Ophiophagus hannah (Cantor, 1836) s. str.

Figs 2–5, 7, 9A–C, 10A, 11A, 12A

Hamadryas hannah Cantor, 1836: 87 .

Hamadryas ophiophagus Cantor, 1838: 187 (replacement name for Hamadryas hannah).

Naia Vittata Elliot, 1840: 41 (sic, for vittata).

Dendraspis hannah sinensis Deraniyagala, 1960: 62 .

Dendraspis hannah nordicus Deraniyagala, 1961: 229 .

Dendraspis hannah brunnea Deraniyagala, 1961: 229 .

Common name

Northern king cobra.

Diagnosis (redefined herein)

A species of Ophiophagus inhabiting the Sub-Himalayas, eastern India, Myanmar, Indochina, southwards to the Isthmus of Kra, showing the following combination of characters: dark-edged yellow bands along body of adults (vs unbanded in adult O. salvatana Gowri Shankar, Das & Wüster sp. nov.; lacking dark edges to the pale bands in O. kaalinga Gowri Shankar, Das & Ganesh sp. nov. and, if present, in O. bungarus). Further, O. hannah differs from all other congeners in having a high pterygoid tooth count of 18–21 (vs 11 in O. bungarus and O. salvatana and 12 in O. kaalinga). Finally, juveniles of O. hannah, with 27–48 body bands, differ from O. salvatana (85–86) and O. bungarus (57–87); relative tail length ranging 21.7–26.4% with a mean of 24.05% (vs 19.3–25.1% [22.2%] in O. bungarus; vs 18.0–19.9% [18.95%] in O. kaalinga; vs 18.7–23.0% [20.85%] in O. salvatana).

Etymology

The source of the epithet hannah was not mentioned by Cantor (1836, 1838). More recently, Adler (2016) argued the same to be an eponym for Hannah Sarah Wallich (1820–1893), eldest daughter of Cantor’s host and uncle, the celebrated botanist (and physician), Nathaniel Wallich (1786–1854), during his time in Calcutta, and at the time of description of the species.

Type material

Neotype (designated herein)

INDIA • ♀; West Bengal State, Kolkata, Howrah, Acharya Jagadish Chandra Indian Botanical Gardens; 22.55° N, 88.29° E; John Anderson leg.; ZSI 8292.

Remarks

The type series comprising four syntypes, of Hamadryas hannah, reported from “Sunderbuns” and “jungle not far from Calcutta”, being lost or untraceable now (not mentioned in Boulenger 1896; Das et al. 1998), along with most of Cantor’s zoological specimens (see Adler 2016), is the cause of some confusion, both in the past and presumably in the future. Indeed, there is no mention of the location of the types in other works that enumerate repositories of Asian snake types (e.g., Smith 1943; Golay 1985; Toriba 1993; David & Ineich 1999; McDiarmid et al. 1999; Iskandar & Colijn 2000).

Since it is evident that the type series of Cantor’s Hamadryas hannah is currently not extant (Article 75.3.4), we invoke Article 75 of the Code (ICZN 1999) to designate ZSI 8292, an adult collected by John Anderson, Superintendent of the Indian Museum from the Indian Botanical Gardens, Howrah, Kolkata, India, as the neotype of Ophiophagus hannah, in the context of the revision of the genus, in order to clarify the taxonomic status of the species (Article 75.3.1) and describe the characters of the specimen (Article 75.3.2), the data and description of the same unequivocally diagnosing the taxon from its congeners (Article 75.3.3). The neotype matches the original description in all observed morphological characteristics (Article 75.3.5), and is drawn from the general geographical area of the type locality, in the north-western suburb of the township of Shibpur, ca 10 km from Kolkata (Article 75.3.6) and preserved in a globally-recognised systematic collection that dates back to the late 1800s (Article 75.3.7).

The members of the genus are widespread and appear morphologically conservative, with incorrect application of names for over a century and the possibility of parapatry or sympatry (such as in fragmented areas of central India, as well as in southern Thailand), and more intense sampling may reveal the existence of additional cryptic lineages within the complex. This is likely within the vast geographical range of O. hannah, as restricted here, as well as in the extreme eastern range of the group (Sulawesi) and the southern islands of the Philippines archipelago, as new material, including genetic samples, becomes available. Despite the long history of collections and research in the southern Bengal region (Das 2004), it is remarkable that no other topotypical specimen exists in a systematic institution, further necessitating the need to draw attention to this significant specimen (ZSI 8292).

On the identity and provenance of Naia vittata

Elliot (1840) erected the nomen Naia vittata based on a stuffed adult (see Günther 1858), a non-skeletal specimen sourced from southern India (Fig. 5). This nomen was however short-lived, as Cantor (1847) went on to synonymise Elliot’s nomen under his own Hamadryas ophiophagus Cantor, 1838: the synonymy has been accepted by subsequent authors. Elliot (1840) in his original description states that the type specimen was given to him by a fisherman who found the live snake contained inside a floating box carried by a catamaran in mid-sea, off the coast of “St. Thome” (currently, Santhome, within Chennai City, Tamil Nadu State, southern India: 13.0303° N, 80.2788° E, 7 m a.s.l.). Thus, its true precise provenance remains unknown. The only other locality mentioned in the original description is Guindy (also in Chennai City: 13.0105° N, 80.2156° E; 14 m a.s.l.). By all evidence, there has never been any wild population(s) of Ophiophagus in either of the two localities mentioned in Elliot’s description (Murthy 1978; Whitaker & Captain 2004). When Deraniyagala (1960) reinstated this nomen as a trinomial for a subspecies, he conferred it to the Western Ghats population. Our studies show that the description and figure of the holotype of Naia vittata are not in conformity with the Western Ghats population, in that the two-scale wide pale bands, distinct black borders of the pale bands and the progressive darkening of the dorsum posteriorly are unique to this nominal species. Thus, taking into account the morphology and the most plausible geographic scenario (see discussion), we treat Naia vittata Elliot, 1840 as a junior subjective synonym of Hamadryas hannah .

Material examined

BHUTAN • 1 ♀; Ronglong, Manas Valley; 26.79° N, 90.94° E; ZSI 22483 .

CHINA (including Hong Kong) • 1 spec., holotype of Dendraspis hannah sinensis Deraniyagala, 1961; Fujian Province “ Guling-Gu Shan ”, near Fuzhou Shi; 26.10° N, 119.28° E; AMNH 29944 • 1 ♀; New Territory, Hong Kong, Tai Po; 22.30° N, 113.95° E– 22.45° N, 114.17° E; BMNH 1983.273 • 1 spec.; Guizhou Province Kowloon; 22.32° N, 114.18° E; BMNH 1955.1.4.69 • 2 specs; Xiuwen; 26.83° N, 106.59° E; CIB 21001, CIB 210002 • 1 ♂; Guizhou Province Xingyi; 25.09° N, 104.89° E; CIB unregistered • 3 specs; Hong Kong Lantau Island i.e., Tai Yue Shan; 22.25° N, 113.93° E; MCZ 176539, MCZ 176540, MCZ 177105 • 1 spec.; Hong Kong New Territories, W slope of Tai Mo Shan; 22.42° N, 114.12° E; MCZ 176365 .

INDIA • 1 ♀; Meghalaya State, Shillong’; 25.58° N, 91.89° E; BMNH 1907.12.16.21 • 1 ♀; same data as for preceding; ZSI 21703 • 1 ♂, holotype of Naia vittata Elliot, 1840; Tamil Nadu State, southeastern India in Mylapore, coastal Chennai “St. Thome”; 13.03° N, 80.28° E; BMNH 1996.451 • 1 ♂; “ Assam ” [no further locality]; BNHS 2274 • 1 spec.; Assam State, Kokrajhar, Kachugaon; 26.38° N, 90.903° E; BNHS 2275 • 1 ♀, paratype of Dendraspis hannah brunnea Deraniyagala, 1961; West Bengal State, Tindharia; 26.85° N, 88.33° E; BNHS 2276 • 1 spec.; Sikkim State, Gangtok; 27.33° N, 88.612° E; BNHS 2270 • 1 spec.; Uttarakhand State, Nainital, Patwadangar; 29.38° N, 79.50° E; BNHS 2278 • 1 spec.; West Bengal State, Duars, ca 30 km area of floodplains in Himalayan foothills north of Brahmaputra; BNHS 2267 • 1 spec; South Andaman Island; ca 11.61° N, 92.61° E; CSPT/S-56 • 1 spec.; Odisha State, Bhitarkanika National Park; ca 20.75° N, 87.00° E; MCBT 152886 • 1 spec.; same data as for preceding; NKBP unreg . • 1 spec.; West Bengal State, Kalimpong, Tarkhola; 27.07° N, 88.48° E; MCZ 58258 • 1 spec.; Arunachal Pradesh, Itanagar; 27.07° N, 93.59° E; SFRI A-4 • 1 spec.; South Andaman Island, Port Blair [currently, Sri Vijaya Puram]; 11.62° N, 92.72° E; USNM 129748 • 1 ♂; South Andaman Island, Mathura; 11.72° N, 92.69° E; ZSIP 366 • 1 spec.; Uttarakhand State, Mussoorie; 30.45° N, 78.08° E; ZSI 12556 • 1 ♂, holotype of Dendraspis hannah brunnea Deraniyagala, 1961; West Bengal State, Darjeeling; 27.05° N, 88.3° E; ZSI 8294 .

MYANMAR • 1 ♀; Bago Division, Bago or Pegu; 17.33° N, 96.48° E; BMNH 68.4.3.31 • 1 spec.; Shan State, Shweli, in a river; 23.67° N, 96.42° E; collection locality presumably in basin; BMNH 1925.12.22.18 • 1 ♀; Shan State, Mogok; 23.00° N, 96.42° E; BMNH 1900.9.20.19 • 1 spec.; Kachin State, Sumprabum Triangle; 26.55° N, 97.57° E; BMNH 1940.6.5.63 • 1 spec.; Magway Division, Thayet-myo; 19.32° N, 95.18° E; BNHS 2271 • 1 ♀; Sinlum Kaba or Sinlumkaba; ca 24° N, 97° E; BNHS 2273 • 1 spec.; Ayeyarwaddy Division, vicinity of Mwe Hauk village; 16.28° N, 94.77° E; CAS 206601 • 1 ♂; Mandalay Division, Mount Popa; 20.92° N, 95.25° E; MCZ 44699 • 1 ♀; Mandalay Division, Popa Mountain Park, Kyauk Pan Tawn; 20.87° N, 95.24° E; CAS 214017 .

NEPAL • 1 spec.; Bagmati Province, Kathmandu; 27.72° N, 85.32° E; BNHS 2268 • 1 spec.; Bagmati Province, ca 14.5 km W of Hetauda, Rapti Bridge; ca 27.43° N, 85.00° E; BNHS 2269 .

THAILAND • 1 spec.; Amphoe Pak Thong Chai, Nakhon Ratchasima Province [= Khorat Province], Sakaerat Environmental Research Station; 14.51° N, 101.93° E; FMNH 180215 • 1 spec.; Prachuap Khiri Khan Province, Koh Lak; 11.82° N, 99.80° E; BMNH 1969.1927 • 1 spec.; same data as for preceding; NRM 2532 • 1 spec.; Lampang Province, northern Thailand, Lakon, Lampang; 18.25° N, 99.50° E; BMNH 1921.4.1.44 • 1 spec.; Pak Nam Pho; 15.71° N, 100.13° E; BMNH 1968.836 • 1 spec.; north-western Thailand, Mae Hong Son Province; 19.25° N, 98.00° E; BMNH 1987.1160 • 1 spec.; Kanchanaburi Province, Sai Yok; 14.12° N, 99.14° E; BMNH 1987.1162 • 1 spec.; northern Thailand, Chiang Mai Province, Doi Suthep; 18.82° N, 98.89° E; FMNH 178405 • 1 spec.; Bangkok Province, Near Bangkok; ca 13.75° N, 100.52° E; MCZ 20331 • 1 spec.; Chiang Mai Province, Mount Angka, possibly Doi Angkei or Doi Angkhang; 19.00° N, 98.67° E; MCZ 20331 • 1 spec.; Khon Kaen Province, Khon Kaen, near Pong Neep Dam; 16.77° N, 102.59° E; TNRC 1120 • 1 spec.; Tak Province, Doi Hua Mod; 15.97° N, 98.85° E; USNM 101040 • 1 spec.; Nakhon Ratchasima Province, Pak Chong or Pak Jong; 14.75° N, 101.42° E; USNM 72726 .

VIETNAM • 1 ♀; Dac Lak Province, Buon Ma Thuot City or Lac Giao; 12.67° N, 108.05° E; FMNH 26475 • 1 ♀; same data as for preceding; MCZ 43744 • 1 spec.; Thuan Pho Ho Chi Minh Province, Ho Chi Minh City or Saigon; 10.75° N, 106.67° E; LSUMZ 34904; 1 spec.; same data as for preceding; MNHN 1975.127 • 2 specs; Ba Ria-Vung Tau Province, Vung Tàu or Cap Saint-Jacque; 10.35° N, 107.07° E; MNHN 1911.66, MNHN 1920.213 • 1 ♀; Lao Cai Province, Cha Pa or Chapa; 22.33° N, 103.83° E; MNHN 1935.108 • 1 spec.; “Cochinchine”, [ currently interpreted as south of Gianh River (Sông Gianh), southern Vietnam, no further data]; MNHN 1912.75 • 3 ♂♂; same data as for preceding; MNHN 3189, MNHN 7671, MNHN 5205 .

Skeletal material

THAILAND • 1 spec.; Nakhon Ratchasima or Khorat Province, Pak Chong or Pak Jong; 101.42° E, 14.75° N; USNM 72726 .

VIETNAM • 1 spec.; Lam Dong Province, Bao Loc; 11.42° N, 107.67°E; USNM 95105 .

Description of neotype (ZSI 8292)

Venter with an incision, appears to be the specimen mentioned in Anderson (1871).

MEASUREMENTS. SVL 2085 mm, TL 462 mm, total 2547 mm.

HABITUS. Body relatively robust (midbody width 38.3 mm, 1.8% SVL), triangular in cross-section; transverse scales on body: DSR1 17; DSR2 15; DSR3 15; ventrals 240; subcaudals 91; cloacal 1; dorsal scales smooth, the vertebral and outer two rows enlarged; ventral scales smooth; first four subcaudal scutes entire; tail short (22.1% SVL), cylindrical, tapering posteriorly.

HEAD. Head relatively large, head length 46.3 mm, head width 29.4 mm; head depth 25.8 mm; distinct from neck, flattened in the orbital region, rounded in the sagittal region, with a slight depression medially, snout projecting slightly beyond mandible; canthus rostralis sharply defined; eye diameter 7.4 mm; interorbital distance 19.3 mm; cephalic scales juxtaposed, smooth-edged, except parietals and occipitals, which are slightly imbricate; rostral trapezoid in shape, distinctly visible from above, over twice as long as wide, concave ventrally, rostral width 5.2 mm; rostral length 12.2 mm; eye to snout distance 15.0 mm; eye to nostril distance 7.0 mm; nostril diameter 5.3 mm; internasals large, subtrapezoidal, wider than long; internasal suture width 19.3 mm; preocular squarish, wider than high, separated from internasal by prefrontal; prefrontals trapezoid, wider than long; frontal trapezoidal, contacting prefrontals, supraoculars and parietals; frontal edge sinuous, short-sided posteriorly; supraocular subtrapezoidal, contacting prefrontal, frontal, parietal, orbit, preocular, upper postocular and two temporals; large paired occipitals, occipital length 13.0 mm; interoccipital scale present behind the suture between parietals; temporals 2/2 (L/R); in anterior pair, lower temporal larger than upper; in posterior pair, upper temporal longer than lower; first row of nuchals slightly enlarged compared to rest of dorsals; supralabials 7/7; III–IV (L/R) touching the eye; III (L/R) contacting preocular and I (L/R) contacting posterior nasal; supralabial I low; II high; III tallest; II and IV subequal; and VI and VII low, narrow and elongate; nostril lateral at posterior of a single concave nasal, horizontally elliptical, its greatest diameter along vertical axis; one preocular and three postoculars; eye large, contained in head length 0.16 times and head depth 0.29 times; pupil rounded; ocular ring (sensu Marx et al. 1987) comprising seven scales: one preocular, three postoculars, one supraocular and two supralabials; suboculars absent; mental small, triangular, wider than deep (mental width 7.8 mm; mental depth 2.4 mm); infralabials 8/8 (L/R), first (L/R) contacting anterior genials; infralabial I contact anterior genial; infralabial IV (L/R) contact posterior genials; infralabial IV largest; cuneate scute on lower jaw absent; two elongate gular scales follow posterior mental; the anterior longer than posterior.

DENTITION. Maxillary teeth recurved and stout, not compressed, gradually increasing in size posteriorly; fang length 4.0 mm; fang width at base 2.0 mm; tooth count obscured by gingivae.

COLOURATION. After over a century in preservative, neotype dorsum mahogany brown, almost band-less anteriorly with traces of mottled reticulation of dark-edged pale bands that become more visible from midbody to near tip of tail; venter similarly coloured as of dorsal ground colour, except posterior edge of each scute edged with pale grey; scutes of head midbrown, those on forehead distinctly edged with dark grey.

HEMIPENIS. Examined in SFRI A-4. Organ everted; distinctly bilobed, rather long and slender, its lengths (L/R) 97/ 85 mm and widths (L/R) 5.5/ 4.5 mm; extending up to 15/10 subcaudal scale; divided at level of first subcaudal scale; lobe head flat and rather circular near apex, with numerous, tiny spinules and flounces; asulcate side relatively less spinose compared to sulcate side; basal part of pedicel with dense congregation of large spines; bilobed tubular length of organ with series of circular whorls of spiny protuberances; sulcal lips not evident, sheath-like, completely surrounding apex base; sulcus spermaticus not evident.

Morphological variation

Six of 43 specimens (14%) examined have an interoccipital scute. The occurrence of divided and undivided subcaudals in members of the complex has been reported in the literature (e.g., Smith 1943: 436). All 43 specimens examined for this character have some undivided subcaudals, typically, the first and in succeeding subcaudals up to subcaudal 26, in what appears to be an irregular pattern, including about a third undivided subcaudals from Subcaudal 18 to 32, again without an apparent pattern that cannot be linked to either sex or geographical distribution. The total pale body band count made on 41 individuals was 27–48 (mean 39.6). Dorsum of more recently collected material examined is a shade of midbrown, Fawn Colour #25, forehead paler, Drab #27, interscale areas darker along body; distinct pale bands, Buff #124, numbering 27–48, each 2–3 scales wide, edged with darker grey, Blackish Neutral Gray #82 areas, about a scale wide, bands separated from each other by five scales; on nape region, the first two pale bands form distinct chevrons, the rest of the body bands less angular in adults; bands sometimes broaden on lower flanks; the largest scales of forehead, frontal, parietals, occipitals, upper temporals and postoculars with a distinct dark edge. Tail and posterior fourth of body nearly black, with 1–2 ivory-coloured scales on tail, separated from each other by 5–6 scales. Some large individuals are grey-black, the interscale regions appearing paler. Mandible and genial region Cream Colour, #54; the gular region to until Ventral 13–27 bright yellow, Warm Buff, #118, sometimes reported as a shade of orange, with two dark areas peripherally, comprising somewhat indistinct bands meeting Ventrals 13–27; and noticeably in larger, hooding individuals. Abdominal region as in gular region, but progressively darkening. Subcaudal region with pale scales, obscurely darkened throughout and with dark edges. Pupil rounded, black, iris brownish-red, Kingfisher Rufous #240, darkening peripherally, with a narrow, yellow ring. Tongue is blackish-grey, the oral cavity pink. Juveniles have ivory white bands, Buff, #124, including four on the head region, the first one in rostral position, a broad one covering rostral, as well as part of Supralabial 1, nasals, and internasals; the second in preocular position, narrower, and covering part of the prefrontals, preoculars and Supralabial III; the third one in postocular position, narrowest of the four bands, and covering part of the frontal and parietals, and comprising large, elongated marks arranged in a transverse series; and the fourth one on cephalic position, broader than the previous one, and covering part of the parietals and occipitals, and comprising large oval spots arranged in a transverse series. Body bands are of the same colour, about two scales in width, and number 35–48 between head bands and the caudal region above vent, and have 4–5 scale wide interband areas. Bands on tail relatively thick, with darker edges, both dorsally and ventrally; and the posterior third of body appears darker than the rest of the body.

Sample size and condition do not permit analysis of ontogenetic change of body bands, although the adult bands do appear to show reduced contrast on the dorsum, compared to hatchlings, and pale markings are absent on the forehead of adults. Individuals (n = 12) that permitted counts of tooth sockets have dentition thus: 15–16 dentaries; 18–21 pterygoids and 3 palatines.

Distribution

The geographical distribution of the nominate species, as restricted here, extends from extreme eastern Pakistan, across the sub-Himalayan region of Kashmir, northern India, Nepal, Bhutan, Tibet, and south to the Godavari-Mahanadi-Ganges deltas of the Circar Coast in eastern India, east to the eastern coast of China, including Hong Kong, the range extending south to Indo-China, including Myanmar, Laos, Vietnam, Cambodia and parts of Thailand, presumably north of the Isthmus of Kra (Pope 1935; Das 1999; Leviton et al. 2003; Whitaker & Captain 2004 [in part]; Ahmed et al. 2009; Nguyen et al. 2009; Sharma et al. 2013; Chandra et al. 2014; Ahsan et al. 2015; Faiz et al. 2017; Dolia 2018; Tshewang & Letro 2018; Francis 2021; Koirala et al. 2021). The records from the extreme west of the distributional range (including Lahore and Palanpur, in Pakistan) were considered by Wall (1928) to be introductions by snake charmers or the result of transport along rivers. However, old records of the occurrence of the snake in the extreme eastern part of Pakistan, discussed in this paper, appear to be reliable, although it is unknown if populations persist at present.