The first riparian skink (Genus: Sphenomorphus Strauch, 1887) from Peninsular Malaysia and its relationship to other Indochinese and Sundaic species

Abstract

Recently discovered populations of skinks of the genus Sphenomorphus from central Peninsular Malaysia represent a new species, S. sungaicolus sp. nov., and the first riparian skink known from Peninsular Malaysia. Morphological analyses of an earlier specimen reported as S. tersus from the Forest Research Institute of Malaysia (FRIM), Selangor indicate that it too is the new riparian species S. sungaicolus sp. nov. Additionally, two specimens from the Tembat Forest Reserve, Hulu Terengganu, Kelantan and another from Ulu Gombak, Selangor have been diagnosed as new the species. The latter specimen remained unidentified in the Bernice Pauahi Bishop Museum, Honolulu, Hawaii since its collection in June 1962. Morphological and molecular analyses demonstrate that S. sungaicolus sp. nov. forms a clade with the Indochinese species S. maculatus, S. indicus, and S. tersus and is the sister species of the latter. Sphenomorphus sungaicolus sp. nov. can be differentiated from all other members of this clade by having a smaller SVL (66.5–89.6 mm); 39–44 midbody scale rows; 72–81 paravertebral scales; 74–86 ventral scales; a primitive plantar scale arrangement; and 20–22 scale rows around the tail at the position of the 10 th subcaudal.

Full text

Accepted by C. Siler: 16 Aug. 2016; published: 3 Oct. 2016
ZOOTAXA
ISSN 1175-5326 (print edition)
ISSN
1175-5334
(online edition)
Copyright © 2016 Magnolia Press
Zootaxa 4173 (1): 029
–
044
http://www.mapress.com/j/zt/
Article
29
http://doi.org/10.11646/zootaxa.4173.1.3
http://zoobank.org/urn:lsid:zoobank.org:pub:E10676C4-F087-4559-B98F-43A51DFBBA4E
The first riparian skink (Genus: Sphenomorphus Strauch, 1887) from Peninsular
Malaysia and its relationship to other Indochinese and Sundaic species
ALEXANDRA SUMARLI
1
, L. LEE GRISMER
1
, PERRY L. WOOD, JR.
2
, AMIRRUDIN B. AHMAD
3
, SYED
RIZAL
3
, LUKMAN H. ISMAIL
3
, NUR AMALINA MOHD IZAM
4
, NORHAYATI AHMAD
4
& CHARLES W.
LINKEM
5
1
Department of Biology, La Sierra University, 4500 Riverwalk Parkway, Riverside, California 92515 USA. E-mail:
sumarli.alex@gmail.com, lgrismer@lasierra.edu
2
Department of Biology, Brigham Young University, 150 East Bulldog Boulevard, Provo, Utah, 84602 USA. E-mail: pwood@byu.edu
3
Pusat Pengajian Sains Marin dan Sekitaran, Universiti Malaysia Terengganu, 21030 Kuala Terengganu, Malaysia.
E-mail: ameyahmad@yahoo.com, syedrizal@umt.edu.my
4
School of Environment and Natural Resource Sciences, Faculty of Science and Technology, University Kebangsaan Malaysia, 43600
Bangi, Selangor, Malaysia. E-mail: amalinanurizam@gmail.com
5
Department of Biology, University of Washington, Box 351800, Seattle, Washington, 98195 USA. E-mail: cwlinkem@gmail.com
Abstract
Recently discovered populations of skinks of the genus Sphenomorphus from central Peninsular Malaysia represent a new
species, S. sungaicolus sp. nov., and the first riparian skink known from Peninsular Malaysia. Morphological analyses of
an earlier specimen reported as S. tersus from the Forest Research Institute of Malaysia (FRIM), Selangor indicate that it
too is the new riparian species S. sungaicolus sp. nov. Additionally, two specimens from the Tembat Forest Reserve, Hulu
Terengganu, Kelantan and another from Ulu Gombak, Selangor have been diagnosed as new the species. The latter spec-
imen remained unidentified in the Bernice Pauahi Bishop Museum, Honolulu, Hawaii since its collection in June 1962.
Morphological and molecular analyses demonstrate that S. sungaicolus sp. nov. forms a clade with the Indochinese spe-
cies S. maculatus, S. indicus, and S. tersus and is the sister species of the latter. Sphenomorphus sungaicolus sp. nov. can
be differentiated from all other members of this clade by having a smaller SVL (66.5–89.6 mm); 39–44 midbody scale
rows; 72–81 paravertebral scales; 74–86 ventral scales; a primitive plantar scale arrangement; and 20–22 scale rows
around the tail at the position of the 10
th
subcaudal.
Key words: Integrative taxonomy, New species, Scincidae, Southeast Asia, Sundaland
Introduction
The genus Sphenomorphus Fitzinger, 1843, is a diverse, polyphyletic genus of scincid lizards that is in the
beginning stages of a major taxonomic revision (see Linkem et al. 2011; Linkem 2013; Linkem, personal
observation). Nineteen of at least 109 species of Sphenomorphus recognized currently (Uetz 2016) occur in
Peninsular Malaysia where they exhibit diverse lifestyles in a wide array of habitats ranging from upland cloud
forest to small, arid, virtually barren islands to lowland and hill dipterocarp forests (Grismer 2011; Grismer & Quah
2015). Some species are diurnal and are often observed foraging along the forest floor or basking on rocks and
trees in sun spots several meters above the ground, whereas others are more secretive, leaving much of their natural
history unknown with some even hypothesized as being nocturnal (Grismer 2011). Despite the ecological diversity
of Sphenomorphus and that of other skink lineages in Peninsular Malaysia, no exclusively riparian species has ever
been reported from this region.
Sphenomorphus tersus (Smith) was first reported from Peninsular Malaysia on the basis of a single specimen
collected along a stream at the Forest Research Institute Malaysia (FRIM), Selangor (Leong et al. 2002). However,
Grismer (2011) questioned the identification of that specimen noting several morphological differences between it
and Taylor’s (1963) account of S. tersus from Thailand and from a specimen of S. tersus collected from Perlis in

SUMARLI E T AL.
30
·
Zootaxa 4173 (1) © 2016 Magnolia Press
northern Peninsular Malaysia along the Thai-Malaysian border. Additionally, we have gained access to a heretofore
unexamined specimen collected by John R. Hendrickson on 13 June 1962 from alongside a small river at Ulu
Gombak, Selangor, 27 km northwest of FRIM. Based on morphological and color pattern characteristics, we
determined this specimen from Ulu Gombak to be conspecific with the FRIM specimen and neither are conspecific
with S. tersus. We recently collected four additional specimens from Peninsular Malaysia––two adults from the
east side of the Banjaran (=Mountain Range) Titiwangsa in the state of Terengganu (LSUHC 11722 from Hutan
Lipur Sekayu and LSUHC 11780 from Hutan Lipur Chemerong 104 km south of Sekayu [Fig. 1]) and one juvenile
(UKMHC 0707) and one adult (UKMHC 0629) from the Tembat Forest Reserve, Hulu Terengganu, Terengganu
approximately 130 km to the west of Hutan Lipur Sekayu. All specimens were collected alongside streams and
closely match the color pattern and morphological characteristics of the FRIM and Gombak specimens from the
western side of the Banjaran Titiwangsa. Given this overlap in morphological characteristics, we consider all six
specimens conspecific. All are similar in general morphology and body stature to S. tersus, S. indicus (Gray), and
S. maculatus (Blyth), which also occur in Peninsular Malaysia (see Grismer 2011 for remarks concerning S.
maculatus). However, the six specimens in question depart from the former species on the basis of several other
morphological and color pattern characters. Previous phylogenetic analyses indicate that S. tersus, S. indicus, and
S. maculatus form a well-supported monophyletic group (Honda et al. 2006; Linkem et al. 2011; Linkem 2013)
into which we hypothesize, based on morphology, this undescribed, riparian species will belong. To test this
hypothesis and the hypothesis of Grismer (2011) that the undescribed species from FRIM is not conspecific with S.
tersus as purported by Leong et al. (2002), we constructed Maximum Likelihood and time-calibrated Bayesian
inference phylogenies based on the mitochondrial 12S and 16S ribosomal RNA (rRNA) genes using the four
recently collected specimens.
Material and methods
Molecular methods. Genomic DNA was isolated from liver or muscle tissues stored in 95% ethanol using the
animal tissue protocol in the Qiagen DNeasyTM tissue kit (Valencia, CA, USA). The mitochondrial genes 12S and
16S were amplified using a double-stranded Polymerase Chain Reaction (PCR) under the following conditions: 1.0
m l (10–33 µg) genomic DNA, 1.0 µl (10 µm M) forward primer and 1.0 µl (10 µM) reverse primer L1091 and
H1478 (Kocher et al. 1989) for 12S rRNA, and, L2606, H3056 (Hedges et al. 1993) for 16S rRNA, 1.0 l
dinucleotide pairs (1.5 µM), 2.0 m l 5x buffer (1.5 µM), 1.0 MgCl 10x buffer (1.5 µM), 0.18 m l Taq polymerase
(5u/µl), and 7.5 m l H2O, primers are from Macey & Verma (1997). All PCR reactions were executed on an
Eppendorf Mastercycler gradient theromocycler under the following conditions: initial denaturation at 95ºC for 2
min, followed by a second denaturation at 95ºC for 35 s, annealing at 55ºC for 35 s, followed by a cycle extension
at 72ºC for 35 s, for 33 cycles. PCR products were visualized on a 1% agarose gel electrophoresis. Successful
targeted PCR products were vacuum purified using MANU 30 PCR plates Millipore plates and purified products
were resuspended in DNA grade water. Purified PCR products were amplified using the PCR primers with the ABI
Big-Dye Terminator v3.1 Cycle Sequencing Kit in an ABI GeneAmp PCR 9700 thermal cycler. Cycle sequencing
reactions were purified with Sephadex G-50 Fine (GE Healthcare) and sequenced on an ABI 3730xl DNA
Analyzer at the BYU DNA Sequencing Center (DNASC). All new sequences produced from this study are
deposited in GenBank under the following accession numbers (Table 1). All sequences were aligned in Geneious
v6.1.8 (Drummond et al. 2012) using 8 iterations and default settings.
Phylogenetic, time calibration, and sequence divergence analyses. The phylogenetic analyses applied two
model-based methods, Maximum Likelihood (ML) and Bayesian Inference (BI). A time calibrated BI analysis was
used to date the cladogenic events within the tree. The Bayesian Information Criterion (BIC) implemented in IQ-
TREE (Nguyen et al. 2015) was used to calculate the best-fit model of evolution for each gene for the ML analysis
(Table 2). Maximum Likelihood analyses using IQ-TREE employed 1000 bootstrap pseudoreplicates via the
ultrafast bootstrap approximation algorithm. Nodes having ML bootstrap support values of 90 and greater and BI
posterior probabilities of 0.95 and greater were considered strongly supported (Huelsenbeck et al. 2001; Nguyen et
al. 2015; Wilcox et al. 2002). Scincella lateralis (Say), Scincella cherriei (Cope), Scincella rupicola (Smith),
Isopachys anguinoides (Boulenger), Lipinia vittigera (Boulenger), Sphenomorphus stellatus (Boulenger),
Sphenomorphus praesignus (Boulenger), Tropidophorus sinicus (Boettger), Tropidphorus brookei (Gray), and
Lamprolepis smaragdina (Lesson) were chosen as outgroups based on their relationships to S. maculatus and S.
indicus as demonstrated by Linkem (2013).

Zootaxa 4173 (1) © 2016 Magnolia Press
·
31
RIPARIAN SKINK IN PENINSULAR MALAYSIA
TABLE 1. Specimens used in this study with locality data and GenBank Accession numbers. FMNH refers to Field Museum of Natural History Collection;
IEBR refers to Institute of Ecology and Biological Resources, Vietnam; KU refers to Kansas University Museum of Natural History collection; KUZ and KUZR
refer to Kyoto University Museum; and SMF refers to Senckenberg Museum of Frankfurt am Main
Voucher no.
Species Name
Locality
GenBank Accession #
12S
16S
KUZ 35371
Isopachys anguinoides
Kaeng Krachan, Thailand
AB028803
AB028815
KUZ 35004
Lamprolepis smaragdina
Mariana Islands, Saipan
AB028774
AB028831
KUZ 32857
Lipinia vittigera
Chantaburi, Thailand
AB028804
AB028816
KUZ 45017
Scincella lateralis
Txler, Texas, USA
AB057387
AB057402
KUZ 40458
Scincella rupicola
Phu Wua, Thailand
AB057388
AB057403
SMF 79812
Scincella cherriei
Bartola, Nicaragua
AB057377
AB057392
FMNH 239867
Sphenomorphus cyanolaemus
Sabah, Malaysia
JF497954
JF498084
KUZ 37239
Sphenomorphus indicus
Yunlin, Taiwan
AB028808
AB028820
KUZ 37239
Sphenomorphus maculatus
Kaeng Krachan, Thailand
AB028809
AB028821
FMNH 243828
Sphenomorphus multisquamatus
Sabah, Malaysia
JF497961
JF498091
KUZ 21251
Sphenomorphus praesignis
Bukit Larut, Malaysia
AB028810
AB028822
FMNH 239881
Sphenomorphus sabanus
Sabah, Malaysia
JF497962
JF498092
FMNH 267739
Sphenomorphus stellatus
Koh Kong Province, Cambodia
KX398015
KX398012
IEBR FN39484
Sphenomorphus stellatus
Vietnam
N/A
HM 773221
LSUHC 11722
Sphenomorphus sungaicolus sp. nov.
Hutan Lipur Sekayu, Terengganu, Malaysia
KX398019
KX398013
LSUHC 11780
Sphenomorphus sungaicolus sp. nov.
Hutan Lipur Chemerong, Terengganu, Malaysia
KX398020
KX398014
LSUHC 9041
Sphenomorphus tersus
Perlis, Malaysia
KX398020
KX398015
KU 309900
Sphenomorphus variegatus
Philippines
JF497966
JF498096
KUZR 19008
Tropidophorus brookei
Sarawak, Malaysia
AB222933
AB222949
KU 311515
Tropidophorus sinicus
Dashang Nature Reserve, China
N/A
KX398016
KU 311520
Tropidophorus sinicus
Dashang Nature Reserve, China
KU311520
KX398017

SUMARLI E T AL.
32
·
Zootaxa 4173 (1) © 2016 Magnolia Press
TABLE 2. Models used for each codon position.
The time calibrated BI analysis was implemented in BEAST v1.8.0 (Drummond et al. 2012). The input file was
constructed in BEAUti v1.8.0 and was run with unlinked substitution models and unlinked clock models.
Sphenomorphus sungaicolus sp. nov., S. tersus, S. indicus, and S. maculatus were chosen as the ingroup and the
same outgroups from the ML analysis based on Linkem (2013) were used. A lognormal relaxed clock and speciation
Yule Process were selected using a 0.65% mutation rate as reported in Honda et al. (2006). MCMC chain lengths
were set to run for 1 million generations per taxon and sampled at every 1000 trees. Log files were checked in Tracer
v1.6 (Rambaut et al. 2014) to ensure ESS (Effective Sample Size) values were above 200. We discarded the first
10% of the trees as burnin after visualization in TreeAnnotater v 1.8.0 (Rambaut & Drummond 2013).
A separate BI analysis was carried out in MrBayes 3.2.3. on XSEDE (Ronquist et al. 2012) using CIPRES
(Cyberinfrastructure for Phylogenetic Research; Miller et al. 2010) employing default priors. Two simultaneous
Markov Chain Monte Carlo (MCMC) runs were performed with four chains per run (three hot and one cold) using
default priors. The analysis was run for 10 million generations, sampled every 1000 generations, and halted after
the average standard deviation split frequency was below 0.01 and convergence was verified in Tracer v1.6
(Rambaut et al. 2014). The first 25% of the trees were discarded as burnin. Nodes having BI posterior probabilities
of 0.95 and greater were considered strongly supported (Huelsenbeck et al. 2001; Wilcox et al. 2002). The same
ingroup and outgroup taxa used in the ML and Time Calibrated BEAST analysis were included. Uncorrected
pairwise sequence divergences were calculated in MEGA v5.2.2 (Tamura et al. 2011).
Morphological and color pattern analyses. Color pattern characters were taken from digital images of living
specimens cataloged in the La Sierra University Digital Photo Collection (LSUDPC) and from living specimens in
the field and include the presence or absence of white spots on dorsum, lateral stripes on body, and the color of
palmer and plantar surfaces relative to the venter.
The following measurements on the type series were taken with Mitutoyo dial calipers to the nearest 0.1 mm
under a Nikon SMZ 1500 dissecting microscope: snout–vent length (SVL); tail length (TaL); axilla–groin length
(AG), distance form posterior junction of fore-limb and body wall to anterior junction of hind limb and body wall
(with limbs held at right angles to the body); head length (HL), tip of snout to posterior margin of parietal or
interparietal, depending on the longest distance; head width (HW), at the widest portion of temporal region; head
height (HH), at the deepest portion of temporal region; snout length (SL), from anterior corner of eye to tip of
snout; snout–tympanum distance (STL), distance from snout to anterior border of tympanum (STL); snout-fore-
limb length (SFIL), from tip of snout to anterior junction of fore-limb and body wall, with the limb held at right
angles to the body; eye–nostril distance (END), distance from anterior margin of eyeball to posterior border of
nostril; eye length (EL), distance between anterior and posterior corners of eyelid; eye tympanum distance (ETL),
distance from anterior border of tympanum to posterior margin of eyeball; tympanum diameter (TYD), maximum
diameter of tympanum; fore-limb length (FIL), from anterior junction of fore-limb and body wall to the tip of
fourth finger; hind limb length (HIL), from anterior junction of hind limb and body wall to the tip of fourth finger.
Variation in foot scalation following Shea (2012) was analyzed and photographs of foot scalation were taken with a
Nikon SMZ800 with attached Nikon Digital Sight DS-Fi1C. Z-stack images were merged using the software
Helicon Focus 6 (Helicon Soft Ltd.). Standard scale counts follow Grismer (2011) and Lim (1998) and include the
number of supraoculars, suboculars, loreals, supralabials, infralabials, lamellae beneath the fourth digit beginning
with the first scale whose posterior margin extends into the body of the foot, enlargement of posterior supraciliary
scales, and degree of contact between adpressed limbs. Additional scale counts were supplemented from Taylor
(1963) and Grismer (2011).
Institutional abbreviations are BPBM (Bernice P. Bishop Museum, Honolulu, Hawaii), LSUHC (La Sierra
University Herpetological Collection, La Sierra University, Riverside, California, USA); LSUDPC (La Sierra
University Digital Photo Collection, La Sierra University, Riverside, California); USMHC (University Sains
Malaysia Herpetology Collection), ZRC (Zoological Reference Collection at the Raffles Museum of Biodiversity
ML BI
Gene Model selected Model selected
12s TIM2+I+Γ GTR+ Γ
16s TIM2+I+Γ GTR+ Γ

Zootaxa 4173 (1) © 2016 Magnolia Press
·
33
RIPARIAN SKINK IN PENINSULAR MALAYSIA
Research, National University of Singapore), and UKMHC (Universiti Kebangsaan Malaysia Herpetological
Collection). The letter “R” next to measurements denotes a regenerated tail, “/” indicates sex is not available, and
“sm” denotes that scales are smooth.
FIGURE 1. Localities of Sphenomorphus sungaicolus sp. nov. from Peninsular Malaysia. Star symbol denotes the type
locality.
Results
The phylogenetic analyses indicate that the new species forms a distinct lineage and is part of a strongly supported
group in the BI analysis (BI/ML = 1.00/79) that includes Sphenomorphus maculatus, S. tersus, and S. indicus
wherein the new species is strongly supported (1.00/98) as being the sister species of S. tersus (Fig. 2). Our time-
calibrated BEAST analysis suggests that the Sekayu and Chemerong populations diverged from one another
approximately 2.6 million years ago (0.4424,5.6266) and S. sungaicolus sp. nov. and S. tersus diverged from one
another approximately 8 million years ago (2.8652,13.7661).

SUMARLI E T AL.
34
·
Zootaxa 4173 (1) © 2016 Magnolia Press
FIGURE 2. BEAST topology inferring the phylogenetic relationships of Sphenomorphus sungaicolus sp. nov. from Peninsular
Malaysia with other members of its clade with Bayesian posterior probabilities and Maximum Likelihood bootstrap values (BI/
ML, respectively) at the nodes. 95% HPD (Highest Posterior Density) bars are displayed for the S. sungaicolus and S. tersus
populations.
TABLE 3. Diagnostic characters differentially separating Spehonomorphus sungaicolus sp. nov. from S. tersus, S.
indicus, and S. maculatus.
* Additional scale counts were supplemented from Taylor (1963) and Grismer (2011). For specimens examined see
Grismer (2011).
The sister species relationship between S. tersus and the new species is also supported by morphological and
color pattern characters in that both share brown dorsal surfaces and darker palmar and plantar surfaces relative to
the venter—character states not found in the other species in the clade (Table 3). Given the phylogenetic
S. indicus S. maculatus S. sungaicolus sp. nov. S. tersus
SVL in mm (Preserved) 49–79 58.6–62.5 66–89.6 90.5–96
Prefontals in contact (1) or seperated posteriorly (0) 0 0 0,1 0,1
Parietals in contact (1) or seperated posteriorly (0) 1 0,1 0,1 1
Superciliaries 8–11 10–12 8 7–12
Scale rows at position of tenth subcaudal row 20–21 17 16–22 16–19
Dark colored palmar and plantar surfaces no no yes yes
Dorsal scales extend onto ventral surface of pes no yes no yes
Lateral stripe absent yes no yes yes
Sample size (n) n=3n=14 n=6n=1*

Zootaxa 4173 (1) © 2016 Magnolia Press
·
35
RIPARIAN SKINK IN PENINSULAR MALAYSIA
relationships of the four specimens in the molecular analyses and the fact that all the specimens bear a unique
combination of scale and color pattern characteristics that unite them with each other but separate them from all
other members of their clade, they are all considered conspecific and described below as the new species:
Systematics
Sphenomorphus sungaicolus sp. nov.
Malaysian Riparian Skink
(Figs. 3–7. Tables 1, 3, 4)
Sphenomorphus tersus — Part: Leong et al. 2002:149; Grismer 2011:673.
Holotype. Adult male (LSUHC 11722) collected from Hutan Lipur Sekayu, Hulu Terengganu, Peninsular
Malaysia (Fig. 3) (4°59'N, 102°55'E) on 1 May 2014 by Syed A. Rizal.
Paratypes. Adult female (LSUHC 11780) collected from Hutan Lipur Chemerong, Terengganu, Peninsular
Malaysia (4°39'N, 103°00'E) on 4 April 2014 by Syed A. Rizal. Adult female (BPBM 43794) collected from Ulu
Gombak, Selangor, Peninsular Malaysia (3°18'N, 101°47'E) on 13 June 1962 by John R. Hendrickson. Adult
female (ZRC. 2.4915) collected from the Forest Research Institute Malaysia (FRIM), Kepong, Selangor,
Peninsular Malaysia (3°14'N, 101°38'E) on 27 February 2001 by Tzi Ming Leong.
Additional referred specimens. Adult female (UKMHC 0629) collected from the Tembat Forest Reserve,
Hulu Terengganu, Terengganu, Peninsular Malaysia (5°00152'N, 102°30987'E) on 4 March 2015 by Nur Amalina
Mohd Izam. Juvenile (UKMHC 0707) collected from the Tembat Forest Reserve, Hulu Terengganu, Terengganu,
Peninsular Malaysia (5°03279'N, 102°56030'E) on 30 October 2014 by Nur Amalina Mohd Izam
Diagnosis (Fig. 5). Adults reach at least 89.6 mm SVL; body slender; tail long (SVL/TL = 66.5–89.6 mm);
limbs not overlapping when adpressed; dorsal scales smooth, 39–44 rows at midbody; 72–81 paravertebral scales;
74–86 ventral scale rows; four supraoculars; prefrontals widely separated or in contact; two loreal scales;
supranasal absent; 18–21 lamellae beneath Toe IV; enlarged precloacal scales; dorsal body bands and lateral stripes
absent; numerous thin, faded, light-colored transverse markings on back continuing onto tail to form rectangular
markings; non-descript small, dark speckles on back, flanks, and tail; venter beige; palmar and plantar surfaces
dark-grey; foot scalation on the postaxial margin of Toe IV exhibits a distinct line of demarcation between smooth,
imbricate scales on dorsal surface and rough, tuberculate scales on ventral surface.
Comparisons. Sphenomorphus sungaicolus sp. nov. can be differentiated from other scincid genera in
Peninsular Malaysia by having limbs bearing five digits unlike those of Larutia (Bleeker); by lacking dorsal scales
with keels as in Dasia (Gray) and Eutropis (Fitzinger); lacking an enlarged central disk on the lower eyelid as in
Emoia (Gray) and Lipinia (Gray); having adpressed limbs meeting unlike those of Lygosoma (Hardwicke & Gray)
and Larutia; and lacking supranasal scales unlike Larutia, Dasia, Eutropis, Emoia, Lipinia, and Lygosoma from
Peninsular Malaysia (Grismer 2011).
Sphenomorphus sungaicolus sp. nov. most closely resembles its sister species S. tersus but can be separated
from it by having a smaller body size (SVL 66.5–89.6 mm versus 90.5–96 mm); 72–81 paravertebral scales versus
70; 74–86 ventral scales versus 68; 20–22 scale rows at position of 10th subcaudal scale row versus 19; and dorsal
scales on the posterior margin of Toe IV of the foot approaching the ventral surface versus terminating in a distinct
line of demarcation between smooth dorsal scales and tuberculate ventral scales at the posterior margin of Toe IV
(Fig. 4). The dorsal surface of S. sungaicolus sp. nov. is earthy brown in color versus reddish brown and its tail
color is uniform with the rest of the body versus being darker towards the end of the tail compared to the rest of the
body in S. tersus (Fig. 3). Paravertebral and ventral scale counts were not included by Taylor (1963) for Thai
specimens of Sphenomorphus tersus.
Sphenomorphus sungaicolus sp. nov. bears less resemblance to S. maculatus and S. indicus and is
differentiated from them based on color pattern, SVL (Table 3), and plantar scalation (Fig. 4). Sphenomorphus
sungaicolus sp. nov. can be distinguished from S. maculatus in being significantly larger (SVL = 66.5–86 mm
versus 59.6–62.5 mm). Sphenomorphus maculatus has a distinct wide, dark, dorsolateral stripe extending from the
nostril to the base of the tail that is bordered dorsally by a thin white stripe that is lacking in S sungaicolus sp. nov.
Sphenomorphus maculatus also differs by having a brown ground color with faint black markings on the top of the

SUMARLI E T AL.
36
·
Zootaxa 4173 (1) © 2016 Magnolia Press
head and exhibits a derived plantar scale arrangement in that the dorsal scales on the posterior margin of the foot
extend onto the ventral surface to Toe III. Sphenomorphus sungaicolus sp. nov. has a larger maximum SVL than S.
indicus (SVL = 89.6 mm versus 79 mm) and can be distinguished from it in lacking a golden ground color with a
distinct dark, dorsolateral stripe extending from nostril to the base of the tail. However, both S. sungaicolus sp. nov.
and S. indicus have the same plantar scale morphology despite not being each others closest relatives. Both S.
indicus and S. maculatus have beige colored palmar and plantar surfaces that match the color of the venter. S.
sungaicolus sp. nov. and S. tersus have darker, brown palmar and plantar surfaces that are darker than the venter.
FIGURE 3. (A) Holotype of Sphenomorphus sungaicolus sp. nov. (LSUHC 11722) from Hutan Lipur Sekayu, Terengganu,
Peninsular Malaysia. (B) S. tersus (LSUHC 9041 from Perlis State Park, Perlis, Peninsular Malaysia. (C) Dorsal view of S.
tersus (LSUHC 9041) on left and S. sungaicolus sp. nov. (LSUHC 11722) on right. (D) Ventral view of S. tersus (LSUHC
9041) on left and S. sungaicolus sp. nov. (LSUHC 11722) on right. Photographs by L. Grismer.
Description of holotype. Adult male, (SVL = 77.6 mm); Tal 156.5 mm; head longer than wide; upper head
scales plate-like, smooth; snout pointed, rounded anteriorly in ventral view; rostral wider than high, visible from
above; supranasals absent; frontonasal wider than long, in contact with rostral, anterior loreals, frontal, and
prefrontals; postnasal fused with nasal scale; prefrontals moderately sized, separated from each other by frontals
and frontonasals; frontal narrow posteriorly, in contact with frontonasal, prefrontals, first and second supraoculars,
and frontoparietals; frontoparietal divided and in contact anteriorly, bordered anteriorly by frontal and
anterolaterally by second, third and fourth supraoculars, parietals, and interparietals; interparietal small, with small
transparent eye spot; parietals in contact posteriorly, posterolateral border surrounded by four scales on each side;
five nuchal scales with the same size as dorsal scale rows bordering parietals.
Nostril in center of fused nasal and postnasal scale; loreals two, anterior loreal same level as posterior loreal,
but narrower; anterior loreal single, in contact with posterior loreal, fused nasal and postnasal scale, frontonasal,
and prefrontal scale; posterior loreal single, in contact with anterior loreal, preocular scales, first supercilliary, and
second supralabial; superciliaries eight, first largest; superciliary row complete along entire length of lateral edge
of supraoculars; supraoculars four, second widest, followed by a small postsupraocular; postoculars single, two
pretemporals; seven postsuboculars in contact with fourth to sixth supralabials; two primary temporals, lower one

Zootaxa 4173 (1) © 2016 Magnolia Press
·
37
RIPARIAN SKINK IN PENINSULAR MALAYSIA
in contact with sixth and seventh supralabials; two secondary temporals, upper one very large and in contact with
parietal; seven supralabials, fifth largest and below eye; a shallow loreal-labial groove extends from posterior
corner of nasals across suboculars obliquely downward to end of fourth supralabial; external ear opening ovoid,
without lobules; tympanum recessed, diameter much smaller than eyeball diameter; mental wider than long,
rounded posteriorly, in contact with first infralabial on each side and one postmental; seven infralabials, second and
third equal in length; postmental single, in contact with mental and second infralabials and anterior pair of chin
shields; three pairs of chin shields, first pair enlarged, in contact anteriorly, second pair enlarged and separated by
one gular scale, third pair small and separated by five gular scales; first pair of chin shields in contact with
infralabials 1 and 2, second pair in contact with infralabials 2 and 3, and third pair in contact with infralabials 3 and
4.
Midbody scales in 40 longitudinal rows; dorsal scales smooth, slightly larger or equal to ventral scales, dorsal
scales in 10 rows transversely across back; 76 undifferentiated paravertebral scales; flank scales smooth; 82 rows
of smooth ventral scales; two enlarged precloacal scales; tail base thick; 22 scale rows around tail at the 10th
subcaudal scale position; median subcaudals enlarged, approximately 1.5 times wider than neighboring scales.
Limbs well-developed, pentadactyl; third and fourth fingers equal in length; scales on dorsal surface of Finger
I in two rows and in three rows in Fingers II–IV; subdigital lamellae smooth, 11/11 under Finger IV and 16/16
under Toe IV; foot scalation on the postaxial margin of Toe IV exhibits distinct line of demarcation between
smooth, imbricate scales on dorsal surface and rough, tuberculate scales on ventral surface. There is no overlap
between the ventral and dorsal scale planes.
FIGURE 4. (A) Ventral view of foot showing scalation of Sphenomorphus sungaicolus sp. nov. (LSUHC 11722), (B) S.
indicus (LSUHC 7445), (C) S. tersus (LSUHC 9041), and (D) S. maculatus (LSUHC 7800). Black arrows indicate the line of
demarcation separating smooth imbricating dorsal scales from the rough tuberculate ventral scales. Photographs by A. Sumarli.

SUMARLI E T AL.
38
·
Zootaxa 4173 (1) © 2016 Magnolia Press
FIGURE 5. Head scalation of the holotype of Sphenomorphus sungaicolus sp. nov. Photograph by A. Sumarli.

Zootaxa 4173 (1) © 2016 Magnolia Press
·
39
RIPARIAN SKINK IN PENINSULAR MALAYSIA
FIGURE 6. Type series of Sphenomorphus sungaicolus sp. nov. From the left: BPBM 43794, ZRC 2.4915, LSUHC 11722,
and LSUHC 11780. Photograph by L. Grismer.
Variation (Fig. 6). The paratype BPBM 43794 closely resembles the holotype in head scale arrangement
except that the prefrontal scales are in point contact. UKMHC 0629 is the largest specimen (SVL = 89.6). BPBM
43794 is slightly more yellow in color likely due to its longer time in alcohol and has minute, dark specks that
extend onto the flanks and its body is more robust compared to the holotype. ZRC. 2.4915 is the smallest (SVL =
66.5 mm) and has a lighter tan color compared to the holotype. UKMHC 0707 is a juvenile (SVL = 34.9).
Differences in scale counts and measurements are listed in Table 4.
Coloration in life of holotype. Ground color of dorsal surface of head, body, limbs, and tail earthy brown; top
of head generally uniform brown with non-descript, small, dark flecks; dorsolateral surface overlain with numerous
thin, faded, transverse, light-colored markings extending from nape to distal end of tail and edged by small, dark
speckles; similar alternating paravertebral markings on dorsum; dark, banding pattern on lips; tail colored same as
dorsum; diffuse, light markings with dark speckles on fore-limbs; diffuse light speckles on hind limbs; all ventral
surfaces except for palmar and plantar regions beige; and palmar and plantar surfaces dark-grey.
Coloration of juvenile specimens (Fig. 7). We obtained an in situ photograph of a hatchling Sphenomorphus
sungaicolus sp. nov. from the Korbu Forest Reserve, Perak (Fig. 7) that matches the color pattern of the hatchling
from UKMHC 0707. The ground color of dorsal surface of head, body, and limbs greyish brown; top of head
generally grey-brown with non-descript, small, dark flecks; dorsolateral surface bearing a row of cream to orange
colored elongate spots extending from nape to end of tail and edged by thin dark borders; similar alternating
paravertebral markings on dorsum; dark banding pattern on lips; tail light orange-brown with dark brown zigzag
markings along the length of the tail; light markings with dark speckles on fore-limbs; diffuse, light-orange to
cream speckles on hind limbs; all ventral surfaces except palmar and plantar regions beige; palmar and plantar
surfaces brown; and ventral surface of tail orange to red color.
Coloration in alcohol. Closely resembles coloration in life except overall ground color of dorsal surface of
head, body, limbs, and tail range from darker brown to tan. Tan coloration may have resulted from a longer time in
alcohol for BPBM 43794 and ZRC. 2.4915.
Etymology. “Sungai” is the Malaysian word for river and “colus” is derived from the Latin meaning “dweller
in”. The specific epithet sungaicolus refers the obligate riparian nature of this new species.

SUMARLI E T AL.
40
·
Zootaxa 4173 (1) © 2016 Magnolia Press
TABLE 4. Selected scale and color pattern characteristics of Sphenomorphus sungaicolus sp. nov. provided in
millimeters (mm). Abbreviations listed in the Materials and Methods.
LSUHC
111722
LSUHC
11780
BPBM
43794
ZRC.
2.4915
UKMHC
0629
UKMHC
0707
Holotype Paratype Paratype Paratype
Sex M F F F F /
SVL 77.6 85.8 80.1 66.5 89.6 34.9
Tal 156.5 79.5R 183.5 125.1 127.2 32.7
AG 36.3 41.1 34.1 34.5 47.7 14
SL 6.7 11.7 9 8.7 6.2 3.1
SFIL 53.3 60 55.2 45.9 56.8 25.6
HL 11.9 12.7 10.5 10.4 15.3 8.2
HW 11 11.6 9.5 9 9.3 4.1
HH 7.3 8.2 7.2 6.8 8.9 3.7
STL 15.9 17.4 13.8 13.9 16.7 8.8
EL 3.9 5.1 3.9 3.4 5.1 3.2
END 4.5 4.9 4.5 3.7 4.7 2.4
ETL 6.3 7.3 6 5.4 6 2.7
TYD 1.5 1.5 1.5 1.3 1.4 1
FIL 27 27.9 24.5 22.3 24.3 10.7
HIL 40.7 40.7 43 33.5 37.3 16.5
Upper head scales sm sm sm sm sm sm
Frontonasal 1 1 1 1 1 1
Prefrontal separation wide none none wide none none
Interparietal with a small transparent
spot
yes yes yes yes yes yes
Number of scales bordering parietals
posterolaterally
544444
Parietals in contact posteriorly yes yes no yes yes yes
Nuchals (L/R) 2/2 2/2 2/2 2/2 2/2 2/2
Supraoculars 4 4 4 4 4 4
Loreals 1 1 1 1 2 2
Loreals seperated from supralabials by
small scales
no no no no no no
Lower eyelid scaly scaly scaly scaly scaly scaly
Superciliaries (L/R) 8/8 8/8 8/8 8/8 9/9 9/9
Supraciliary row interrupted by fourth
Supraocular
no no no no no no
Supralabials (L/R) 7/7 7/7 7/7 7/7 7/7 7/7
Infralabials (L/R) 7/7 7/7 7/7 7/7 7/7 7/7
Primary temporal 2 2 2 2 2 2
Secondary temporal 2 2 2 2 2 2
Midbody scale rows 40 39 41 43 44 41
Doral scale rows across the back 10 10 10 8 9 9
Paravertebral scales 76 72 78 81 80 77

Zootaxa 4173 (1) © 2016 Magnolia Press
·
41
RIPARIAN SKINK IN PENINSULAR MALAYSIA
FIGURE 7. (A) Type locality of Sphenomorphus sungaicolus sp. nov. at Hutan Lipur Sekayu, Terengganu, Peninsular
Malaysia. Photograph by L. Grismer. (B) Juvenile S. sungaicolus sp. nov. (not collected) from the Korbu Forest Reserve,
Perak. Photograph by Z. Dzulkaf.
Distribution. Sphenomorphus sungaicolus sp. nov. is known from Hutan Lipur Sekayu, Hutan Lipur
Chemerong, and the Tembat Forest Reserve, Hulu Terengganu, Terengganu, Peninsular Malaysia—localities east
of the Banjaran Titiwangsa. Localities on the western side of the Banjaran Titiwangsa are FRIM and Ulu Gombak,

SUMARLI E T AL.
42
·
Zootaxa 4173 (1) © 2016 Magnolia Press
Selangor and the Korbu Forest Reserve, Perak to the north (Fig. 1). It is likely this species has a greater distribution
throughout Peninsular Malaysia similar to what has been reported for other species of lizards whose distribution
wraps around the southern end of the Banjaran Titiwangsa (Grismer 2011).
Natural history. Sphenomorphus sungaicolus sp. nov. is a lowland species not known to occur above 300 m in
elevation and found only in riparian areas coursing through lowland dipterocarp forest. All specimens were found
along the edges of watercourses. ZRC.2.4915 from FRIM was found on boulders next to a large stream (Leong et
al. 2002) and the holotype was found at night running in water at the edge of a small stream amongst rocks at
Hutan Lipur Sekayu. The Hutan Lipur Chemerong and Ulu Gombak specimens were collected from along
riverbanks. The Hulu Terengganu specimens were collected from pitfall traps located approximately 2.5–3 meters
from the edge of a river. Sphenomorphus sungaicolus sp. nov. is the first obligate riparian skink known from
Peninsular Malaysia. A hatchling S. sungaicolus sp. nov. from the Korbu Forest Reserve, Perak (Fig. 7) was
photographed along the sandy edge of a rocky stream at an elevation of approximately 300 m (Zaharil Dzulkafly in
litt. 2015).
Discussion
The discovery of a new Sphenomorphus in Peninsular Malaysia along with several other descriptions of new
species from this genus in recent years (Grismer 2006; Grismer 2007; Grismer 2008; Grismer et al. 2009a,b;
Grismer & Quah 2015) brings the total number of species in Peninsular Malaysia to 20. Sphenomorphus
sungaicolus sp. nov. is the first confirmed report of an obligate riparian skink from Peninsular Malaysia, although
riparian proclivities have been observed in both lowland S. maculatus (Diong et al. 1995; Ibrahim et al. 2006;
Grismer 2011) and S. tersus (Grismer 2011) but not in the upland S. indicus (Grismer 2011).
Between Sphenomorphus sungaicolus sp. nov. and S. tersus, there is an uncorrected pairwise sequence
divergence of 4.5%. and between S. sungaicolus from Hutan Lipur Sekayu and Hutan Lipur Chemerong there is a
modest degree (1.0%) of sequence divergence as well. This is not uncommon in other riparian species we have
examined such as the bufonid species in the genus Ansonia Stoliczka, 1870 (Davis et al., 2016; Grismer et al.
2016). We attribute this intraspecific genetic variation as a result of being restricted to specific riparian systems for
extended periods of time and only being able to share genes by way of relatively infrequent episodes of stream
capture. This stands in opposition to many other species capable of dispersing through forest habitats that are
genetically very similar yet separated by hundreds of kilometers of uninhabitable terrain (Loredo et al. 2013;
Grismer et al. 2014a,b; Grismer et al. 2015).
As noted by Grismer et al. (2015), the annual rate of new lizard species being discovered in Peninsular
Malaysia since 2003 exceeds that of all other Southeast Asian countries and highlights the continued need for
exploration-driven field work in biodiversity hotspots. As new species are discovered, so will the attendant new
ecologies and adaptations of many of them such as the first riparian species of skink reported here for Peninsular
Malaysia. Such discoveries underscore the need for forest management techniques that take into account small
watershed systems in order to protect endemic riparian microhabitat specialists alongside management plans that
protect broader expanses of forest.
Acknowledgements
We thank The Edmund C. Jaeger Undergraduate Endowment and The Ryckman Undergraduate Research
Endowment for funding this research for AXS. Funding in part to LLG was provided by the College of Arts and
Sciences, La Sierra University. We thank Z. Dzulkafly for providing the picture of the juvenile skink from the
Korbu Forest Reserve and we thank Raul E. Diaz for allowing AXS to use his microscope and Helicon 6 software
for Fig 4. Funding for field work and molecular costs for PLWJ was supported by an NSF EF-1241885 grant issued
to J.W. Sites, Jr. Lastly, we thank Jesse L. Grismer and Chan Kin Onn for their advice and assistance with the
phylogenetic analyses.

Zootaxa 4173 (1) © 2016 Magnolia Press
·
43
RIPARIAN SKINK IN PENINSULAR MALAYSIA
Literature cited
Diong, C.H., Kiew, B.H. & Lim, B.L. (1995) An annotated checklist of the lizard fauna in the Temengor Forest Reserve, Hulu
Perak, Malaysia. Malaysia Nature Journal, 48 (3–4), 353–356.
Drummond, A.J., Suchard, M.A., Xie, D. & Rambaut, A. (2012) An annotated checklist of the lizard fauna in the Temengor
Forest Reserve, Hulu Perak, Malaysia. Molecular Biology and Evolution, 29 (8), 1969–1973.
http://dx.doi.org/10.1093/molbev/mss075
Fitzinger, L. (1843) Systema Reptilium. Fasciculus Primas Amblyglossae. Braumüller et Seidel, Vienna, 106 pp.
Grismer, L.L. (2007) A new specie of small montane forest floor skink (Genus Sphenomorphus Fitzinger, 1843) from southern
Peninsular Malaysia. Herpetologica, 64 (4), 544–551.
http://dx.doi.org/10.1655/0018-0831(2007)63[544:ansosm]2.0.co;2
Grismer, L.L. (2008) A new species of insular skink (Genus Sphenomorphu Fitzinger, 1843) from the Langkawi Archipelago,
Kedah, West Malaysia with the first report of the herpetofauna of Pulau Singa Besar and an updated checklist of the
herpetofauna of Pulau Langkawi. Zootaxa, 1691, 53–66.
Grismer, L.L. (2011) Lizards of Peninsular Malaysia, Singapore, and their adjacent archipelagos. Edition Chimaira, Frankfurt,
36 pp. [pp. 641–676]
Grismer, L.L., Ahmad, N. & Onn, C.K. (2009a) A new diminutive, upland Sphenomorphus Fitizinger, 1843 (Squamata:
Scincidae) from the Belum-Tememgor Forest Complex, Peninsular Malaysia. Zootaxa, 2312, 27–38.
Grismer, L.L., Wood, P.L. Jr. & Grismer, J.L. (2009b) A New Insular Species of Skink of the Genus Sphenomorphus Strauch,
1887 (Squamata: Scincidae) from Pulau Perhentian Besar, Terengganu, Peninsular Malaysia. Tropical Life Sciences
Research, 20 (1), 51–69.
Grismer, L.L., Quah, E.S.H, Anuar, M.S., S., Muin, M.A., Wood, P.L. Jr. & Nor, A.M. (2014a) A diminutive new species of
cave-dwelling Wolf Snake (Colubridae: Lycodon Boie, 1826) from Peninsular Malaysia. Zootaxa, 3815 (1), 051–067.
http://dx.doi.org/10.11646/zootaxa.3815.1.3
Grismer, J.L., Onn, C.K., Quah, E.S.H. & Pauwels, S.A. (2014b) Systematics and natural history of Southeast Asian Rock
Geckos (genus Cnemaspis Strauch, 1887) with descriptions of eight new speices from malaysia, Thailand, and Indonesia.
Zootaxa, 3880 (1), 1–147.
http://dx.doi.org/10/11646/zootaxa.3880.1.1
Grismer, L.L. & Quah, E.S.H. (2015) The Rediscovery of Sphenomorphus malayanus Doria, 1888 (Squamata: Scincidae) from
the Titiwangsa Mountain Range of Peninsular Malaysia and its redescription as S. senja sp. nov. Zootaxa, 3931 (1), 63–70.
http://dx.doi.org/10.11646/zootaxa.3931.1.4
Grismer, L.L., Wood, P.L. Jr., Aowphol, A., Cota, M., Grismer, M.S., Murdoch, M.L., Aguilar, C. & Grismer, J.L. (2016) Out
of Borneo again and again: biogeography of the Stream Toad genus Ansonia Stoliczka (Anura: Bufonidae) and the
discovery of the first limestone cave dwelling species. Biological Journal of the Linnean Society, in press.
Grismer, L.L, Wood, P.L. Jr., Lee, C.H., Quah, E.S.H., Anaur, S., Ngadi, E. & Sites J.W., Jr. (2015) An integrative taxonomic
review of the agamid genus Bronchocela (Kuhl, 1820) from Peninsular Malaysia with descriptions of new montane and
insular endemics. Zootaxa, 3948 (1), 001–023.
http://dx.doi.org/10.11646/zootaxa.3948.1.1
Hedges, S.B., Nussbaum, R.A. & Maxson, L.R. (1993) Caecilian phylogeny and biogeography inferred from mitochondrial
DNA sequences of the 12S rRNA and 16S rRNA genes (Amphibia: Gymnophiona). Herpetological Monographs, 7, 64–
76. Available from: http://www.jstor.org/stable/1466952 (accessed 24 July 2016)
Honda, M., Ota, H., Murphy, R.W. & Hikida, T. (2006) Phylogeny and biogeography of the water skinks of the genus
Tropidophorus (Reptilia: Scincidae): a molecular approach. Zoological Scripta, 35 (1), 85–95.
http://dx.doi.org/10.1111/j.1463-6409.2005.00215.x
Huelsenbeck, J., & Ronquist, F. (2001) MRBAYES: Bayesian inference of phylogeny. Bioinformatics, 17 (8), 754–755.
http://dx.doi.org/10.1093/bioinformatics/17.8.754
Ibrahim, J., Shahrul, M.S., Norhayati, A., Shukor, M.N., Shahriza, S., Nurul’ Ain, E., Nor Zalipah, M. & Rayan, D.M. (2006)
An annotated checklist of the herpetofauna of Langkawi island, Kedah, Malaysia. Malayan Nature Journal, 57, 369–382.
Kocher, T.D., Thomas, W.K., Meyer, A., Edwards, S.V., Pääbo, S., Villablanca, F.X. & Wilson, A.C. (1989) Dynamics of
mitochondrial DNA evolution in animals: Amplification and sequencing of conserved primers. Proceedings of the
National Academy of Sciences, USA, 86 (16), 6196–6200.
Leong, T.M., Yaakob, S.Y. & Das, I. (2002) Geographic distribution. Sphenomorphus tersus. Herpetological Review, 33 (2),
149.
Lim, L.J. (1998) The taxonomy of West Malaysia and Singapore Scincidae (Reptilia: Sauria). Unpublished M.S. Thesis,
National University for Singapore, Kent Ridge, 234 pp.
Linkem, C.W., Diesmos, A.C. & Brown, R.M. (2011) Molecular systematics of the Philippine forest skinks (Squamata:
Scincidae: Sphenomorphus): testing morphological hypotheses of interspecific relationships. Zoological Journal of the
Linnean Society, 163 (4), 1217–1243.
http://dx.doi.org/10.1111/j.1096-3642.2011.00747.x
Linkem, C.W. (2013) Molecular Phylogenetics and Biogeography of Sphenomorphini (Squamata: Scincidae). Ph.D. Thesis.
Department of Ecology and Evolutionary Biology, University of Kansas. Available from: https://kuscholarworks.ku.edu/

SUMARLI E T AL.
44
·
Zootaxa 4173 (1) © 2016 Magnolia Press
handle/1808/10436 (accessed 24 July 2016)
Loredo, A.I., Wood, P.L. Jr., Quah, E.S.H., Anuar, S.H., Greer, L., Norhayati, A. & Grismer, L.L. (2013) Cryptic speciation
within Asthenodipsas vertebralis (Boulenger, 1900) (Squamata: Pareatidae), the description of a new species from
Peninsular Malaysia, and the resurrection of A. tropidonotus (Lidth de Jude, 1923) from Sumatra: an integrative taxonomic
analysis. Zootaxa, 3664 (4), 505–524.
http://dx.doi.org/10.11646/zootaxa.3664.4.5
Macey, J.R. & Verma, A. (1997) Homology in phylogenetic analysis: Alignment of transfer RNA genes and the phylogenic
position of snakes. Molecular Phylogenetics and Evolution, 7 (2), 272–279.
http://dx.doi.org/10.1006/mpev.1997.0379
Maddison, W.P. & Maddison, D.R. (2015) Mesquite: a modular system for evolutionary analysis. Version 3.04. Available from:
http://mesquiteproject.org (accessed 12 December 2015)
Miller, M.A., Pfeiffer, W. & Schwartz, T. (2010) Creating the CIPRES Science Gateway for inference of large phylogenetic
trees. Proceedings of the Gateway Computing Enviroments Workshop (GCE), 14 November 2010, New Orleans, LA, 1–8.
Nguyen, L.-T., Schmidt, H.A., von Haeseler, A. & Minh, B.Q. (2015) IQ-TREE: A fast and effective stochastic algorithm for
estimating maximum likelihood phylogenies. Molecular Biology and Evolution, 32 (1), 268–274.
http://dx.doi.org/10.1093/molbev/msu300
Rambaut, A., Suchard, M.A., Xie, D. & Drummond, A.J. (2014) Tracer v1.6. Available from: http://beast.bio.ed.ac.uk/Tracer
(accessed 21 December 2015)
Rambaut, A. & Drummond, A.J. (2013) TreeAnnotator v1.8.0 MCMC Output Analysis. Available from:http://
beast.bio.ed.ac.uk/TreeAnnotator (accessed 15 February 2015)
Ronquist, F., Teslenko, M., van der Mark, P., Ayres, D.L., Darling, A., Höhna, S., Larget, B., Liu, L., Suchard, M.A. &
Huelsenbeck, J.P. (2012) MrBayes 3.2: Efficient Bayesian Phylogenetic Inference and Model Choice across a Large
Model Space. Systematic Biology, 61 (3), 539–542.
http://dx.doi.org/10.1093/sysbio/sys029
Shea, G.M. (2012) On the identity of the type species of Sphenomorphus (Squamata: Scincidae): Lygosoma melanopogon
Dúmeril and Bibron 1839, with a note on a new scalation character on the pes in Sphenomorphus. Zootaxa, 3490, 1–29.
Tamura, K., Peterson, D., Peterson, N., Stecher, G., Nei, M. & Kumar, S. (2011) MEGA5: Molecular Evolutionary Genetics
Analysis using Maximum Likelihood, Evolutionary Distance, and Maximum Parsimony Methods. Molecular Biology and
Evolution, 28 (10), 2731–2739.
http://dx.doi.org/10.1093/molbev/msr121
Taylor, E.H. (1963) The lizards of Thailand. University of Kansas Science Bulletin, 44, 687–1077.
Wilcox, T.P., Zwickl, D.J., Heath, T.A. & Hillis, D.M. (2002) Phylogenetic relationships of the Dwarf Boas and a comparison
of Bayesian and bootstrap measures of phylogenetic support. Molecular Phylogenetics and Evolution, 25 (2002), 361–371.
http://dx.doi.org/10.1016/s1055-7903(02)00244-0
Uetz, P. (2016) The Reptile Database. Available from: http://www.reptile-database.org (accessed 24 July 2016)