Craspedocephalus gramineus (Shaw, 1802), 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 gramineus (Shaw, 1802)

Figure 19

Coluber gramineus Shaw, 1802

Coluber viridis (nec Trimeresurus viridis Lacépède, 1804) Bechstein, 1802

Vipera viridis (nec Trimeresurus viridis Lacépède, 1804) Daudin, 1803

Trigonocephalus fario Jan, 1859 (nomen nudum)

Colubar gramineus - Mason, 1860 (nomen incorrectum)

Bothrops viridis fario Jan, 1863 (nomen nudum)

Bothrops viridis genei Jan, 1863 (nomen nudum)

Lachesis graminea - Wall, 1919 part

Trimeresurus gramineus - Whitaker & Captain, 2004 part

Craspedocephalus gramineus - Wallach et al., 2014 part

Taxonomic history.

Russell (1796) described ' Boodroo Pam ' based on a specimen from Vizagapatam (now Vishakhapatnam, Andhra Pradesh state, India) in the northern Eastern Ghats. Shaw (1802) provided a nominate description of that taxon as Coluber gramineus , based on the specimen depicted by Russell (1796). Bechestein (1802) erected the nomen Coluber viridis and Daudin (1803) erected the nomen Vipera viridis based on the same specimen depicted in Russell (1796). These nomina are connected by an objective synonymy and therefore share the same type specimen (see Russell, 1796) and type locality, Vizagapatam. Notwithstanding these, Lacépède (1804) described an Indonesian species as Trimeresurus viridis . This nomen Trimeresurus viridis Lacépède, 1804 has now been superseded by its junior subjective synonym Trimeresurus insularis Kramer, 1977 that is now in prevailing usage (see David et al. 2011). Three unavailable nomina Trigonocephalus fario Jan, 1859, Bothrops viridis fario Jan, 1863 and Bothrops viridis genei Jan, 1863 are present.

Jerdon (1854) discusses this taxon under the nomen Trigonocephalus (Cophias) viridis and miscredits it to Merrem. Beddome (1862) discusses this taxon in part (specimen from Condipalli hills) under the nomen Trimeresurus viridis (see below). Günther (1864) discusses unrelated East Asian species from Penang, Mergui, Laos, Khasya, Sikkim, Ladakh and Ningpo. Günther (1864) questioned if Andaman and Nicobar populations are conspecific with Tri. gramineus s. auct. but concludes that it could not be ascertained based on Blyth’s (1846) brief notes. Theobald (1876) also followed a similar view and refers populations from Sylhet, Burma and Malacca under this name. Subsequently, he maintained the same stance when he mentioned specimens from Sikkim, Khasi hills, Assam, Pegu, Andamans, Bengal and Nepal (Theobald, 1876).

Later Boulenger (1890), perhaps prompted by earlier works (e.g. Theobald), lumped together several S.E. Asian nomina such as Trimeresurus viridis Lacépède, 1804 (see David et al. 2011 for its proper explanation), Trigonocephalus erythrurus Cantor, 1839, Trimeresurus albolabris Gray, 1842, Trimeresurus elegans Gray, 1853 and even a non-green taxon Trimeresurus mutabilis Stoliczka, 1870 (see Vogel et al. 2014 for its current status). Stejneger (1907) also followed Boulenger (1890). This diluted and over-circumscribed concept of ' Tri. gramineus ' (sic) encompassing almost all green Trimeresurus species, gave rise to the false notion that it is a very widespread and variable species.

Remarks.

Coluber viridis Bechestein, 1802 and Vipera viridis Daudin, 1803 are both objective junior synonyms of Coluber viridis Shaw, 1802, all being described on the basis of the very same type specimen described and illustrated by Russell (1796) as " Boodroo Pam ". Trimeresurus viridis Lacépède, 1804 is a nomen oblitum superseded by Trimeresurus insularis Kramer, 1977 that is now in prevailing usage for this Indonesian species ( Wallach et al. 2014). Two female paratype specimens from the Eastern Ghats and one male specimen from Matheran, northern Western Ghats of a former subjective junior synonym (fide Wallach et al. 2014) Trimeresurus occidentalis Pope & Pope, 1933 is herein referred to Tri. gramineus (Shaw, 1802) with respect to its phylogenetic and morphological affinities.

Type.

Holotype (iconotype), a 762 mm specimen described and illustrated in Russell (1796: 13-14, pl. 9, titled " Boodroo Pam ").

Type locality.

"hills in the vicinity of Vizagapatam, coast of Coromandel, India" [= Vishakhapatnam, Andhra Pradesh, India, 17°41'N; 83°13'E].

Etymology.

Craspedocephalus gramineus , ‘grassy’ in Latin, alluding to its grass-green dorsal colouration.

Material examined.

NHMUK 1946.1.19.87 from Cuddapa Hills, Andhra Pradesh, by R.H. Beddome paratype of Trimeresurus occidentalis Pope & Pope, 1933; CESS033 from Gandagan, Orissa by Ashok Kumar Mallik; CESS100, CESS101, CESS102 from Naneghat, Maharashtra by Ashok Kumar Mallik in 2010; AFS96.13 from Masinagudi, and AFS96.14 Gingee, Tamil Nadu by Anita Malhotra in 1997; BNHS 2627 from Khandala, Maharashtra by Charles Mc Cann in 1941, BNHS 2764 from Ahwa, Dangs, Gujarat by Dr. E.M. Shull in 1963 and BNHS 3275 from Sirumalai hills, Tamil Nadu by S.P. Vijayakumar, 2001; CESS526 from Shevroys, Tamil Nadu by Achyuthan Srikanthan and M.V. Shreeram in 2019.

Lineage diagnosis.

A cryptic lineage (L1) belonging to the C. gramineus complex, it is distinguished from C. occidentalis (L2) as follows: lower ventral scale count 158-179 (vs.142-154). It is genetically divergent from C. occidentalis by 8.1 % and 1.0 % at cyt b and 16S respectively. This lineage is far more widespread than its parapatric sister taxon C. occidentalis , occuring almost throughout peninsular India from Odisha in the east, southern Gujarat to the north-west and as far south as the Srivilliputhur hills.

Description of referred material (n=11).

A species with a slender, cylindrical body of snout to vent length (SVL) up to 679 mm and a prehensile tail; dorsal scales keeled with anterior dorsal scale rows (DSR) 19-21, mid body scale rows (MSR) 19-21 and posterior scale rows (PSR) 15; head prominent, clearly distinguished from the neck with small, juxtaposed scales on the head; rostral scale sub triangular with the upper side roughly one fourth the size of the lower side with the tip visible from above; supraoculars separated by 8-11 scales, between the posterior edge of the supraocular scales; canthus rostralis distinct with three to four canthal scales on the ridge; two to three preoculars, two to three postoculars and a thin elongated crescent shaped subocular; eye with a distinct elliptical pupil; temporal scales mildly keeled; aperture of the nostril completely covered by the nasal scale, undivided and subrectangular; nasal scale bordering the first supralabial; loreal pit present in contact with the second supralabial with two scales between the nasal and the second supralabial; 10-12 supralabials and 11-13 infralabials, with 9-13 scales between the last supralabial, including the last supralabial up to the start of the ventral scales; ventrals 158-179, laterally separated from the dorsal scale rows by a slightly broader row of dorsal scales anal scale undivided, divided subcaudals 54-67; terminal scale on the tail larger than the previous scale, blunt at the tip.

Colour in life.

Head and dorsum colour from verdant green to leaf green, sometimes bluish green with black, alternating saddle shaped markings on the dorsum; preocular/ temporal stripe in black, sometimes green fading into black; ventrals with bright yellow, creamish yellow to a dirty white colour; the region where the ventrals meet the dorsum alternating with the ventral colour once every 2-3 scales.

Habitat.

A highly arboreal species found in dry scrub, dry deciduous, semi evergreen and lowland riparian forests from 100-1600 m asl.

Distribution.

Endemic to Peninsular India. Found in the Eastern Ghats (Simlipal hills, Chota Nagpur plateau southwards upto Sirumalai hills) and hills of Central India (Pachmahri, Seoni hill ranges) as well as the Western Ghats from Surat Dangs, southwards till the Shencottah gap.

Figures

Figure 19. Craspedocephalus gramineus in life, showing: (A) entire lateral view (Thamini, Pune), (B) head and forebody dorsal view (Belapur, Navi Mumbai), (C) head and forebody lateral view, (D) entire ventrolateral view; from Kharghar, Navi Mumbai, Maharashtra.