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Sinomicrurus macclellandi (REINHARDT, 1844)

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Higher TaxaElapidae, Colubroidea, Caenophidia, Alethinophidia, Serpentes, Squamata (snakes)
SubspeciesSinomicrurus macclellandi macclellandi (REINHARDT 1844)
Sinomicrurus macclellandi nigriventer (WALL 1908)
Sinomicrurus macclellandi univirgatus (GÜNTHER 1858) 
Common NamesE: MacClelland’s Coral Snake
G: MacClellands Korallenotter
Chinese: 中华珊瑚蛇
E: Black-bellied Coral Snake [nigriventer] 
SynonymElaps macclellandi REINHARDT 1844: 532
Elaps personatus BLYTH 1855 (fide SMITH 1943)
Callophis macclellandii — ANDERSON 1871: 189
Calliophis macclellandii — STEJNEGER 1907: 391
Callophis maclellandii [sic] — WALL 1908: 780
Calliophis macclellandii var. gorei WALL 1909: 842
Callophis macclellandi (var. typica) — VENNING 1910: 775
Calliophis swinhoei VAN DENBURGH 1912
Callophis [sic] formosensis THOMPSON 1912
Callophis macclellandi — SMITH 1943: 423
Hemibungarus macclellandi — GOLAY et al. 1993
Micrurus macclellandi — WELCH 1994: 85
Calliophis macclellandii — ZUG & MITCHELL 1995
Calliophis macclellandi — COX et al. 1998: 32
Sinomicrurus macclellandi — SLOWINSKI, BOUNDY & LAWSON 2001
Hemibungarus macclellandii — GRUBER in SCHLEICH & KÄSTLE 2002
Hemibungarus macclellandii — ORLOV et al. 2003
Sinomicrurus macclellandi — LEVITON et al. 2003
Sinomicrurus macclellandi — ZIEGLER et al. 2007
Sinomicrurus macclellandi — WALLACH et al. 2014: 676
Sinomicrurus macclellandi — SMART et al. 2021

Sinomicrurus macclellandi macclellandii (REINHARDT 1844)
Elaps macclellandii REINHARDT 1844
Micrurus macclellandi macclellandi — WELCH 1994: 85
Hemibungarus macclellandi macclellandi — CHAN-ARD et al. 1999: 209
Calliophis macclellandi macclellandi — MURTHY 2010

Sinomicrurus macclellandi nigriventer (WALL 1908)
Calliophis macclellandi nigriventer WALL 1908: 266
Sinomicrurus nigriventer — MIRZA et al. 2020
Calliophis macclellandi — SMART et al. 2021

Sinomicrurus macclellandi univirgatus (GÜNTHER 1858)
Elaps univirgatus GÜNTHER 1858: 231
Calliophis maclellandi univirgatus — KRAMER 1977
Micrurus macclellandi univirgatus — WELCH 1994: 87
Sinomicrurus macclellandi univirgatus — NGUYEN et al. 2009 
DistributionIndia (Assam, Sikkim, Darjeeling; Arunachal Pradesh [A. Captain, pers. comm.], Mizoram), Nepal, Bangladesh, Bhutan,
N Myanmar (= Burma), Thailand, Vietnam,
China (incl. Hong Kong, Hainan, north to Gansu and Shaanxi), Japan (Ryukyu Islands), Taiwan

Type locality: Assam, India

univirgatus: Nepal, India (Sikkim), 1000-2000 m elevation

nigriventer: India (Himachal Pradesh, Uttarakhand); Type locality: “Kasauli”  
TypesHolotype: Unknown (fide LEVITON et al. 2003), ZMUC 65399 (H. V. Kristensen, pers. comm.)
Holotype: CAS 18864 [formosensis]
Holotype: BNHS 342, male (fide Smart et al. 2021); BMNH 1948.1.7.7 (fide Mirza et al. 2020), BMNH 1946.1.17.82 (fide BMNH catalogue), but see Smart et al. 2021 for a discussion [nigriventer]
Syntypes: BMNH 1946.1.17.84, BMNH 1946.1.17.90, collected by B.H. Hodgson [univirgatus] 
DiagnosisDiagnosis. Based on 113 individuals (44 females, 69 males);
comparatively large compared to most species of Sinomicrurus (1) SVL 174–662 mm (mean 436.5 mm) in males (1000 mm total length recorded from Nepal by Shrestha & Majapuri, 1977), 173–714 mm (mean 415.4) in females; (2) temporals 1+1; (3) no teeth behind fangs, palatine teeth 5–8, pterygoid teeth 2–6, and dentary teeth 7–11; (4) supralabials 7/7; (5) infralabials 6/6; (6) postoculars 2; (7) nuchal band width 2–5 scales in males, 2–6 scales in females; (8) black body bands 0–29 in males, 0–35 in females; (9) no blotching between body bands; (10) ventrals 188–233 in males, 196–252 in females; (11) anal divided; (12) paired subcaudals 21–36 in males, 22–34 in females; (13) tail bands 0–5 in males, 0–4 in females; (14) relative tail length (TL/SVL) 0.08–0.15 (mean 0.10) in males, 0.07–0.12 (mean 0.09) in females (Smart et al. 2021).

Comparisons. Sinomicrurus macclellandi, a banded temperate Asian coralsnakes with 13 dorsal scale rows (with S. annularis S. iwasakii, S. peinani, S. macclellandi, S. swinhoei; Table 2); most closely resembles its sister species S. peinani, but differs on the extent of black bands present on head and neck; from S. peinani by black rostralband spilling into frontal (vs. not covering frontal), nuchal-band beginning at parietals or at their edge, usually pointing forward (vs. nuchal-band beginning at middle or posterior margin of mid-dorsal scale located after parietals, curved forward, Figure 8), a more rostral posterior edge of parietal-band at midline (0.44–1.89 vs. 1.86– 2.90, PPB-M-Ratio) and at level of eye (0.67–2.19 vs. 2.04–3.03, PPB-S-Ratio), and by black rostral-band either spilling onto periphery (specimens from Nepal and Sikkim) or extending halfway down frontal (most populations from Assam to Vietnam) (vs. black rostral band only skirting upper boundary of frontal shield, without spilling into it, Figure 8); from all other banded temperate Asian coralsnakes with 13 dorsal scale rows (S. annularis S. iwasakii, and S. swinhoei) differs by a larger overall size (mean SVL >400 mm vs. <400 mm); additionally from S. iwasakii, by having 1+1 temporals (vs. 1+2, at least on one side); less subcaudals (females 22–34 vs. 32–37, males 21–36 vs. 35–40) and tail black bands (females 0–4 vs. 4–6, males 0–5 vs. 4–6); additionally from S. swinhoei, by having less subcaudals (females 22–34 vs. 32–36, males 21–36 vs. 36–41), less body black bands (females 0–35 vs. 26–38, males 0–29 vs. 27–37) and tail black bands (females 0–4 vs. 4–6, males 0–5 vs. 4–7), and a more rostral posterior edge of parietal-band at midline (1.59–2.01 vs. 1.86–2.90, PPB-M-Ratio).
Sinomicrurus macclellandi can be distinguished from other congeners (Tables 4 and 5), from S. kelloggi in having fewer dorsal scale rows (13 vs. 15) and 1+1 temporals (vs. 1+2); from S. boettgeri, S. japonicus, and S. sauteri in lacking mid-dorsal stripe (vs. present); from S. boettgeri and S. sauteri in lacking lateral longitudinal stripes (vs. present); from S. boettgeri and S. sauteri in lacking lateral longitudinal stripes (vs. mid-dorsal and lateral longitudinal stripes present); from S. boettgeri and S. japonicus, in having a pale parietal-band (yellow/ cream/white; vs. absent); from S. kelloggi, S. boettgeri, S. japonicus, and S. sauteri, in lacking maxillary teeth behind fang (0 vs. 2–5); from S. kelloggi and S. boettgeri, in having fewer palatine teeth (5–8 vs. 10–14); from S. boettgeri, S. japonicus, and S. sauteri, in having fewer pterygoid teeth (2–6 vs. 7–16); from S. kelloggi, S. boettgeri, and S. japonicus, in having fewer dentary teeth (7–11 vs. 13–18).
Sinomicrurus macclellandi differs from other congeners in hemipenial morphology (Table 6, Figure 11c,d); from S. annularis in having lobes attenuate (vs. subcylindrical), spines around main body 18–30 (vs. 11–16), organ medium length—organ length/TL 0.21–0.23 (vs. 0.19), lobe length/organ length 0.11–0.15 (vs. 0.23); from S. iwasakii in having lobes attenuate (vs. subcylindrical), main body bulge moderately to highly enlarged (vs. slightly enlarged), organ medium length— organ length/TL 0.21–0.23 (vs. 0.4), basal pocket short—pocket length/organ length 0.17–0.22 (vs. 0.37), retractor muscle anterior vertebral insertion 21–24 (vs. 31–31; subcaudals); from S. peinani in having lobes attenuate (vs. bulbous), spines on main body thin (vs. thick), base with many small spines (vs. many thick but short spines); from S. swinhoei in having sulcus spermaticus centripetal (vs. centrolineal), spines around main body 18–30 (vs. 14–16), organ medium length—organ length/TL 0.21–0.23 (vs. 0.08–0.15); from S. kelloggi in having lobes attenuate (vs. attenuate and horn-like), sulcus spermaticus centripetal (vs. centrolineal), base with many small spines (vs. few small spines), basal pocket shallow to moderately deep (vs. very deep) with bordering lip moderately thin to thick (vs. very thick), main body bulge complete and moderately to highly enlarged (vs. restricted to asulcate side and moderately enlarged) with groove above (asulcate side) shallow to deep and narrow (vs. shallow and broad), organ medium length—organ length/TL 0.21–0.23 (vs. 0.18), length to organ furcation/organ length 0.85–0.86 (vs. 0.56), base length/ organ length 0.37–0.39 (vs. 0.12), lobe length/organ length 0.11– 0.15 (vs. 0.44); from S. boettgeri in having lobes attenuate (vs. subcylindrical), sulcus spermaticus centripetal (vs. centripetal-revolute), base with many small spines (vs. smooth), lip bordering basal pocket moderately thin to thick (vs. very thin), main body bulge complete and moderately to highly enlarged (vs. absent), length to organ furcation/organ length 0.85–0.86 (vs. 0.67–0.75), base length/organ length 0.37–0.39 (vs. 0.44–0.53), basal pocket short—pocket length/ organ length 0.17–0.22 (vs. 0.41–0.47), lobe length/organ length 0.11–0.15 (vs. 0.25–0.33); from S. japonicus in having lobes attenuate (vs. subcylindrical), spines around main body 18–30 (vs. 14), base with many small spines (vs. smooth), basal pocket shallow to moderately deep (vs. extremely shallow to absent) with bordering lip moderately thin to thick (vs. very thin), main body bulge complete and moderately to highly enlarged (vs. absent), length to organ furcation/organ length 0.85–0.86 (vs. 0.68), base length/organ length 0.37–0.39 (vs. 0.56), basal pocket short—pocket length/organ length 0.17–0.22 (vs. 0.39), lobe length/organ length 0.11–0.15 (vs. 0.32); from S. sauteri in having base with many small spines (vs. smooth, very few tiny spines), basal pocket short and shallow to moderately deep (vs. very long and very deep), main body bulge complete and moderately to highly enlarged (vs. absent), organ medium length— organ length/TL 0.21–0.23 (vs. 0.30–0.31), base length/organ length 0.37–0.39 (vs. 0.43–0.53), basal pocket short—pocket length/organ length 0.17–0.22 (vs. 0.36–0.36). The hemipenes of S. macclellandi has been illustrated and described by Lalremsanga and Zothansiama (2015), where they mention no calyces, as we do, but mention that Smith (1943) mentions calyces in the description. We agree with Lalremsanga and Zothansiama (2015) and do not regard as calyces the semi-honeycomb configuration at the base of spines. Other features in these descriptions agree with ours (Smart et al. 2021).

Color. Medium sized snakes with at least one geographic and several chromatic variants, each one with significantly different patterns of black nuchal and muzzle bands, as well as body and tail ornamentation: 1) var typica—black eye band beginning at anterior edge of internasals, completely covering prefrontals and ending more than half way through supraoculars and at half point mark of frontal; black nuchal-band narrowing mesially anteriorly stopping short of making contact with frontal, while extending back the extent of about three scale rows; 2) var. univirgatus—Same as above except lacks black body bands and displays medial black line running along length of the body, usually stopping close to vent; tail with 0–3 bands; 3) var. gorei—differs from both aforementioned variants in absence of black body bands and black dorsal stripe, latter replaced by diminutive black spots down length of the spine; black nuchal-band concave anteriorly (as in S. annularis); 4) var. concolor—differs from above-mentioned variants in lacking dorsal bands or traces of them; 4) var. nigriventer—as var. univirgatus, lacking black body bands, displays a medial black line running along length of the body, having a tail with 3–4 bands, and being characterized by a longitudinal black band encompassing the body venter (Figure 10g,h) (Smart et al. 2021).

Description (nigriventer): Morphologically, it differs from its congeners in having a reddish brown dorsum and a mid-dorsal black vertebral strip extending from the nape to the vent and with three thin bands on the tail, belly white with black smear up to the vent, tail dorsum with a few band-like blotches, and males with 230–235 ventral scales (probably from Mirza et al. 2020).

Original description (nigriventer): “It constitutes a very distinct colour variety for which I propose the name nigriventer. It differs from univirgatus, the variety peculiar to the Eastern Himalayas, in having no trace of black rings, except on the tail where there are three, each being wholly imperfect on the left side. Further there is a continuous broad black irregularly outlined band, running down the middle of the belly. The snake is otherwise exactly similar to univirgatus in colour and lepidosis. The ventrals are 232, and the subcaudals 28.” (Wall 1908: 266). 

Conservation: Calliophis macclellandi iwasakii is classified as “near threatened” in Japan (Ota 2000).

Type species: Elaps macclellandii Reinhardt 1844 is the type species of the genus Sinomicrurus SLOWINSKI, BOUNDY & LAWSON 2001.

Synonymy: for a more extensive and commented synonymy see Smart et al. 2021: 2238. Kaiser et al. 2013 considered the generic name Funkelapidus Hoser 2012 invalid and rejected its use instead of Sinomicrurus. Sinomicrurus nigriventer is embedded within a clade comprising S. peinani, but not S. macclellandi s. l., with a genetic distance of 7–16% from its congeners, hence Mirza et al. 2020 elevated it to full species. However, Smart et al. 2021 argued that molecular data does not support this elevastion and synonymize nigriventer with macclellandi. Given the small number of specimens apparently known from nigriventer, there may not be enough data to fully support either position, so we treate nigriventer as subspecies for the time being.

Subgenera: Wood et al. (2020) established more subgeneric clade names than there are species within Sinomicrurus, hence we do not use them here: Taiwanosinomicrurus, Ravdotosinomicrurus, Ryukyusinomicrurus, Nanyuesinomicrurus, Zonisinomicrurus, Microsinomicrurus, Imalaiosinomicrurus, Megalosinomicrurus, but see Smart et al. 2021: 2259ff for a discussion and definitions.

Subspecies: Sinomicrurus macclellandi iwasakii (MAKI 1935) and Sinomicrurus macclellandi swinhoei VAN DENBURGH 1912 have been elevated to full species by Smart et al. 2021.

NCBI taxonomy ID: 2731268 [nigriventer]

Distribution: for a map of localities see Smart et al. 2021: 2214 (Fig. 1). 
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