Taxonomic revision of the king cobra Ophiophagus hannah (Cantor, 1836) species complex (Reptilia: Serpentes: Elapidae), with the description of two new species Author Das, Indraneil 85A2D637-43A8-4422-B42E-A23B81592326 Institute of Biodiversity and Environmental Conservation, Universiti Malaysia Sarawak, 94300 Kota Samarahan, Sarawak, Malaysia. Author Shankar, P. Gowri FD48E857-17FC-4D01-B727-66BE3BBA76FB Maharaja Shrirama Chandra Bhanja Deo University, Baripada, Takatpur, Odisha, India. & Centre for Ecological Sciences, Indian Institute of Science, Bangalore 560 012, Karnataka, India. & Department of Ecology and Genetics, Evolutionary Biology Centre, Uppsala University, SE- 751 05 Uppsala, Sweden. & Kālinga Foundation, Guddakere, Agumbe, Shivamogga 577 411, Karnataka, India. Author Swamy, Priyanka Centre for Ecological Sciences, Indian Institute of Science, Bangalore 560 012, Karnataka, India. Author Williams, Rhiannon C. Molecular Ecology and Evolution at Bangor, School of Environmental and Natural Sciences, Bangor University, Environment Centre Wales, Bangor LL 57 2 UW, Wales, UK. Author Lalremsanga, Hmar Tlawmte Developmental Biology & Herpetology Laboratory, Department of Zoology, Mizoram University, Aizawl 796 004, Mizoram, India. Author Prashanth, P. Kālinga Foundation, Guddakere, Agumbe, Shivamogga 577 411, Karnataka, India. Author Sahoo, Gunanidhi Department of Zoology, Utkal University, Bhubaneswar 751 004, Odisha, India. Author Vijayakumar, S. P. Kālinga Foundation, Guddakere, Agumbe, Shivamogga 577 411, Karnataka, India. Author Höglund, Jacob Department of Ecology and Genetics, Evolutionary Biology Centre, Uppsala University, SE- 751 05 Uppsala, Sweden. Author Shanker, Kartik Centre for Ecological Sciences, Indian Institute of Science, Bangalore 560 012, Karnataka, India. Author Dutta, Sushil K. Department of Zoology, Assam Don Bosco University, Tapesia 782 402, Assam, India. Author Ganesh, S. R. Kālinga Foundation, Guddakere, Agumbe, Shivamogga 577 411, Karnataka, India. Author Wüster, Wolfgang Molecular Ecology and Evolution at Bangor, School of Environmental and Natural Sciences, Bangor University, Environment Centre Wales, Bangor LL 57 2 UW, Wales, UK. text European Journal of Taxonomy 2024 2024-10-16 961 1 1 51 https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2681/12413 journal article 10.5852/ejt.2024.961.2681 2118-9773 13944155 8E064900-1289-4648-BE9A-F17461CCF25C Ophiophagus kaalinga Gowri Shankar, Das & Ganesh sp. nov. urn:lsid:zoobank.org:act: 84BAA541-67F1-4CE1-B72A-9832157B0267 Figs 8 , 9G–I , 10C , 11C , 14 Common name Western Ghats king cobra. Diagnosis A species of Ophiophagus endemic to the Western Ghats of the Indian peninsula, defined by the following combination of characters: possessing pale bands that lack darker edges, along body of adults (vs unbanded in adult O. salvatana sp. nov. and many O. bungarus ; with dark edges to the pale bands in adult O. hannah ). Ophiophagus kaalinga sp. nov. differs from O. hannah through a lower pterygoid teeth count (12 vs 18–21). Finally, juveniles of O. kaalinga have 28–48 body bands fewer than O. salvatana (85–86) and O. bungarus (57–87); relative tail length ranging 18.0–19.9% with a mean of 18.95% (vs 21.7–26.4% [24.05%] in O. hannah ; vs 19.3–25.1% [22.2%] in O. bungarus ; vs 18.7–23.0% [20.85%] in O. salvatana sp. nov. ). Fig. 8. Plate 8, from Fayrer (1872) depicting the ‘Dusky variety’ of Ophiophagus elaps ( Günther, 1858 ) , and corresponding to Ophiophagus kaalinga Gowri Shankar, Das & Ganesh sp. nov. , showing entire body of a hooding adult in mostly a dorsolateral view. 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. Etymology The specific epithet kaalinga is derived from Kannada language of Karnataka , India , alluding to the snake’s dark colouration (‘Kali’ / ‘Kari’ = dark / black), an abbreviated form of ‘Kaalinga Havu’ / ‘Sarpa’ (see Das 1998 ), associated with Lord Shiva, as a demigod; here coined as a noun in apposition and hence invariable. Type material Holotype INDIA • ♂ ; Karnataka State , Shivamogga District , Agumbe ; 13.57° N , 75.10° E ; 21 Apr. 2021 ; P. Gowri Shankar leg.; BNHS 3655 . Paratypes INDIA • 1 ♀ ; Karnataka State , Uttara Kannada District , Dandeli Wildlife Sanctuary ; 15.25° N , 74.57° E ; ZSIC ( ex-MCBT 152882 ) • 1 spec. ; Kerala State , Quilon ; 8.88° N , 76.6° E ; BNHS 2280 • 1 spec. ; Tamil Nadu State , Anaimalai or Annamalai; 10.30° N , 77.00° E ; BMNH 61.12.30.83 • 1 spec. ; Karnataka , Karwar District , Sirsi ; 14.60° N , 74.90° E ; MCBT 152884 • 1 spec. ; Kerala State , Vanjikadavu , Kundurmadu ; 10.39° N , 76.34° E ; ANSP 35312 . Fig. 10. View of dorsum of entire body of respective type specimens of species of Ophiophagus Günther, 1864 . A . Ophiophagus hannah ( Cantor, 1836 ) , neotype, ♀ (ZSI 8292). B . Ophiophagus bungarus ( Schlegel, 1837 ) comb. nov. , lectotype, ♂ (RMNH 1334). C . Ophiophagus kaalinga Gowri Shankar, Das & Ganesh sp. nov. , holotype, ♂ (BNHS 3655). D . Ophiophagus salvatana Gowri Shankar, Das & Wüster sp. nov. , holotype, ♂ (CAS 61329). Description of holotype ( BNHS 3655) Laterally incised to access liver tissue; three ventral scales and two rib tips clipped as tissue samples. MEASUREMENTS . SVL 2475 mm , TL 530 mm , total 3005 mm . HABITUS . Body relatively robust (midbody width 43.2 mm , 1.6% SVL), triangular in cross-section; Transverse body rows: DSR1 19; DSR2 15; DSR3 15; ventrals 241; subcaudals 89; 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 1–8 and 16–22 entire, the rest divided; tail short (20.4% SVL), cylindrical, tapering posteriorly. HEAD . Head relatively large, head length 72.0 mm; head width 52.5 mm ; head depth 30.0 mm; distinct from neck, flattened in the orbital region, rounded in the sagittal region, with a slight depression medially; eye diameter 8.0 mm; interorbital distance 27.0 mm; canthus rostralis sharply defined; 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 7 mm ; rostral length 16 mm ; Eye to snout distance 25 mm ; eye to nostril distance 11.5 mm ; nostril diameter 6.0 mm; internasals large, subtrapezoidal, wider than long; preocular squarish, wider than high, separated from internasal by prefrontal; prefrontals trapezoid, wider than long; frontal trapezoidal, contacting prefrontals, supraoculars and parietals; frontal edge straight, short-sided posteriorly; supraocular subtrapezoidal, contacting prefrontal, frontal, parietal, orbit, preocular, upper postocular but not temporals; large paired occipitals; occipital length 10.6 mm ; interoccipital scute present; temporals 2/2 (L/R); in anterior pair, upper temporal longer than lower; 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; IV (L/R) contacting preocular and I and II (L/R) contacting posterior nasal; supralabial I low; II and III progressively higher; III tallest; V and VI subequal; and VI and VII low, narrow and elongate; Supralabial III contacts preoculars; nostril lateral at posterior of a single concave nasal, horizontally elliptical, its greatest diameter at a vertical plane; one preocular and three postoculars; orbit large, pupil rounded; ocular ring comprises seven scales- one preocular, three postoculars, one supraocular and two supralabials; suboculars absent; mental small, triangular, wider than deep; 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. 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. HEMIPENES . Organ partially everted on left side, fairly short and thick, length 24 mm and width 6 mm ; extending to 3 rd subcaudal scale; divided at level of 2 nd subcaudal scale; divided lobe length 7 mm ; divided lobe width 5 mm ; lobe head forked near apex; lobe head with numerous tiny spinules and flounces; asulcate side rather smooth and less spinose; sulcate side with dense congregation of larger spines; sulcal lips not evident, sheath-like, completely surrounding apex base; sulcus spermaticus groove-like and rather straight than convoluted; pedicel more or less conical and tapering down towards its base. DENTITION . Maxillary teeth recurved and stout, gradually enlarged posteriorly; fang length 9 mm ; fang width at base 2 mm ; a reserve fang present on left side. COLOURATION . Dorsum dark grey, the ventrals paler. In life, the dorsal surface of the body Dark Brownish Olive (#129), darkening at the posterior third of body to Blackish Neutral Gray (#82); forehead Drab (#27), the largest scales of forehead, including frontal, parietal and occipitals and upper temporals with distinct dark edges; interscale areas of dorsum darker along body, distinctly yellowish when within the pale bands; dorsum and flanks with narrow, oblique, chevron-shaped Straw Yellow (#56) bands, covering a single scale on vertebral region, expanding to cover 4–5 scales on the lower flanks, and separated by 7–8 scales; mandible and genial region Straw Yellow (#56); gular region Buff-Yellow (#53), extending to 20 th ventral scale, intervened by a 2-scale wide dark band at the level of 12–13 ventrals; abdominal region as in gular region, but progressively darkening, from Ventral 20 initially from the peripheral region of the scales. Subcaudal scales grey throughout, and with dark edges. Pupil rounded, black, iris brownish, darkening peripherally, with a narrow, yellow ring. Tongue dark red. Body bands Yellow Ocher (#123C), 1–2 scales in width, widening on lower flanks to 3–4 scales, 29 such bands present between head bands and the caudal region above vent, with 4–5 scale wide interband areas. The first two bands in nuchal region chevron-like. Bands on tail with relatively thick, dark, edges, both dorsally and ventrally; and the posterior third of body is darker than rest of body. In preservation, after over two years in alcohol, colouration generally the same, except the eye becoming dirty white to grey and venter becoming ivory coloured anteriorly to light ashy grey posteriorly. Morphological variation Two ( ZSIC ex-MCBT 152882 and MCBT 152884) of four specimens examined had an interoccipital scute. Three of the specimens had intact tails to permit examination of subcaudal fusion pattern. One showed undivided subcaudals 1–7 ( ZSIC ex-MCBT 152882), a second, 1–6 and 12–13 ( BMNH 61.12.30.83) and the third, 1–8 and 10–11 ( MCBT 152884). Wall (1919) reported slightly lower subcaudal (85) and ventral (239 and 241) counts, although it is unclear how these were made. Only two specimens permitted counts of body bands that number 28 and 48. Specimens that permitted counts of tooth and / or sockets had dentition thus: 12 dentary sockets and 3–4 palatine sockets. Juveniles have ivory white bands, Buff, #124, including four on the head region, the first 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. Distribution Ophiophagus kaalinga sp. nov. is endemic to the Western Ghats of south-western India , covering parts of Tamil Nadu , Kerala , Karnataka , Goa and the adjacent border of Maharashtra States. The species is known from the Ashambu hills near Kanyakumari (formerly, Cape Comorin), through the Agasthyamalai and Devarmalai ranges, the Cardamom hills, the Meghamalai mountains, the Anamalai-Palni ranges, across the Palghat Gap, through Nilgiri-Waynad, on to the Malnad regions (Coorg-Agumbe-Sharavathi-Anshi), until about the Goa Gap, abutting Maharashtra State ( Ishwar et al. 2001 ; Whitaker & Captain 2004 [part]; Khaire 2006 ; Hutton & David 2009 ; Ganesh et al. 2013 , 2014 ; Yadav & Yankanchi 2015 ). It is essentially a hill-dwelling species, that is prevalent in mid-elevation (ca 500–900 m a.s.l.) rainforests, while extending lower to the very foothills (ca 100 m a.s.l.), in mesic windward western versant, or lower slopes (ca 300 m a.s.l.) in the drier leeward eastern versant, reaching up to the high elevation plateaus (ca 1800 m a.s.l.) covered with montane forests (unpubl. data).