Lycodon zoosvictoriae, Neang, Thy, Hartmann, Timo, Hun, Seiha, Souter, Nicholas J. & Furey, Neil M., 2014

Neang, Thy, Hartmann, Timo, Hun, Seiha, Souter, Nicholas J. & Furey, Neil M., 2014, A new species of wolf snake (Colubridae: Lycodon Fitzinger, 1826) from Phnom Samkos Wildlife Sanctuary, Cardamom Mountains, southwest Cambodia, Zootaxa 3814 (1) : -

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, 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.

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, 1st SL ventrally, posterior nasal in contact with internasal and prefrontal dorsally, loreal posteriorly, 1st and 2nd SL ventrally; loreal large, elongated, subrectangular, in contact with prefrontal dorsally, preoculars posteriorly, so not in contact with orbit, 2nd & 3rd 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 3rd SL, allowing 3rd SL to broadly enter orbit, 2 preoculars on the right side, upper vertically elongated, subrectangular, lower much smaller, subrectangular, separating 3rd 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 5th& 6th 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 laterally beneath parietal on the right side, subrectangular on the left side. SL 8/8, 1st SL in contact with rostral anteriorly, nasal dorsally, 2nd SL in contact with nasal anteriorly, loreal dorsally, 3rd 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, 4th& 5th SL in contact with orbit, 5th SL in contact with lower postocular dorsally, 6th SL in contact with lower postocular, anterior temporal dorsally, 7th SL in contact anterior temporal and posterior temporal anteriorly, 8th SL in contact posterior temporal; mental moderate, triangular; InL 10/10, 1st pair in broad contact with each other, anterior chin shield posteriorly, 1–5th in contact with first chin shield, 5th InL in contact with both first and second chin shield; 6th largest in contact with second chin shield centrally and anteriormost gular scale, 7–10th 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 ).

Dentition. On maxilla: nine maxillary teeth, 1–5th 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 14th ventral scale; DSM 17, at level of 108th ventral scale; DSV 15, at level of 206th ventral scale; VS 213, venter keeled, angulate laterally; SC 85, divided; anal plate single; scales rhomboid, 1–4th scale row smooth, 4th scale row indistinctly keeled, 5–8th 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 95th 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).

literature and comparative material. Data were taken from Anderson (1879); Smith (1943); Taylor (1965); Saint Girons (1972); Daltry & Wüster (2002); Pauwels et

. (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 L. capucinus L. cardamomensis L. davidi L. fasciatus sp. nov.

SVL 411 258–684 436–709 308 358–717 Dorsal surface of head Dark brown Dark Black Olive-brown Dark

o of transverse blotches or 31 Reticulate 11–13 84 19–49

bands on body

o of transverse blotches or 26 – 7 38 7 –21

crossbands on tail

Color of blotches or Dark brown Mixed reticulate Whitish Pale tan brown Whitish grey

crossbands

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 White with dark blotches White with dark transverse

dark blotches blotches ……continued on the next page Venter coloration Cream, speckled White Cream to beige- Cream White Grayish

posteriorly brown

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 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).

Figures

FIGURE 2. Holotype (CBC 02238) of Lycodon zoosvictoriae sp. nov. in life. Photo: Thy Neang.

FIGURE 3. Head of the holotype (CBC 02238) 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 (CBC 02238) of Lycodon zoosvictoriae sp. nov. in preservative. Photos: Thy Neang.

FIGURE 5. Submontane habitat of the holotype (CBC 02238) of Lycodon zoosvictoriae sp. nov. Photo: Thy Neang.

FIGURE 6. Striking position of (CBC 02238) Lycodon zoosvictoriae sp. nov. during defensive behavior. Photo: Thy Neang.