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Trimeresurus nebularis VOGEL, DAVID & PAUWELS, 2004

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Higher TaxaViperidae, Crotalinae, Colubroidea, Alethinophidia, Serpentes, Squamata (snakes) 
Common NamesCameron Highlands pitviper, Clouded Pit viper 
SynonymTrimeresurus nebularis VOGEL, DAVID & PAUWELS 2004
Trimeresurus gramineus — BOULENGER 1912: 217 (part.) (non Coluber gramineus Shaw, 1802)
Trimeresurus gramineus — SMITH 1930: 90 (part.)
Trimeresurus gramineus — SMEDLEY 1932: 123
Trimeresurus gramineus — HOGE & ROMANO HOGE 1981: 257 (part.)
Trimeresurus popeiorum — TWEEDIE 1954: 117 (part.)
Trimeresurus popeiorum — TWEEDIE 1957: 121 (part.)
Trimeresurus popeiorum — TWEEDIE 1983: 139 (part.) (non Trimeresurus popeiorum SMITH 1937)
Trimeresurus popeiorum — LIM 1982: cover, 20 & 21: Fig. 22)
Trimeresurus popeiorum — LIM 1990: 393, 394: Fig. 7
Trimeresurus popeiorum — LIM 1991: cover, 23 (Fig. 25)
Trimeresurus popeiorum — LIM et al. 1995: 361 (part.)
Trimeresurus popeiorum — MANTHEY & GROSSMANN (1997: 409: Fig. 316).
Trimeresurus popae — TWEEDIE 1940: 131 (part.)
Trimeresurus popeorum — LIM 1982: 21 [partim].
Trimeresurus popeiorum ssp. — CHAN-ARD et al. 1999: 199
Trimeresurus popeiorum ssp. — CHAN-ARD et al. 1999: 199 [bottom], 200 [both pictures])
Trimeresurus nebularis — DAVID et al. 2009
Trimeresurus (Popeia) nebularis — DAVID et al. 2011
Trimeresurus nebularis — LIVIGNI 2013: 396
Popeia nebularis — WALLACH et al. 2014: 575
Popeia nebularis — CHAN-ARD et al. 2015: 292 
DistributionWest Malaysia (Cameron Highlands, endemic), Thailand

Type locality: Gunung Brinchang [now Gunung Batu Berinchang], Cameron Highlands, State of Pahang, West Malaysia.

Popeia inornata: West Malaysia; Type locality: Cameron Highlands, Pahang, Malaysia. Map legend:
TDWG region - Region according to the TDWG standard, not a precise distribution map.

NOTE: TDWG regions are generated automatically from the text in the distribution field and not in every cases it works well. We are working on it.
TypesHolotype: USNM 142425; Paratypes in ZFMK, IRSNB, MNHN, ZRC

Holotype (P. inornata): ZMB (Institut für systematische Zoologie, Museum für Naturkunde der Humboldt-Universität zu Berlin, Berlin, Germany) 66281, adult male, collected by indiginous collectors, July 2002. Paratypes: BMNH, CAS 
CommentVenomous! For synonymy and references see VOGEL, DAVID & PAUWELS 2004. This paper was published 15 Nov 2004 and thus T. nebularis may be a junior synonym of Popeia inornata.

Diagnosis. —A species of Trimeresurus characterized by (1) hemipenes long, without spines; (2) 1 st supralabial distinct from nasal; (3) 21 DSR at midbody, moderately keeled; (4) overall bright green coloration with blue tones and blue upper lips in males and females, and yellowish green chin and throat; (5) large size, with a TL up to about 1000 mm in males and at leat 950 mm in females; (6) postocular streak absent in males and females; (7) ventrolateral stripes often missing, or white or pale blue; (9) upper lips bluish-green; (10) eye green in males and females; (11) tail dark rusty brown vertebrally, green laterally with a sharp border between the colours (similar to Trimeresurus albolabris or T. macrops); (12) a low number of scales between the supraoculars (9–10); (13) tail average in females, with a ratio TaL/TL 0.165–0.172; (14) VEN 147–153; SC: 50–65 (after VOGEL et al. 2004).

Comparison with other species. —Trimeresurus nebularis differs from all other pitvipers of the Trimeresurus popeiorum complex by the combination of the following characters: (1) tail with a sharp border between the green and the brown colour; (2) eye colour green at both sexes; (3) postocular streak absent in both sexes; (4) a larger size; (5) the lowest number of scales between the supraoculars; (6) the lowest number of subcaudals; (7) a low number of ventrals; (8) the lack of sexual dimorphism; (9) upper lips bluish- green; (10) ventrolateral stripes often pale blue. Main characters separating Trimeresurus nebularis from other taxa of the group are given in Tables 12–13 (in VOGEL et al. 2004). Trimeresurus nebularis differs from T. popeiorum by (1) the colour of the eye (never red in T. nebularis); (2) the number of Cep between the supraoculars; (3) the presence of a postocular streak in males of T. popeiorum; (4) the presence of a ventro-lateral streak in females of T. popeiorum; (5) a stronger keeling of the Occ and Tem in T. popeiorum; (6) a lower number of VEN in females (147–153 [x = 150.4, s = 2.3] vs. 154– 168 [x = 161.6, s = 4.2] in T. popeiorum). Trimeresurus nebularis differs from Trimeresurus fucatus by (1) the absence of dorsal crossbands in males; (2) a white or blue ventrolateral stripe in males, vs. a bicolor ventro-lateral stripe in males of T. fucatus, and the absence of this stripe in females vs. a conspic-uous white stripe in females of T. f ucat us; (3) the absence of a postocular streak in males; (4) the pattern of the tail; (5) a lower number of ventral scales (149–153 [x = 151.7, s = 2.3] in males of T. nebularis vs. 156–171 [x = 164.0, s = 3.5] in males of T. fucat us; in females 147–153 [x = 150.4, s = 2.3] vs. 157–170 [x = 163.5, s = 3.4] in T. fucatus; (6) a lower number of subcaudal scales (61–65 [x = 63.0, s = 2.8] in males of T. nebularis vs. 69–84 [x = 75.9, s = 3.5] in males of T. fucatus; in females 50–60 [x = 55.2, s = 3.8] vs. 59– 73 [x = 63.8, s = 3.6] in T. fucatus). Trimeresurus nebularis differs from Trimeresurus sabahi by: (1) a larger size; (2) the colour of the eyes (green vs. bright red in T. sabahi); (3) the pattern of the tail; (4) the presence of ventrolateral stripes in both sexes of T. sabahi; (5) a lower number of SC in both sexes (in males 61–65 [x = 63.0, s = 2.8] vs. 69–76 [x = 71.6, s = 2.5] in T. sabahi; in females 50–60 [x = 55.2, s = 3.8] vs. in T. sabahi 59–65 [x = 62.2, s = 2.7]); (6) the contact of 3 rd SL with the subocular, in contact in all examined specimens of T. nebularis, sepa-rated in 14/20 occurrences in T. sabahi; (7) by proportionnally wider supraocular scales in T. nebularis (ratio L–SpOc/W–SpOc: 2.4–2.8 ([x = 2.5, s = 0.1] in T. nebul ari s, vs. 2.8–3.3 [x = 3.0, s = 0.2] in T. sabahi). Lastly, Trimeresurus nebularis differs from T. barat i by: (1) a larger size; (2) the pat-tern of the tail; (3) the colour of eyes in both sexes; (4) the ventrolateral stripe in males (bicolor in T. barati); (4) the number of MSR (21 vs. usually 17–19 in T. barati); (5) a snout shorter in T. nebularis than in T. barat i (ratio DEP/DEN: 0.52–0.57 [x = 0.53, s = 0.03] vs. in T. barati 0.56–0.62 [x = 0.59, s = 0.02]) (after VOGEL et al. 2004).

Named after the Latin adjective nebularis, meaning “from the clouds”, in allusion to the cloudy montane rainforests, or cloud forests, inhabited by this species.

Abbreviations: SVL = Snout-vent length, TaL = Tail length, TL = Total length, HL = Head length, SnL = Snout length (from the tip of rostral to a line connecting the ante-rior, eye margins), HED = Eye diameter (horizontal), VED = Eye diameter (vertical), DEL = Distance lower eye margin–edge of the lip, DEN = Distance between the anterior eye margin and the nostril, DEP = Distance between the anterior eye margin and the loreal pit, WInN = Width of internasals (means), LSupOc = Length of supraoculars, WSupOc = Width of supraoculars, L3SL = Length of 3 rd supralabial, H3SL = Height of 3 rd supralabial, H4SL = Length of 4 th supralabial, TaL/TL = Ratio tail length/Total length, SnL/HL = Ratio snout length/head length, DEP/HL = Ratio distance eye–pit/head length, DEN/HL = Ratio distance eye–nostril/head length, DEP/DEN = Ratio distance eye–pit/distance eye–nostril, WInN/WSupOc = Ratio width of internasals/width of supraoculars, L3SL/HL = Ratio length of 3 rd supralabial/head length, VED/DEL = Ratio: vertical eye diameter/distance eye margin–edge of the lip, LSupOc/WsupOc = Ratio of the length of supraocular/width of the supraoculars, DSR = Dorsal scale rows, MSR = Dorsal scale rows at midbody, VEN = Ventral plates, SC = Subcaudal plates, SL = Supralabial scales, HeSc = Head scales (scales on a longitudinal row between the interna-sals, and the limit of the neck), SnSc = Snout scales (scales on a line between the internasals and a line, connecting the anterior margin of eye), InN = Internasal scale(s), Can = Canthal scales (scales between the internasal and the subocular), Cep = Cephalic scales (scales on a line between the middle of supraoc-ulars), Tem = Temporal scales, IL = Infralabials, NVEN = Number of ventral plates, NSC = Number of subcaudal plates, NASR = Number of dorsal scale rows behind head, NMSR= Number of dorsal scale rows at midbody, NPSR = Number of dorsal scale rows before vent, KMSR = Keeling of dorsal scale rows at midbody, TNSL = Total number of supralabial scales, C3SL = Number of scales between 3 rd supralabial and subocular, C4SL = Number of scales between 4 th supralabial and subocular, C45SL = Number of scales between 4 th and 5 th supralabial and subocular, NCep = Number of cephalic scales on a line between the supraoculars, KOcc = Keeling of the occipital scales, KTem = Keeling of temporal scales, NInN = Number of scales separating the internasals, CSupOC = Number of scales directly in contact with supraocular, NSnSc = Number of snout scales (defined as above), NHeSc = Number of longitudinal head scales (defined as above), TNIL = Total number of infralabial scales, DBB = Presence of darker bands on body, VSB Presence of a line of white vertebral spots, POSTM = Presence of a postocular streak in males, POSTF= Presence of a postocular streak in females, CPOST = Coloration of the postocular streak, VELSM = Presence of a ventrolateral stripe in males, VELSF = Presence of a ventrolateral stripe in females, CVEL = Coloration of the ventrolateral stripe, COLEM = Colour of eyes in males, COLEF = Colour of eyes in females, TAP = Pattern of the tail.

Diagnosis (P. inornata): The new species is distinguished from other species of the Trimeresurus (sensu lato) complex, with the exception of other members of the genus Popeia and some members of the genus Cryptelytrops Cope, 1860, by a long calyculate hemipenis without papillae (as described in Malhotra and Thorpe, 2004b). Popeia inornata is distinguished from members of Cryptelytrops by full separation of the first supralabial and nasal scales. It is distinguished from other members of Popeia by the absence of a red or orange lateral stripe in adult males. Both sexes of P. inornata are separated from P. popeiorum sensu stricto by low ventral scale counts (no higher than 156, compared to no lower than 158), and from both P. popeiorum and the southern clade species by green eye coloration, with the exception of West Malaysian representatives of the southern clade, from which P. inornata can be distinguished by low ventral scale counts alone (149 – 156 vs. 157 – 172). With respect to the classification of Regenass and Kramer (1981), P. inornata is distinguished from the Borneo population (T. p. sabahi sensu Regenass and Kramer, 1981) by the absence of a red or orange lateral stripe in adult males and by green eye coloration in both sexes. It differs from snakes from the Barisan range of western Sumatra (T. p. barati sensu Regenass and Kramer, 1981) in having a longer and less compact head, a coloration of interstitial skin and dorsal scale bases that forms black bands with narrow light blue interspaces (vs. no dark bands on uniformly bluish gray to blue interstitial skin), and lower and upper margins of supralabials and infralabials, respectively, that are distinctly blue in life (vs. bluish white to lighter blue). Popeia inornata is further distinguished from west Sumatran specimens by the dorsal coloration of the tail which in life is a well-defined and solid brick red with light margins on red dorsal scales (vs. a light brown to golden hue which is less clearly demarcated laterally). While Regenass and Kramer (1981) distinguished T. p. barati on the basis of lower midbody dorsal scale counts (17 – 19 vs. 21 in the rest of their T. popeiorum group), an additional specimen examined by Sanders, Malhotra, and Thorpe (unpublished) from west Sumatra has 21 scale rows. Additional morphological characters that can be used in combination to diagnose the new species are listed in Table 1; diagnostic molecular characters are given above (3.2).
Popeia inornata is not known to share its geographic range with any other green pitviper species. It is easily distinguished from juvenile and adult male Tropidolaemus wagleri (Boie, 1827) by the distinct red and white pre- and postocular stripe, red and white dorsal bands or spots, and keeled scales on the lower side of the head possessed by the latter species. Adult females of T. wagleri are even more distinct from P. inornata due to their highly divergent, largely black and yellow coloration and relatively massive girth. The new species can be distinguished from Parias hageni (Lidth de Jeude, 1886) by the dorsal coloration and pattern of the tail, which in Pa. hageni is green with a few irregular reddish-brown spots, and solid reddish-brown only in about the distal half of the tail (vs. laterally well-defined and solid brick red from about the anal region to the tip in P. inornata). 
EtymologyEtymology (P. inornata): The specific epithet inornata is an adjective in the feminine singular gender. The scientific name of the new species alludes to the fact that adult males of P. inornata lack ornamentation by a distinctive orange to brick red ventrolateral stripe, which is typically present in adult males of sexually dimorphic members of the genus Popeia, and that both sexes show reduction of the white lateral stripe. 
  • Chan-ard, T., Parr, J.W.K. & Nabhitabhata, J. 2015. A field guide to the reptiles of Thailand. Oxford University Press, NY, 352 pp. [see book reviews by Pauwels & Grismer 2015 and Hikida 2015 for corrections] - get paper here
  • Chan-ard,T.; Grossmann,W.; Gumprecht,A. & Schulz,K. D. 1999. Amphibians and reptiles of peninsular Malaysia and Thailand - an illustrated checklist [bilingual English and German]. Bushmaster Publications, Würselen, Gemany, 240 pp. [book review in Russ. J Herp. 7: 87] - get paper here
  • Das, I. 2012. A Naturalist's Guide to the Snakes of South-East Asia: Malaysia, Singapore, Thailand, Myanmar, Borneo, Sumatra, Java and Bali. Oxford J, ohn Beaufoy Publishing - get paper here
  • David, P., Petri, M., Vogel, G. & G. Doria 2009. A new species of pitviper of the genus Trimeresurus (Popeia) from northern Sumatra (REPTILIA, SQUAMATA, VIPERIDAE). Annali del Museo Civico di Storia Naturale "G. Doria", Genova 100: 323-346
  • DAVID, PATRICK; GERNOT VOGEL & ALAIN DUBOIS 2011. On the need to follow rigorously the Rules of the Code for the subsequent designation of a nucleospecies (type species) for a nominal genus which lacked one: the case of the nominal genus Trimeresurus Lacépède, 1804 (Reptilia: Squamata: Viperidae). Zootaxa 2992: 1–51 - get paper here
  • Lim, Boo Liat 1982. Poisonous snakes of peninsular Malaysia, 2.ed. Malayan Nature Soc., Kuala Lumpur 73 pp.
  • LiVigni, F. (ed.) 2013. A Life for Reptiles and Amphibians, Volume 1. Chimaira, Frankfurt, 495 pp. - get paper here
  • Sanders, K. L., A. Malhotra, A. Gumprecht, R. S. Thorpe & U. Kuch 2004. Popeia inornata, a new species of pitviper from West Malaysia (Squamata: Viperidae: Crotalinae). Russ. J. Herpetol. 11(3): 171-184 - get paper here
  • Sumontha,M., Kunya,K., S. G. Pauwels,O., Nitikul,A., and Punnadee,S. 2011. Trimeresurus (Popeia) phuketensis, a New Pitviper (Squamata: Viperidae) from Phuket Island, Southwestern Thailand. Russ. J. Herpetol. 18 (3): 11-17 - get paper here
  • Thompson, C. & Thompson, T. 2008. First contact in the Greater Mekong - new species discoveries. WWF, 40 pp. - get paper here
  • Vogel, G.; David, P. & Pauwels, O.S.G. 2004. A review of morphological variation in Trimeresurus popeiorum (Serpentes: Viperidae: Crotalinae), with the description of two new species. Zootaxa 727: 1–63 - get paper here
  • Vogel, Gernot 2010. Zur Systematik der Schlangen der orientalischen Region: Forschungsdefizite führen zur Unterschätzung der Artenvielfalt. Ophidia 4 (2): 18-26 - get paper here
  • Wallach, Van; Kenneth L. Williams , Jeff Boundy 2014. Snakes of the World: A Catalogue of Living and Extinct Species. Taylor and Francis, CRC Press, 1237 pp.
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