Ophiophagus bungarus ( Schlegel, 1837 ) Das & Shankar & Swamy & Williams & Lalremsanga & Prashanth & Sahoo & Vijayakumar & Höglund & Shanker & Dutta & Ganesh & Wüster, 2024

Das, Indraneil, Shankar, P. Gowri, Swamy, Priyanka, Williams, Rhiannon C., Lalremsanga, Hmar Tlawmte, Prashanth, P., Sahoo, Gunanidhi, Vijayakumar, S. P., Höglund, Jacob, Shanker, Kartik, Dutta, Sushil K., Ganesh, S. R. & Wüster, Wolfgang, 2024, Taxonomic revision of the king cobra Ophiophagus hannah (Cantor, 1836) species complex (Reptilia: Serpentes: Elapidae), with the description of two new species, European Journal of Taxonomy 961 (1), pp. 1-51 : 19-24

Ophiophagus bungarus ( Schlegel, 1837) comb. nov.

Figs 6 , 9D–F , 11B , 12B , 13B , 14

Naja bungaroides Boie, 1828a: 250 (nom. nud.).

Naja bungaroidea [sic] Wagler, 1830: 342.

Hoplocephalus bungaroides Wagler, 1830: 342 , by implication.

Naja bungarus Schlegel, 1837: 184 .

Naja ingens van Hasselt, 1858: 141 .

Dimeresurus boiei Bleeker, 1858: 263 (nom. nud.).

Dendraspis hannah borneensis Deraniyagala, 1960: 61 .

Nomenclatural comment

Due to the impending Case before the International Commission of Zoological Nomenclature and to future-proof the binomen of this species, we have not used Wagler’s name for this species, but opted for the next-oldest available name, Naja bungarus Schlegel, 1837 .

Common name

Sunda king cobra.

Diagnosis

A species of Ophiophagus inhabiting Malesia, south of the Isthmus of Kra (southern Thailand, peninsular Malaysia, Sunda Archipelago, east to parts of southern and central Philippines) and showing the following combination of characters: mostly unbanded in large individuals, occasionally with narrow pale bands, lacking darker edges, along body of adults (vs unbanded in adult O. salvatana sp. nov.; dark-edged pale band in O. hannah ). It differs from O. kaalinga sp. nov. in having a brownish-yellow to mahogany dorsum, with or without (in large adults) slightly paler, narrow bands (vs dorsum dark grey with yellow bands that expand on lower flanks); dorsum ground colour showing little to no contrast at the meeting point of ventral scales at gular region (vs with a clear line of separation from ventrals in gular region). The higher body band counts (57–87) of juvenile O. bungarus separate it from those of O. hannah (27–48), and O. kaalinga (28–48). The pale body bands of juvenile O. bungarus are more rounded, the dark interband areas covering 4–9 scales ( Fig. 12B ), whereas they are extremely angular in O. salvatana , with the dark interband areas covering 2–3 scales ( Fig. 12D ); relative tail length ranging vs 19.3–25.1% with a mean of 22.2% (vs 21.7–26.4% [24.05%] in O. hannah ; vs 18.0–19.9% [18.95%] in O. kaalinga ; vs 18.7–23.0% [20.85%] in O. salvatana ). Finally, O. bungarus differs from O. hannah in having a lower pterygoid tooth count of 11 (vs 18–21).

Etymology

The specific epithet probably alludes to either morphological (partially undivided subcaudals) or behavioural (ophiophagous) characters of kraits (genus Bungarus ).

Material examined

Lectotype

INDONESIA • ♂, hatchling; Java, Java Barat Province, Tjihao or Cihoe or Tji Hoe ; 6.43° S, 107.13° E; Apr. 1827; Heinrich Boie (1784–1827) and Heinrich Christian Macklot (1799–1827) leg. [see Troelstra 2016]; RMNH 1334 .

Schlegel (1837) mentioned two specimens in his description of Naja bungarus : Boie’s specimen (extant as RMNH 1334), and a specimen from Sumatra of unconfirmed whereabouts, but possibly RMNH 1338 (Wolfgang Denzer and Esther Dondorp in litt.; Wolfgang Wüster pers. obs.). Both of these are currently syntypes of the name. We hereby designate the extant specimen RMNH 1334 as the lectotype of Naja bungarus Schlegel, 1837 as per Art. 74 of the Code.

Other material examined

BRUNEI DARUSSALAM • 1 spec.; Dewan Museum , Kota Batu, Bandar Seri Begawan; 4.89° N, 114.97° E; BMNH 2.1979 • 1 ♂; Bandar Seri Begawan , Kampung Pinto Halim; 4.89° N, 114.97° E; BMNH 224.1991 • 1 ♂; Bandar Seri Begawan , Kampung Beribi, Gadong; 4.89° N, 114.89° E; BMNH 6.1998 • 1 spec.; Bandar Seri Begawan , Jalan Muara, Forest Hill; 4.90° N, 115.00° E; LSUMZ 55839 • 1 spec.; Bandar Seri Begawan , Kota Batu; 4.88° N, 114.97° E; BMNH 1984.80 .

INDONESIA • 1 spec.; Jawa Barat, Pasir Banggoer Krawang; 6.42° S, 107.67° E; MZB 376 • 1 spec.; Jawa Barat, Batavia or Jakarta; 6.17° S, 106.78° E; MZB 368 • 1 spec.; Jawa Barat, Muara Karang; 6.12° S, 106.78° E; MZB 2206 • 1 spec.; Jawa Barat, Jakarta, Antjol or Ancol; 6.13° S, 106.83° E; MZB 402 • 1 spec.; Jawa Barat, Indramayu or Indramaju; 6.32° S, 108.32° E; MZB 399 • 1 spec.; Borneo , Kalimantan, Timur Province, Balikpapan; 1.24° S, 116.85° E; BMNH 1912.6 .26.21 • 1 spec.; Borneo, Berau , Kalimantan Timur Province, Sambulajan, Sungei Segah; 2.17° N, 117.50° E; MZB 1339 • 1 spec.; Borneo , Central Kalimantan Province, Purukcahu or Puruktyau or Purukt Jahu; 0.58° S, 114.58° E; RMNH 7546 • 1 spec.; Borneo , East Kalimantan Province, Sebulu; 0.27° S, 117.00° E; USNM 200500 • 2 ♂♂; Sumatra, Bukit Tinggi or Fort de Kock ; 0.32° S, 100.37° E; BMNH 1928.2 .8.43, BMNH 1928.2 .18.43 • 1 spec.; Sumatra, Siulakderas, Kerinchi or Korinchi; 1.92° S, 101.30° E; BMNH 1915.12 .2.39 • 1 spec.; Sumatra, Langkat; 3.87° N, 98.31° E; RMNH 6351 • 1 spec.; Sumatra, Sinabang, Simeulue Island ; 2.47° N, 96.37° E; RMNH 5192 • 1 spec.; Sumatra, Tarussan Bay or Teluk Tarusan; 1.2167° S, 100.42° E; USNM 35763 • 2 ♂♂; Sumatra, Labuhandeli or Deli Belawan; 3.75° N, 98.68° E; ZMA 13487 , ZMA 17476 • 1 ♂; Sumatra, Klein Sungei, Karang Galang; 1.15° N, 104.18° E; ZMA 11235 • 1 spec.; Sumatra, Aceh Special District , Padang Miangatas Gunung Jago; 4.00° N, 97.06° E; MZB 401 • 1 spec.; Sumatra, Bengkulu Province, Kepahiang; 3.65° S, 102.57° E; USNM 070972 • 1 ♂; Sumatra, Bengkulu Province, Kepahiang; 3.65° S 102.57° E; ZRC 2.3212 • 1 spec.; Riau Archipelago , Pulau Galang; 0.75° N, 104.23° E; ZRC 2.3254 - 6 • 1 spec.; Bangka Belitung Province, Pulau Bangka, Pangkal Pinang; 2.12° S, 106.12° E; MZB 369 • 1 spec.; Bangka Belitung Province, Belitung, Kampung Rembikang, Tanjung Pandan; 2.75° S, 107.65° E; MZB 373 .

MALAYSIA • 1 spec.; West Malaysia , Perak, Pulau Dinding; 4.23° N, 100.56° E; BMNH 1903.6 .13.3 • 1 spec.; West Malaysia, Perak, in Gunung Korbu ; 4.68° N, 101.30° E; BMNH 1957.1 .11.40 • 1 ♂; West Malaysia , Selangor; 3.51° N, 101.50° E; IMR uncatalogued • 2 ♂♂; West Malaysia, Selangor, Kuala Langat, Bukit Mandol Forest Reserve ; 2.95° N, 101.53° E; IMR 110487 , IMR 5063 • 1 spec.; West Malaysia , Pulau Pinang; 5.40° N, 100.23° E – 5.73° N, 103.00° E; MCZ 944 • 1 ♂; West Malaysia , Kuala Lumpur; 3.17° N, 101.70° E; MNHN 1899.172 • 1 spec.; East Malaysia ( Borneo ), Sarawak, Kuching; 1.55° N, 110.33° E; FMNH 67282 • 1 spec.; same data as for preceding; SM uncatalogued • 2 ♂♂; same data as for preceding; SM 5.45 .25, SM 5.45 .2 • 1 ♂; East Malaysia (Borneo) Sarawak, Subis Forest Reserve , Niah National Park ; 3.80° N, 113.80° E; FMNH 128275 • 1 ♂; East Malaysia (Borneo), Sarawak, Bintulu Division , Labang Camp on Sungei Serin ; 1.27° N, 110.45° E; FMNH 150891 • 1 spec.; East Malaysia (Borneo), Sarawak, Sungei Baram ; 4.58° N, 113.97° E [collection locality presumably upriver]; FMNH 71659 • 1 spec.; East Malaysia (Borneo), Sarawak, Sibu ; 2.30° N, 111.81° E; SM 5.45.2C • 1 ♂; East Malaysia (Borneo), Sabah, Lahad Datu , Danum Valley Field Centre; 5.02° N, 117.78° E; FMNH 233152 • 1 ♂; East Malaysia ( Borneo ), Sabah, Sandakan; 5.87° N, 118.07° E; FMNH 63566 • 2 ♂♂; East Malaysia , Sabah; 5.02° N, 117.78° E; SSM 0109 , SSM 0096 • 1 ♂; East Malaysia ( Borneo ), Sabah, Sandakan; 5.87° N, 118.07° E; SSM 0360 • 1 ♂; “ Malaysia ” accidentally imported with logs; MCBT 152883 .

PHILIPPINES • 1 ♂; 10 km SSW of Iwahig ; 9.740° N, 118.66° E; CAS 129629 • 1 spec.; same data as for preceding; CAS 157468 • 1 spec.; Palawan, Tagup River, Brooke’s Point, Borangbato , Mainit ; 8.82° N, 117.78° E; NMP 2231 .

SINGAPORE • 1 ♂; “Singapore”, no further data; BMNH 82.11 .29.1 • 1 ♂: “Singapore”, no further data; CAS 16785 • 1 spec.; Mandai Lake Road ; 1.42° N, 103.75° E; ZRC 2.2301 .

THAILAND • 1 spec.; Yala Province, Betong ; 5.75° N, 101.08° E; BMNH 1938.8 .7.60 • 1 spec.; Pattani Province, Pattani Na Prado ; 6.68° N, 101.14° E; CUZM unreg . • 1 spec.; Nakhon Si Thammarat Province, Nakhon Si Thammarat; 8.43° N, 99.97 °E; TNRC 1124 • 3 specs; Pattani Province, Pattani Na Prado ; 6.68° N, 101.15° E; TNRC 1126 , TNRC 1128 , TNRC unreg . • 1 spec.; Trang Province, Trang; 7.56° N, 99.60° E; USNM 023014 • 1 spec.; Songkhla Province, Singora ; 7.08° N, 100.50° E; USNM 079533 • 1 spec.; Trang Province, Trang [presumably also south of Isthmus of Kra ]; 7.56° N, 99.60° E; USNM 220395 • 2 specs; “ Southern Thailand ”; AMNH 10046.1 , AMNH 10046.2

Skeletal material

THAILAND • 1 spec.; Trang; 7.58° N, 90.50° E; USNM 220395 .

Description of the lectotype ( RMNH 1334)

MEASUREMENTS. SVL 496 mm, TL 123 mm, total 619 mm.

HABITUS. Body relatively robust (midbody width 10.4 mm, 2.1% SVL), triangular in cross-section; Transverse body rows: DSR1 19; DSR2 15; DSR3 15; ventrals 253; subcaudals 108; supralabials 7; infralabials 8; anterior temporals 2; posterior temporals 2; cloacal 1; dorsal scales smooth, the vertebral and outer two rows enlarged; ventral scales smooth; subcaudals 1–28 entire, the rest divided; tail short (24.8% SVL), cylindrical, tapering posteriorly.

HEAD. Head relatively large, distinct from neck, head length 17.5 mm, head width 9.8 mm; head depth 7.3 mm; 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 width 4.3 mm; interorbital distance 8.1 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 0.8 mm; rostral width 1.5 mm; rostral length 4.4 mm; eye to snout distance 5.7 mm; eye to nostril distance 2.1 mm; nostril diameter 1.5 mm; internasals large, subtrapezoidal, wider than long; internasal suture width 1.0 mm; preocular squarish, wider than high, making a narrow contact with internasal; prefrontals trapezoid, wider than long; frontal trapezoidal, contacting prefrontals, supraoculars and parietals; frontal edge slightly sinuous, short-sided posteriorly; supraocular subtrapezoidal, contacting prefrontal, frontal, parietal, orbit, preocular, upper postocular but not the temporals; large paired occipitals; occipital length 3.5 mm; interoccipital scale present behind the suture between parietals; temporals 2/2 (L/R); in anterior pair, lower temporal broader but shorter 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) contact spectacle; III (L/R) contacting preocular and I and II (L/R) contacting posterior nasal; Supralabial I low; II high; III tallest; II and IV subequal; and VI and VII low, narrow and elongate; Supralabial IV does not contact preoculars; 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 0.24 times and head depth 0.6 times; pupil rounded; ocular ring comprises seven scales- one preocular, three postoculars, one supraocular and two supralabials; suboculars absent; mental small, triangular, wide than deep (mental width 0.43 mm; mental depth 0.14 mm); infralabials 8/8 (L/R), first (L/R) contacting anterior genials; infralabial IV (L/R) contact posterior genials; infralabial IV largest; one pair of genials; 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 decreasing in size posteriorly; fang length 1.7 mm; fang width at base 0.5 mm; tooth count obscured by gingivae.

COLOURATION. The dorsum of the lectotype, a hatchling, is brownish-grey; cephalic and body bands ivory yellow, numbering 65 up to beginning of tail; throat and rest of venter unpatterned pale yellow, except the slightly darker edges of the ventral scales.

Morphological variation

Fourteen of 59 specimens (23.7%) examined have an interoccipital scute. Divided into geographical regions, the lowest occurrence of specimens with interoccipital scales were from Borneo and Palawan (11.5%; n = 26), followed by the Malay Peninsula (28.6%; n = 18) and Sumatra and Java (46.6%; n = 15).

Of 59 specimens examined for subcaudal imbrication character, 26 have one series of undivided subcaudals, in what appears to be an irregular pattern, the rest with up to eight series of undivided subcaudals, ranging from a single scute to 26 scutes, without an apparent pattern that cannot be linked to either sex, ontogeny or geographical distribution. Hatchlings and young adults bear multiple, narrow pale bands on body, pale body bands occasionally retained in presumed adult (> 2000 mm) individuals. Specimens with intact dentition or those permitting counts of tooth sockets have dentition thus: 11–15 dentary sockets; 11–11 pterygoid sockets and 3 palatine sockets.

Dorsum is a shade of brown, dorsal surface of head darker, Kingfisher rufous #240, forebody ranging from Cinnamon, #139 to Buff, #124, interscale areas darker along body; the largest scales of forehead, frontal, parietal and occipitals and upper temporals with a distinct dark edge. Some large individuals are grey-black, the interscale regions appearing paler, while others appear to show pale transverse body bands, one scale wide, with 6–8 scales in interband areas. Isolated, irregular dark smudges may be present in such darker individuals on anterior part of gular scales. Mandible and genial region Buff, #124; the gular region to until Ventrals 12–28 bright yellow, Warm Buff, #118, with at least two dark areas peripherally, comprising somewhat indistinct bands meeting Ventrals 7–8 and 14–15; and noticeably in larger, hooding individuals, the first 5–6 nuchal scales are darker. Abdominal region as in gular region, but progressively darkening, initially from the peripheral region of the scales. 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, 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 comprise 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 comprise large oval spots arranged in a transverse series. Body bands are of the same colour, about two scales in width, and number 57–87 (65 in the lectotype), between head bands and the caudal region above vent, and have 4–5 scale wide interband areas. Bands on tail with relatively thick, dark, edges, both dorsally and ventrally; and the posterior third of body appears darker than the rest of the body.

Excluding hatchlings, which bear multiple, narrow pale bands on body, no pale body bands were seen in presumed adult (total length> 2,000 mm) individuals.

Distribution

The range of the species extends from south of the Isthmus of Kra, across the Malay Peninsula (including extreme southern Thailand, West Malaysia, Singapore, and offshore islands), Sumatra, Borneo, Java, Bali, and some of the islands of the southern Philippines Archipelago ( David & Vogel 1996; David et al. 2006; Chanhome et al. 2011 [in part]; de Lang 2011, 2017; Stuebing et al. 2014; Das et al. 2015; Ismail et al. 2018).

Remarks

The king cobra genome sequenced by Vonk et al. (2013) came from a specimen of this species from Bali, Indonesia. Card et al. (2021) documented the genomic mechanisms of facultative parthenogenesis in this species (mitochondrial sequences included in Gowri Shankar et al. 2021).

Figures

Fig. 6. Plate XVII, figures 8–9 from Schlegel (1837), showing head in dorsal and lateral views of lectotype of Naja bungarus Schlegel, 1837 (RMNH 1334).

Fig. 9. Head in dorsal, ventral and lateral views of respective type specimens of species of Ophiophagus Günther, 1864. A–C. Ophiophagus hannah (Cantor, 1836), neotype, ♀ (ZSI 8292). A. Dorsal view of head. B. Ventral view of head.C. Left lateral view of head.D–F. Ophiophagus bungarus (Schlegel, 1837) comb. nov., lectotype, ♂ (RMNH 1334). D. Dorsal view of head. E. Ventral view of head. F. Left lateral view of head. G–I. Ophiophagus kaalinga Gowri Shankar, Das & Ganesh sp. nov., holotype, ♂ (BNHS 3655). G. Dorsal view of head. H. Ventral view of head. I. Left lateral view of head. J–L. Ophiophagus salvatana Gowri Shankar, Das & Wüster sp. nov., holotype, ♂ (CAS 61329). J. Dorsal view of head. K. Ventral view of head. L. Left lateral view of head.

Fig. 11. Variation in dorsal bands in adults of the four species of Ophiophagus Günther, 1864 recognised in this study. A. O. kaalinga Gowri Shankar, Das & Ganesh sp. nov. B. O. hannah (Cantor, 1836). C. O. bungarus (Schlegel, 1837) comb. nov. D. O. salvatana Gowri Shankar, Das & Wüster sp. nov.

Fig. 12. Variation in dorsal body bands in juveniles of the four species of Ophiophagus Günther, 1864 recognised in this study.A. O. hannah (Cantor, 1836) (photo: H.T. Lalremsanga).B. O. bungarus (Schlegel, 1837) comb. nov. (photo: T. Charlton). C. O. kaalinga Gowri Shankar, Das & Ganesh sp. nov. (photo: P. Gowri Shankar). D. O. salvatana Gowri Shankar, Das & Wüster sp. nov. (photo: Bernard Tarun).

Fig. 13. Live unvouchered adults of Ophiophagus hannah (Cantor, 1836) showing variations in colour pattern. A. Mizoram, India (photo: H.T. Lalremsanga). B. China (Adam Francis). C. Thailand (photo: P. Gowri Shankar). D. Andaman Islands, India (photo: Aaron Fernandes).

Fig. 14. Live unvouchered adults of Ophiophagus bungarus (Schlegel, 1837) comb. nov. showing variations in colour pattern. A. Peninsular Malaysia (photo: Ahmad Khaldun Ismail). B. Java, Indonesia (photo: Nathan Rusli). C. Bali, Indonesia (photo: Shinta Sukum). D. East Malaysia (photo: P. Gowri Shankar).