A diminutive new species of cave-dwelling Wolf Snake (Colubridae: Lycodon Boie, 1826) from Peninsular Malaysia Author Grismer, L. Lee Author Quah, Evan S. H. Author Anuar, Shahrul Author Muin, Mohd Abdul Author Wood, Perry L. Author Nor, Siti Azizah Mohd text Zootaxa 2014 3815 1 51 67 journal article 36835 10.11646/zootaxa.3815.1.3 61e18952-5077-4bcd-bfc1-d281b2ff4088 1175-5326 286258 FADC54BE-301E-40E7-9029-159082652822 Lycodon cavernicolus sp. nov. Gua Wang Burma Wolf Snake Figs. 3 –5 Holotype . Adult female ( LSUHC 9985) collected on 12 March 2011 by Evan S.H. Quah and Shahrul Anuar M.S. from Gua Wang Burma , Perlis State Park, Perlis, Peninsular Malaysia (6° 41.594N 100° 11.400E at 175 m in elevation). Paratype . Juvenile male ( LSUHC 10500) has the same data as the holotype except for being collected on 23 May 2010 . Diagnosis. Lycodon cavernicolus sp. nov. is separated from all other species of the L. ruhstrati and L. fasciatus groups by having the combination of an elongate loreal scale that enters the orbit; 245 (male) and 232 (female) ventral scales; 113 (male) and 92 (female) paired subcaudal scales; a single precloacal plate; nine or 10 supralabials; 10 or 11 infralabials; a maximum total length of 508 mm for the single female; relative tail length 0.25–0.27; venter immaculate as juveniles and with dark edging on the posterior margins of the ventral scales in adults; and bands in juveniles that are white (Tables 2,3). Description of holotype ( Figs. 3 –5). Head flattened anteriorly, distinct from the neck; snout elongate; nostril oval, large, in the middle of the nasal; eye large, with a vertically elliptic pupil; rostral triangular, hardly visible from above; nasal vertically divided by a furrow along posterior margin of nostril; two square internasals, in wide, medial contact, and in contact with two large, subrectangular prefrontals posteriorly; single, azygous, subpentagonal frontal, longer than wide; two large, elongate parietals, contacted laterally by upper anteriror and posterior temporals and a larger paraparietal; 1/1 wide, triangular supraocular; 1/1 small preocular, located above the posterior portion of loreal; 2/2 postoculars of similar size; 1/1 narrow, elongate loreal entering orbit, in contact with second, third, and fourth supralabials ventrally, the prefrontal and preocular dorsally, the nasal anteriorly; 9/10 supralabials all higher than wide except last scale in the series; first and second supralabials in contact with nasal; fourth, fifth, and sixth supralabials entering orbit; seventh supralabial largest; upper row of two (long anterior and short posterior) temporals; lower row of three (two anterior and one posterior) temporals; a middle posterior temporal ventral to upper posterior temporal and dorsal to lower posterior temporal; posterior temporals smaller than anterior temporals; 11/11 infralabials; first pair of infralabials separated medially by deep, medial groove; first five infralabials in contact with first pair of chinshields; anterior and posterior pair of chinshields elongate, generally same size and shape, and bearing a deep, medial groove that is confluent with groove separating first pair of infralabials. Body elongate, somewhat laterally compressed; SVL 406 mm ; TaL 102 mm ; TL 508 mm . 232 ventrals (plus two preventrals), 92 paired, subcaudals; anal single; dorsal scales in 17–17–15 rows, the eight medial rows weakly keeled; vertebral row not enlarged; no apical pits. FIGURE 3. Dorsal (upper) and lateral (lower) head views of the holotype of Lycodon cavernicolus sp. nov. LSUHC 9985 showing the location of head scales. F = frontal; IF = infralabial; IN = internasal; L = loreal; LAT = lower anterior temporal; LPT = lower posterior temporal; M = mental; MPT = middle posterior temporal; N = nasal; P = parietal; PF = prefrontal; PO = postocular; PP = postparietal; PrO = preocular; R = rostral; SL = supralabial; SO = suprocular; UAT = upper anterior temporal; and UPT = upper posterior temporal. FIGURE 4. Left: ventral pattern of the holotype of Lycodon cavernicolus sp. nov. LSUHC 9985. Right: ventral pattern of the paratype LSUHC 10500. Photos by L.L. Grismer Coloration in life ( Figs. 4 , 5). Body and tail nearly uniformly light-brown; body bearing 36 faint, lighter colored bands; tail bearing 29 similarly colored bands; head coloration same as that of the faint bands; venter ground color beige; posterior edges of ventral scales edged in light-brown, generally beginning with ventral scale 40; subcaudals mottled with light-brown. Variation ( Figs. 4 , 5). Differences in scalation and color pattern between the holotype , the paratype , and 12 individuals of Lycodon butleri from throughout its range are listed in Table 3. The paratype is a hatchling and bears a bold, contrasting, dorsal color pattern similar to that of juvenile Lycodon butleri (Fig. 5). Its venter however, is nearly immaculate unlike the holotype whose ventrals are edged posteriorly with dark-brown ( Fig. 4 ). It also has 45 irregularly shaped, whitish bands with darkened centers on the body and 41 similarly colored caudal bands. A wide, white band covers the occipital and posterior temporal regions. The anterior 11 bands on the body are more widely separated and distinct than the posterior body and caudal bands. Presumably, the banding pattern fades considerably with maturity as in L. butleri but not to the extent observed in L. cavernicolus sp. nov. (Figs. 5, 7). Etymology. The specific epithet “ cavernicolus ” is an adjective derived from the Latin caverna meaning “cave” and the Latin cola meaning “dweller of” and refers to this species being a cave-dweller. Natural history. Both the holotype and paratype were found deep within Gua Wang Burma cave approximately 200 m from the cave entrance ( Fig. 6 ). Both specimens were found at approximately 1100 hrs. The holotype was observed scaling a vertical wall approximately 2 m above the ground while exploring nooks and crevices. She was gravid but expelled two eggs soon after capture. The paratype was found crawling over a slanting cave wall approximately 3 m above the ground in a more exposed area. Other species of amphibians and reptiles observed in the cave or near the cave entrance were the bufonid Phrynoides aspera , the gekkonids Cnemaspis biocellata , Cyrtodactylus astrum , and C. macrotuberculatus and the colubrid Othriophis taeniurus ridleyi .

FIGURE 5. Upper: holotype of Lycodon	cavernicolus sp. nov. LSUHC 9985. Middle: paratype of L.	cavernicolus sp.	nov.
LSUHC 10500. Photos by E.S.H. Quah.	Lower: juvenile L. butleri LSUHC 9421 from Bukit Larut,	Perak. Photo by	L.L.
Grismer.

FIGURE 6. Microhabitat inside the Gua Wang Burma cave at the type locality in Perlis State Park, Perlis. Photos by L.L. Grismer. The gravid holotype indicates that the reproductive season of Lycodon cavernicolus sp. nov. extends through March. The only potential food source we found deep within cave is Cyrtodactylus astrum (juvelies) which are also known to occur on the cave walls ( Grismer et al. 2012 ). Comparisons. Lycodon cavernicolus sp. nov. is distinguishable from all species of the ruhstrati group by having a single loreal scale that enters the orbit as opposed to the loreal scale not entering the orbit. From the species of the fasciatus group it differs by having more ventral scales (232–245 vs. 182–227 collectively); more subcaudal scales in the male (113 vs. 65–92 collectively); a much smaller adult female total length (508 vs. 615–762 collectively); more caudal bands (29–41 vs. 7–23 collectively); and a belly pattern that lacks wide, dark bands or dark spots ( Table 2 ). Lycodon cavernicolus sp. nov. is further separated from its closest relative L. butleri by the loreal and internasals being separat as opposed to contacting and having an uncorrected p -distance of 9.3% (Table 4).