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Trimeresurus popeiorum SMITH, 1937

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Higher TaxaViperidae, Crotalinae, Colubroidea, Caenophidia, Alethinophidia, Serpentes, Squamata (snakes)
Subspecies 
Common NamesE: Pope’s Tree Viper, Pope's Bamboo Pit Viper, Pope’s Green Pit Viper
G: Popes Bambusotter, Popes Lanzenotter
Chinese: 坡普竹叶青蛇
E: Yingjiang green pitviper [yingjiangensis]
Chinese: Yíng jiāng zhú yè qīng (盈江竹叶青) [yingjiangensis] 
SynonymTrimeresurus popeiorum SMITH 1937
Trimeresurus popeorum — GRANDISON 1972: 94
Trimeresurus popeiorum — COX et al. 1998: 21
Trimeresurus popeorum — HOFFMANN 1998
Trimeresurus popeorum — MCDIARMID, CAMPBELL & TOURÉ 1999: 340
Trimeresurus popeiorum — TU et al. 2000
Trimeresurus popeiorum — VOGEL et al. 2004
Popeia popeiorum — MALHOTRA & THORPE 2004
Popeia popeiorum — GRISMER et al. 2006
Trimeresurus popeiorum — DAVID et al. 2009
Trimeresurus (Popeia) popeiorum — DAVID et al. 2011
Popeia popeorum — WALLACH et al. 2014: 575
Popeia popeiorum — GUO et al. 2015
Popeia popeiorum — CHAN-ARD et al. 2015: 291
Trimeresurus popeiorum — MULCAHY et al. 2017
Popeia popeiorum — GUO et al. 2018
Trimeresurus yingjiangensis CHEN, ZHANG, SHI, TANG, GUO, SONG & DING 2019
Popeia yingjiangensis — WANG 2022
Trimeresurus popeiorum — MIRZA et al. 2023 
DistributionNE India (Mizoram etc.), Nepal, Myanmar (= Burma), N Thailand, N Laos, Malaysia (Pulau Tioman? [LIM & LIM 1999]); China (Yunnan)

Type locality: Khasi Hills, Assam (State of Meghalaya), India (designated by TAYLOR & ELBEL 1958)

yingjiangensis: China (Yunnan), Myanmar (Htamanthi wildlife sanctuary, Sagaing Division); Type locality: Heihe Village, Kachang Town, Yingjiang County, Yunnan Province (24.782° N, 97.878° E, 1112 m elevation)  
Reproductionovovivparous 
TypesHolotype: BMNH 72.4.17.137 (Lectotype)
Holotype: CIB DL2017070101, adult male, collected by Li DING on 19 July 2017 (Figure 1). Paratypes: ZL-tspynglg-2018- 01(allotype, GIABR) adult female, collected from Heping Village, Tongbiguan Town, Yingjiang County, Yunnan Province (24.584° N, 97.738° E, 1 200 m) by Jian XU. DL201070102 (CIB), adult male, and DL201070103 (CIB), adult male, collected by Li DING at the same time as Holotype. OV2671 (IVPP), collected from the Yingjiang County (24.734° N, 97.843° E, 1074 m) by Jingsong SHI on 6 September 2017. [yingjiangensis] 
DiagnosisDIAGNOSIS (DIAGNOSTIC CHARACTERS): Scales in 21 (rarely 23) longitudinal rows at midbody; 9–11 upper labials, first upper labials separated from nasals by a distinct suture; a single supraocular; above green, below pale green to whitish, the two separated by a bright bicolored orange or brown (below) and white (above) (males) or white (females) ventrolateral stripe, which occupies the whole of the outermost scale row and a portion of the second row; ventrals 155–169; subcaudals 52–76, in males the base of the tail enlarged to the level of subcaudals 20–25; hemipenes long and slender, smooth, without spines. Total length 770 mm, tail length 170 mm. [after LEVITON 2003]

Diagnosis (popeiorum). A Trimeresurus bearing 21 (rarely 19) dorsal scale rows at midbody with an overall bright green colour, lacking bands; males may bear a bicoloured postocular stripe and a bicoloured ventrolateral stripe is always present. SVL 414–692 mm in males, 417–710 mm in females. Hemipenis deeply forked, reaching the 25th subcaudal. Ventrals 158–170 in males, 161–169 in females; subcaudals 62–76 in males, 55–66 in females; nasal and first supralabial separate. TaL 107–194 mm in males, 75–139 mm in females; TaL/TL 0.18–0.22 in males, 0.14–0.19 in females. Palatine with four teeth; pterygoid with eight teeth; 10–12 dentary teeth. Maxilla with one functional and 5–6 replacement fangs. (Mirza et al. 2023)

Description based on examined specimens (popeiorum, n = 19). Body long and moderately stout, SVL 414–600 mm; head triangular and elongate, head length 20–24.3 mm (HL/SVL 0.04–0.05); head width 13.4–21 mm; (HW/ HL 0.64–0.88) clearly distinct from neck; distance between nostrils 3.8–7 mm; distance between preoculars 8.6–12.8 mm; distance between the tip of snout and anterior border of eye 6.6 mm; distance between nostril to eye 5.1–6.7 mm. Canthus rostralis distinct; a single large scale between the nasal and supraocular. Rostral subtriangular, slightly visible when viewed from above; nasal and first supralabial separate, wider than tall; three internasals, the outer pair of internasals larger than the one in the middle, and the one in the middle less than half the width of the outer ones. bordered by six scales on its posterior margin; two small scales separate the third supralabial from the nasal; second and third supralabial and three preoculars encompass the loreal pit; the lower preocular forms the lower margin of the loreal pit; one elongate and narrow supraocular; cephalic scales (CEP) small, irregular, subimbricate, smooth; longitudinal cephalic scales 27–31, gradually increasing in size towards the posterior part of the head; 10–11 CEP between anterior edge of the supraoculars and 14–17 at the posterior edge; occipital scales smooth; seven rows of scales between the internasals and anterior border of the supraoculars flat and irregular in their shape; the rows towards the posterior part of the head gradually show a feeble keel; temporals feebly keeled and subequal; subocular crescent shaped; 9–13 supralabials; SL1 not fused with nasal scale, 2nd much higher than 1st, 3rd highest among the supralabials; 4th widest, separated from the subocular by a single row of smooth scales and the lower loreal scale; 4th supralabial separated from the subocular by two rows of smooth scales; the remaining supralabials slightly decreasing in size posteriorly and in contact with temporal scales; 10–13 infralabials, the first pair in contact with each other; the first three pairs in contact with anterior chin shields; six pairs of chin shields, each pair in contact medially; separated from infalabials by 1–5 scale rows.
Body scalation. 19, 21 or 23 dorsal scales one head length behind the head (rarely 17); 21 dorsal scales at midbody, rarely 19; 13–15 dorsal scales one head length anterior to the vent; dorsal scales rhomboid, moderately keeled except for the first row which is smooth; 0–3 preventrals; ventrals 158–170 in males, 161–169 in females ventral scales; subcaudals 62–76 in males, 55–66 in females subcaudal scales; paired; single cloacal plate. Eye large, with VED/DEL ratio 0.85; tail short; ventrally depressed; TaL 107–194 mm in males, 75–139 mm in females; TaL/TL 0.18–0.22 in males, 0.14–0.19 in females. Tail prehensile. Hemipenis long and deeply forked at the 5–6th subcaudal, extending to the 23–25th subcaudal scales, calyculate throughout the arms of the fork, lacking spines. (Mirza et al. 2023)

Colouration in life (Fig. 3). Males overall in a shade of bright green throughout, slightly lighter ventrally. Scales on the dorsum are green with cyan edges. Bicoloured postocular strip, usually red and white, runs from the posterior part of the subocular to the angle of the jaw. The line extends all the way up to the tail; the outermost dorsal scale row bears a white spot at the tip and is blood red for the rest of the part, bordered by the second row, which is white in its lower half and green on its upper half. The tail in both sexes may be rusty, red or brown. The ventrolateral aspect of the tail in males may bear a discontinuous stripe. The females lack the postocular and ventrolateral stripes. The ventral scales bear a slight yellowish tinge. The males may lack the postocular stripe. The eyes of both sexes are blood red. (Mirza et al. 2023)

Diagnosis (yingjiangensis). Trimeresurus yingjiangensis sp. nov. is assigned to Popeia group by hemipenes morphology (Malhotra and Thorpe, 2004), differ with its congeners by a combination of following characters: (1) dorsal body olive drab,without cross bands on the scales; (2) a conspicuous bicolor ventrolateral stripe present on each side of males, first row of dorsal scales firebrick with a white ellipse dot on posterior upper part in male, these strips absent in females; (3) the eyes firebrick in both gender; tail red, mottled with green laterally, and the ventrolateral stripes discontinuous on the tail; (4) hemipenes long, reaching 23rd to 25th SC, forked opposite 5-6th SC (n = 4), bifurcated near the base and the sulcus spermaticus split from the apex to basal without spines; (5) 21 DSR at middle body, moderately keeled; VEN = 164–168 (n = 5), SC = 60–76 (n = 5); Sexual dimorphism, the female has more ventrals and fewer subcaudals than males; (6) tail long, with ratios of TaL/TL between 0.199 and 0.219 in male.

Comparison with other species (yingjiangensis). T. yingjiangensis sp. nov. is distinct from T. popeiorum (Lectotype, Pope and Pope, 1933) by the following characters: (1) First row of dorsal scales on each side firebrick with a white ellipse dot on posterior upper part in males vs. “brown with yellow tip” in males; (2) lowest quarter of second row white in males vs. “yellow below keel”; (3) suboculars separated from 3rd upper labial by one scale on each side vs. “two scales on each side”; (4) upper part of second upper labial separated from nasal by two small scales smaller than nostril on either side vs. “separated from nasal by two large scales on either side”.
T. yingjiangensis sp. nov. is distinct from T. nebularis by the following characters: (1) body olive drab, upper lips green with white tip, vs. body bright green with blue tones and blue upper lips; (2) eyes firebrick vs. pale green; (3) ventrolateral stripes bicolor in males vs. uniformly white or blue; (4) in males: ventrals 164–167 (n = 4), vs. 149–153 (n = 8), subcaudals 71–76 (n = 4) vs. 61–65 (n = 8).
T. yingjiangensis sp. nov. is different from T. sabahi, T. cf sabahi (from Sumatra), T. fucatus, T. barati and T. toba by following characters: (1) MSR 21 (vs. MSR 19 in T. barati); (2) dorsal color was olive drab with conspicuous bicolor ventrolateral stripes and no crossbars (vs. dorsal color was green with irregular rusty or reddish-brown dorsal crossbands and white dots on the vertebral scales in T. fucatus); (3) the ventrolateral stripe bicolor in males (vs. white in males of T. cf sabahi from Sumatra, and white in males of T. toba); (4) the temporal scales smaller than those in T. toba; (5) no ventrolateral stripe present on females (vs. white or yellow ventrolateral stripe present on females of T. sabahi); (6) the eyes firebrick (vs. yellow in T. sabahi, and orange in T. barati); (7) VEN 164–168, n = 5 (vs. VEN 142–158, n = 17 in T. barati, 147–157, n = 14, in T. sabahi, and 153–155, n = 3, in Sumatra T. cf sabahi). The comparisons of main morphological characters are summarized in Table 2 in Chen et al. 2019. 
CommentVenomous!

Distribution: Possibly in Bhutan (Lenz 2012).

Synonymy: "Contrary to Hoge & Romano Hoge (1981: 257) and Welch (1988: 137), we consider that the specific name gramineus must be restricted to the form inhabiting the Peninsular India, and that the correct name for the species distributed in Sumatra and other parts of southeastern Asia is Trimeresurus popeiorum Smith, 1937." (from David & Vogel 1996). Mirza et al. 2023 synonymized T. yingjiangensis with T. popeiorum.

Diversity: Mulcahy et al. 2017 found populations of T. popeiorum that are paraphyletic with T. nebularis but are morphologically indistinguishable from T. popeiorum and hence appear to be a cryptic species.

Similar species: Most often confused with T. stejnegeri (q.v.), the two have quite distinct hemipenes, which does not make identification of individuals in the field or in the laboratory any easier without recourse to (a) male individuals and (b) an examination of the hemipenes. However, the two species are not known to have overlapping distributions, at least based on available materials. Also, closely allied to T. popeiorum is T. yunnanensis (q.v.); ordinarily, the two are more easily be told apart by the number of midbody scale rows, 21 in T. popeiorum, 19 in T. yunnanensis (from LEVITON et al. 2003).

Type species: Trimeresurus popeiorum is the type species of the genus Popeia MALHOTRA & THORPE 2004. Popeia may not be valid fide Li et al. 2020.

For abbreviations see T. nebularis.

Habitat: partly arboreal (Harrington et al. 2018).

NCBI taxonID: 2653482 [yingjiangensis] 
Etymology"This species was named in honour to Clifford H. Pope and Sarah H. Pope. The original spelling of the specific epithet, popeiorum, was corrected into popeorum by Smith (1943: 518) on the basis that it was indeed a clerical error. Unfortunately, according to the Art. 32 (c, ii) of the Code (ICZN 1985), such a change does not fall into the category of a „correction of an incorrect original spelling" According to the Art. 33 (d), the use of a termination -orum in a subsequent spelling of a species-group name that is a genitive based upon a personal name in which the correct original spelling terminates with -iorum, is an incorrect subsequent spelling, even if the change is deliberate. The original spelling, popeiorum, must therefore be conserved." (from David & Vogel 1996)

This issue is adressed again in Biology of the Vipers (Schuett et al, ed., 2002). On page 353 Orlov et al. state that "The specific epithet is most commonly spelled popeorum. [...] William W. Lamar (in litt.) has communicated to us, however, that Smith (1943) was clearly correcting a copyist's error, that such a change was allowed at the time, and that it was specifically allowed under the first published ICZN. While the 1985 and subsequent Codes prohibit this, they apply only to actions taken post-1985. Thus, the correct name is popeorum."

The specific name yingjiangensis refers to the location of type specimens, Yingjiang Country, Yunnan Province, China. 
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