Ophiophagus hannah, s. str.

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 •