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Oligodon arenarius VASSILIEVA, 2015

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Higher TaxaColubridae, Colubrinae, Colubroidea, Caenophidia, Alethinophidia, Serpentes, Squamata (snakes) 
Subspecies 
Common Names 
SynonymOligodon arenarius VASSILIEVA 2015 
DistributionS Vietnam (Ba Ria–Vung Tau)

Type locality: Binh Chau–Phuoc Buu Nature Reserve, Xuen Moc District, Ba Ria–Vung Tau Province, southern Vietnam, coordinates 10°29'46"N, 107°27'54"E, elevation 5 m elevation.  
Reproduction 
TypesHolotype. ZMMU R-14503 (field ID ZMMU ABV–00813), adult male, collected by Anna B. Vassilieva on 14 November 2014.
Paratypes. ZMMU R-14002 (field ID ZMMU NAP-03137), adult male from type locality: Binh Chau–Phuoc Buu Nature Reserve, Xuen Moc Disctrict, Ba Ria–Vung Tau Province, southern Vietnam (10°32'18"N, 107°28'37"E, 43 m a.s.l.), collected on 8 November 2010 by Anna B. Vassilieva; ZMMU R-14002 (field ID ZMMU NAP-03138), adult female from type locality (10°29'50"N, 107°27'58"E, 5 m a.s.l.) collected on 9 November 2010 by Anna B. Vassilieva; VNMN 04724 (field ID NAP-03884), adult female from type locality, environs of the resort Ho Coc (10°29'54"N, 107°28'08"E; 8 m a.s.l.), captured on 10 July 2012 by local people and collected by Anna B. Vassilieva and Nikolay Poyarkov; ZMMU R-14504, adult female from type locality (10°29'41"N, 107°27'46"E; 7 m a.s.l.), collected on 16 November 2014 by Anna B. Vassilieva. 
CommentDiagnosis. The species is allocated to the genus Oligodon based on the following characters considered to be diagnostic for the genus (Smith, 1943): posterior maxillary teeth strongly enlarged and compressed; palatine teeth well developed or vestigial; head short, not distinct from neck, usually displaying a specific coloration pattern; head scalation complete or reduced; eye moderate, with rounded pupil; rostral scale enlarged, extending on to the upper surface of the snout and partly separating the internasals; body cylindrical; scales smooth; ventrals rounded or obtusely keeled laterally; subcaudals paired.
The new species is distinguishable from other species of the genus by a combination of the following features: (1) medium size in adults (TL 336–389 mm); (2) 17(19)–17–15 dorsal scale rows; (3) 6–8 maxillary teeth, the posterior three being enlarged; (4) head scalation lacking a loreal but including a presubocular (except one specimen), (5) nasal divided; (6) two postoculars; (7) 131–144 ventrals; (8) 36–60 subcaudals (36–40 in females, 58–60 in males); (9) anal plate entire; (10) tail quite long for the genus in males (TaL/TL = 0.26–0.28) and moderate in females (TaL/TL = 0.13–0.17); (11) unforked hemipenes without spines; (12) head coloration pattern including ocular band, temporal bands and chevron-shaped mark on nape; (13) dorsal coloration without specific pattern, uniform or with dark speckling; (14) ventrals pinkish in life, immaculate; (15) iris bicolored with silvery upper third and dark brown lower two thirds.

Comparisons. Oligodon arenarius sp. nov. differs from all other species of the genus by the unique combination of the following characters: 17 DSR, the absence of a loreal scale, unforked hemipenis without spines or obvious papillae, the absence of a specific dorsal coloration pattern (blotches, crossbars or longitudinal lines) and the expressed sexual dimorphism in the relative tail length and number of subcaudals.
The absence of a loreal is a rather rare feature in kukri snakes. In particular, among the 22 Oligodon species known from Indochina, this element of head scalation is consistently absent in only three species: O. annamensis Leviton (Vietnam, Cambodia), O. catenatus (Blyth) (Vietnam, Thailand, Myanmar) and O. lacroixi Angel et Bourret (Vietnam, China); also, two Indochinese species occurring in Vietnam, O. mouhoti (Boulenger) and O. macrurus (Angel), lack the loreal scale facultatively. Oligodon arenarius sp. nov. differs from them by the following features: from O. annamensis, O. catenatus and O. lacroixi, by a greater DSR count (17 vs. 13, 13 and 15, respectively); usually present presubocular; 8(7) supralabials vs. 6(5), 6 and 5, respectively; 9(8) infralabials vs. 6, 7 and 6, respectively; a lesser number of ventrals (131–144 vs. 148–170, 179–212 and 162–178, respectively) and a dorsal coloration pattern (uniform or with dark speckling vs. thin light crossbars in O. annamensis, chained vertebral line in O. catenatus and four indistinct longitudinal lines in O. lacroixi); additionally, Oligodon arenarius sp. nov. differs from O. annamensis and O. lacroixi by a greater number of subcaudals (58–60 vs. 44–46 in males, 36–40 vs. 30 in females of O. annamensis, 29–34 in O. lacroixi) and from O. catenatus and O. lacroixi by an entire anal plate instead of divided one; Oligodon arenarius sp. nov. differs from O. mouhoti by a longer tail in males (0.26–0.28 vs. 0.17–0.19) and a greater number of subcaudals in males (58–60 vs. 39–43), as well as by dorsal coloration features (vertebral stripe and two large dark blotches on tail in O. mouhoti) and the hemipenial morphology (see below); from O. macrurus, by a relatively shorter tail in males (TaL/TL 0.26–0.28 vs. 0.34–0.37, calculated after the data of Geissler et al. 2011, or 0.31 in one specimen, P. David, pers. comm.), a markedly lesser number of subcaudals in males (58–60 vs. 73–94), a slightly lesser number of ventrals in males (131–134 vs. 139– 145) and the hemipenial morphology (see below).
Oligodon arenarius sp. nov. also differs from many congeners by its hemipenial morphology (unforked hemipenis devoid of spines and papillae). In particular, it differs from all the members of the informal O. taeniatus group (sensu Smith 1943, David et al. 2008b) with Indochinese distribution, namely O. taeniatus (Günther), O. pseudotaeniatus David, Vogel & Van Rooijen, O. deuvei David, Vogel & Van Rooijen, O. barroni (Smith), O. moricei David, Vogel & Van Rooijen and O. mouhoti discussed above, which possess deeply forked hemipenes. Additionally, Oligodon arenarius sp. nov. differs from them externally by the dorsal coloration (all members of the O. taeniatus group possessing vertebral stripes or blotches) and by the following characters: from O. taeniatus, by 17 vs. 19 DSR and a greater subcaudals count in males (58–60 vs. 38–48); from O. pseudotaeniatus, by 9 vs. 8 infralabials and a greater subcaudals count in males (58–60 vs. 44–46); from O. deuvei, by a slightly lesser ventrals count (131–134 vs. 140–147 in males, 142–144 vs. 147–155 in females) and a greater subcaudals count in males (58–60 vs. 36–47); from O. barroni, by a relatively longer tail in males (0.26–0.28 vs. 0.17–0.19), a greater subcaudals count in males (58–60 vs. 36–48) and females (36–40 vs. 28–35) and the immaculate belly coloration; from O. moricei, by a lesser ventrals count (131–144 vs. 175), 6–8 maxillary teeth vs. 12 and a usually present presubocular.
Oligodon arenarius sp. nov. also differs by its hemipenial morphology from all the members of the informal Oligodon cyclurus group (sensu Smith 1943, David et al. 2008a, Green et al. 2010). The species of Oligodon cyclurus group are distributed in Indochina, China and India, and possess deeply forked hemipenes without spines but sometimes with papillae; these features were documented for O. cattienensis Vassilieva, Geissler, Galoyan, Poyarkov, Van Devender & Böhme, O. chinensis (Günther), O. cyclurus (Cantor), O. fasciolatus (Günther), O. formosanus (Günther), O. juglandifer (Wall), O. kampucheaensis Neang, Grismer & Daltry, O. ocellatus (Morice), O. saintgironsi David, Vogel & Pauwels, and O. macrurus (discussed above). Additionally, from the species of the Oligodon cyclurus group occurring in Indochina, Oligodon arenarius sp. nov. differs externally by the dorsal coloration features (all members of the Oligodon cyclurus group except O. macrurus possess blotches, reticulation or crossbars) and by the following characters: from O. cattienensis, by the immaculate belly and a lesser ventrals count (131–144 vs. 167–178); from O. chinensis, by a much lesser ventrals count (131–144 vs. 170–206); from O. fasciolatus, by DSR 17 vs. 21–23 and by a lesser ventrals count (131–144 vs. 147–210); from O. formosanus, by DSR 17 vs. 19 and a lesser ventrals count (131–144 vs. 154–189); from O. kampucheaensis, by DSR 17 vs. 15 and a lesser ventrals count (131–144 vs. 165); from O. ocellatus, by DSR 17 vs. 19, a relatively longer tail in males (TaL/TL 0.26–0.28 vs. 0.11–0.14) and a greater subcaudals count (58–60 vs. 32–44 in males, 36–40 vs. 26–33 in females); from O. saintgironsi, by a lesser ventrals count (131–144 vs. 166–184), a lesser subcaudals count in females (36–40 vs. 53) and a relatively longer tail in males (TaL/TL 0.26–0.28 vs. 0.19–0.20).
From the members of the informal Oligodon cinereus group (sensu Smith 1943, Green et al. 2010, David et al. 2011, 2012, Neang et al. 2012), which are distributed in Indochina, China, India, Andaman and Nicobar islands, and are characterized by the unforked hemipenes, not spinose but with papillae, Oligodon arenarius sp. nov. differs by the absence of a loreal (present in all group members except O. melanozonatus Wall from India) and by the following features: from O. cinereus (Günther), by a lesser ventrals count (131–144 vs. 155–186) and by the dorsal coloration pattern (from all subspecies having blotches, reticulations or crossbars except the uniformly colored O. c. cinereus); from O. albocinctus (Cantor), by a lesser ventrals count (131–144 vs. 177–207) and by the dorsal coloration pattern (light crossbars in O. albocinctus); from O. inornatus (Boulenger), by DSR 17 vs. 15 and a lesser ventrals count (131–144 vs. 171–174); from O. joynsoni (Smith), by a lesser ventrals count (131–144 vs. 186–195) and the dorsal coloration pattern (blotches and reticulated crossbars in O. joynsoni); from O. maculatus (Taylor), by a lesser ventrals count (131–144 vs. 156–164) and the dorsal coloration pattern (dark blotches in O. maculatus); from O. melanozonatus, by a lesser ventrals count (131–144 vs. 171–173) and the dorsal coloration pattern (black- edged or black crossbars in O. melanozonatus); from O. nagao David, Nguyen, Nguyen, Jiang, Chen, Teynié & Ziegler, by a lesser ventrals count (131–144 vs. 184–193), a relatively longer tail in males (TaL/TL 0.26–0.28 vs. 0.14–0.15) and the dorsal coloration (butterfly-shaped blotches in O. nagao); from O. splendidus (Günther), by DSR 17 vs. 21, a single pair of both prefrontals and internasals (up to four pairs of each in O. splendidus), a lesser ventrals count (131–144 vs. 169–193) and the dorsal coloration (large spots on dorsum in O. splendidus); from O. woodmansoni (Sclater), by 8(7) supralabials vs. 6, a lesser ventrals count (131–144 vs. 180–190) and pale belly coloration (dark brown with white edges in O. woodmansoni).
The members of the informal O. dorsalis group (sensu Smith 1943) are distributed in Indochina, India, Nepal and Myanmar; the species with documented hemipenial morphology are characterized by unforked and usually spinose hemipenes. From all congeners belonging to this group, except O. lacroixi and O. catenatus discussed above, Oligodon arenarius sp. nov. differs by the dorsal coloration (longitudinal stripes or crossbars in all members of the O. dorsalis group) and an entire anal plate vs. divided, as well as by the following features of the pholidosis: from O. dorsalis (Gray & Hardwicke), by DSR 17 vs. 15, the absence of a loreal and a lesser ventrals count (131–144 vs. 162–188); from O. eberhardti Pellegrin, by DSR 17 vs. 13, 8(7) supralabials vs. 6 and a lesser ventrals count (131–144 vs. 165–191); from O. erythrogaster Boulenger, by a lesser ventrals count (131–144 vs. 163–186); from O. hamptoni Boulenger, by DSR 17 vs. 15, 8(7) supralabials vs. 5 and a lesser ventrals count (131– 144 vs. 160–175); from O. mcdougalli Wall, by DSR 17 vs. 13 and a lesser ventrals count (131–144 vs. 199).
In the species of the informal O. torquatus group (sensu Smith 1943, Green et al. 2010) with the distribution in India, Myanmar and Thailand, hemipenes are unforked with apical papillae. Oligodon arenarius sp. nov. differs from the species of this group by the dorsal coloration features (longitudinal stripes in the O. torquatus group except O. planiceps), an entire anal plate vs. divided and additionally by the following external features: from O. cruentatus (Günther), by a lesser ventrals count (131–144 vs. 148–173); from O. planiceps Boulenger, by DSR 17 vs. 13, 8(7) supralabials vs. 5(4), a greater subcaudals count (36–60 vs. 22–27) and the immaculate ventrals (ventrals with paired dark blotches in O. planiceps); from O. theobaldi (Günther), by the absence of a loreal and a lesser ventrals count (131–144 vs. 164–180); from O. torquatus (Boulenger), by DSR 17 vs. 15 and the absence of a loreal.
In the species of the informal O. venustus–O. taeniolatus group (sensu Smith 1943, Green et al. 2010) with distribution in India, Sri Lanka, Afghanistan, Pakistan, Nepal and Bangladesh, hemipenes are unforked and spinose. Oligodon arenarius sp. nov. differs from the species of this group by the absence of spines on the hemipenis, the dorsal coloration pattern (crossbars or longitudinal stripes or blotches in all species of the O. venustus–O. taeniolatus group), an entire anal plate vs. divided, and the following external characters: from O. affinis Günther, by the immaculate belly (with black subrectangular spots in O. affinis); from O. arnensis (Shaw), by a lesser ventrals count (131–144 vs. 164–202); from O. brevicauda Günther, by the presence of internasals, DSR 17 vs. 15, a lesser ventrals count (131–144 vs. 158–173) and a greater subcaudals count (36–60 vs. 25–29); from O. calamarius (Linnaeus), by the absence of a loreal and a greater subcaudals count (36–60 vs. 20–34); from O. erythrorachis Wall, by DSR 17 vs. 15, 9(8) infralabials vs. 4 and a lesser ventrals count (131–144 vs. 154); from O. melaneus Wall, by DSR 17 vs. 15, the absence of a loreal and a lesser ventrals count (131–144 vs. 152–160); from O. sublineatus Duméril, Bibron & Duméril, by DSR 17 vs. 15 and the absence of a loreal; from O. taeniolatus (Jerdon), by DSR 17 vs. 15, the absence of a loreal and a lesser ventrals count (131–144 vs. 158–218); from O. travancoricus Beddome, by a lesser ventrals count (131–144 vs. 154–155); from O. venustus (Jerdon), by the immaculate belly (with large black spots in O. venustus).
From other species of the genus, which are not clearly assigned to any specific groups of species, Oligodon arenarius sp. nov. differs by the following characters: from O. ancorus (Girard) (the Philippines), by the dorsal coloration (large dorsal spots in O. ancorus), the absence of a loreal and a lesser ventrals count (131–144 vs. 147– 173); from O. annulifer (Boulenger) (Malaysia: Borneo), by the dorsal coloration (crossbands in O. annulifer), DSR 17 vs. 15 and the absence of a loreal; from O. bitorquatus Boie (Indonesia), by the dorsal coloration (red and yellow dots in O. bitorquatus) and the immaculate belly (with black spots in O. bitorquatus); from O. booliati Leong & Grismer (Peninsular Malaysia), by the dorsal coloration (crossbands in O. booliati) and 9(8) infralabials vs. 7; from O. everetti Boulenger (Malaysia and Indonesia), by DSR 17 vs. 15 and the absence of a loreal; from O. forbesi (Boulenger) (Indonesia), by the absence of a loreal, a lesser ventrals count (131–144 vs. 150–170) and the immaculate belly (ventrals with brown spots in O. forbesi); from O. jintakunei Pauwels, Wallach, David & Chanhome (Thailand), by the dorsal coloration (crossbars in O. jintakunei), DSR 17 vs. 15, an entire anal plate and a lesser ventrals count (131–144 vs. 189); from O. kheriensis Achardji & Ray (Nepal and India), by the absence of a loreal and a lesser ventrals count (131–144 vs. 196); from O. lungshenensis Zheng & Huang (China), by DSR 17 vs. 15, 8(7) supralabials vs. 6, 9(8) infralabials vs. 6 and a lesser ventrals count (131–144 vs. 163–179); from O. meyerinkii (Steindachner) (the Philippines and Malaysia), by the absence of a loreal and a lesser ventrals count (131–144 vs. 154–169); from O. modestus Günther (the Philippines), by DSR 17 vs. 15, 8(7) supralabials vs. 6 and a lesser ventrals count (131–144 vs. 158–176); from O. nikhili Whitaker & Dattatri (India), by DSR 17 vs. 15, the dorsal coloration (longitudinal stripes in O. nikhili) and the immaculate belly (ventrals with black spots in O. nikhili); from O. notospilus Günther (the Philippines), by the absence of a loreal and the dorsal coloration (large rhombic yellow spots in O. notospilus); from O. octolineatus (Schneider) (Malaysia and Indonesia), by the dorsal coloration (longitudinal stripes in O. octolineatus), the absence of a loreal and a lesser number of ventrals (131–144 vs. 155–197); from O. ornatus Van Denburgh (China and Taiwan), by the dorsal coloration (crossbars in O. ornatus), DSR 17 vs. 15 and a lesser number of ventrals (131–144 vs. 156–182); from O. perkinsi (Taylor) (the Philippines), by the absence of a loreal and a lesser number of ventrals (131–144 vs. 183–188); from O. petronellae Roux (Indonesia), by the dorsal coloration (dark spots in O. petronellae), DSR 17 vs. 15 and the absence of a loreal; from O. praefrontalis Werner (Indonesia), by the presence of internasals, an entire anal plate vs. divided and a lesser number of ventrals (131–144 vs. 183–193); from O. propinquus Jan (Indonesia), by the absence of a loreal and DSR 17 vs. 15; from O. pulcherrimus Werner (Indonesia), by the dorsal coloration (elongate spots on dorsum in O. pulcherrimus), the absence of a loreal and a lesser number of ventrals (131–144 vs. 152–176); from O. purpurascens (Schlegel) (Thailand, Malaysia and Indonesia), by the absence of a loreal, DSR 17 vs. 19–21 and a lesser number of ventrals (131–144 vs. 150–210); from O. signatus (Günther) (Indonesia and Malaysia), by the dorsal coloration (yellow spots on dorsum in O. signatus) and the absence of a loreal; from O. trilineatus (Duméril, Bibron & Duméril) (Indonesia), by the dorsal coloration (longitudinal stripes in O. trilineatus) and the absence of a loreal; from O. unicolor (Kopstein) (Indonesia), by the absence of a loreal and a lesser number of ventrals (131– 144 vs. 162); from O. vertebralis Günther (Indonesia and Malaysia), by the dorsal coloration (white or yellow dorsal spots in O. vertebralis), DSR 17 vs. 15 and the absence of a loreal; from O. waandersi (Bleeker) (Indonesia), by DSR 17 vs. 15 and a greater number of subcaudals (36–60 vs. 18–29); from O. wagneri David & Vogel (Indonesia), by DSR 17 vs. 15 and the dorsal coloration (white crossbars in O. wagneri).

Variation: O. arenarius is quite variable in color, with beige, light brown, greyish, reddish variants (Vassilieva 2015, Figs. 5-6).

Sympatry: O. deuvei, O. fasciolatus 
Etymology The new species name is an adjective in the nominative case, masculine gender, derived from the Latin word "arena" meaning "sand", "sandy land". The name is intended to reflect the evident preference by the new species for sandy coastal habitats and the snake's remarkable ability to plunge into the sand and move under it. 
References
  • VASSILIEVA, ANNA B. 2015. A new species of the genus Oligodon Fitzinger, 1826 (Squamata: Colubridae) from coastal southern Vietnam Zootaxa 4058 (2): 211–226 - get paper here
 
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