You are on page 1of 17

See discussions, stats, and author profiles for this publication at: https://www.researchgate.

net/publication/274219762

A New Species Of The Genus Tropidophorus Duméril & Bibron, 1839


(Squamata: Sauria: Scincidae) From Vietnam

Article  in  Zootaxa · April 2010


DOI: 10.5281/zenodo.194925

CITATIONS READS

5 159

5 authors, including:

Truong Nguyen Tao Thien Nguyen


The Institute Of Ecology and Biological Resources Vietnam Academy of Science and Technology
277 PUBLICATIONS   1,968 CITATIONS    65 PUBLICATIONS   832 CITATIONS   

SEE PROFILE SEE PROFILE

Andreas Schmitz
Natural History Museum of Geneva
151 PUBLICATIONS   2,682 CITATIONS   

SEE PROFILE

Some of the authors of this publication are also working on these related projects:

Save Vietnam Biodiversity View project

biodiversity View project

All content following this page was uploaded by Truong Nguyen on 29 August 2015.

The user has requested enhancement of the downloaded file.


Zootaxa 2439: 53–68 (2010) ISSN 1175-5326 (print edition)
www.mapress.com / zootaxa/ Article ZOOTAXA
Copyright © 2010 · Magnolia Press ISSN 1175-5334 (online edition)

A new species of the genus Tropidophorus Duméril & Bibron, 1839


(Squamata: Sauria: Scincidae) from Vietnam

TRUONG QUANG NGUYEN1,5,7, TAO THIEN NGUYEN2, ANDREAS SCHMITZ3,


NIKOLAI L. ORLOV4 & THOMAS ZIEGLER6
1
Institute of Ecology and Biological Resources, 18 Hoang Quoc Viet, Hanoi, Vietnam. E-mail: nqt2@yahoo.com
2
Vietnam National Museum of Nature, 18 Hoang Quoc Viet, Hanoi, Vietnam. E-mail: nguyenthientao@gmail.com
3
Muséum d'histoire naturelle, C.P. 6434, CH-1211 Geneva 6, Switzerland. E-mail: andreas.schmitz@ville-ge.ch
4
Zoological Institute, Russian Academy of Sciences, Universitetskaya nab., 1, St. Petersburg 199034, Russia.
E-mail: azemiops@zin.ru
5
Zoologisches Forschungsmuseum Alexander Koenig, Adenauerallee 160, D-53113 Bonn, Germany
6
AG Zoologischer Garten Köln, Riehler Strasse 173, D-50735 Cologne, Germany. E-mail: ziegler@koelnerzoo.de
7
Corresponding author

Abstract

A new species of the genus Tropidophorus is described from Hoang Lien Mountain, Sa Pa and Van Ban districts, Lao Cai
Province, northern Vietnam. Tropidophorus boehmei sp. nov. is characterized by the combination of the following
features: size relatively large (maximum SVL 103.5 mm); upper head scales smooth; frontonasal undivided; prefrontals
very small, widely separated; loreals 2, anterior loreal undivided; supralabials 6; supraciliaries 7–8, supraciliary row
completed along the entire length of the lateral edge of supraoculars; external ear opening present, tympanum superficial,
tympanum diameter smaller than eye length; midbody scales in 30–32 rows; dorsal and lateral scales on body and tail
smooth; paravertebral scales 60–69, not widened; scale rows at position of tenth subcaudal 17–18; ventral scales 56–66;
medial subcaudals divided from first to fifth, remaining ones about 1.5 times wider than neighboring scales; 16–19
smooth lamellae under fourth toe; dorsum and tail base dark grey with light transverse bands and two dorsolateral rows
of white dots from neck to tail base; body sides dark grey with irregular small white dots from behind the eye to tail base.
In addition, previous records of Tropidophorus berdmorei from northern Vietnam are discussed.

Key words: Tropidophorus boehmei sp. nov., T. berdmorei, taxonomy, phylogeny, Vietnam: Lao Cai Province: Hoang
Lien Mountain

Introduction

The genus Tropidophorus Duméril & Bibron, 1839 is a skink group with a high level of local endemism, with
15 of 27 species being restricted to a single country (Greer & Biswas 2004; Chuaynkern et al. 2005; Ziegler et
al. 2005; Honda et al. 2006; Nguyen et al. 2010). In Vietnam, a total of eight water skink species of the genus
Tropidophorus have been recorded and three of them are endemic to the country (Nguyen et al. 2009). From
the Indochinese Peninsula, 15 species of Tropidophorus are currently known: T. baviensis Bourret, 1939; T.
berdmorei (Blyth, 1853); T. cocincinensis Duméril & Bibron, 1839; T. guangxiensis Wen, 1992; T. hainanus
Smith, 1923; T. hangnam Chuaynkern, Nabhitabhata, Inthara, Kamsook & Somsri, 2005; T. laotus Smith,
1923; T. latiscutatus Hikida, Orlov, Nabhitabhata & Ota, 2002; T. matsuii Hikida, Orlov, Nabhitabhata & Ota,
2002; T. microlepis Günther, 1861; T. murphyi Hikida, Orlov, Nabhitabhata & Ota, 2002; T. noggei Ziegler,
Vu & Bui, 2005; T. robinsoni Smith, 1919; T. sinicus Boettger, 1886; and T. thai Smith, 1919 (Hartmann et al.
2009; Nguyen et al. 2009; Nguyen et al. 2010). With the exception of T. berdmorei and T. laotus, all
aforementioned species have keeled dorsal and lateral scales. During recent field work in northern Vietnam,
we found a collection of the water skinks from Hoang Lien (Fansipan) Mountain, which resembled

Accepted by S. Carranza: 15 Mar. 2010; published: 28 Apr. 2010 53


Tropidophorus berdmorei by having smooth dorsal and lateral scales and an undivided frontonasal. However,
closer examination of morphology and mitochondrial DNA revealed them to represent a distinct taxon that
herein is described as a new species of Tropidophorus.

Materials and methods

Field surveys were conducted in May 2004 in Van Ban and in June 2009 in Sa Pa, Lao Cai Province, northern
Vietnam. Specimens referred to in this paper are deposited in the collections of the American Museum of
Natural History (AMNH), New York; Institute of Ecology and Biological Resources (IEBR), Hanoi; Vietnam
National Museum of Nature (VNMN), Hanoi; Muséum d'histoire naturelle (MHNG), Geneva; Royal Ontario
Museum (ROM), Toronto; Zoologisches Forschungsmuseum Alexander Koenig (ZFMK), Bonn; and
Zoological Institute (ZISP), Russian Academy of Sciences, St. Petersburg. Beside the recently collected
specimens, we also examined a specimen from Bourret’s collection which is deposited at the Faculty of
Biology, Vietnam National University (VNUH), Hanoi, Vietnam.
Molecular analysis. Molecular data were collected to examine sequence variation between the new
species and its morphologically closest relatives. A portion of the mitochondrial 16S rRNA gene of two
specimens of the new species (IEBR A.0923, VNMN 822; GenBank accession numbers: GU550104–
GU550105) were sequenced. These sequences were then compared to sequences of other Tropidophorus
sequences from earlier published papers (Honda et al. 2006; Ziegler et al. 2007).
DNA was extracted using QiAmp tissue extraction kits (see Wagner et al. 2009a). The primers 16sar-L
(light chain; 5’ – CGC CTG TTT ATC AAA AAC AT – 3’) and 16sbr-H (heavy chain; 5’ – CCG GTC TGA
ACT CAG ATC ACG T – 3’) of Palumbi et al. (1991) were used to amplify a portion of the mitochondrial
16S ribosomal RNA gene. PCR cycling procedure was as described in Schmitz et al. (2005). PCR products
were purified using Qiaquick purification kits (Qiagen). Sequences were obtained using an automatic
sequencer (ABI 377). Sequences were aligned using ClustalX (Thompson et al. 1997; default parameters) and
manually checked using the original chromatograph data in the program BioEdit (Hall 1999); this resulted in
a total of 500 bp for the chosen section of the 16S gene. Phylogenetic trees were calculated in the program
Paup* 4.0b10 (Swofford 2002) using the neighbor-joining (NJ), and the maximum parsimony (MP: heuristic
searches on parsimony-informative characters only, with random addition of taxa for 100 replications, and
tree bisection-reconnection (TBR) branch-swapping) algorithms and bootstrapping with 20000 and 2000
pseudo-replicates respectively, to estimate node support. Further, we used the online version of PhyML
(Montpellier bioinformatics platform; Guidon & Gascuel 2003) to do a maximum likelihood (ML) analysis
with 1000 bootstraps. The optimal model for the ML analysis (GTR+I+Г) was calculated with Modeltest 3.7
(Posada & Crandall 2001). For NJ, ML and MP we considered clades with values ≥ 70% as strongly
(significantly) supported (Hillis & Bull 1993). Paup* 4.0b10 was also used to compute the uncorrected
pairwise distances for all sequences. Finally, we used the Bayesian (PP) reconstruction method as described in
Schmitz et al. (2005) and Wagner et al. (2009b). All Bayesian analysis were performed with MrBayes 3.12
(Huelsenbeck & Ronquist 2001). For the Bayesian analyses parameters of the model were estimated from the
data set using MrModeltest 2.2 (Nylander 2002). Clades with PP ≥ 95% were considered significantly
supported.
Morphological analysis. Measurements were taken with a digital caliper to the nearest 0.1 mm. The
following abbreviations were used: SVL: Snout-vent length, TaL: Tail length, AG: Distance from posterior
junction of forelimb and body wall to anterior junction of hind limb and body wall (with the limbs held at
right angles to the body), HL: Head length (from tip of snout to posterior margin of parietal or interparietal,
depending on the longest distance), HW: Head width (at the widest portion of temporal region), HH: Head
height (at the deepest portion of temporal region), SL: Snout length (from anterior corner of eye to tip of
snout), STL: Distance from snout to anterior border of tympanum, SFlL: Snout-forelimb length (from tip of
snout to anterior junction of forelimb and body wall to the tip of fourth finger, with the limb held at right
angles to the body), END: Distance from anterior corner of eye to posterior border of nostril, EL: Eye length

54 · Zootaxa 2439 © 2010 Magnolia Press NGUYEN ET AL.


(distance between anterior and posterior corners of eyelid), ETL: Distance from anterior border of tympanum
to posterior corner of eye, TYD: Maximum diameter of tympanum, FlL: Forelimb length (from anterior
junction of forelimb and body wall to the tip of fourth finger, with the limb held at right angles to the body),
HlL: Hind limb length (from anterior junction of hind limb and body wall to the tip of fourth finger, with the
limb held at right angles to the body). Scalation: nuchals: enlarged scales behind parietals, paravertebral
scales: number of scales in a line from posterior edge of parietals to dorsal point opposite posterior margin of
the medial precloacals, ventral scales: number of scales from first gular to anterior margin of precloacals,
scale rows at position of tenth subcaudal on tail including subcaudal. Bilateral scale counts were given as left/
right. Other abbreviations: SD: Standard deviation, a.s.l.: above sea level.

FIGURE 1. Phylogram based on 477 bp of the mitochondrial 16S ribosomal RNA gene sequences. Values above the
nodes represent bootstrap values in percent for the neighbor-joining (NJ), and maximum parsimony (MP) analyses.
Values below the nodes represent the corresponding values for maximum likelihood (ML) and Bayesian (PP)
calculations. Significant values (≥70 for NJ, MP and ML; ≥ 0.95 for PP) are written in bold.

A NEW TROPIDOPHORUS FROM VIETNAM Zootaxa 2439 © 2010 Magnolia Press · 55


56 · Zootaxa 2439 © 2010 Magnolia Press NGUYEN ET AL.
Results

Molecular results

In their recent publication, Honda et al. (2006) could clearly show a basal split within Tropidophorus which
separates a clade of continental Indochinese species (exclusive of T. cocincinensis and T. microlepis) from the
species from Borneo, Sulawesi, and the Philippines. Therefore, we used the published sequences of the
species of the former clade (Honda et al. 2006; Ziegler et al. 2007) to compare it with the sequences we
gained from the newly collected species from Vietnam. From the 500 bp gained for each sequence, a small
hypervariable part (23 bp) needed to be excluded from the analyses because it was to variable to be
unambiguously aligned. As expected for the used 16S rRNA gene and its comparatively short sequences
compared, it lacks the power to resolve all the basal nodes within the recovered phylogenetic tree (Fig. 1), but
it clearly shows Tropidophorus sp. nov. to be embedded deep in the Indochinese subclade of Tropidophorus
sensu Honda et al. (2006).
More specifically, not counting the new taxon, all used phylogenetic reconstruction methods show 6
terminal clades (several of them significantly supported) and two of them even give some comparatively weak
support (NJ: 58/ML: 55) for the new taxon to be part of a specific clade containing T. matsuii, T. latiscutatus
(known only from northeastern and eastern Thailand) and the recently described T. noggei (Ziegler et al.
2007), a species endemic to Vietnam.
Table 1 summarizes the uncorrected p-distances for the used gene fragment. It becomes immediately clear
that the new species shows large and very similar genetic differences from all other member of this subclade
as these latter species show to each other. The full range within this Indochinese subclade ranges from ca.
3.4% (between T. matsuii and T. latiscutatus) to a maximum of ca. 9.8% (between T. sinicus to T. noggei), and
therefore shows a similar range of sequence divergence as known from other studies on Asian scincid lizards
(e.g., Mausfeld & Schmitz 2003). The distances of the newly identified Tropidophorus species to the already
described species ranges from ca. 5.3% (to T. matsuii) to ca. 9.2% (to T. sinicus) and therefore lie perfectly
within the genetic range expected for the species rank within this genus. We also compared the new species to
Sphenomorphus cryptotis (another species of aquatic skinks with a superficial tympanum) and, as expected
because of the allocation of this species to a different genus, found a much higher distance of ca. 13.1%.
The identified genetic structure with more or less equidistant inter-specific genetic distances confirms the
specific distinctness of Tropidophorus sp. nov., and together with the clear morphological evidence (see
below) warrants the recognition of this taxon as new valid species.

Tropidophorus boehmei sp. nov.


(Figs. 2, 3)

Holotype. VNMN 822, adult male, collected on 10 June 2009 by Nguyen Thien Tao and Nguyen Van Quang
in the forest of Hoang Lien Mountain, near Ban Khoang (22o24’N and 103o47’E), Sa Pa District, Lao Cai
Province, northern Vietnam, at an altitude of 1200–1300 m a.s.l.
Paratypes. IEBR A.0922 (adult male), IEBR A.0923 (adult female), VNMN 820 (adult female), VNMN
909 (adult male), MHNG 2721.49 (adult male), ZFMK 89136 (adult male), ZFMK 89137 (adult female), the
same data as the holotype; ZISP 25745 (adult female) collected on 6 May 2004 by Nikolai Orlov in the forest
of Hoang Lien Mountain, near Na Nheo (21o59’N, 104o20’E), Khanh Yen Ha Commune, Van Ban District,
Lao Cai Province, Vietnam, at an altitude of 1000 m a.s.l.; and VNUH S.162 (Bourret’s collection in Hanoi),
collected from “Chapa, Tonkin” (now Sa Pa, Lao Cai Province, northern Vietnam).
Diagnosis. Size large, maximum SVL up to 103.5 mm; upper head scales smooth; frontonasal undivided;
prefrontals small, widely separated from each other; nuchals 1–4; supralabials 6; supraciliaries 7–8,
supraciliary row completed along the entire length of the lateral edge of supraoculars; external ear opening
present, tympanum diameter smaller than eye length; midbody scale rows 30–32; dorsal and lateral scales

A NEW TROPIDOPHORUS FROM VIETNAM Zootaxa 2439 © 2010 Magnolia Press · 57


smooth; paravertebral scales 60–69, not widened; ventral scales 56–66; scale rows at position of tenth
subcaudal 17–18; medial subcaudals divided from first to fifth, remaining ones approximately 1.5 times wider
than neighboring scales; limbs well developed, dorsal surface covered by smooth scales; lamellae smooth,
16–19 under fourth toe; dorsum and tail base dark grey with light transverse bands and two dorsolateral rows
of white dots from neck to tail base; body sides dark grey with small white dots from behind the eye to tail
base.

FIGURE 2. Dorsolateral (A) and ventral (B) views of the holotype (VNMN 822) of Tropidophorus boehmei sp. nov.
from Sa Pa, Lao Cai Province, northern Vietnam. Photo T. T. Nguyen.

Description of holotype. Size large (SVL 99.1 mm, TaL 138.8 mm), for further measurements see Table
2; head longer than wide; upper head scales smooth; snout obtuse, rounded anteriorly; rostral wider than high,
visible from above; supranasals absent; frontonasal as long as wide, in contact with rostral, nasals, anterior
loreals, frontal, and prefrontals; prefrontals very small, widely separated from each other; frontal narrowing
posteriorly, 1.5 times longer than the distance to the snout, in contact with frontonasal, prefrontals, first to
third supraoculars, and frontoparietals; frontoparietals in contact anteriorly, and bordered by frontal, third and

58 · Zootaxa 2439 © 2010 Magnolia Press NGUYEN ET AL.


A NEW TROPIDOPHORUS FROM VIETNAM Zootaxa 2439 © 2010 Magnolia Press · 59
FIGURE 3. Dorsal (A), ventral (B), and lateral (c) views of the head of the holotype (VNMN 822) of Tropidophorus
boehmei sp. nov. from Sa Pa, Lao Cai Province, northern Vietnam. Photo T. Ziegler.

60 · Zootaxa 2439 © 2010 Magnolia Press NGUYEN ET AL.


FIGURE 4. Map showing the distribution (red circle) of Tropidophorus boehmei sp. nov. in Lao Cai Province, northern
Vietnam.

A NEW TROPIDOPHORUS FROM VIETNAM Zootaxa 2439 © 2010 Magnolia Press · 61


62 · Zootaxa 2439 © 2010 Magnolia Press NGUYEN ET AL.
A NEW TROPIDOPHORUS FROM VIETNAM Zootaxa 2439 © 2010 Magnolia Press · 63
fourth supraoculars, parietals and interparietal; interparietal small, without small transparent spot; parietals in
contact posteriorly, posterolateral border surrounded by 4 scales on each side; nuchal scales in 2 pairs; nostril
in centre of nasal; loreals 2, anterior loreal higher but narrower than posterior one; anterior loreal undivided,
posterior loreal in contact with anterior loreal, prefrontal, first supraciliary, preocular, upper presubocular, and
second supralabial; preoculars 2; presuboculars 2, anterior large, posterior one in contact with third and fourth
supralabials; supraciliaries 8, first largest, supraciliary row completed along the entire length of the lateral
edge of the supraoculars; supraoculars 4, second widest, followed by a small postsupraocular; postocular
single; pretemporal single; postsuboculars 4, lower one in contact with fourth and fifth supralabials; primary
temporals 2, lower one in contact with fifth and sixth supralabials; secondary temporals 3, upper one very
large and in contact with parietal; lower eyelid moveable, with four enlarged opaque scales, separated from
supralabials by two rows of small scales; supralabials 6, fourth largest and below the eye; a shallow groove on
loreal-labial border, from posterior corner of nasal across subocular obliquely downward to the end of fourth
supralabial; external ear opening ovoid, without lobules; tympanum superficial, tympanum diameter much
smaller than eye length; mental wider than long, rounded anteriorly, in contact with first infralabial on each
side and postmental; infralabials 5, first longest; postmental undivided, in contact with mental, first infralabial
on both sides, and anterior pair of chinshields; chinshields in 3 pairs, first pair in contact anteriorly, second
pair separated by a gular scale, third pair very small and separated by three scales; first and second pairs of
chinshields in contact with infralabials, third pair separated from infralabials by small scales.
Midbody scales in 30 rows; dorsal scales smooth, subequal to lateral and ventral scales, in 8 rows across
the back; paravertebral scales 64, not widened; lateral scales smooth; ventral scales smooth, in 65 rows;
precloacals 2+2, inner precloacals overlapping outer ones, central two enlarged, right one overlapped by left
one; partially everted hemipenis bifurcated apically; tail thick at base, 17 scale rows at position of tenth
subcaudal; medial subcaudals divided from first to third, remaining ones about 1.5 timer wider than
neighboring scales.
Limbs well developed, pentadactyl; third and fourth fingers equal in length; scales on dorsal surface of
basal digit IV in two rows and terminal three scales in single row; subdigital lamellae smooth, numbering 12/
12 under fourth finger and 16/17 under fourth toe.
Coloration in alcohol. Dorsal head, back, and tail base dark grey with indistinct light transverse bands,
faded from anterior to posterior: one between anterior corners of eye, two on neck, seven between limb
insertions, three on tail base; dorsum with two rows of white dots from neck to the point above vent; body
sides dark grey with irregular white dots from behind the eye to groin; supralabials and infralabials with light
dots; limbs dorsally dark grey with white dots; chin and throat grey; venter and underside of tail base
brownish cream; underside of tail tip dark grey. For coloration in life see Fig. 2.
Variation of paratypes. Measurements, pholidosis and color pattern of the paratypes are summarized in
Tables 2 and 3. The specimen from Van Ban (ZISP 25745) has parietals separated by a small scale posteriorly,
and midbody scales in 32 rows. Males are somewhat larger than females (SVL 89.7 ± 14.3 mm, n = 6 versus
72.9 ± 9.4 mm, n = 4, respectively).
Distribution. Tropidophorus boehmei sp. nov. is currently known only from Hoang Lien Mountain in Sa
Pa and Van Ban districts, Lao Cai Province, Vietnam (Fig. 4).
Natural history. Specimens were collected in the late afternoon and in the evening (1600–2100 h) in and
nearby rocky forest streams. When disturbed they disappeared in the water or in rock holes. The stomach
content of the female (IEBR A.0923) contained remains of a spider (Arachnida). Stomachs were empty in
VNMN 820, 909, and ZFMK 89137. The holotype had white testes (7.1 mm in length and 3.4 mm in
diameter). The females contained small white follicles.
Etymology. We name this new species in honor of Prof. Dr. Wolfgang Böhme, vice director of the
Zoological Research Museum Alexander Koenig in Bonn, head of the Vertebrate Section and curator for
herpetology, in recognition of his outstanding contributions towards lizard systematics and training of many
herpetologist generations. For the common names we suggest Boehme’s Water Skink (English), Thằn lằn tai
boê-me (Vietnamese), and Böhmes Wasserskink (German).
Comparisons. Based on data from the literature (e.g., Blyth 1853; Boulenger 1887; Smith 1919, 1923,
1935; Taylor 1963; Wen 1992; Ngo et al. 2000; Hikida et al. 2002; Greer & Biswas 2004; Chuaynkern et al.
2005; Ziegler et al. 2005, 2007; Bourret 2009; Nguyen et al. 2010) and specimen-based research (Appendix

64 · Zootaxa 2439 © 2010 Magnolia Press NGUYEN ET AL.


1), we compare Tropidophorus boehmei sp. nov. in the following with congeners from the Indochinese
Peninsula:
Tropidophorus boehmei sp. nov. differs from T. cocincinensis, T. hainanus, T. guangxiensis, T. microlepis,
T. robisoni, T. sinicus, and T. thai by having smooth dorsal head scales (versus striated in latter species).

TABLE 4. Comparisons of diagnostic characters in Tropidophorus with smooth dorsal head scales from the Indochinese
Peninsula (* = data obtained from literature).
baviensis boehmei berdmorei hangnam laotus latiscutatus matsuii murphyi noggei
Frontonasal divided (2) 1 1 1 1 2 1 2 1 1
or undivided (1)
Prefontals in contact 1, rarely 0 0 0 or 1 1 0 or 1 0 1 1 1or 0
(1) or separated (0)
Parietals in contact (1) 0 1, rarely 0 1, rarely 0 0 1 0 0 0 0
or separated (0)
posteriorly
Loreals 2 2 2 1 2 2 2 2 2
Anterior loreal divided yes no yes, no no yes yes yes yes no
Supraciliaries 5–7 7–8 8 4 8–9 6–7 8 5–8 2–5
Supraciliary row no no yes yes yes no no no no
interrupted by fourth
supraocular
Supralabials 6 6 6 7–8 6–7 6 6 6 6
Infralabials 5–6 5 5–7 5-6 5–6 4–6 6 5–6 4–5
Midbody scale rows 28–30 30–32 32–40 28–31 30–36 28–30 34 30–32 22–24
Paravertebrals 47–53 60–69 64 52 – 58–63 65 55–67 43–49
Paravertebral scales 0 0 – 0 – 1 0 0 0
widened (1) or not (0)
Dorsal body scales kl sm sm (kl in kl sm kl kl kl kl
smooth (sm) or keeled juveniles)
(kl)
Ventral scales 44–52 56–66 53 – – – – – 40–44
Scale rows at position 11 17–18 – – – 13–15 15 13 9–10
of tenth subcaudal
Subcaudals divided on 1 1 1 2 1 1 1 1 1
anterior part of tail (1)
or on whole tail (2)
Lamellae under fourth 18–22 16–19 22 18–19 18–22 18–22 22–23 20–25 18–20
toe
White spots on dorsum absent present present absent present absent absent absent absent
Sample size (n) n = 5, * n = 10 * * * * * n = 2, * n=8

Tropidophorus boehmei sp. nov. differs from remaining water skink species, that also have smooth dorsal
head scales (Table 4) as follows: from T. baviensis, T. hangnam, T. latiscutatus, T. matsuii, T. murphyi, and T.
noggei in having smooth dorsal body scales (versus being keeled in the latter species). The new species
further differs from T. baviensis, T. latiscutatus, T. matsuii, T. murphyi, and T. noggei by having a higher
number of scale rows at position of tenth subcaudal (17–18 versus 9–15). Tropidophorus boehmei sp. nov. is
most similar to Tropidophorus berdmorei and T. laotus due to the presence of smooth body scales. However,
Tropidophorus boehmei sp. nov. differs from both latter species by having the supraciliary row completed
along the entire length of the lateral edge of the supraoculars. The new species is further distinguished from T.
berdmorei by having lower counts of midbody scale rows (30–32 versus 32–40 in T. berdmorei), lower counts
of lamellae under the fourth toe (16–19 versus 22 in T. berdmorei), and a higher number of ventral scales (56–

A NEW TROPIDOPHORUS FROM VIETNAM Zootaxa 2439 © 2010 Magnolia Press · 65


66 versus 53 in T. berdmorei). Tropidophorus boehmei sp. nov. furthermore differs from T. laotus by having
the tympanum diameter (1.7–2.9 mm) smaller than the eye length (3.6–5.2 mm) (versus tympanum as large as
eye in T. laotus) and frontonasal undivided (versus frontonasal divided in T. laotus).

Discussion

Tropidophorus boehmei sp. nov. is most similar to T. berdmorei, a species which is widely distributed from
China through Myanmar and Vietnam to Thailand (Nguyen et al. 2009; Nguyen et al. 2010). The holotype of
T. berdmorei, collected from Mergui (Myanmar), has 8 distinct rows of keeled scales along the back as was
stated in Blyth’s (1853) description. Hence Tropidophorus boehmei sp. nov. differs from the holotype of T.
berdmorei by having dorsal scales smooth (versus keeled). Although Blyth (1853) had a single type specimen,
Smith (1923) noted three types from Myanmar in the Indian Museum in Calcutta (one of which is probably
Blyth's type). According to Smith (1923) these three specimens have 32–34 midbody scale rows and the
dorsal scales are distinctly keeled in two small specimens but smooth in the largest one of the series.
Therefore, Tropidophorus boehmei sp. nov. is further distinguishable from T. berdmorei by having a lower
number of midbody scale rows (30–32 versus 32–34). Boulenger (1887) mentioned a collection of T.
berdmorei from Pegu and Tenasserim (Myanmar) which obviously differs from Tropidophorus boehmei sp.
nov. by having 38–40 midbody scale rows; 6–7 supraciliaries and the supraciliary row being interrupted by
the fourth supraocular. Boulenger (1887) used the name T. yunnanensis for three specimens from Yunnan,
which were originally labelled as T. berdmorei by Anderson (1878). Subsequently, T. yunnanensis was
synonymized with T. berdmorei by Smith (1923). According to Smith (1923), the Yunnan specimens are lost,
however, Tropidophorus boehmei sp. nov. also differs from these by having lower numbers of midbody scale
rows (30–32 versus 34 rows) and dorsal scales being smooth (versus dorsal scales keeled in all three Yunnan
specimens) (Smith 1923). Finally, according to Taylor (1963) and Greer & Biswas (2004), the specimens of T.
berdmorei from Thailand have the supraciliary row interrupted by the fourth supraocular. This character is
absent in Tropidophorus boehmei sp. nov., revealed by the examination of nine specimens recently collected
during our field work in Vietnam and two specimens of Bourret's collection (see Bourret 2009: Fig. 97b, S.88;
per. obs., S.162). Therefore, the previous records of T. berdmorei from Sa Pa (e.g., Tran et al. 1981; Bobrov
1998; Bourret 2009; Nguyen et al. 2009) should be replaced by Tropidophorus boehmei sp. nov. and the
records of T. berdmorei published from other localities in Vietnam must be carefully re-examined in the
future.

Acknowledgements

We thank the directorate of the Hoang Lien National Park and the Forest Protection Department of Lao Cai
Province for issuing relevant permits and support our field work. Nguyen Thien Tao is grateful to Nguyen Van
Quang (Lao Cai), Nikolai Orlov thanks Ho Thu Cuc (Hanoi) and Rustam K. Berdiev (St. Petersburg) for
assistance in the field. Nguyen Quang Truong thanks Thomas Giang (Cologne) for assistance in analysing the
stomach contents. For the loan of specimens, we acknowledge Le Xuan Canh, Pham Van Luc, Vu Ngoc
Thanh (Hanoi), Wolfgang Böhme (Bonn), and Natalia Ananjeva (St. Petersburg). We thank Eleanor Sterling
(New York) and Kevin Koy (Berkeley) for providing the map; Tsutomu Hikida (Kyoto), Hidetoshi Ota
(Hyogo), and Yodchaiy Chuaynkern (Pathum Thani) kindly provided literature. This research was partially
funded by the Vietnam National Museum of Nature (VNMN); Zoological Institute, Russian Academy of
Sciences, St. Petersburg (ZISP, Grant No. RFBR08-04-00041); and German Academic Exchange Service
(DAAD).

66 · Zootaxa 2439 © 2010 Magnolia Press NGUYEN ET AL.


Literature cited

Anderson, J. (1878) Anatomical and zoological researches: comprising an account of the zoological results of the two
expeditions to western Yunnan in 1868 and 1875; and a monograph of the two Cetacean genera, Plantanista and
Orcella. Bernard Quaritch, London, 15, xxv + 985 pp.
Blyth, E. (1853) Notices and descriptions of various reptiles, new or little known. The Journal of the Asiatic Society of
Bengal, Calcutta, 22, 639–655.
Bobrov, V.V. (1998) Lizards of Fan Si Pan mountain region and Sa Pa vicinity. In: Korzun, L. P & Kalyakin, M. V.
(Eds.). Materials of zoological and botanical studies in Fansipan Summit area (North Vietnam). Moscow – Hanoi,
pp. 227–232 (in Russian).
Boulenger, G.A. (1887) Catologue of the lizards in the British Museum (Natural History). Volume III, Trustees, London,
xii + 575 pp. + XL pls.
Bourret, R. (2009) Les lézards de l’Indochine. Edition Chimaira, Frankfurt am Main, 346 pp.
Chuaynkern, Y., Nabhitabhata, J., Inthara, C., Kamsook, M. & Somsri, K. (2005) A new species of the water skink
Tropidophorus (Reptilia: Squamata: Scincidae) from northeastern Thailand. The Thailand Natural History Museum
Journal, 1, 165–176.
Greer, A.E. & Biswas, S. (2004) A generic diagnosis for the Southeast Asian scincid lizard genus Tropidophorus
Duméril & Bibron, 1839 with some additional comments on its morphology and distribution. Journal of
Herpetology, 38, 426–430.
Guindon S. & Gascuel O. (2003) A simple, fast, and accurate algorithm to estimate large phylogenies by maximum
likelihood. Systematic Biology, 52(5), 696–704.
Hall, T.A. (1999) BioEdit: a user–friendly biological sequence alignment editor and analysis program for Windows 95/
98/NT. Nucleic Acids Symposium Series, 41, 95–98.
Hartmann, T., Chipich, H., Handschuh, M., Nguyen, Q.T. & Böhme, W. (2009) First record of Tropidophorus
cocincinensis Duméril & Bibron, 1839 from Cambodia. Herpetology Notes, 2, 87–89.
Hikida, T., Orlov, N. L., Nabhitabhata, J. & Ota, H. (2002) Three new depressed-bodied water skinks of the genus
Tropidophorus (Lacertilia: Scincidae) from Thailand and Vietnam. Current Herpetology, 21, 9–23.
Hillis, D.M. & Bull, J.J. (1993) An empirical test of bootstrapping as a method for assessing confidence in phylogenetic
analysis. Systematic Biology, 42, 182–192.
Honda, M., Ota, H., Murphy, R.W. & Hikida, T. (2006) Phylogeny and biogeography of water skinks of the genus
Tropidophorus (Reptilia: Scincidae): a molecular approach. Zoologica Scripta, 35, 85–95.
Mausfeld, P. & Schmitz, A. (2003) Molecular phylogeography, intraspecific variation and speciation of the Asian scincid
lizard genus Eutropis Fitzinger, 1843, (Squamata: Reptilia: Scincidae): taxonomic and biogeographic implications.
Organisms, Diversity and Evolution, 3(3), 161–171.
Ngo, A., Murphy, R.W., Orlov, N.L., Darevsky, I.S. & Nguyen, V.S. (2000) A redescription of the Ba Vi water skink
Tropidophorus baviensis Bourret, 1939. Russian Journal of Herpetology, 7, 155–158.
Nguyen, Q.T., Nguyen, V.S., Nikolai, O., Hoang, N.T., Böhme, W. & Ziegler, T. (2010) A review of the genus
Tropidophorus (Squamata: Scincidae) from Vietnam with new species records and additional data on natural history.
Zoosystematics and Evolution, 86(1), 5–19.
Nguyen, V.S., Ho, T.C. & Nguyen, Q.T. (2009) Herpetofauna of Vietnam. Edition Chimaira, Frankfurt am Main, 768 pp.
Palumbi, S.R., Martin, A., Romano, S., McMillan, W.O., Stice, L. & Grabowski, G. (1991) The simple fool's guide to
PCR. Department of Zoology and Kewalo Marine Laboratory, Hawaii, 47 pp.
Posada, D. & Crandall, K.A. (2001) Selecting the best-fit model of nucleotide substitution. Systematic Biology, 50, 580–
601.
Schmitz, A., Ineich, I. & Chirio, L. (2005) Molecular review of the genus Panaspis sensu lato in Cameroon, with special
reference to the status of the proposed subgenera. Zootaxa, 863, 1–28.
Smith, M.A. (1919) The lizards of the genus Tropidophorus in Siam, with description of two new species. Journal of
Natural History Society of Siam, 3, 223–228.
Smith, M.A. (1923) A review of the lizards of the genus Tropidophorus on the Asiatic mainland. Proceedings of the
Zoological Society of London, 1923, 775–781.
Smith, M.A. (1935) The fauna of British India, including Ceylon and Burma: Reptilia and Amphibia. Vol. 2–Sauria.
Taylor and Francis, London, xii + 440 pp.
Swofford, D.L. (2002) PAUP*: Phylogenetic analysis using Parsimony (*and other methods), version 4.0b10.
Sunderland, MA: Sinauer Associates.
Taylor, E.H. (1963) The lizards of Thailand. The University of Kansas Science Bulletin, 44, 687–1077.
Thompson, J.D., Gibson, T.J., Plewniak, F., Jeanmougin, F. & Higgins, D.G. (1997) The ClustalX windows interface:
flexible strategies for multiple sequence alignment aided by quality analysis tools. Nucleic Acids Research, 24,
4876–4882.

A NEW TROPIDOPHORUS FROM VIETNAM Zootaxa 2439 © 2010 Magnolia Press · 67


Tran, K., Nguyen, V.S. & Ho, T.C. (1981) [Baseline survey on the herpetofauna in the North of Vietnam (1955-1976)].
In: Dao, V. T., Nguyen, D. T. Nguyen, K. L. & Vu, D. T. (Eds). [Baseline survey on the fauna in the North of
Vietnam]. Science and Technology Publishing House, Hanoi, pp. 365–427 (in Vietnamese).
Wagner, P., Barej, M. & Schmitz, A. (2009a) Studies on African Agama VII. A new species of the Agama agama –group
(Linnaeus, 1758) (Sauria: Agamidae) from Cameroon & Gabon, with comments on Agama mehelyi Tornier, 1902.
Bonner zoologische Beiträge, 56(4), 285–297.
Wagner, P., Böhme, W., Pauwels, O.S.G. & Schmitz, A. (2009b) A review of the African red-flanked skinks of the
Lygosoma fernandi (Burton, 1836) species group (Squamata: Scincidae) and the role of climate change in their
speciation. Zootaxa, 2050, 1–30.
Wen, Y.-T. (1992) A new species of the genus Tropidophorus (Reptilia: Lacertilia) from Guangxi Zhang Autonomous
Region, China. Asiatic Herpetological Research, 4, 18–22.
Ziegler, T., Heidrich, A., Schmitz, A. & Böhme, W. (2007) Phylogenetic position, morphology and natural history of the
Vietnamese water skink Tropidophorus noggei Ziegler, Vu & Bui, 2005 (Sauria: Scincidae). Revue suisse de
Zoologie, 114, 599–614.
Ziegler, T., Vu, N.T. & Bui, N.T. (2005) A new water skink of the genus Tropidophorus from the Phong Nha–Ke Bang
National Park, central Vietnam (Squamata: Sauria: Scincidae). Salamandra, 41, 137–146.

Appendix 1. Comparative specimens examined

Tropidophorus baviensis: Nghe An Province: IEBR A.0744, A.0753, IEBR A.0754, ZFMK 87588; Phu Tho Province:
VNMN 181.
Tropidophorus cocincinensis: Thua Thien–Hue Province: IEBR 387–390, 393–395, 397, 400, 402, 411–413, 415, 417–
418, 443, 445, 391–392, 396, 398–399, 401, 403–404, 414, 416, 420, 441–442, 444, A.0815–A.0818; Quang Tri
Province: IEBR 2258; Quang Nam Province: IEBR 3125.
Tropidophorus hainanus: Ha Tinh Province: AMNH 147123; Bac Giang Province: IEBR A.0711, 505, 563; Lao Cai
Province: IEBR 1219; Lang Son Province: IEBR A.0710, ZFMK 84618; Ha Giang Province: IEBR A.73, ZFMK
84619.
Tropidophorus microlepis: Dong Nai Province: IEBR A.0756, ZFMK 88042, IEBR A.0846– A.0847.
Tropidophorus murphyi: Cao Bang Province: ROM 41225, 41229 (paratypes).
Tropidophorus noggei: Quang Binh Province: ZFMK 83668 (holotype), ZFMK 83669, VNUH 18605.1 (paratypes),
MHNG 2683.99, Collection of the Phong Nha - Ke Bang National Park: PNNP 111–112, 174–175.
Tropidophorus sinicus: Cao Bang Province: IEBR A.0759; Bac Giang Province: IEBR 501–503, 3642, 3662–3663,
3667; Quang Ninh Province: IEBR A.0810; Hai Duong Province: IEBR A.0811.

68 · Zootaxa 2439 © 2010 Magnolia Press NGUYEN ET AL.

View publication stats