus group based on mitochondrial markers, and
present a key to identification of the known Bornean species in the genus.
Key words: Borneo, new species, Scincidae, Tropidophorus sebi sp. nov.
Introduction
The water skink genus Tropidophorus Duméril & Bibron, 1839 (type species: T. cocincinensis Duméril & Bibron,
1839) is diagnosed by the exposure of the tympanum and presence of a single scale at the anterior and posterior
corners of the eyelid (see Hikida et al. 2002; Greer & Biswas 2004). Remarkable amongst members of its family,
these lizards are predominantly aquatic (Barbour 1921a; Bauer & Jackman 2008) and obligates of lowland forests.
Currently, the genus comprises 28 nominal species that range from Bangladesh and north-eastern India, through
mainland south-east Asia and southern China, southwards to Indo-China, the Malay Peninsula and Borneo, as well
as the southern Philippines and Sulawesi (Brown & Alcala 1980; Honda et al. 2006; Ziegler et al. 2005; Nguyen et
al. 2010a). On Borneo, six species of Tropidophorus have been recorded—T. beccarii Peters, 1871, T. brookei
(Gray, 1845), T. iniquus van Lidth de Jeude, 1905, T. micropus van Lidth de Jeude, 1905, T. mocquardii Boulenger,
1894 and T. perplexus Barbour, 1921 (de Rooij 1915; Das 2004).
In this communication, we describe a new species of Tropidophorus from a previously unsurveyed site in
central Sarawak, East Malaysia, in northern Borneo. We allocate this species to this nominal genus for showing the
following characters that are currently associated with the genus (see Smith 1923, Mittleman 1952; Taylor 1963;
Manthey & Grossmann 1997; Hikida et al. 2002; Greer & Biswas 2004), in addition to genetic data provided
herein: lack of palatine teeth; limbs and eyelids well developed and moveable; lower eyelid scaly; supranasals
absent; prefrontals well developed; frontoparietal distinct from interparietal; tympanum large and superficial; and
2–3 large preanals. The new Tropidophorus differs from congeners in a combination of size, colour pattern, body
proportion and squamation characters. We define a species using the General Lineage Concept of de Queiroz
(1998, 1999) as an extension of the Evolutionary Species Concept of Wiley (1978), in which phenotypically
divergent populations represent distinct and separately evolving lineages. For the new Tropidophorus, we consider
the possession of non-overlapping and diagnostic morphological characters relative to its Sundaic congeners to be
indicative of speciation.
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Materials and methods
The type specimens were collected between 19–20 May 2015. Specimens were photographed in life, euthanized,
fixed in formalin, and subsequently washed in water and transferred to 70% ethanol about a week after collection.
Specimens were deposited at the Museum of the Institute of Biodiversity and Environmental Conservation,
Universiti Malaysia Sarawak. Sexes of the type specimens were determined through dissection. The following
measurements were taken with Mitutoyo
TM
dial vernier callipers (to the nearest 0.1 mm) approximately two months
after collection: snout–vent length (SVL, from snout tip to cloaca); tail length (TaL, from cloaca to tail tip); total
length (TL, from snout tip to tail tip); head length (HL, distance between angle of jaws and snout-tip); head width
(HW), measured across retroarticular process of the mandible; head depth (HD), greatest transverse depth of head,
taken posterior of the orbital region; tibia length (TBL), distance between surface of knee to surface of heel, with
the knee flexed; eye diameter (ED), horizontal diameter of the eyes-measured across the anterior and posterior
margins of the palpebral margin; internarial distance (IN), distance between nostrils; eye–snout distance (E–S),
distance between anteriormost point of eyes and tip of snout; eye–nostril distance (E–N), distance between
anteriormost point of eyes and posterior-edge of nostrils; nostril–snout distance (N–S), distance between
anteriormost point of nostril and tip of snout; axilla–groin distance (A–G), distance between posterior edge of
forelimb at its insertion to body to anterior edge of hind limb at its insertion to body and body width (BW), greatest
width of body. Scalation: nuchals (enlarged scales behind parietals); paravertebral scales (number of dorsal scales
from posterior edge of parietals to a point above vent); ventrals in transverse rows (number of scales from first
gular to precloacal scales), and scale rows at position of 10th subcaudal on tail including subcaudal.
Sources of data on character states and distribution include material in Appendix I as well as the following
works: Barbour (1921b); Boulenger (1894); Bourret (2009); Bacon in Brown & Alcala (1980); Chuaynkern et al.
(2005); Das (2004, 2010); de Rooij (1915); Fei et al. (2010); Guo et al. (2010); Hartmann et al. (2009); Hikida &
Ota (1994); Hikida et al. (2002); Malkmus et al. (2002); Manthey & Grossmann (1997); Ngo et al. (2000); Nguyen
et al. (2010a; 2010b); Rao et al. (2011); Smith (1919; 1923); Taylor (1963); van Lidth de Jeude (1905); Waiprom et
al. (2013); Wen (1992); and Ziegler et al. (2005; 2007).
Museum abbreviations follow Sabaj (2016), except for the Lee Kong Chian Museum of Natural History
(formerly, the Raffles Museum of Biodiversity Research, National University of Singapore, Singapore) for which
we use the acronym ZRC, following widespread usage; BM = Brunei Museum, Bandar Seri Begawan, Brunei
Darussalam; BMNH = The Natural History Museum London; FMNH = Field Museum of Natural History,
Chicago, U.S.A.; MCZ = Museum of Comparative Zoology, Harvard University, Cambridge, U.S.A.; RMNH =
Naturalis, the Nationaal Natuurhistorisch Museum, Leiden, the Netherlands; SBC = Sarawak Biodiversity Centre
Museum, Jalan Semenggoh, Malaysia; SM = Sarawak Museum, Kuching, Malaysia; UBD = Museum of Brunei
Darussalam, Bandar Seri Begawan, Brunei Darussalam; USNM = National Museum of Natural History,
Smithsonian Institution, Washington, D.C., U.S.A.
Genomic DNA was extracted as described in Aljanabi & Martinez (1997). Polymerase chain reactions (PCR)
was performed on a Eppendorf Mastercycler nexus gradient thermocycler and PCR product was cleaned using a
home-made magnetic bead solution (Rohland & Reich 2012). All PCR reactions began with an initial 2 min
denaturation at 95°C, followed by 95°C for 35 s, annealing for 35 s at 50°C and extension at 72°C for 1 min 35 s
for 34 cycles. Cycle sequencing was performed using Big Dye v3.1 chemistry, followed by an additional magnetic
bead cleanup and analysis on an ABI3730xl. We sequenced the mitochondrial ribosomal 12s (395 bp) and
ribosomal 16s (537 bp) genes for comparison with previously sequenced Tropidophorus. For amplifications of 12s,
we used the primers L1019 (5’-AAAAAGCTTCAAACTGGGATTAGATACCCCACTAT-3’) and H1478 (5’-
TGACTGCAGAGGGTGACGGGCGGTGTGT-3’) of Kocher et al. (1989) and for 16s we used the primers 16sbr-
L (5’- CGCCTGTTTATCAAAAACAT-3’) and 16sbr-H (5’- CCGGTCTGAACTCAGATCACGT-3’) of Palumbi
et al. (1991). All newly generated sequences are available on GenBank (GB Accession Numbers KY488450,
KY488451, KY488452, KY488453, KY488454, KY488455, KY488456 and KY488457). DNA reads were
manually checked and corrected for errors. Consensus sequences were aligned by eye to all Tropidophorus
sequences available on GenBank: for 12s (AB028811, AB222933, AB222934, AB222935, AB222936, AB222937,
AB222938, AB222939, AB222940, AB222941, AB222942, AB222943, AB222944, AB222945, AB222946,
AB222947, AB222948, AY308473, AY308474) and for 16s (AB028823, AB222949, AB222950, AB222951,
AB222952, AB222953, AB222954, AB222955, AB222956, AB222957, AB222958, AB222959, AB222960,
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AB222961, AB222962, AB222963, AB222964, AY308322, AY308323, EF611186, GU550104, GU550105).
These data were further aligned to the 12s and 16s genes from the existing dataset of Sphenomorphus group skinks
of Linkem et al. (2011) downloaded from Dryad (http://datadryad.org/resource/doi:10.5061/dryad.30064) and
trimmed to include a single specimen from each species. Pairwise sequence divergences were assessed using
Geneious v9.0.4 (http://www.geneious.com, Kearse et al. 2012).
We concatenated the two genes and performed maximum likelihood (ML) and Bayesian inference (BI)
phylogenetic analyses using RAxML v8.1.15 (Stamatakis 2006) and MrBayes v3.2.1 (Ronquist et al. 2012). For
each analysis, we determined the optimal partitioning scheme using PartitionFinder v.1.1.1 (Lanfear et al. 2012),
with the BIC and greedy algorithm supporting separate partitions for each mitochondrial gene and the GTR+I+G
evolutionary model in both partitions. For the maximum likelihood analysis, we ran RAxML with both partitions
under the GTRCAT model of evolution with 1000 rapid bootstrap replicates. For Bayesian inference, we ran the
Markov Chain Monte Carlo in MrBayes for 20 million generations for two runs with four chains each, specifying
the GTR+I+G model for both gene partitions. We assessed adequate burn-in of the Markov Chains in Tracer v1.6
(Rambaut & Drummond 2013) by eye and by ensuring all ESS values were greater than 200.
Systematics
Tropidophorus sebi sp. nov.
(Figs. 1–2)
Holotype. Museum of the Institute of Biodiversity and Environmental Conservation, UNIMAS P1167, from a
small stream located at the First Count Logging Camp (01.35.644°N; 113.47.377°E; datum WGS84), Putai, upper
Baleh, Kapit district, Sarawak, East Malaysia (Borneo), altitude 117 m ASL, coll. Pui Yong Min and Mohamad
Paisal bin Wahab, 19 May 2015. Adult female. The type locality is shown in Fig. 3.
Paratype. UNIMAS P1166, same locality as holotype. Collected 20 May 2015. Adult female.
Diagnosis. A large-sized (SVL 83.8 mm and 85.9 mm in the two specimens known, both adult females); upper
head shields, dorsal and lateral scales smooth; parietal scales in two pairs; presuboculars three; supraciliaries eight,
first largest; supraoculars four, fourth contacting orbit; supralabials seven (supralabial five contacting orbit);
infralabials four, second longest; postmental undivided; longitudinal scale rows 58; ventrals 53; transverse scale
rows at midbody 34; subcaudals 98; preanals enlarged, single; and subdigital lamellae on Toe IV 19.
Etymology. The species name, sebi derives from the acronym ‘SEB’, for Sarawak Energy Berhad, with a
genitive ending. We are pleased to name the new species after this organisation in acknowledgement of support
received for our long-term research of the herpetofauna of the Upper Baleh region. Suggested common name:
Baleh Water Skink.
Description of holotype. Adult female. Large size, SVL 83.8 mm, TL 121.4 mm; snout acute (IN:IO ratio
0.37), projecting beyond lower jaws; nostril laterally oriented; oval, situated closer to snout-tip than to orbit; eye–
nostril distance 4.83 mm, eye–snout distance 6.58 mm (E-N:E-S ratio 0.73); head long, much longer than wide, HL
13.60 mm, HW 12.48 mm (HL:HW ratio 1.09); head shape slightly flattened, HD 8.91 mm (HL:HD ratio 1.53);
upper head shields smooth; supranasals absent; frontonasal trapezoidal, as long as wide; prefrontals trapezoidal, in
narrow contact medially; frontal elongated, trapezoidal, wider anteriorly; frontoparietals joined; interparietal
single, with small transparent spot; parietal scales in two pairs; large posterior parietals, in contact posteriorly;
small anterior parietal, in contact with frontoparietal and supraocular IV; nuchal scales in two pairs; nostrils on
nasal located closer to anterior loreal; postnasal absent; supranasals absent; loreals two, anterior lower than
posterior; preoculars two; presuboculars three, anterior large, second in contact with third and fourth supralabials,
and third in contact with fourth and fifth supralabials; supraciliaries eight, first largest; supraoculars four, fourth
contacting orbit; postsupraocular present; two postocular; pretemporal single; six postsuboculars, lower one
contact with fifth and sixth supralabials; six primary temporals, lower one in contact with sixth and seventh
supralabials; five secondary temporals; seven supralabials (supralabial five contacting orbit); four infralabials,
second longest; mental smaller than rostral; rostral broad, projecting onto snout; posterior border of rostral straight;
postmental undivided; genials in three pairs, first and second pair in broad contact, and third pair separated by three
scales; auricular opening scaleless, ovoid and smaller than orbit of eye, its location indicated by a shallow
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depression; eyes relatively small; pupil discernable in preserved specimen; moveable eyelids; upper palpebrals 14;
lower palpebrals 14; tongue short; undivided anteriorly, tip obtuse, not pointed; teeth relatively small and
somewhat pointed.
Body slender, BW 12.52 mm (BW:SVL ratio 0.15); head distinct from neck and body; 58 longitudinal scale
rows from parietal to above level of anterior margin of hind limb; dorsal and lateral scales smooth; ventrals 53,
counted from first postgular to last scale before preanals; transverse scale rows at midbody 34; subcaudals 98;
preanals enlarged, single; tail laterally compressed, relatively long, longer than snout-vent length (TL:SVL ratio
1.45); tail tip acute; tail base wider than rest of tail; tail gradually tapering to a point; median row of subcaudals
enlarged. Visceral fat bodies absent in abdomen.
Limbs short and pentadactyl, digits short and clawed; lamellae smooth, enlarged; adpressed limbs touching;
lamellae under Finger I-8; II-12; III-11; IV-9; V-5 and lamellae under Toe I-7; II-11; III-15; IV-19; V-12.
FIGURE 1. Female holotype of Tropidophorus sebi sp. nov. (UNIMAS P1167) in life. (SVL = 83.8 mm).
Colouration in life. Dorsum chocolate brown, with dark greyish-brown transverse bars on trunk and tail; bars
on nape fused to form collar-like pattern; blackish-grey postocular band, starting from posterior corner of orbit and
broadening to axillary region, and narrowing thereafter along the flanks of torsi; forehead unpatterned chocolate
brown; temporal region grey with cream-coloured flecks; axilla and lower flanks with scattered yellow flecks;
labials brown with greyish-black bars; dorsal surface of tail greyish-brown with darker areas forming incomplete
bands; dorsal surface of fore- and hindlimbs chocolate brown with darker variegation; digits brown with darker
bands; gular region pale grey, the chin with dark grey variegation, forming about 10 lines that extend to the pectoral
region; upper two-thirds of pectoral region pale yellow with indistinct grey stripes; lower third orange, each scale
edged with yellow; cloacal region deep orange; lowest edge of anal scale dark grey; tail base grey with large,
orange-coloured scales; rest of tail dark grey; lower parts of forearm unpatterned pale grey; hindlimbs unpatterned
pale orange; manus and pes dark grey; pupil rounded, black, with a narrow, orange ring; iris grey with an orange
cast; oral cavity pale cream-coloured; tongue dark grey, not pale apically.
Measurements (in millimeters; holotype with paratype, in parentheses). SVL 83.8 (85.9); HL 13.6 (14.7);
HW 12.5 (13.6); HD 8.9 (9.8); BW 12.5 (13.5); TBL 11.3 (11.9); ED 4.1 (4.3); IN 3.1 (3.2); E–S 6.9 (6.9); E–N 4.8
(5.3); N–S 1.7 (1.7); A–G 36.7 (41.5) and TL 121.4 (87.2).
Squamation (holotype with paratype in parentheses). Transverse scale rows at midbody 34 (33);
longitudinal scale rows 58 (57); ventral scale rows 53 (53); supralabials 7 (7); infralabials 4 (4); subcaudals 98 (22
+; regenerated tail) and lamellae under toe IV 19 (19).
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FIGURE 2. Female holotype of Tropidophorus sebi sp. nov. (UNIMAS P1167), showing head in a. lateral; b. ventral; and c.
dorsal views (scale markers = 10 mm); and d. pes and e. manus. (scale markers = 1 mm).
Ecological Notes and Distribution. The type series was collected from narrow crevices of rocky banks of
small streams at the headwaters of Sungei Baleh. Currently, the new species is only known from its type locality,
Putai, Upper Baleh, central Sarawak (Fig. 4).
Phylogenetic relationships. The ML and BI topologies were identical with respect to relationships within
Tropidophorus. Phylogenetic analysis of mitochondrial l2s and 16s provide strong support for Tropidophorus
beccarii as the closest relative of the new species (see Fig. 5). This sister relationship suggests a single origin of
smooth dorsal scales among Tropidophorus on Borneo (see Comparison), assuming an ancestral state of keeled
dorsal scales for the genus. The phylogenetic results presented here are generally concordant with results of Honda
et al. (2006), though we recover much weaker support for the paraphyly of Tropidophorus with respect to the
Sphenomorphus group (ML bootstrap 36 vs. 73). The ML and BI topologies were identical with respect to
relationships within Tropidophorus.
Raw pairwise sequence divergence between T. beccarii and the new species is 6.8–7.0% for 12s and 5.4–5.6%
for 16s. Genetic divergence between the two type specimens of the new species is low, with raw distances of 0.2%
and 0.3% for 12s and 16s, respectively. However, this measure of genetic diversity within the new species may be
an underestimate as the two type specimens were collected from the same locality, and increased genetic
divergence would likely exist across a larger spatial area.
Comparison. We initially compare the new species with Bornean congeners here. Tropidophorus sebi sp. nov.
differs from T. brookei (distribution: widespread on Borneo), T. iniquus (distribution: upper Sungei Kajan, central
Kalimantan, Indonesia), T. micropus (distribution: Long Bloe in upper Sungei Mahakkam, Kalimantan, Indonesia
and Putai, Sarawak, Malaysia) and T. perplexus (distribution: Sungei Tinjar, Sarawak, Malaysia) in having smooth
dorsal scales (versus keeled in these four species). The new species is further distinguished from another two
smooth dorsal scales congeners, T. beccarii (widespread on Borneo) and T. mocquardii (distribution: northern
Sabah, Malaysia; treated as a synonym of T. beccarii by some authors but considered valid here; see below) from
Borneo, in showing a higher count of midbody scale rows (33–34) versus lower in T. beccarii (28–30), in having
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four supraoculars versus five supraoculars, and in lacking a thick, dark, dorsolateral stripe along flanks, that are
with light transverse bands in T. mocquardii.
FIGURE 3. Map showing type locality of Tropidophorus sebi sp. nov., at Putai, Upper Sungei Baleh, Kapit District, Sarawak,
Malaysia (Borneo). Inset: Map of the Indo-Pacific, showing the location of Borneo.
The presence of smooth dorsal scales distinguishes the new species from the following extra-Bornean
congeners that possess keeled dorsal scales: T. assamensis Annandale, 1912 (distribution: north-eastern
Bangladesh and north-eastern India, including Mizoram and Assam states); T. baconi Hikida, Riyanto & Ota, 2003
(distribution: Patunuang Natural Reserve, southern Sulawesi, Indonesia); T. baviensis Bourret, 1939 (distribution:
Mount Ba Vi, Ha Tay Province, northern Vietnam and possibly, adjacent Thailand); T. berdmorei (Blyth, 1853)
(distribution: Yunnan in southern China, Myanmar, Thailand and Vietnam); T. cocincinensis Duméril & Bibron,
1839 (distribution: Thailand, southern Vietnam and Cambodia); T. davaoensis Bacon, 1980 (distribution: south-
central Mindanao, Philippines); T. gr ayi Günther, 1861 (distribution: Panay, Luzon, Polillo, Leyte, Negros,
Mastabe, Cebu, Philippines); T. guangxiensis Wen, 1992 (distribution: Guangxi and Hunan, China); T. hainanus
Smith, 1923 (distribution: Hainan, Guangxi, Jiangxi, Hunan, Guangdong and Guizhou in China, and Bac Kan, Cao
Bang, Dak Lak, Ha tay, Hai Duong, Hoa Binh, Kon Tum, Lai chau, Ninh binh, Phu Tho, Quang Ninh, Vinh Phuc
and Nam Dinh in Vietnam); T. hangnam Chuaynkern, Nabhitabhata, Inthara, Kamsook & Somsri, 2005
(distribution: Chaiyaphum Province, north-easten Thailand), T. latiscutatus Hikida, Orlov, Nabhitabhata & Ota,
2002 (distribution: Phu Wua Wildlife Sanctuary, Nong Kai Province, eastern Thailand); T. matsu i i Hikida, Orlov,
Nabhitabhata & Ota, 2002 (distribution: Phu Pa Namtip, Roi Et Province, eastern Thailand); T. microlepis Günther,
1861 (distribution: southern Thailand, Vietnam, Laos and Cambodia); T. misaminius Stejneger, 1908 (distribution:
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FIGURE 4. Habitat of Tropidophorus sebi sp. nov. at Putai, Upper Sungei Baleh, Sarawak, Malaysia (Borneo).
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FIGURE 5. Concatenated ML phylogram of the mitochondrial 12s and 16s genes for Tropidophorus and close relatives with
ML bootstraps (left number) and BI posterior probabilities (right number) shown at the nodes. Nodes with ML bootstrap ≥ 70
and BI posterior probability ≥ 0.95 are denoted with a black dot. The new species is indicated in bold.
Basilan, Camiguin and Mindanao, Philippines); T. murphyi Hikida, Orlov, Nabhitabhata & Ota, 2002 (distribution:
Cao Bang Province, northern Vietnam); T. noggei Ziegler, Thanh & Thanh, 2005 (distribution: Phong Nha - Ke
Bang National Park, Quang Binh Province, Vietnam); T. pa rtel loi Stejneger, 1910 (distribution: Mataling River
Falls, Cotabato Province, Mindanao, Philippines); T. robinsoni Smith, 1919 (distribution: Myanmar and Thailand);
T. sinicus Boettger, 1886 (distribution: Guangxi, Guangdong, Hong Kong in China and Vietnam) and T. t h ai Smith,
1919 (distribution: Myanmar and northern and north-eastern Thailand).
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Two extra-Bornean congeners possess smooth scales: T. boehmei Nguyen, Nguyen, Schmitz, Orlov & Ziegler,
2010 (distribution: Hoang Lien Mountain, Lao Cai Province, northern Vietnam) and T. laotus Smith 1923
(distribution: Pak Lai district in Laos and Thailand). Tropidophorus boehmei differs from the new species in
showing tiny, reduced prefrontals vs. trapezoidal prefrontals that are in contact, supralabials six vs. seven,
longitudinal scale rows 60–69 vs. 58, and transverse scale rows at midbody 30–32 vs. 34 in the new Bornean
species. Tropidophorus laotus differs from the new species in showing a frontonasal that is divided vs. fused,
supralabials six vs. seven, and anterior loreal divided vs. undivided in the new species.
Finally, to aid field identification, we present below a dichotomous identification key to the Bornean species of
the genus Tropidophorus.
Key to Bornean species of Tropidophorus
1. Dorsal scales smooth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
- Dorsal scales keeled . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
2. Midbody scale rows 28–30; thick dark dorsolateral stripe along flanks with light transverse bands . . . . . . . . . . . . . . . . beccarii
- Midbody scale rows 33–34; no dark dorsolateral stripe along flanks. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ..6
3. Four supraoculars . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . micropus
- Five supraoculars . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
4. Dorsal scales bicarinate, grooved . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . iniquus
- Dorsal scales unicarinate; not grooved . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
5. Supraciliaries 14; midbody scale rows 32 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. brookei
- Supraciliaries 5–6; midbody scale rows 30. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ... perplexus
6. Four supraoculars. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .sebi sp. nov.
- Five supraoculars. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .mocquardii
Acknowledgements
We thank the Institute of Biodiversity and Environmental Conservation, Universiti Malaysia Sarawak, for
supporting our researches, which received financial support from Sarawak Energy Berhad, under the Hydropower
Environmental Sustainability Program (grant no. GL(I01)SEB/2014/02[03] to ID). Karen Lee served as liaison
between these two institutions. AMB was supported by Grant EF 1241885 (subaward 13-0632) from the National
Science Foundation, USA. Permission and advice to conduct research in Sarawak was provided by the Sarawak
Forest Department and the Sarawak Forestry Corporation (permit no. NCCD.907.4.4(Jld.11–68 and 508/2014), and
we are grateful to the staff of these organisations, especially Haji Wan Shardini bin Wan Salleh, Oswald Braken
Tisen, Engkamat anak Lading, Mohamad bin Kohdi and Nur Afiza binti Umar. We thank the curatorial staff of the
following institutions for permission and facilities at their respective institutions: AMNH (D. R. Frost and L. Ford),
BM (M. Wong and Hj. Bakariah), BMNH (E. N. Arnold, D. Gower and C. J. McCarthy), FMNH (H. K. Voris, R. F.
Inger and A. Resetar), MCZ (J. E. Cadle, J. Hanken and J. P. Rosado), MZB (Ir. Boeadi and P. Mumpuni), SM (C.
Leh), USNM (W. R. Heyer, R. I. Crombie and G. R. Zug), ZRC (P. K. L. Ng and K. K. P. Lim) and ZSI (J. R. B.
Alfred and B. H. C. Murthy). We are also grateful to R. de Ruiter, RMNH for the image of the holotype of
Tropidophorus micropus (RMNH 4452). Our colleague, Jiang Jian-Ping from the Chinese Academy of Sciences
supplied several critical references. At Putai, we thank Mr. Lee and his staff of the First Count Camp. Finally, we
are grateful to Andrew Alek Tuen, Rahah binti Mohamad Yakup, Mohd. Hasri Al-Hafiz, Mohamad Paisal bin
Wahab, Pang Sing Tyan, Attiqqah Fadziliah and Melinda Cheok for assistance in the field.
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APPENDIX I. Comparative material examined.
Tropidophorus assamensis Annandale, 1912: ZSI 17029 (holotype), ‘Haraigaj range, 550 ft., Sylhet hills, Assam’ (at present
Sylhet District, north-eastern Bangladesh).
Tropidophorus beccarii Peters, 1871: AMNH 111914. Tubau Camp on Sungei Pesu, Miri Division, Sarawak, Malaysia; ZRC
2.1848. Sungei Baram, Sarawak, Malaysia; SBC R.30. Bukit Meraja, Bau, Sarawak, Malaysia; BMNH 72.2.19.15–16.
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Gunung Matang, Sarawak, Malaysia; SM c.c.6.3.1 [two specimens]. Gunung Penrissen, Sarawak, Malaysia; FMNH
221481. Gunung Sungei Segaham, Kapit, Sarawak, Malaysia; USNM 197818. Labang Camp at Sungei Pesu, Sarawak,
Malaysia; FMNH 248113; 248525. Tawau Hills Park, Tawau District, Sabah, Malaysia; FMNH 248526. Maliau Basin,
Kinabatangan/Pensiangan Districts, Sabah, Malaysia; BM 1992.90. Sungei Ingei, Belait District, Brunei; UBD 84, 235,
295, 313, 538. Batu Apoi, Temburong District; Brunei; ZRC 2.4572. Sintang, along road at km 463 to Pontianak,
Kalimantan Barat, Indonesia; ZRC 2.5801. Sungei Moleng at Baloi, Kalimantan Tengah, Indonesia; ZRC 2.5802.
Mahakam basin, Kalimantan Tengah, Indonesia; MZB 2941–43; 18081; 18185; 18217. Maruwai, Kalimantan Tengah
Propinsi, Indonesia.
Tropidophorus berdmorei (Blyth, 1853): ZRC 2.4618. Nam Tok Natee Rachan, Mae Hong Son Province, Thailand; ZSI 2270–
72 (syntypes of Aspris Berdmorei Blyth, 1853), ‘Mergui’ (Myeik or Beik), Myanmar.
Tropidophorus brookei (Gray, 1845): AMNH 111915–18. Nanga Tekalit on Sungei Mengiong, Kapit Division, Sarawak,
Malaysia; MZB 2391. Sungei Ulu Matai, Taman Nasional Bentuang Karimun, Kalimantan Barat Propinsi, Indonesia;
MZB 3740–42. Sungei Menggulang, Taman Nasional Bentuang Karimun, Kalimantan Barat Propinsi, Indonesia; ZRC
2.1850. Sungei Baram, Sarawak, Malaysia; ZRC 2.3395, 2.4663. Bako National Park, Sarawak, Malaysia; ZRC 2.3447.
Sarikei, Sarawak, Malaysia; ZRC 2.3522. Sungai Stum Muda near Lundu, Sarawak, Malaysia; ZRC 2.5739. waterfall at
base of Gunung Pueh, Sarawak, Malaysia; ZRC 2.1852–58. Lawas, Sarawak, Malaysia; ZRC 2.4943. Nanga Ulai near
Betong, Sarawak, Malaysia; ZRC 2.5749–51; 2.5845–46. Kapit, Sungai Belajau, Kapit, Sarawak, Malaysia; ZRC 2.4937,
ZRC 2.4943. Ranchan Pool, Serian, Sarawak, Malaysia; ZRC 2.4617. Rajang basin near Kapit, Sarawak, Malaysia; ZRC
2.5337. Long Asap, Sarawak, Malaysia; ZRC 2.5338. Lambir Hills National Park, Miri, Sarawak; ZRC 2.3710. Danum
Valley, Sungai Taliwas, Danum Valley, Sabah, Malaysia; ZRC 2.1850. Kabayo near Gunung Kinabalu, Sabah, Malaysia;
ZRC 2.5805. Sungai Sepan, Sungei Sepan, Brunei Darussalam; ZRC 2.5266–67. Kuala Belalong Field Studies Centre,
Temburong District, Brunei Darussalam; ZRC 2.5803. Sanggau, Kalimantan Barat, Indonesia.
Tropidophorus micropus van Lidth de Jeude, 1905: UNIMAS P1168. First Count Logging Camp, Putai, upper Sungei Baleh,
Kapit District, Sarawak, Malaysia; RMNH 4452 (holotype of Tropidophorus micropus; digital image). “Long Bloe Upper
Mahakkam River” (= Long Blu, also spelt Long Bloéoe, upper reaches of Sungei Mahakam, Kalimantan Timur Propinsi,
Indonesia).
Tropidophorus perplexus Barbour, 1921: MCZ 14632 (holotype) “..near the Fort at Long Loba, Tinjar River, Sarawak” (=
vicinity of Fort Hose, Marudi, Miri Division, Malaysia); FMNH 138551–56. Nanga Tekalit Camp on Sungei Mengiong,
Kapit, Sarawak, Malaysia; FMNH 235156. Mendolong, Sipitang District, Sabah, Malaysia.
Tropidophorus sinicus Boettger, 1886: ZRC 2.4814. Lam Tseun, Pak Ngau Shek, New Territories, Hong Kong, China; ZRC
2.3478. Tai Po Kau Forest Reserve, New Territories, Hong Kong, China.
