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A new species of wolf snake (Colubridae: Lycodon Fitzinger, 1826) from Phnom Samkos Wildlife Sanctuary, Cardamom Mountains, southwest Cambodia

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  • National Meanchey University

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A new species of the genus Lycodon Fitzinger, 1826 is described from the Cardamom Mountains of southwest Cambodia. Lycodon zoosvictoriae distinctly differs from all other species of Lycodon in Southeast Asia by a combination of its morphometric characters and unique coloration. The new species has 17 dorsal scales at midbody; 2+2 temporals; 8 supralabials; 10 infralabials; loreal separated from internasal and orbit; 213 ventrals; 85 subcaudals; pale tan brown ground color; irregular dark brown blotches on anterior part, 31 transverse blotches on posterior part of body and 26 blotches on tail. Given its submontane type locality, the new species could prove to be endemic to the Cardamom Mountains of southwestern Cambodia and probably Southeast Thailand.
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Accepted by Z. T. Nagy: 24 Apr. 2014; published: 6 Jun. 2014
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http://dx.doi.org/10.11646/zootaxa.3814.1.3
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A new species of wolf snake (Colubridae: Lycodon Fitzinger, 1826) from Phnom
Samkos Wildlife Sanctuary, Cardamom Mountains, southwest Cambodia
THY NEANG
1,4
, TIMO HARTMANN
2
, SEIHA HUN
1,3
, NICHOLAS J. SOUTER
1,3
& NEIL M. FUREY
1,3
1
Fauna & Flora International, Cambodia Program.# 19, Street 360, Boeng Keng Kong 1, Chamkarmorn, Phnom Penh, Cambodia
2
Zoological Research Museum Alexander Koenig (ZFMK), Adenauerallee 160, D-53113 Bonn, Germany
3
Centre for Biodiversity Conservation, Room 415, Faculty of Science, Royal University of Phnom Penh, Confederation of Russia Bou-
levard, Phnom Penh, Cambodia
4
Corresponding author. E-mail: thy.neang@fauna-flora.org
Abstract
A new species of the genus Lycodon Fitzinger, 1826 is described from the Cardamom Mountains of southwest Cambodia.
Lycodon zoosvictoriae distinctly differs from all other species of Lycodon in Southeast Asia by a combination of its mor-
phometric characters and unique coloration. The new species has 17 dorsal scales at midbody; 2+2 temporals; 8 suprala-
bials; 10 infralabials; loreal separated from internasal and orbit; 213 ventrals; 85 subcaudals; pale tan brown ground color;
irregular dark brown blotches on anterior part, 31 transverse blotches on posterior part of body and 26 blotches on tail.
Given its submontane type locality, the new species could prove to be endemic to the Cardamom Mountains of southwest
Cambodia and probably Southeast Thailand.
Key words: Diagnosis, herpetofauna, Pursat, Indochina, systematics, morphology, etymology, Lycodon
Introduction
The colubrid wolf snakes of the genus Lycodon Fitzinger, 1826 are characterized by (1) their strongly arched
maxillary bone bending inwards anteriorly, having three to six enlarged and fang-like anterior maxillary teeth
without venomous grooves, increasing in size posteriorly, separated by a diastema from the posterior seven to
fifteen teeth, posteriorly increasing in size, the last two being enlarged; (2) rounded orbit with vertical elliptic
pupil; (3) smooth or weakly keeled dorsal scales in rows of 15–21 (Smith 1943; Taylor 1965; Lanza 1999; Stuebing
et al. 1999; Daltry & Wüster 2002; Das 2010; Vogel & David 2010; Vogel et al. 2009, 2012); (4) most wolf snakes
have a distinct or indistinct white or pale brown nuchal collar followed posteriorly by a black to blackish-brown
ground color with various white, cream or yellow bands, blotches and speckles (some without pale coloration) on
body and tail; (5) the anal plate varies from divided to undivided (Lanza 1999; Daltry & Wüster 2002; Das 2010).
The most recent phylogenetic revisions by Siler et al. (2013) and Guo et al. (2013) indicate that the genus Dinodon
Duméril, 1853 is morphologically and genetically a closely related genus and a junior synonym of Lycodon, a view
we follow herein.
Herpetofaunal field investigations and examination of museum specimens in recent years have yielded
descriptions of 14 new species of Lycodon, six of which were reported from the Philippines: L. alcalai Ota & Ross,
1994; L. bibonius Ota & Ross, 1994, L. chrysoprateros Ota & Ross, 1994, L. solivagus Ota & Ross, 1994, L.
ferroni Lanza, 1999 and L. fausti Gaulke, 2002; three from China: L. synaptor Vogel & David, 2010, L. gongshan
Vogel & Luo, 2011 and L. liuchengchaoi Zhang, Jiang, Vogel & Rao, 2011; two from India: L. zawi Slowinski,
Pawar, Win, Thin, Gyl, Oo & Tun, 2011 and L. flavicollis Mukherjee & Bhupathy, 2007; one from Cambodia: L.
cardamomensis Daltry & Wüster, 2002; one from southern Thailand: L. ophiophagus Vogel, David, Pauwels,
Sumontha, Norval, Hendrix, Vu & Ziegler, 2009; and one from Laos: L. davidi Vogel, Nguyen, Kingsada &
Ziegler, 2012. As a consequence, the genus Lycodon has now become one of the most diverse genera within the
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A NEW SPECI ES OF WOL F SNAKE FROM SOUTHWEST CAMBODIA
Colubridae worldwide. At present, the genus contains 47 species (including eight species formerly assigned to
Dinodon) which occur from eastern Iran and central Asia to southern China and Japan, southward to the
Philippines and the Indo-Australian Archipelago (Ota & Ross 1994; Lanza 1999; Vogel & David 2010; Vogel &
Luo 2011; Zhang et al. 2011; Uetz 2013). Of the 47 known species, 33 are reported from Southeast Asia, 21 from
the Indo-Burmese Region, 14 from Indochina (Uetz 2013) and five species from Cambodia. In the latter country,
the genus is represented by Lycodon capucinus (Boie, 1827), L. cardamomensis Daltry Wüster, 2002, L. laoensis
Günther, 1864, L. septentrionalis (Günther, 1875) and L. subcinctus Boie, 1827 (Saint Girons 1972; Daltry &
Wüster 2002; Stuart et al. 2006; Grismer et al. 2007, 2008, 2011; Bezuijen et al. 2009; Das 2010).
During a field survey in 2013 at Phnom Samkos Wildlife Sanctuary (Fig. 1) in southwest Cambodia, 22 reptile
species were collected. Of these, a single wolf snake specimen could not be assigned to any of the nominal
Lycodon species currently recognized. Due to significant differences in its morphology and coloration, we describe
the specimen as a new species.
FIGURE 1. Type locality of Lycodon zoosvictoriae sp. nov.: Phnom Samkos Wildlife Sanctuary, Cardamom Mountains,
southwest Cambodia.
Material and methods
Intensive sampling of amphibians and reptiles was undertaken during day and night times from 11–16 June 2013 at
Phnom Samkos Wildlife Sanctuary, Cardamom Mountains, southwest Cambodia. The field work was conducted by
Thy Neang, Seiha Hun and four assistants from a nearby village. The above-mentioned specimen was caught using
snake tongs and kept in a cloth bag for one night. The specimen was photographed prior to euthanasia after which
liver tissue was taken and stored in 97 % ethanol for future genetic analysis. Following this, the specimen was
preserved in 10% formalin for 36 hours before being soaked in water for 24 hours to avoid dehydration and then
immediately transferred to 70% ethanol for storage in the zoological museum of the Centre for Biodiversity
Conservation at the Royal University of Phnom Penh, Cambodia.
Measurements and counts of the morphological characters of the type specimen were carried out using a poly
dial caliper (SPI 31–415–3) to the nearest 0.1 mm under a Nikon SMZ 645 dissecting microscope. Morphological
data measured included: snout to vent length (SVL)–measured from the tip of the snout to the vent; tail length
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(TaL)–from the vent to the tip of the tail; TaL/SVL–ratio of TaL/SVL; head width (HW)–the widest section of the
head at posterior margin of parietals; head length (HL)–from the tip of the snout to the posterior margin of the
mandible; head depth (HD)–vertical height between upper and ventral surfaces of head measured at HW;
interorbital distance (IO)–the distance between outer margins of supraoculars; eye diameter (ED)–horizontal
diameter of eye; snout length (SnL)–distance between the tip of the snout and anterior edge of eye; eye to nostril
(EN)–distance between posterior margin of nostril and anterior margin of eye; nostril distance (ND)–horizontal
distance between nostrils; dorsal scale rows at neck (DSN)–the number of scale rows at one head length behind the
head; mid-body scale rows (MSR)–the number of scale rows at mid-body; dorsal scale rows anterior to the vent
(DSV)–the number of dorsal scale rows at one head length prior to the vent; dorsal scale rows (DSR)–a general
scale formula in the form “DSN–MSR–DSV”. Ventral scales were excluded from the dorsal scale count.
Ventral scales (VS) were counted according to Dowling (1951); the number of scales from the third ventral
scales leaving two preventrals immediately posterior to gulars to the vent excluding anal plate; anal plate (AP)–the
number of single or divided terminal ventral scales immediately anterior to the vent; subcaudal scales (SC)–the
number of paired subcaudal scales excluding the terminal scute; supralabials (SL)–the number of scales on upper
lips; SL-orbit contact–number of SL entering orbit; infralabials (InL)–the number of scales on lower lips; maxillary
teeth (MT)–the number of maxillary teeth on both sides of upper jaw; temporals (TP)–the number of scales
immediately behind postoculars and between posterior SL and parietals; number of transverse blotches on
body–the number of blotches transversely extending across the back and downward to the sides but not encircling
the body; number of blotches on tail–the number of blotches on the tail. Values of paired characters are given in left
and right order.
Species morphological data used for comparison were taken from literature (see Table 1) and examination of
museum specimens (see Appendix 1). Museum acronyms used are: CBC–Centre for Biodiversity Conservation,
Royal University of Phnom Penh, Cambodia; ZFMK–Zoologisches Forschungsmuseum Alexander Koenig, Bonn,
Germany.
Systematics
The new species described below is placed in the genus Lycodon, since it possesses the characteristically arched
maxillary bones bearing fang-like teeth, posteriorly increasing in size and rounded orbits with vertical elliptic
pupils (see introduction).
Lycodon zoosvictoriae sp. nov.
(Fig. 2–6; Table 1)
Holotype (Fig. 2). CBC02238, an adult female, collected by Thy Neang and Seiha Hun, 15 June 2013 at
N12º09'22.0", E102º59'18.7", 1,284 m above sea level (a.s.l.) in Phnom Samkos Wildlife Sanctuary, Cardamom
Mountains, Pursat Province, southwest Cambodia.
Diagnosis. Lycodon zoosvictoriae sp. nov. is differentiated from all of its congeners by the following unique
combination of morphological characters: ratio of TaL/SVL 0.27; temporals 2+2; maxillary teeth 9 (5 anterior and
4 posterior); supralabials 8; preoculars 1L/2R; postoculars 2; Lo-orbit contact absent; Lo-internasal contact absent;
infralabials 10; DSR 17–17–15; ventral scales 213; subcaudal scale 85; anal plate entire; dorsal surface of head
dark brown with pale tan brown on edges and between scales; irregularly scattered dark brown blotches on anterior
dorsum, 31 regular dark brown transverse blotches on posterior dorsum and 26 on tail; color of blotches dark
brown; body coloration light tan brown; dorsal scale weakly keeled; venter coloration white with dark pigments or
spots.
Description of holotype. Adult female, SVL 411 mm; tail long, much slender posteriorly, TaL 109.7 mm, ratio
of TaL/SVL 0.27 or 27% of SVL; head elongate, distinct from neck, HL 14 mm, twice as long as wide (HW 7 mm),
narrower anteriorly and somewhat truncate at the end of the snout tip, larger posteriorly reaching maximum width
at level of posterior parietals, forming a subtrapezoid shape, rather depressed, HD 4.1 mm; IO 3.5 mm, twice
smaller than head width; snout elongate, SnL 3.7 mm; rostral shield moderate, sub-convex crescentic shape,
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A NEW SPECI ES OF WOL F SNAKE FROM SOUTHWEST CAMBODIA
leaving a small median groove for the tongue to protrude, invisible from above, 1.9 mm in width, about twice as
long as depth, 1 mm; internasals subrectangular, 1 mm in width and 1 mm in depth, forming a narrow suture, 1
mm; prefrontal rather large, subpentagonal, posterior sharper point slightly intruding toward anterior suture
between supraoculars and sides of frontal, 1.9 mm in width, same length as depth 1.9 mm, forming a suture 1.5 mm
in length; frontal large, triangular, its length from anterior side to posterior sharper point 3.3 mm, longer than its
width 2.5 mm, measuring the anterior transverse side of maximum frontal contacting with posterior prefrontals;
supraoculars narrow, elongated, subrectangular, 2 mm in length; parietals large, butterfly-like in shape, bordered by
supraoculars, frontal, upper postoculars anteriorly, upper anterior and posterior temporals, paraparietal laterally and
three nuchal scales posteriorly.
FIGURE 2. Holotype (CBC02238) of Lycodon zoosvictoriae sp. nov. in life. Photo: Thy Neang.
Nostrils rather large, located at EN 2 mm, at ND 2.3 mm, valve on upper edge of nostril visible from lateral
view; nasals completely divided by nostrils into two scales about equal in size, anterior nasal in contact with rostral
anteriorly, internasal dorsally, 1
st
SL ventrally, posterior nasal in contact with internasal and prefrontal dorsally,
loreal posteriorly, 1
st
and 2
nd
SL ventrally; loreal large, elongated, subrectangular, in contact with prefrontal
dorsally, preoculars posteriorly, so not in contact with orbit, 2
nd
& 3
rd
SL ventrally, separating from internasal by
broad contact between posterior nasal and prefrontal and from orbit by preoculars; preoculars 1L/2R, one on the
left side rather large, vertically elongated, subrectangular, in contact with prefrontal anteriorly, supraocular
dorsally, orbit posteriorly, lower preocular fused with upper edge of 3
rd
SL, allowing 3
rd
SL to broadly enter orbit, 2
preoculars on the right side, upper vertically elongated, subrectangular, lower much smaller, subrectangular,
separating 3
rd
SL from orbit; orbit moderate, rounded, ED 1.7 mm, almost twice shorter from its distance to the tip
of the snout, pupil vertically elliptic; postocular 2/2, subrectangular (except the lower one on the right side
pentagonal), upper smaller in contact with supraocular dorsally, parietal, anterior temporal posteriorly, lower
pentagonal in contact with 5
th
& 6
th
SL ventrally, both anterior temporal posteriorly; TP 2+2, on the left side anterior
temporals horizontally elongated, hexagonal, larger and longer than posterior temporals, upper one pentagonal,
lower one subrectangular, on the right side upper anterior temporal much longer than lower, in contact with parietal
dorsally, paraparietal posteriorly and upper posterior temporal ventrally, lower anterior temporal subrectangular,
posterior lower temporal pentagonal, anterior half of its size locating beneath posterior part of anterior temporal,
posterior half locating beneath anterior part of paraparietal; paraparietal, large, elongated, irregular shape, locating
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laterally beneath parietal on the right side, subrectangular on the left side. SL 8/8, 1
st
SL in contact with rostral
anteriorly, nasal dorsally, 2
nd
SL in contact with nasal anteriorly, loreal dorsally, 3
rd
SL on the left side fused with
lower preocular, in contact with orbit, on the right side in contact with loreal, lower preocular dorsally, separating
from orbit by lower preocular, 4
th
& 5
th
SL in contact with orbit, 5
th
SL in contact with lower postocular dorsally, 6
th
SL in contact with lower postocular, anterior temporal dorsally, 7
th
SL in contact anterior temporal and posterior
temporal anteriorly, 8
th
SL in contact posterior temporal; mental moderate, triangular; InL 10/10, 1
st
pair in broad
contact with each other, anterior chin shield posteriorly, 1–5
th
in contact with first chin shield, 5
th
InL in contact
with both first and second chin shield; 6
th
largest in contact with second chin shield centrally and anteriormost gular
scale, 7–10
th
in contact with gular scales; two rather large chin shields, in broad contact, the first pair larger than the
second, both forming a distinct suture of 3 mm, second chin shield bordered posteriorly by five gular scales, first
pair of gular scales at central region intruding posterior concave part of second chin shield; first pair of gular scale
followed posteriorly by second pair of gular scales, first preventral, second preventral, and first ventral scale (Fig. 3).
FIGURE 3. Head of the holotype (CBC02238) of Lycodon zoosvictoriae sp. nov. A. lateral view of the right side of head; B.
lateral view of the left side of head; C. ventral view of head (all in preservative); D. dorsal view of head (in life). Photos: Thy
Neang.
FIGURE 4. Dorsal (A) and ventral (B) views of the holotype (CBC02238) of Lycodon zoosvictoriae sp. nov. in preservative.
Photos: Thy Neang.
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A NEW SPECI ES OF WOL F SNAKE FROM SOUTHWEST CAMBODIA
Dentition. On maxilla: nine maxillary teeth, 1–5
th
anterior, increasing in size, the last one the largest fang-like
tooth without venom groove, separated posteriorly by a diastema from two enlarged teeth, anterior similar in size to
the third anterior teeth, posterior larger, again interspaced by a gap which is smaller than that of the diastema,
ending at the two posteriormost teeth similar in size of fourth anterior teeth, locating next to one another; on
palatine of lower jaw: five anterior teeth, increasing in size, followed by a diastema, then one large fang-like tooth,
again interspaced by a small gap, followed by 11subsequent teeth similar in size.
Body scalation: DSN 17, corresponding at level of 14
th
ventral scale; DSM 17, at level of 108
th
ventral scale;
DSV 15, at level of 206
th
ventral scale; VS 213, venter keeled, angulate laterally; SC 85, divided; anal plate single;
scales rhomboid, 1–4
th
scale row smooth, 4
th
scale row indistinctly keeled, 5–8
th
and vertebral scale row weakly
keeled; scale apical pit absent.
Coloration. In life, the specimen possesses a pale tan brown background color on dorsum with darker brown
speckles on scales; large irregular dark brown spots or blotches on anterior one third of dorsum, followed by more
or less 31 distinct, discontinuous, transverse dark brown blotches on posterior two-thirds of dorsum; the first
regular blotches starting at the level of the 95
th
ventral scale; similar indistinct dark brown transverse blotches with
distinct “flying bat” shaped dark brown blotches on three vertebral scales wide between the larger transverse
blotches; blotches on the back transversely extending downwards to the edges of ventral scales; 26 dark brown
transverse blotches on tail, the third anterior blotch encircling the venter; an indistinct vertebral dark brown stripe,
more distinct anteriorly at nuchal region and less distinct posteriorly (washed out in preservative); top and lateral
sides of head with dark brown on scales and pale tan brown speckles between scales, a pale tan brown spot on each
parietal, posterior parietals less darker brown; transverse dark brown nuchal blotches; eye pale tan brown with dark
brown pupil. In preservative, eye creamy gray, pupil milky white; pale tan brown color becomes more creamy
white except on top of head; dark brown color becomes lighter dark brown; anterior and posterior ventral and
subcaudal surface white with irregular dark pigments, mid-ventral surface reddish-orange with irregular dark
pigments (Fig. 4).
Natural history. The holotype of Lycodon zoosvictoriae sp. nov. was found at night at 21:18 hours while it
was slowly moving on a 40 cm diameter tree trunk about 2 m above ground during light rain. The encounter
occurred on 15 June 2013 in submontane forest where most parts of tree trunks and their branches are covered with
mosses (Fig. 5). Hence we carefully assume that L. zoosvictoriae is a nocturnal, arboreal and probably terrestrial
species, which as with other species of Lycodon, may feed on small lizards and frogs. Lizards and frogs known to
inhabit the area include Hemiphyllodactylus sp., Sphenomorphus indicus, Limnonectes kohchangae, Microhyla
heymonsi, Chiromantis samkosensis, Raorchestes cardamonus, R. parvulus, Kurixalus bisacculus and
Rhacophorus rhodopus. When threatened, the reptile protected itself aggressively from danger by repeatedly
striking (Fig. 6) and shaking its tail like a rattlesnake or by coiling and hiding its head inside the coiled body.
Etymology. The specific epithet zoosvictoriae refers to “Zoos Victoria” – the Zoological Parks and Gardens
Board of the southeastern Australian State of Victoria. “Zoos Victoria” has funded the Cardamom Mountains
Research Group in collaboration with the Cambodian Ministry of Environment and Fauna & Flora International
since 2006. Their support has built the capacity of Cambodian researchers and conservationists and greatly
improved understanding of Cambodian herpetofauna. For the vernacular name, we propose Zoos Victoria’s Wolf
Snake (in English).
Comparison. The new species is easily recognized by its unique coloration within the genus Lycodon in
combination with distinct morphometric and meristic characters. Detailed comparisons with its congeners
occurring in Cambodia and neighboring countries between 19° N latitude in the north and the Isthmus of Kra in the
south are provided below and in Table 1.
First, Lycodon zoosvictoriae sp. nov. can be distinguished from L. cardamomensis Daltry & Wüster, 2002
(characters of the latter are given in parenthesis) from Cardamom Mountains in Cambodia and southeastern
Thailand by having TP 2+2 (vs. 2+3); fewer MT 9: 5A & 4P [vs.11–12 (6–7)A/5P]; fewer DSR 17–17–15 (vs.
19–17–15); fewer VS 213 (vs. 215–225) and fewer SC 85 (vs. 92–93); dorsal surface of head dark with pale tan
brown on scale edges and between scales (vs. purely black); irregularly scattered dark brown blotches on anterior
dorsum and 31 relatively regular blotches on posterior body (vs. having 11–13 distinct crossbands on body); 26
blotches on tail (vs. 7 crossbands); color of body blotches dark brown (vs. cream crossbands) and color of body
pale tan brown (vs. black); ventral coloration white with scattered dark pigments (vs. white with encroached dark
blotches).
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TABLE 1. Measured and counted morphological characters of Lycodon zoosvictoriae sp. nov. and characters of relevant Southeast Asian species taken from
literature and comparative material. Data were taken from Anderson (1879); Smith (1943); Taylor (1965); Saint Girons (1972); Daltry & Wüster (2002); Pauwels et
al. (2005); Grismer et al. (2008); Vogel et al. (2009, 2012); Das (2010); Vogel & David (2010); and Zhang et al. (2011). Abbreviations are defined in the methods
and materials and additional abbreviations are: maxillary teeth (MT), anterior=A, posterior=P; preocular (PreOc), present=1 or 2, absent=0, left (L) or right (R);
postocular (PoC); loreal (Lo), present=1, absent=0; lo-internasal in contact=1, lo-internasal not in contact=0; lo-orbit in contact=1, lo-orbit not in contact=0; anal
plate (AP): single=1, divided=2; number of blotches or bands, absent= –; nuchal band, absent= –; rare or exceptional characters are given in parenthesis.
Characters L. zoosvictoriae
sp. nov.
L. capucinus L. cardamomensis L. davidi L. fasciatus
SVL 411 258–684 436–709 308 358–717
TaL 109.7 73–132 109–187 81.5 95–201
TaL/SVL 0.27 0.19–0.28 0.25–0.28 0.26 0.23–0.29
TP 2+2/2+2 2+3/2+3 2+3/2+3 2+2/2+3 2+2
MT (5A/4P) ? 6–7A/4–5P 6A/5P ?
SL 8/8 9–10/9–10 8/8 8/8 8/8
SL-orbit in contact (3–5th)L/(4–5
th
)R 3–5th 3–5
th
3–5th 3–5th
PreOc 1L/2R 1 1 1 1
PoC 2/2 2/2 2–3/2–3 2/2 2/2
Lo 1 1 1 1 1
Lo-orbit in contact 0 0 0 0 1
Lo-internasal in contact 0 1 0 0 0
InL 10/10 9–10/9–10 10/10 10/10 9/9 (8,10)
DSN 17 17 19 17 17
DSM 17 17 17 17 17
DSV 15 15 15 15 15
VS 213 182–211 215–225 224 182–225
SC 85 59–74 92–93 99 65–94
AP 1 2 1 1 1
Dorsal surface of head Dark brown Dark Black Olive-brown Dark
N
o
of transverse blotches or
bands on body
31 Reticulate 11–13 84 19–49
N
o
of transverse blotches or
crossbands on tail
26 – 7 38 7–21
Color of blotches or
crossbands
Dark brown Mixed reticulate Whitish Pale tan brown Whitish grey
Color of body Light tan brown Mixed brownish grey Black Brownish black Blackish
Dorsal scales Weakly keeled Weakly keeled Weakly keeled Weekly keeled Weakly keeled
Venter coloration White with dark pigments Dirty white White withencroached
dark blotches
White with dark blotches White with dark transverse
blotches
……continued on the next page
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A NEW SPECI ES OF WOL F SNAKE FROM SOUTHWEST CAMBODIA
TABLE 1. (Continued)
Characters L. futsingensis L. laoensis L. paucifasciatus L. ruhstrati abditus L. septentrionalis L. subcinctus
SVL up to 663 333.4–391 536–640 320–761 354.4–990 209–750
TaL up to 187 75–107 137 84–203 90–190 173
TaL/SVL 0.2–0.23 0.21–0.30 0.20 0.21–0.24 0.19–0.25 0.23–0.30
TP 2(1)+3(2) 2+3(4) 2+3 2(1)+3(2) 2+2 (3) 1+2/1+2
MT 7–8A/5–7P ? 6A/5–6P 6–7A/5–6P 4A/3P ?
SL 8(7) 9–10/9–10 8 8 8/8 8/8
SL-orbit in contact 3(2–4) –5
th
(4–6) 3–5th 3–5
th
3–5
th
3–5th 3(4)–5th
PreOc 1 1 1 1 1 0
PoC 2 (3) 2(3)/2(3) 2 2 2/2 2(3)/2(3)
Lo 1 1 1 1 1 1
Lo-orbit in contact 0 0 0 0 1 1
Lo-internasal in contact 0 0(1) 0 0 0 0
InL 10(9,11) 10/10 10 10 (9,11) 7–8/7–8 8–9/8–9
DSN 17 17 19 17 17 17
DSM 17(16) 17 19–17 17 17 17
DSV 15 15 15 15 15 15
VS 193–208 163–192 221–222 214–229 202–217 192–230
SC 72–85 60–76 92 90–103 80–92 60–91
AP 1 2 1 1 1 2 (1)
Dorsal surface of head Grayish brown Dark Dark Dark Light Dark Largely white
N
o
of transverse blotches or bands
on body
19–33 19–36 14–25 19–43 23–35 9–15
N
o
of transverse blotches or
crossbands on tail
9–18 11–23 8–11 10–23 11–17 –
Color of blotches or crossbands Brownish-speckled
white
Yellowish white Dirty cream Whitish White Whitish
Color of body Dark brown Black Blackish brown Blackish brown Black Black
Dorsal scales Smooth Smooth Distinctly keeled Distinctly keeled Weakly keeled Weakly keeled
Venter coloration Cream, speckled
posteriorly
White Cream to beige-
brown
Cream White Grayish
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FIGURE 5. Submontane habitat of the holotype (CBC02238) of Lycodon zoosvictoriae sp. nov. Photo: Thy Neang.
FIGURE 6. Striking position of (CBC02238) Lycodon zoosvictoriae sp. nov. during defensive behavior. Photo: Thy Neang.
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Lycodon zoosvictoriae sp. nov. differs from L. capucinus (Boie, 1827) which widely occurs in Southeast Asia
by having TP 2+2 (vs. 2+3); SL 8 (vs. 9–10); greater number of SC 85 (vs. 59–74); dorsal surface of head dark
brown with pale tan brown on scale edges and between scales (vs. purely dark); irregularly scattered dark brown
blotches on anterior dorsum and 31 blotches on posterior body (vs. reticulate with pale lines); 26 blotches on tail
(vs. absent); color of body blotches dark brown (vs. mixed reticulate); color of body pale tan brown (vs. mixed
brownish-grey); ventral coloration white with dark pigments (vs. dirty white).
Morphologically, the new species is separated from L. davidi Vogel, Nguyen, Kingsada & Ziegler, 2012 from
Laos by having fewer MT 9 [5A & 4P][vs. 11 (6A & 5P)]; fewer VS 213 (vs. 224); fewer SC 85 (vs. 99); dorsal
surface of head dark brown with light tan brown on scale edges and between scales (vs. olive-brown); irregularly
scattered dark brown blotches on anterior part and 31 blotches on posterior part of body (vs. 84 crossbands on
body); 26 blotches on tail (vs. 38 crossbands on tail); color of body blotches dark brown (vs. pale tan brown); color
of body pale tan brown (vs. brownish black); ventral coloration white with dark pigments (vs. white with dark
blotches).
In having a ratio of TaL/SVL 0.27, TP 2+2, SL 8, loreal present and not in contact with internasals, InL 10,
DSR 17–17–15, VS 213, SC 85, and dorsal scale weakly keeled, L. zoosvictoriae sp. nov. mostly resembles L.
fasciatus (Anderson, 1879) from Southwest China, India, Myanmar, northern Thailand, northern Laos and
Vietnam, although its coloration pattern is completely different. The new species distinctly differs from L. fasciatus
by having its loreal not in contact with orbit (vs. Lo-orbit in contact); dorsal surface of head dark brown with pale
tan brown on scale edges and between scales (vs. purely dark brown); color of body blotches dark brown (vs.
whitish-grey crossbands); color of body pale tan brown (vs. blackish-brown), venter coloration white with dark
pigments (vs. white with dark transverse blotches, see Vogel & David 2010).
The new species differs from L. futsingensis (Pope, 1928) recorded from Southeast China and North to Central
Vietnam by having fewer MT 9 [5A & 4P] [vs. 12–15 (7–8A & 5–7P)]; a distinctly higher TaL/SVL of 0.27 (vs.
0.2–0.23); 26 blotches on tail (vs. 9–18 crossbands ); color of body blotches dark brown (vs. brownish-speckled
white); color of body pale tan brown (vs. dark brown); dorsal scales weakly keeled (vs. smooth).
Lycodon zoosvictoriae sp. nov. is distinguished from L. laoensis Günther, 1864 which occurs in India,
Thailand, Laos, Vietnam, Cambodia, West Malaysia and China by having TP 2+2 (vs. 2+3); SL 8 (vs. 9–10);
higher VS 213 (vs. 163–192); higher SC 85 (vs. 60–76); dorsal surface of head dark brown with pale tan brown on
scale edges and between scales (vs. purely dark); higher number of blotches 26 (vs. 11–23); color of body blotches
dark brown (vs. yellowish-white crossbands); color of body pale tan brown (vs. black); dorsal scales weakly keeled
(vs. smooth ); ventral coloration white with dark pigments (vs. white).
Lycodon zoosvictoriae sp. nov. can easily be distinguished from L. paucifasciatus (Rendahl, 1943) from
Central Vietnam by its lower number of dorsal scales at the neck 17 (vs. 19); only weakly keeled dorsal scales (vs.
distinctly keeled); its proportionally longer tail, TaL/SVL 0.27 (vs.0.20); having 2 posterior temporals (vs. 3);
fewer MT 9 (vs. 11–12); more blotches on body 31 (vs. 14–25) and tail 26 (vs. 8–11); color of body blotches dark
brown (vs. mottled cream) and color of body pale tan brown (vs. blackish brown). Since L. ruhstrati (Fischer,
1886) is a polytypic species with its nominotypic subspecies restricted to Taiwan, we confine our comparison to L.
ruhstrati abditus (Vogel et al. 2009) from which Lycodon zoosvictoriae sp. nov. can be differentiated by its weakly
keeled dorsal scales (vs. distinctly keeled); its proportionally shorter tail, TaL/SVL 0.27 (vs. 0.21–0.24); fewer MT
9 (vs. 11–13); slightly lower number of subcaudals 85 (vs. 90–103); more blotches on tail 26 (vs. 11–23); color of
body blotches dark brown (vs. whitish) and color of body pale tan brown (vs. blackish brown).
The new species differs from Lycodon septentrionalis (Günther, 1875) which occurs from India through
Myanmar, Thailand, Cambodia, Laos, Vietnam to China by having a longer tail, TaL/SVL 0.27 (vs. 0.19–0.25);
loreal not in contact with orbit (vs. Lo-orbit in contact ); irregularly scattered dark brown blotches on anterior part
and 31 blotches on posterior part of body (vs. 23–35 crossbands on body); 26 blotches on tail (vs. 11–17); color of
body blotches dark brown (vs. white bands) and color of body pale tan brown (vs. black); ventral coloration white
with dark pigments (vs. white).
Lastly, Lycodon zoosvictoriae sp. nov. is distinguished from Lycodon subcinctus Boie, 1827 which occurs
from Brunei, West Malaysia, Indonesia, India, Philippines, China, Vietnam, Laos, Thailand to Cambodia by having
TP 2+2 (vs. 1+2); preoculars 1L/2R (vs. preocular absent); loreal not in contact with orbit (vs. Lo-orbit in contact);
InL 10 (vs. 8–9); dorsal surface of head dark brown with pale tan brown on scale edges and between scales (vs.
largely white); irregularly scattered dark brown blotches on anterior body and 31 on posterior body (vs. 9–15
NEANG ET AL.
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Zootaxa 3814 (1) © 2014 Magnolia Press
crossbands on body); color of body blotches dark brown (vs. whitish); color of body pale tan brown (vs. black);
ventral color pattern white with dark pigments (vs. grayish).
Discussion
Morphologically, wolf snakes of the genus Lycodon were traditionally diagnosed by their inwardly bent maxillary
bone with 3–6 anterior teeth, and, following a diastema, 7–15 posterior teeth (Smith, 1943; Taylor, 1965). Although
the diagnostic value of teeth is still questionable (Siler et al. 2013; Guo et al. 2013), many recently described
species, including a Cambodian specimen (CBC00678, see appendix 1), indicate that this is not always correct as
the number of posterior teeth can be as low as four or five (Lanza 1999; Daltry & Wüster 2002; Vogel & David
2010; Vogel et al. 2012).
The new species appears to be a cryptic, arboreal and probably terrestrial snake. It likely benefits by its dull
pale tan brown coloration which may act as camouflage from predation on similarly colored tree bark which are
largely covered with moss at the type locality due to the continually wet conditions that prevail there. As such, it is
unsurprising that the species has eluded the intensive herpetological sampling effort in Cambodia over the last
decade. This peculiar coloration renders Lycodon zoosvictoriae sp. nov. highly distinct from its congeners and
permits its specific recognition on morphological differences alone. Nevertheless, since the species is presently
known only from a single specimen, future molecular studies would be of great interest. Additional samples are
needed to evaluate genotype variation within the species population and its appropriate position in existing
phylogenetic trees (Siler et al. 2013). Further, the fusion of the preocular on the left side causes the third supralabial
to contact the orbit in the existing specimen and this character may prove to be variable when additional specimens
are found. Given its unique coloration, submontane habitat and altitudinal separation (1,284 m a.s.l.) from other
wolf snakes of Lycodon genus in the region, the species could prove to be endemic to the Cardamom Mountains of
Cambodia and probably Thailand.
The discovery of Lycodon zoosvictoriae sp. nov. from Cambodia raises the total number of Lycodon currently
recognized to 48 species and the number described in the last two decades to 15 species (see introduction). It also
increases the number of snake species described from Cambodia in the last 12 years to eight (Daltry & Wüster
2002; David et al. 2008a,b; Malhotra et al. 2011; Murphy et al. 2012; Neang et al. 2012). Our latest finding is an
additional proof of the critical importance of the herpetofauna of Cambodia’s Cardamom Mountains for IUCN Red
List assessments and conservation oriented research. On the basis of our single specimen description, we suggest
inclusion of this species in the IUCN Red List of Threatened Species as Data Deficient (DD) until additional data
are available and thoroughly assessed.
Acknowledgements
The authors are indebted to His Excellency Chay Samith, Delegate of the Royal Government of Cambodia in
charge as General Director of General Department of Administration for Nature Conservation & Protection,
Ministry of Environment, for granting research permission No. 0116 dated 5
th
May 2013. Grateful thanks go to
Patrick David, Gernot Vogel, Lee Grismer and Bryan Stuart for providing literature and to Sovannarun Sim for
producing Figure 1. We also thank our assistants for their very hard work during the field survey. Our research was
made possible by grants from the Zoological Parks and Gardens Board of Victoria (Australia), Darwin Initiative
(DEFRA, UK: EIDPO028), and the John D. and Catherine D. MacArthur Foundation (US: 09-92411-000-GSS).
We also thank Patrick David and one anonymous reviewer for their constructive reviews of the manuscript.
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APPENDIX 1. Comparative material.
Lycodon capucinus: ZFMK 92630 and ZFMK 90370, KPNP, Siem Reap province, Cambodia; ZFMK 54884, Phnom Penh,
Cambodia; ZFMK 84097, W-Flores: Labuan, Indonesia; ZFMK 86832, Da Nang: Son Tra, Vietnam; ZFMK 88296, Son
Tra, Vietnam.
Lycodon cardamomensis: CBC02206, Phnom Samkos Wildlife Sanctuary, southwest Cambodia.
Lycodon fasciatus: Cao Bang: Ha Lang, An Lac, Vietnam; ZFMK 86448-86450, Quang Binh, Phong Nha-Ke Bang National
Park, Vietnam.
Lycodon laoensis: CBC00399, Phnom Samkos Wildlife Sanctuary; ZFMK 54885-54887, Phnom Penh Capital, Cambodia.
Lycodon septentrionalis: CBC00678, Virachey National Park, northeast Cambodia.
Lycodon subcinctus: ZFMK 92631, ZFMK 90371-90372, Kulen Promtep National Park (KPNP), Siem Reap province,
Cambodia.
... The Cardamom Mountains are relatively isolated from other significant mountain blocks in the region ) and contain the highest, wettest and largest tract of evergreen forest remaining in Cambodia (Stuart & Emmett 2006). The Cardamom Mountains on both sides of the international border have been the subject of historical (e.g., Mouhot 1864;Smith & Kloss 1915;Smith 1917;Saint Girons 1972;reviewed in Stuart & Emmett 2006) and especially of contemporary herpetological surveys (e.g., Daltry & Wüster 2002;Ohler et al. 2002;Stuart & Emmett 2006;Grismer et al. 2007aGrismer et al. ,b, 2008Grismer et al. , 2011Neang et al. 2011Neang et al. , 2012Neang et al. , 2014Murdoch et al. 2019). However, to date, the sole record of Leptobrachella from the entirety of the Cardamom Mountains has been a single preserved metamorph of an unknown species that was collected in early 2001 at Phnom Aural Wildlife Sanctuary, Kampong Speu Province, Cambodia (Ohler et al. 2002). ...
... The specific epithet is a patronym honoring Mr. Thy Neang of Wild Earth Allies, Phnom Penh, Cambodia, collector of the new species, and cherished friend and colleague of the authors. As Cambodia's first national with specific expertise in taxonomy of amphibians and reptiles, Thy has made tremendous contributions to the herpetology of his country (e.g., Neang & Holden 2008;Neang & Hun 2013;Neang et al. 2011Neang et al. , 2012Neang et al. , 2014Neang et al. , 2018aNeang et al. ,b, 2020. The first and second authors have had the privilege of working with Thy since 2003 and 2007, respectively, including the collaborative discoveries and descriptions of the other two Leptobrachella species known from Cambodia (Rowley et al. 2010(Rowley et al. , 2015aFig. ...
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... Specimen identification. For taxonomic identification, we referred to Bourret (1936Bourret ( , 1942, Smith (1943), Liu (1950), Taylor (1962), Manthey & Grossmann (1997), Dubois & Ohler (1998), Lathrop et al. (1998), Ziegler & Köhler (2001), Ziegler (2002), Leviton et al. (2003), Bain et al. (2004), Ohler (2003), Ohler & Delorme (2006), , Fei et al. (2008), Stuart & Heatwole (2008), Fei et al. (2009), Vogel et al. (2009), Inger & Stuart (2010), Nguyen et al. (2011), Chen et al. (2013), Hecht et al. (2013), Luu et al. (2013), Nemes et al. (2013), Ostroshabov et al. (2013), Le et al. (2014a, b), Neang et al. (2014), Nguyen et al. (2016a, b), Pham et al. (2016, Nguyen et al. (2018), Janssen et al. (2019), and Yu et al. (2019). Species names follow Nguyen et al. (2009), Frost (2020, and Uetz et al. (2020). ...
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... A tissue sample was collected from the holotype of Lycodon pictus (IEBR 4166) then extracted following formalin protocol developed by Friedman &Desalle (2008) using GTE andNeang et al., 2014). From Vietnam, sixteen species of Lycodon have been reported to date, comprising L. capucinus (Boie, 1827), L. cardamomensis (Daltry & Wü ster, 2002), L. davisonii (Blanford, 1878), L. fasciatus (Anderson, 1879), L. fl avozonatus (Pope, 1928), L. futsingensis (Pope, 1928), L. laoensis Gü nther, 1864, L. meridionalis (Bourret, 1935), L. namdongensis Luu, Ziegler, Ha, Le & Hoang, 2019, L. paucifasciatus Rendahl in Smith, 1943, L. pictus Janssen, Pham, Ngo, Le, Nguyen & Ziegler, 2019, L. rosozonatus (Hu & Zhao, 1972[1975), L. rufozonatus Cantor, 1842, L. ruhstrati abditus Vogel, David, Pauwels, Sumontha, Norval, Hendrix, Vu & Ziegler, 2009, L. septentrionalis (Gü nther, 1875 and L. subcinctus Boie, 1827(Uetz et al., 2020. ...
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The Painted Wolf Snake, Lycodon pictus, was recently described based on the type series from Trung Khanh and Ha Lang districts in Cao Bang Province, northern Vietnam. Herein, we report new findings and a range extension of the recently described species. One individual was collected close by the type locality, in Ha Lang District, Cao Bang Province, Vietnam and another one in Nonggang National Nature Reserve, Longzhou County, Guangxi Zhuang Autonomous Region, China, approximately 60 km apart from the type locality. Both specimens generally accorded with the morphological diagnosis of L. pictus provided in the original description. Molecular analyses supported the morphological findings: the newly collected specimens were approximately 0.3-0.9% (cyt b) genetically divergent from those of the type series. Molecular analyses also revealed that the holotype of L. pictus, for which no sequences were available so far, showed, based on the results of a formalin protocol, about 0.9-1.3% differentiation from the remaining type series, 0.5% from the new record from Cao Bang and 0.9% from the new record from China. Based on the morphological and molecular fi ndings, we herein present the fi rst country record of L. pictus from China, with a detailed morphological description of the specimen from Nonggang National Nature Reserve and slightly extend the original diagnosis of the species.
... Morphological characters of recognized Lycodon species were obtained from examination of museum specimens (see Appendix 1) and from the following references: Günther (1864), Günther (1875), Blanford (1878), Boulenger (1893), Boulenger (1900), Wall (1906), Stejneger (1907), Griffin (1909), Taylor (1922), Pope (1928a, b), Smith (1943), Taylor (1950), Leviton (1965), Hu et al. (1975), Zhao (1981), Ota and Ross (1994), Manthey and Grossmann (1997), Captain (1999), Lanza (1999), Slowinski et al. (2001), Daltry and Wüster (2002), Gaulke (2002), Gaulke et al (2003), Jackson and Fritts (2004), Vijayakumar and David (2005), Zhao (2006), Mukherjee and Bhupathy (2007), Mistry et al. (2007), , Bahuguna and Bhuta (2010), Vogel and David (2010), Vogel and Luo (2011), Zhang et al. (2011), Vogel et al. (2012), Guo et al. (2013), Vogel and Harikrishnan (2013), Grismer et al. (2014), Lei et al. (2014), Neang et al. (2014), Zhang et al. (2015), Gawor et al. (2016), Do et al. (2017), Wostl et al. (2017), Ganesh and Vogel (2018), Luu et al. (2018), Melvinselvan et al. (2018), Janssen et al. (2019), Luu et al. (2019), and Vogel and David (2019). Data shown in Table 1 was modified based on Janssen et al. (2019), with distinguishing characters marked in bold. ...
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