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

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DOI: 10.11646/zootaxa.3814.1.3
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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 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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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
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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.
  • ... Comparisons were mainly based on the data provided by Boulenger (1893), , Smith (1943), Leviton (1965), Ota and Ross (1994), Manthey and Grossmann (1997), Lanza (1999), Vogel et al. (2009), Vogel and David (2010) and Neang et al. (2014), with additional references provided in the comparisons and legends of the tables. Additionally, studied specimens are listed in the Appendix 1. ...
    ... separated) and cream body bands (vs. yellow) (Grismer et al. 2014, Neang et al. 2014. ...
    ... In addition, the new species has cream body bands (vs. pinkish red) (Hu et al. 1975, Neang et al. 2014. ...
    Article
    Full-text available
    A new species of the genus Lycodon is described from Cao Bang Province, Vietnam, based on three individuals with distinct differences in morphology and molecular data. The new species is differentiated from its congeners by a combination of the following characters: dorsal scales in 17-17-15 rows, smooth throughout; supralabials usually eight (rarely nine); infralabials ten; one elongated loreal on each side, in contact with the eye; precloacal plate single; ventral scales 212-218 (plus one or two preventral scales); subcaudals 90 or 91; maxillary teeth 13 or 14; dorsal surface of body with 28 or 29 light body bands; dorsal surface of tail with 13 cream bands, forming a distinct blotch in the vertebral region. Based on phy-A peer-reviewed open-access journal Helen Y. Janssen et al. / ZooKeys 875: 1-29 (2019) 2 logenetic analyses of mitochondrial cytochrome b sequence data, the new species is recovered as the sister species to a clade containing L. multizonatus and L. liuchengchaoi with strong support from the Bayesian analysis. The new species is at least 7.5% divergent from other species within this clade in uncorrected pairwise distance calculated using a fragment of more than 1000 bp of the mitochondrial cytochrome b. This discovery increases the number of Lycodon species known from Vietnam to 16.
  • ... The genus Lycodon Boie is one of the most diverse genera of colubrid snakes, with 59 currently recognized species (Figueroa et al. 2016;Luu et al. 2018;Uetz et al. 2018). The members of this genus have a wide distribution range from eastern Iran to southern China and Japan, southward to the Philippines and the Indo-Australian Archipelago (Lanza 1999;Silver et al. 2013;Neang et al. 2014). Five species of Lycodon have been recently described from Southeast Asian countries, namely L. davidi Vogel, Nguyen, Kingsada & Ziegler, 2012 from Boulenger 1893;Smith 1943;Deuve 1970;Hu et al. 1975;Ziegler et al. 2007;Nguyen et al. 2009;Vogel & David 2010;Geissler et al. 2011;Orlov et al. 2011;Neang et al. 2014;Luu et al. 2013a,b;Jestrzemski et al. 2013;Phan et al. 2014Grismer et al. 2014Ziegler et al. 2014;Gawor et al. 2016;Ngo & Duong 2016;Nguyen et al. 2016;Do et al. 2016;Vassilieva et al. 2016;Pham et al. 2017;Do et al. 2017;Luu et al. 2018). ...
    ... The members of this genus have a wide distribution range from eastern Iran to southern China and Japan, southward to the Philippines and the Indo-Australian Archipelago (Lanza 1999;Silver et al. 2013;Neang et al. 2014). Five species of Lycodon have been recently described from Southeast Asian countries, namely L. davidi Vogel, Nguyen, Kingsada & Ziegler, 2012 from Boulenger 1893;Smith 1943;Deuve 1970;Hu et al. 1975;Ziegler et al. 2007;Nguyen et al. 2009;Vogel & David 2010;Geissler et al. 2011;Orlov et al. 2011;Neang et al. 2014;Luu et al. 2013a,b;Jestrzemski et al. 2013;Phan et al. 2014Grismer et al. 2014Ziegler et al. 2014;Gawor et al. 2016;Ngo & Duong 2016;Nguyen et al. 2016;Do et al. 2016;Vassilieva et al. 2016;Pham et al. 2017;Do et al. 2017;Luu et al. 2018). 1-maximum snout-vent length (in mm); 2-maximum tail length; 3-maximum of total length; 4-number of supralabials; 5-number of infralabials; 6-loreal touching orbit; 7-number of preoculars; 8-number of postoculars; 9-number of loreals; 10-dorsal scale row fomula; 11-number of ventral scales; 12-number of subcaudals; 13-number of cloacal (1 or 2); 14-head pattern; 15-belly banded; 16-dorsal body pattern; 17-number of body bands/blotches; 18-number of tail bands/blotches; 19-color of light body bands/blotches; 20-ground coloration on dorsum; 21-dorsal scales keeled/smooth; 22-belly pattern; 23-vertebral scale row enlarged; 24-first body band position (at VEN); 25-first body band width; 26-ventral tail banded; 27-maxillary teeth, anterior=A, posterior=P; 28-temporals (A+P); 29-nuchal band; 30-distribution. ...
    ... The new species has a loreal which is not in contact with the orbit and thus differs from the following species of the Lycodon fasciatus group which have the loreal entering the orbit: L. butleri Boulenger; L. fasciatus; L. subcinctus. In addition, the new species differs from L. butleri by having fewer dorsal bands (23 versus 28-37), fewer tail bands (14 versus 17-23), belly pattern mostly cream (versus banded & spotted) (Grismer et al. 2014); from L. fasciatus by having dorsal scale rows being completely smooth (versus weakly keeled), dorsal head pattern whitish grey (versus dark), first body band starting at ventral scale 25 (versus 5-18), first body band covered by one vertebral scale (versus 3-12), ventral pattern mostly cream (versus banded) (Vogel & David 2010;Neang et al. 2014); from L. subcinctus by the presence of a preocular scale (versus absent), having dorsal scale rows being completely smooth (versus weakly keeled), single cloacal (versus divided), and more body bands (23 versus 9-15) (Neang et al. 2014) (for more details see Table 1). ...
    Article
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    We describe a new species of the genus Lycodon based on an adult male specimen from Thanh Hoa Province, Vietnam. Lycodon namdongensis sp. nov. is differentiated from its congeners by a combination of the following characters: dorsal scales in 17−17−15 rows, all smooth; supralabials 8; infralabials 10; one large loreal on each side, separated from the eye; cloacal single; ventral scales 218 (plus two preventral scales); subcaudals 85, paired; a total length of 723 mm; tail / total length ratio 0.205; dorsal head pattern whitish grey; dorsal surface of body with 23 narrow greyish cream bands; dorsal surface of tail with 14 cream bands; ventral surface of body mostly cream with a few small dark spots posteriorly; ventral surface of tail dark grey. Based on molecular comparisons, Lycodon namdongensis sp. nov. is recovered as a sister species to L. futsingensis with strong support values from all three analyses. The new species is at least 6.5% genetically divergent from other species within this clade as shown by a fragment of more than 1000 pbs of the mitochondrial cytochrome b. This discovery increases the number of Lycodon species known from Vietnam to 15.
  • ... Keel (keeled dorsal scale rows); PreVEN (preventral scales); VEN notched (present or absent); VEN keeled (present or absent); SC (subcaudal scales); numbers of pattern-units (like crossbars or vertebral blotches) are provided as number on body + number on tail. For comparisons, we referred to the data provided by Boulenger (1893), Smith (1943), Orlov & Ryabov (2004), Neang et al. (2014), and Grismer et al. (2014); studied specimens are listed in the Appendix. Museum abbreviations are as follows: CAS-California Academy of Sciences; GP-Specimens in the collection of Peng Guo; LSUMZ-The Louisiana State University Museum of Natural Science; MNHN-Muséum National d'Histoire Naturelle, Paris, France; VNUF-Vietnam National University of Forestry. ...
    ... Figs 2-5, Table 3 Holotype (Neang et al., 2014). From the remaining species in the fasciatus group, the new species differs as follows: from L. butleri Boulenger by having more ventral scale rows (241 versus 220-227), dorsal pattern blotched (versus banded), and belly pattern Distribution: Lycodon banksi sp. ...
    ... nov. from closely related species and the remaining Lycodon species recorded from Laos (data obtained from Boulenger, 1893;Smith, 1943;Taylor, 1965;Deuve, 1970;Orlov & Ryabov, 2004;Vogel et al., 2009Vogel et al., , 2012Siler et al., 2013;Luu et al., 2013;Teynié et al., 2014;Neang et al., 2014). ...
    Article
    We describe a new species of the genus Lycodon Boie based on an adult male specimen from Khammouane Province, central Laos. Lycodon banksi sp. nov. is distinguished from its congeners by a combination of the following characters: (1) dorsal scales in 17-17-15 rows, dorsal scales on the anterior 2/3 of the body length smooth, the six central dorsal scale rows of the posterior 1/3 of the body length feebly keeled; (2) supralabials 8; (3) infralabials 10; (4) loreal in contact with the eye; (5) cloacal single; (6) ventral scales 241; (7) dorsal surface of body with 87 greyish yellow blotches; (8) ventral surface of body and tail uniformly grey cream. Based on the molecular comparison, Lycodon banksi sp. nov. is placed in a clade with other species previously considered to be members of the genus Dinodon. The new species is at least 9% genetically divergent from other species within this clade as shown by a fragment of the mitochondrial cytochrome b. This discovery increases the number of Lycodon species known from Laos to eleven. © 2018 Museum d'Histoire Naturelle de Geneve. All rights reserved.
  • ... Keel (keeled dorsal scale rows); PreVEN (preventral scales); VEN notched (present or absent); VEN keeled (present or absent); SC (subcaudal scales); numbers of pattern-units (like crossbars or vertebral blotches) are provided as number on body + number on tail. For comparisons, we referred to the data provided by Boulenger (1893), Smith (1943), Orlov & Ryabov (2004), Neang et al. (2014), and Grismer et al. (2014); studied specimens are listed in the Appendix. Museum abbreviations are as follows: CAS-California Academy of Sciences; GP-Specimens in the collection of Peng Guo; LSUMZ-The Louisiana State University Museum of Natural Science; MNHN-Muséum National d'Histoire Naturelle, Paris, France; VNUF-Vietnam National University of Forestry. ...
    ... Figs 2-5, Table 3 Holotype (Neang et al., 2014). From the remaining species in the fasciatus group, the new species differs as follows: from L. butleri Boulenger by having more ventral scale rows (241 versus 220-227), dorsal pattern blotched (versus banded), and belly pattern Distribution: Lycodon banksi sp. ...
    ... nov. from closely related species and the remaining Lycodon species recorded from Laos (data obtained from Boulenger, 1893;Smith, 1943;Taylor, 1965;Deuve, 1970;Orlov & Ryabov, 2004;Vogel et al., 2009Vogel et al., , 2012Siler et al., 2013;Luu et al., 2013;Teynié et al., 2014;Neang et al., 2014). ...
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  • ... Post-conflict herpetological field resurgence mainly concentrated on the Cardamom Mountains of southwest Cambodia (Grismer et al., 2008;Neang et al., 2014;Ohler et al., 2002;Stuart & Emmett, 2006) and north-eastern Cambodia Stuart et al., , 2010. Some sporadic surveys were also carried out along the Mekong River (Bezuijen et al., 2009), Tonle Sap Lake (Brook et al., 2007a(Brook et al., , 2007(Brook et al., , 2009) and around Siem Reap (Hartmann et al., 2013). ...
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    Conservation International, in cooperation with the Forestry Administration of Cambodia, undertook biodiversity surveys of Prey Lang between June 2014 and February 2015. The objectives of the survey were to determine the biodiversity values and conservation priorities within Prey Lang, identify threats, and produce recommendations for the alleviation of these threats. The survey covered vegetation, mammals (including a specific study of bats), birds, amphibians and reptiles. The survey teams recorded evidence of the presence of species, using camera trapping, mist netting, harp traps, direct observation and captures. Sign surveys and interviews were also undertaken. The field surveys aimed to build on existing research undertaken in the area, and the results sections present the aggregation of both primary and secondary data
  • ... With at least 51 currently accepted species, Lycodon Boie in Fitzinger is one of the most diverse Asian snake genera Neang et al. 2014;Uetz et al. 2019). It occurs from central Asia and eastern Iran to southern and eastern China and Japan (Ryukyu Islands), southwards, across the entire Indo-Chinese Peninsula, and into the Indo-Australian Archipelago and the Philippines (Lanza 1999). ...
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    A new species of the genus Lycodon Boie in Fitzinger is described on the basis of three specimens collected on the Khorat Plateau, Nakhon Ratchasima Province, Thailand. Lycodon gibsonae spec. nov. is superficially similar to L. fasciatus (Anderson), it differs by its much larger size, a higher number of ventral and subcaudal scales, and a lower number of bands on the body, the last three characters presenting no overlap with those of L. fasciatus sensu stricto. The new species seems to be endemic to the Khorat Plateau.
  • ... However, with the political reconciliation and integration of the late 1990's, field work was permitted to recommence. In the past two decades, numerous surveys have been made in the Cardamoms and adjacent upland areas of Cambodia resulting in the discovery of no less than 12 new endemic species of amphibians and reptiles (Ohler et al. 2002;Daltry & Wüster 2002;Grismer et al. 2007bGrismer et al. , 2010Wood et al 2010;Neang et al. 2011;Mahony 2011;Neang et al. 2012;Geissler et al. 2014;Neang et al. 2014). The growing number of species from these mountains highlights the need for continued field research and conservation in these unique areas. ...
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    An integrative taxonomic analysis using color pattern, morphology, and 1449 base pairs of the ND2 mitochondrial gene and its five flanking tRNAs demonstrated that eight species-level lineages occur within the Cyrtodactylus intermedius complex (Cyrtodactylus intermedius sensu stricto, C. phuquocensis and related populations) of the Cardamom mountains and associated highlands that have a sequence divergence ranging 3.4–8.9%. Additionally, each lineage is discretely diagnosable from one another based on morphology and color pattern and most occur in specific geographic regions (upland areas, karst formations or islands) that prevent or greatly restrict interpopulation gene flow. Six of these lineages were masquerading under the nomen C. intermedius and are described as the following: Cyrtodactylus auralensis sp. nov. endemic to Phnom Aural, the highest mountain in Cambodia; C. bokorensis sp. nov. endemic to the Bokor Plateau, Cambodia; C. cardamomensis sp. nov. from the main block of the Cardamom mountains; C. thylacodactylus sp. nov. endemic to Phnom Dalai the northernmost peak of the Cardamom mountains; C. laangensis sp. nov. endemic to the Phnom Laang karst formation, Cambodia; and C. septimontium sp. nov. from the Bảy Núi Hills of southwest Vietnam.
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    The Asiatic Wolf Snakes genus Lycodon Boie, 1826 as currently recognized, consists of about 48 species of which 15 have been formally described in the last 15 years (Neang et al. 2014). These snakes have for most of the last 188 years been treated as being within a single genus, namely Lycodon Boie, 1826. Other authors have proposed generic names for given species or species groups, with most if not all invariably being synonymised with Lycodon. By way of example, the most recent phylogenetic revisions by Siler et al. (2013) and Guo et al. (2013) merged the long recognized and used genus Dinodon Duméril, 1853 within a greater Lycodon. This is a position maintained by most other authors including for example Neang et al. (2014). However a comparison of where the line of divergence is drawn to define a genus between snakes within Lycodon sensu lato and other snake genera as currently recognized shows that by any reasonable interpretation Lycodon should be split to be consistent with the majority of other genera. This view is supported by molecular studies of Siler et al. (2013), Guo et al. (2013) and Grismer et al. (2014) if lined up against the comparative order-wide supermatrix constructed by Pyron et al. (2013), which clearly shows Lycodon as paraphyletic. As a result of the evidence within these and other studies, including numerous taxonomic ones based on morphology, Lycodon as currently recognized is split into seven genera, for which names are available for five. The other two genus groups, Snakebustersus gen. nov. and Apollopierson gen. nov. are formally named and defined according to the rules of the Zoological Code (Ride et al. 1999). Within various genera five new subgenera are also formally named for the first time and another genus Cercaspis Wagler, 1830 is resurrected and treated as being a monotypic subgenus within Lycodon. Also recognized are the following associated genera: Lepturophis Boulenger, 1900, Dryocalamus Günther, 1858 and Hydrophobus Günther, 1862, making a grand total of 10 genera within Lycodon sensu-lato. In order to best identify the relationships between all the associated genera a new tribe Snakebustersini tribe nov. is erected to accommodate the genera. Keywords: Taxonomy; Nomenclature; Lycodon; Dinodon; Tytleria; Ophites; Cercaspis; Tetragonosoma; Leptorhytaon; Dryocalamus; Hydrophobus; Sphecodes; Dannyelfakharikukri; new tribe; Snakebustersusini; new genera; Snakebustersus; Apollopierson; new subgenera; Mindanaosnakebustersus; Myanmarelfakhari; Sinoelfakhari; Paralycodon; Kotabilycodon.
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