Ophiophagus salvatana Gowri Shankar, Das & 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 : 30-35

publication ID

https://doi.org/ 10.5852/ejt.2024.961.2681

publication LSID

lsid:zoobank.org:pub:8E064900-1289-4648-BE9A-F17461CCF25C

persistent identifier

https://treatment.plazi.org/id/1D2264EB-DFE1-4A71-AAA3-0EC155480486

taxon LSID

lsid:zoobank.org:act:1D2264EB-DFE1-4A71-AAA3-0EC155480486

treatment provided by

Plazi

scientific name

Ophiophagus salvatana Gowri Shankar, Das & Wüster
status

sp. nov.

Ophiophagus salvatana Gowri Shankar, Das & Wüster sp. nov.

urn:lsid:zoobank.org:act:1D2264EB-DFE1-4A71-AAA3-0EC155480486

Figs 9J–L View Fig , 10D View Fig , 11D View Fig , 15 View Fig

Common name

Luzon king cobra.

Diagnosis

A species of Ophiophagus inhabiting Luzon Island in the northern Philippines and exhibiting the following combination of characters: lacking pale bands along body of adults (vs with dark-edged, pale bands in O. hannah ; lacking dark edges to the pale bands, if present, in O. bungarus , or with a pale band in O. kaalinga sp. nov.). Further, it differs from O. hannah in having fewer pterygoid teeth (11 vs 18– 21). Finally, juveniles of O. salvatana sp. nov. have extremely angular pale body bands, with the dark intervening areas covering 2–3 scales (vs more rounded pale body bands, the intervening areas covering 4–9 scales in congeners). The new species, with 85–86 pale body bands, can be easily separated from O. hannah (27–48) and O. kaalinga (28–48); relative tail length ranging 18.7–23.0% with a mean of 20.85% (vs 21.7–26.4% [24.05%] in O. hannah ; vs 19.3–25.1% [22.2%] in O. bungarus ; vs 18.0–19.9% [18.95%] in O. kaalinga ).

Etymology

The specific epithet salvatana is the Tagalog (a vernacular language spoken in Luzon and adjacent regions of the Philippines, of Austronesian origin) name for the king cobra in the Luzon region (northern Philippines), here coined as a noun in apposition and hence invariable.

Type material

Holotype

PHILIPPINES • ♂; Luzon , Benguet Province, Baguio; 16.40° N, 120.60° E; Edward H. Taylor leg.; CAS 61329 About CAS . GoogleMaps

Paratypes

PHILIPPINES • 1 spec.; Luzon , Isabella; 17.00° N, 122.00° E; BMNH 94.10 .24.15 GoogleMaps 1 ♀; Luzon, Camarines Sur Province, Gota Beach Caramoan ; 13.88° N, 123.70° E; UF 50927 GoogleMaps 1 ♂; Luzon, Camarines Sur Province, Caramoan Municipality, Tarago Base Camp ; ca 13.82° N, 123.80° E; UF 55008 GoogleMaps 1 ♂; Luzon, Pampanga Province, Sapang Tagalog , Tarlac; 15.42° N, 120.59° E; UPM 1692 View Materials GoogleMaps 1 ♂; Luzon, Camarines Sur Province, San Pedro, Iriga ; 13.49° N, 123.47° E; FMNH 53553 About FMNH GoogleMaps .

Description of holotype ( CAS 61329)

MEASUREMENTS. SVL 2060 mm, TL 517+ mm, total 2577+ mm.

HABITUS. Body relatively robust (midbody width 30.0 mm, 1.5% SVL), triangular in cross-section; Transverse body rows: DSR1 19; DSR2 15; DSR3 15; ventrals 252; subcaudals 95+; supralabials 7; infralabials 8; anterior temporals 2; posterior temporals 2; cloacal plate 1; dorsal and ventral scales smooth; subcaudals 1–26 and 31–37 undivided, the rest divided; tail short, cylindrical, tapering posteriorly, tail tip missing.

HEAD. Head relatively large, head length 52.2 mm; head width 31.7 mm; head depth 21.1 mm; distinct from neck, flattened in the orbital region, rounded in the sagittal region, with a depression medially, snout projecting slightly beyond mandible; canthus rostralis sharply defined; eye width 7.5 mm; interorbital distance 22.4 mm; cephalic scales juxtaposed, smooth-edged, except parietals and occipitals, which are slightly imbricate; rostral trapezoid in shape, distinctly visible from above, slightly under twice as high as wide, concave ventrally, rostral width 12.6 mm; rostral length 6.7 mm; eye to snout distance 19.7 mm; eye to nostril distance 9.4 mm; nostril diameter 5.4 mm; internasals large, subtrapezoidal, wider than long; internasal suture length 4.6 mm; internasal width 6.9 mm; prefrontals trapezoid, wider than long; prefrontal length 7.7 mm; prefrontal width 9.3 mm; prefrontal suture 5.8 mm; frontal lanceolate, contacting prefrontals, supraoculars and parietals, rectangular in shape, short-sided anteriorly; supraocular subtrapezoidal, contacting prefrontal, frontal, parietal, orbit, preocular and upper postocular; large paired occipitals; occipital length 12.4 mm; interoccipital scute absent; nuchals undifferentiated; supralabials 7/7; III–IV (L/R) touching the eye; II (L/R) contacting nasal and I, II and III (L/R) contacting posterior nasal; Supralabial I low; II high; III tallest; IV and V subequal; and VI and VII low, narrow and elongate; Supralabial IV does not contact preoculars; nostril lateral at posterior of a single concave nasal, oval in shape, its greatest diameter at a vertical plane; nasal irregularly triangular, one preocular and three postoculars; eye large, contained in head length 0.14 times and head depth 0.36 times; pupil rounded; ocular ring comprises seven scales – one preocular, three postoculars, one supraocular and two supralabials; suboculars absent; temporals 2/2 (L/R) + 3/3 (L/R); anterior upper temporal longer than lower temporal; mental small, triangular, wide than deep (mental width = 6.98 mm; mental depth = 3.64 mm); infralabials 8/8 (L/R); infralabial I–IV contact anterior genial; infralabial IV–V (L/R) contact posterior genials; infralabial IV largest; two pairs of genials, with the anterior larger than posterior; cuneate scute on lower jaw absent; three elongated 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 6.4 mm; fang width at base 1.8 mm; teeth count obscured by gingivae.

COLOURATION. Dorsum of holotype is yellowish-grey, each scale on forehead and body edged with pale grey, lacking distinct pale bands; the venter is paler, scales similarly dark-edged, edges progressively darker posteriorly.

Morphological variation

None of the six specimens examined have an interoccipital scute. All subcaudals were divided in the holotype, while other specimens examined have some undivided subcaudals, typically, the first, and some succeeding subcaudals, up to Subcaudal 26, in what appears to be an irregular pattern that cannot be linked to either sex, ontogeny or geographical distribution. Variations observed include undivided subcaudals 1–17, 20–25, 28, 30–35, 46 ( UPMNH 1692); 1, 4–6, 25 ( UF 50927); 1–18 ( UF 55008); and 1–26, 31–37 ( CAS 61329). In life, dorsal surface of body is Dark Neutral Gray #83, forehead and posterior fourth of body darker, especially on scale periphery, making the posterior of body darker than the rest; interscale areas dark along body; the largest scales of forehead, frontal, parietal and occipitals and upper temporals with a distinct dark edge; ventral scales, from Ventral I suffused with yellow, gradually turning entirely yellow with irregular patterned grey posterior edges; oval scales of the 2–3 lower-most row of dorsal scale rows meeting ventrals yellow; posterior third of body dorsum darkening due to increasing intensity and extent of dark edges of scales; tail dorsum nearly black, individual scales with yellow centres. Mandible and genial region Buff Yellow, #53; the gular region brighter, Orange Yellow, #18, with darker areas peripherally. 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, with pale yellow narrow ring; iris brownish-grey with a yellowish-brown ring. Tongue blackish-grey, oral cavity pink. Images of two hatchlings were examined for colour and pattern. The forehead of hatchlings is Light Neutral Gray (#85), including the posterior cephalic scales (including upper temporal, parietals and occipital), dark edged, Jet Black (#89), the dark areas of the latter pair extensive, separating the pale areas of the centre as eye-like spots; dorsum of body is Jet Black (#89), 2–3 scales across, with 101, extremely angular, Pale Neutral Gray (#86) bands, a single scale across; the pattern on back is undifferentiated posteriorly. Apart from two hatchlings, on which we counted 85–86 narrow pale bands on the body, the distinct pale bands seen in other species of the genus are absent in O. salvatana sp. nov. In preservative, forehead and dorsum of body are greyish-brown, with darker edges of scutes; no other patterns are discernable; venter is similarly coloured but slightly paler. Three individuals that permitted count of tooth and / or sockets have dentition thus: 15 dentaries; 11 pterygoids and 3 palatines.

Distribution

The range of the species is restricted to the Luzon islands in the northern Philippines ( Diesmos et al. 2005; McLeod et al. 2011; Siler et al. 2011; Devan-Song & Brown 2012; Brown et al. 2013; Cruz & Afuang 2018; Cruz et al. 2018). The affinities of king cobras from other islands of the Philippine Archipelago remain to be confirmed.

Comparisons

Early attempts to understand variation in the complex suffered from a variety of shortcomings, due to the variation within ( Figs 11–12 View Fig View Fig ) and among populations ( Figs 13–16 View Fig View Fig View Fig View Fig ) and scarcity of vouchered specimens, especially entire specimens of adults. For instance, all four species recognised here demonstrate dramatic ontogenetic colour and pattern changes, with the banded juveniles losing the pattern in adults consistently in one species ( O. salvatana sp. nov.) and sometimes in another ( O. bungarus comb. nov.), retaining it into adulthood in a third population ( O. kaalinga sp. nov.), and developing dark edges to the band in the fourth ( O. hannah ). Further, the allopatric populations of the four species recognized in this paper do not show categorical differences in the counts of the major scales that have been traditionally employed for taxonomic differentiation of species within snakes. However, a number of characters are useful to unequivocally diagnose the four species recognized here. Character states useful for identification and diagnosis of the members of the complex include dorsal ground colour and pattern, nature of body bands in adults (unbanded and banded, the bands with or without a dark edge) and pterygoid tooth/tooth socket counts.

Means, ranges and sample size of characters differentiating the four lineages identified in this study and supported by our mitochondrial phylogeny (see Gowri Shankar et al. 2021: 5) are shown in Table 1. Measurement data are in Table 2. Furthermore, we provide here a dichotomous identification key for the identification of the species of the genus Ophiophagus recognized here.

Key to the species of Ophiophagus Günther, 1864

1. More than 17 pterygoid tooth sockets; light body bands present at all ages,>1.5 scales wide, with dark edges in most adults .......................................................................... O. hannah ( Cantor, 1836) View in CoL

– 12 or fewer pterygoid tooth sockets, body bands not as above ........................................................ 2

2. Pale body bands always present, fewer than 55,>1.5 scales wide; <90 subcaudals ......................... .......................................................................... O. kaalinga Gowri Shankar, Das & Ganesh sp. nov. – Body bands absent or narrow (<1 scale wide), more than 55; often>90 subcaudals ..................... 3

3. Juveniles with 57–87 regular narrow light body bands, adults with faint narrow bands or unpatterned ......................................................................................... O. bungarus ( Schlegel, 1837) View in CoL

– Juveniles with>80 often irregular, highly angular narrow light bands, adults always without pale bands ................................................................ O. salvatana Gowri Shankar, Das & Wüster sp. nov.

CAS

California Academy of Sciences

UF

Florida Museum of Natural History- Zoology, Paleontology and Paleobotany

Kingdom

Animalia

Phylum

Chordata

Class

Squamata

Family

Elapidae

Genus

Ophiophagus

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