Craspedocephalus malabaricus (Jerdon, 1854), 1822

Mallik, Ashok Kumar, Srikanthan, Achyuthan Needamangalam, Ganesh, Sumaithangi Rajagopalan, Vijayakumar, Seenapuram Palaniswamy, Campbell, Patrick D., Malhotra, Anita & Shanker, Kartik, 2021, Resolving pitfalls in pit viper systematics - A multi-criteria approach to species delimitation in pit vipers (Reptilia, Viperidae, Craspedocephalus) of Peninsular India reveals cryptic diversity, Vertebrate Zoology 71, pp. 577-619 : 577

Craspedocephalus malabaricus (Jerdon, 1854)

Figures 10 , 11 , 12

Trigonocephalus (Cophias) malabaricus Jerdon, 1854

Trigonocephalus (Cophias) wardii Jerdon, 1854

Lachesis coorgensis Rao, 1917

Trimeresurus malabaricus - Smith, 1943

Lachesis anamallensis (non Günther, 1864) - Wall, 1919, 1924 part

Craspedocephalus malabaricus - Wallach et al., 2014

Taxonomic history.

Jerdon (1854) originally described this species as Trigonocephalus (Cophias) malabaricus and the type locality of this species was mentioned as the "West Coast of Peninsular India", restricted to the Western Ghats. The types of this species are currently untraceable (Goley et al. 1993, Gumprecht et al. 2004, Wallach et al. 2014). Other subsequently described nomina of C. malabaricus , that are currently recognized as synonyms include Trigonocephalus (Cophias) wardii Jerdon 1854, Trimeresurus anamallensis Günther, 1864 (now a valid species; see below) and Lachesis coorgensis Rao, 1917. Jerdon (1854) in his original descriptions of Trimeresurus malabaricus and Tri. wardii did not mention any locality names or geographic province. Our perusal of historical collection catalogues (Boulenger, 1896; Das et al. 1998; Sclater, 1891; Theobald, 1868, 1876) in two potential repositories associated with Jerdon’s herpetological specimens (the Natural History Museum London and the Zoological Survey of India Kolkata) all reveal that only three precisely-named places have ever been associated with these nomina: Anamalais, Nilgiris and Wayanad (also see Beddome, 1862; Boulenger, 1890).

The specific epithets, mostly toponyms, also suggest the same: C. malabaricus : from ‘Malabar’ i.e. north Kerala and Coimbatore / Nilgiris, in the Western Ghats; C. anamallensis : from Anamalai hills of the Southern Western Ghats; Lachesis coorgensis : from ‘Coorg’ (=Kodagu) a part of Malnad region in Central Western Ghats. The only remaining nomen in its synonymy is Tri. wardii of Jerdon, 1854. Our perusal of historical literature reveal that this eponym could only be named after Samuel Neville Ward (18th June 1813 - 31st January 1897). Samuel M. Ward was with the Madras Civil Service (1832-63) and had a final appointment as Judge of Koyambatur (=Coimbatore). He had reportedly been corresponding with naturalists such as Charles Darwin, Edward Blyth, as well as T. C. Jerdon, who described Tri. wardii . Ward had been reported to have visited Sirsi, in North Canara for his work on Indian wildlife, mostly butterflies and birds (Pittie, 2016 and references therein). Beddome (1862) explicitly stated that Tri. wardii is from the Nilgiris. The earliest reference of this group from Agasthyamalai is that of Ferguson (1895), who used Günther’s name ( C. anamallensis ) to refer to it. Thus historical collections prior to typifications (till Rao, 1917) of the C. malabaricus complex apparently happened only from Anaimalai, Nilgiris and Wayanad.

Type.

Syntypes lost (after Smith, 1943; also see Wallach et al. (2014:188), ‘Holotype’ lost (after Das et al. 1998). Hence, as per Art. 75.3 of the Code ( ICZN 1999) since we are revising the taxonomy of this group, we designate as neotype the holotype of its junior synonym Lachesis coorgensis Rao, 1917. The selected neotype (read below) is chosen to clarify the identity of this nominal taxon, a well-preserved adult originating from a precise locality, in conformity with the original description of the conferred nomen and housed in a permanent national repository/museum. Thus Craspedocephalus malabaricus and Lachesis coorgensis are now objective synonyms.

Materials examined.

Specimen series: Neotype: ZSI-18161 from Coorg, Karnataka, 12°24.82'N; 75°43.85'E by C.R. Narayan Rao in 1917, the holotype of Lachesis coorgensis Rao , 1917. - Other referred material: BNHS 2609-778 from Kotagiri , Tamil Nadu and BNHS 2069-781 from Coonoor , Tamil Nadu by A. M. Kinloch in 1907. Additional comparative material collected for this study - CESS053, CESS055 from Amboli , Maharashtra ; CESS063, CESS065 from Saklespura Karnataka ; CESS273 from Coorg , Karnataka ; CESS086 from Thirunelli , Kerala by Ashok Kumar Mallik between 2009-11 and CESS141 from Silent valley by Saunak Pal and Mrugank Prabhu in 2010; BNHS 2609A from Kotagiri and BNHS 2609B from Coonor , Nilgiris , Tamil Nadu by A.P Kinloch in 1907; BNHS 2596 from Sirsi , Karnataka by Charles McCann in 1938; BNHS 2599 from Mahabaleshwar , Maharashtra by H. Abdulali in 1953; BNHS 2601 from Castle Rock, Karnataka by Mrs. H. Pearson in 1907.

Type locality.

Originally given as "West Coast, Peninsula of India". Here restricted to Coorg (12°24.82'N; 75°43.85'E) in Central Western Ghats, by neotype designation.

Etymology.

The specific epithet Craspedocephalus malabaricus is a toponym, alluding to its type locality - the Malabar region of the Western Ghats.

Lineage diagnosis (redefined herein).

A lineage of the C. malabaricus complex, C. malabaricus s. str. (L5) is here restricted only to populations north of the Palghat Gap. This nominotypical population is 8.3-9% and 1.2-2.2% divergent at cyt b and 16S respectively, from those south of the Palghat Gap (L3 & L4), recognised here as two nominate taxa: C. anamallensis ( Günther, 1864) and Craspedocephalus travancoricus sp. nov. (see below). These taxa are allopatric with respect to each other and C. malabaricus .

Description.

Neotype in good condition, small lesion near the nape, possibly caused while collecting and euthanizing the individual; specimen with a slender, cylindrical body of snout to vent length (SVL) 481 mm and a prehensile tail of length (TL) 73 mm; dorsal scales keeled with anterior dorsal scale rows (DSR) 21, mid body scale rows (MSR) 21 and posterior scale rows (PSR) 13; head prominent, of length 24.3 mm, clearly distinguished from the neck with small, mildly keeled scales on the head; rostral scale trapezoid, with the lower side roughly more than twice the size of the upper side with the tip visible from above; supraoculars divided, separated by eight cephalic scales between both supraoculars at its posterior border; seven scales bordering each supraocular. Canthus rostralis distinct with four canthal scales; three preoculars, two postoculars and a thin elongated crescent shaped subocular; eye with a distinct elliptical pupil, vertical diameter of the eye 3.31 mm and horizontal diameter 3.68 mm; strongly keeled temporal scales and cephalic scales in the posterior sides above the mandibular joint; aperture of the nostril completely covered by the nasal scale, undivided and sub-pentagonal shaped, in contact with the first two canthal scales and the 1st and 2nd supralabial; loreal pit present in contact with the second supralabial and the 2nd and 3rd preoculars; nine supralabials and 12 infralabials, with eight scales between the last supralabial, including the last supralabial up to the start of the ventral scales; 1st, 2nd and 3rd infralabial scale in contact with the first pair of genials; a gap of four scales including the posterior genials followed by 148 ventrals, laterally separated from the dorsal scale rows by a slightly broader row of dorsal scales; anal scale undivided, followed by 38 divided subcaudals scales; Terminal scale on the tail larger than the previous scale, blunt at the tip.

Variation (n=20).

The referred materials are of SVL up to 670 mm and TL up to 126 mm with colours varying from dark brownish red to light green throughout the specimens in its current preservation state, differs from the holotype with respect to pholidosis by having 19 to 22 DSR, 19 to 23 MSR, 13 to15 PSR, 145-149 ventrals and 52-54 subcaudals; head distinct with supralabials ranging between 8-10 and infralabials ranging between 10-13; one to three preocular scales, one to two scales (some specimens showing an absence of these scales) between the 3rd supralabial scale and the suboculars, seven to nine cephalic scales and seven to eight scales surrounding the supraoculars from the dorsum.

Colour in life.

A highly variable and polymorphic species, with respect to colour, specimens can be found in a variety of colour morphs - greenish blue-cyan, bright yellow, green, rufous brown, bright orange and red coloured specimens have been encountered during this study; head characterized with a thick dark brown to black postocular stripe till the nape, labials sometimes marked with blotches and a highly variable pattern above the head, sometimes fully dark, some individuals with no markings at all, body with alternating zig-zag saddle shaped markings with the last rows of scales on the tail banded with different colour; these markings vary from brick reddish, dark brown to black, sometimes intermixed with spots of other colours such as green, yellow and blue; the base colour of the body varies from light brown in juveniles, light cream, orange, yellow, brick red, bluish green and sometimes morphs mottled with all or some of the aforementioned colours; ventrals sometimes vary from the colour of the dorsum, complementing the variety of vibrant dorsal colours, but often are coloured similar to the dorsum; colour change is also observed to be seasonal ( Kanagavel et al. 2012); juveniles brown with dark brown to black markings, neonates and younger juveniles possess a tail ‘lure’ that is often different from the body’s colouration. Mandibular region and the ventrals in alternating light green, blue, yellow to creamy yellow with speckles, separated from the dorsal scales with a longitudinal lighter irregular stripe, two prominent, labial stripes from the eye and the loreal pit, up to the edge of the lower end of the supralabial region.

Colour in preservative.

Fairly faded specimen with a light brown dorsum, ventrals and tail; scales bordered with dark brown; temporal stripe visible on the right side of the specimen in dark brown, left side with no visible temporal stripe; body with barely distinguishable dark brown saddle shaped markings throughout the body; tail with dark brown stripes from the vent up to the tip.

Habitat.

An arboreal species, commonly found on bushes and in undergrowth in forests and near streams in evergreen forests to moist deciduous and lowland riparian forests. Due to anthropogenic changes to the landscape, this species is also abundantly found in agricultural landscapes such as coffee plantations, from 100-1800m MSL.

Distribution.

Restricted to the central and northern Western Ghats from Mahabaleshwar - Koyna in Maharashtra to the Nilgiris and Elivalmalai hills, north of the Palghat Gap. Known to occur sympatrically with C. occidentalis comb. nov. in Nilgiris and Coorg, with C. gramineus in Amboli and may have some overlap with C. strigatus in the mid to high elevations of the western slopes of the Nilgiris (at the upper limit of its altitudinal distribution).

Figures

Figure 10. Craspedocephalus malabaricus in life, showing: (A) orange morph entire dorsal view (Agumbe), (B) purple morph (Anshi), (C) maroon morph (Agumbe), (D) cream morph (Amboli), (E) green morph (Amboli), (F) yellow morph (Anshi) & (G) brown morph (Anshi).

Figure 11. Craspedocephalus malabaricus neotype (Lachesis coorgensis holotype) (ZSI- 18161), in preservation, showing: (A) head dorsal view, (B) head lateral left view, (C) head lateral right view, (D) head ventral view.

Figure 12. Craspedocephalus malabaricus neotype (Lachesis coorgensis holotype) (ZSI- 18161), in preservation, showing: (A) entire dorsal view, (B) entire ventral view.