Lycodon fasciolatus Shaw, 1802 comb. nov. - Russell's wolfsnake

Deepak, V., Narayanan, Surya, Mohapatra, Pratyush P., Dutta, Sushil K., Melvinselvan, Gnanaselvan, Khan, Ashaharraza, Mahlow, Kristin & Tillack, Frank, 2021, Revealing two centuries of confusion: new insights on nomenclature and systematic position of Argyrogena fasciolata (Shaw, 1802) (auctt.), with description of a new species from India (Reptilia: Squamata: Colubridae), Vertebrate Zoology 71, pp. 253-316 : 253

Lycodon fasciolatus Shaw, 1802 comb. nov. - Russell's wolfsnake

Coluber . - Russell 1796, p. 26 f., pl. 21, "Nooni Paragoodoo", locality not given, collector unknown; Anonymous 1796, p. 1667, "Nuni Paragudu"; Duncan and Duncan 1798, p. 5, "Nooni Paragoodoo"; Gmelin 1798, p. 21 "Nuhni Paraguhduh"; Bechstein 1802, p. 83 f., pl. 9, fig. 3, "Nuhni Paragudih" or "Russelsche Hofnatter"; Wilkes 1810, p. 804 f., 811, "Nooni paragoodo"; Buhle et al. 1835, p. 93, "Hofnatter, oder Nuhnih-Paragudih"; Druitt 1860, p. 166, "Nooni Paragoodo".

Coluber Fasciolatus Shaw, 1802, p. 528, 413, "Fasciolated Snake", type locality “India”, based on Russell’s (1796, p. 26-27) account of "Nooni Paragoodoo" and illustrated on plate 21, holotype lost according to Bauer (2015); Rees 1819, "Nooni Paragoodoo".

Coluber fasciolatus . - Gmelin 1809, p. 590, 649, "Fasciolated Viper"; Gray 1831, Appendix, p. 92; Cantor 1847a, p. 73, 150; Cantor 1847b, p. 919, 1069; Cantor 1848, p. 274; Froriep and Schomburgk 1848, col. 282; Jerdon 1853, p. 529; Blyth 1855b, p. 740.

Coluber hebe Daudin, 1803 p. 385 ff., type locality “Coromandel”, like Coluber fasciolatus Shaw based on Russell’s (1796) account of the "Nooni Paragoodoo", see above; Cuvier and Duméril 1818, p. 190, "La Couleuvre Hébé”, “Nouni-paragoudou”; Gray 1831, Appendix, p. 92; Hoeven 1856, p. 282.

Natrix [ Natrix ] Natrix hebe . - Merrem 1820, p. 95.

Lycodon Hebe. - Boie, H. 1826, p. 238 [partim]; Boie, H. in Boie, F. 1827, col. 551 [partim]; Schlegel 1827, col. 293 [partim]; Wagler 1830, p. 186; Schlegel 1837, pt. Générale, p. 78, 142, 231 [partim], pt. Descriptive, p. 106 [partim]; Filippi 1840, p. 183 [partim]; Fitzinger 1843, p. 27 [partim]; Burmeister 1850, p. 80; Beddome 1862, p. 22.

Lycodon aulicum . - Duméril, Bibron and Duméril 1854, p. 370 [ Varieté B, partim, non Lycodon aulicus (Linneus, 1758)].

Coryphodon fasciolatus . - Günther 1858, p. 109 [partim]; Gray 1863, p. 83 [partim].

Lycodon anamallensis Günther, 1864, p. 318 f., type locality "Anamallai Mountains", holotype BMNH 1946.1.14.92, coll. B.[sic] R.H. Beddome; Ganesh and Vogel 2018, p. 29 ff. [see also further citations in their synonym list].

Lycodon osmanhilli Taylor, 1950, p. 562 ff, pl. 20, type locality "Colombo, Ceylon", holotype KUMNH 24141, coll. W.C. Osman-Hill.

Lycodon fasciolatus .- this work.

Coluber fasciolatus formally named by George Kearsley Shaw (1802) in the third volume of the "General Zoology" series is entirely based on Russell’s (1796) description of a " Coluber " named "Nooni Paragoodoo" by native inhabitants of the "Coromandel Coast", encompassing the lowlands of the today’s Tamil Nadu and Andhra Pradesh states on the southeastern coast region of peninsular India. Besides short remarks on pholidosis, colour, pattern, measurements and brief comments on experiments about possible toxic effects of its bite ( Russell 1796, p. 26-27), the specimen is illustrated on plate 21 [reproduced here in Fig. 3 A ]. A complete German translation of Russell’s account was published by Johann Matthäus Bechstein (1802, p. 83-84) under "Nuhni Paragudih oder Russelsche Hofnatter", accompanied by a rearranged sketch on plate 9, figure 3, illustrating another position and different colouration [reproduced here in Fig. 3 C ]. Additional shortened German versions can be found in Anonymous (1796), Gmelin (1798), and Buhle et al. (1835), while further English versions were published by e.g. Gmelin (1809) and Wilkes (1810).

On the same basis and almost simultaneously as Shaw (1802), François Marie Daudin described Coluber hebe in 1803. Merrem (1820, p. 96, footnote a) was the first to recognise the synonymy of Coluber fasciolatus and Coluber hebe but he preferred to use Daudin’s younger name as valid and combined it with the generic name Natrix Laurenti, 1768. This synonymy went unnoticed by subsequent authors and while Coluber fasciolatus remained in the genus Coluber , Daudin’s Coluber hebe was relegated in the newly erected genus Lycodon by H. Boie (1826). A few years later Gray (1831), and subsequently Schlegel (1837) and Duméril, Bibron and Duméril (1854) recognized the conspecificity and continued to use Lycodon as the preferred genus. Like Merrem (l.c.), Schlegel (1837) also disregarded the priority of the name Coluber fasciolatus Shaw and considered specimens from Java to be identical to the taxon described from India by Shaw (l.c.) and Daudin (l. c.). It has to be noted that some subsequent authors erroneously credited the name Coluber hebe to Schlegel (e.g. Filippi 1840; Burmeister 1850; Günther 1861; Mason 1861; Gray 1863; Steindachner 1867; Müller 1878; Boulenger 1893; Wallach et al. 2014) or Boie (e.g. Fitzinger 1843; Hoeven 1856). The confusion was further increased by the synonymization of Coluber hebe Daudin with Coluber aulicus Linnaeus, 1758 and Lycodon capucinus Boie, H. in Boie, F., 1827 by Duméril, Bibron and Duméril (l.c.), two morphologically almost indistinguishable species with the same lepidosis and variable colour and pattern (see O’Shea et al. 2018).

In his "Catalogue of Colubrine snakes [ …] of the British Museum" Günther (1858, p. 203) listed a "Var. F." under the synonymy of Lycodon aulicus without voucher specimens. This refers to Russell’s (1769, p. 34-35, pl. 29) account on a " Coluber " ("Patza Tutta") from “Casemcottah” which is undoubtedly a specimen of what is commonly regarded as a "Banded racer" today, but not a Lycodon (wolfsnake). With his comment "belongs perhaps to Coryphodon fasciolatus " [= Coluber fasciolatus Shaw] he justified the confusion of two valid colubrid taxa, which continues today. In 1864 (p. 316) Günther noted: "The synonymy [of Lycodon aulicus ] in all preceding herpetological works is confused - C. striatus , C. malignus , C. hebe , C. fasciolatus , and probably C. capucinus belonging to other species". He ignored the previous generic allocation to Lycodon , excluded the aforementioned taxa from the synonymy of L. aulicus , relegated Coluber fasciolatus Shaw, and Coluber hebe Daudin together with the simultaneously synonymized Coluber curvirostris Cantor, 1839 to the genus Zamenis Wagler, 1830, and created a conglomerate of unrelated taxa of racers and wolfsnakes. From that point on, authorities used the name Coluber fasciolatus in combination with different genera as the scientific name for the Banded racer whereas C. hebe together with C. curvirostris were treated as junior synonyms (e.g. Theobald 1876; Boulenger 1890, 1893; Wall 1921; Smith 1943; Wallach et al. 2014; Bauer 2015; Uetz et al. 2019). At the same time, Russell’s "Nooni Paragoodoo" and C. hebe were also recognized as synonyms of Lycodon aulicus or L. capucinus (e.g. Boulenger 1893; Wall 1921; Iskandar and Colijn 2002; Wallach et al. 2014; Uetz et al. 2019).

Smith (1943, p. 531-532) studied the sketches in Russell’s (1796) 'Indian Serpents’ and identified the specimens depicted on plate 21 "Nooni Paragoodoo" together with that on plate 29 "Patza Tutta" as Coluber fasciolatus and continued to confuse the two different taxa. Further misidentifications are done in the list of determinations made from the Thomas Hardwicke collection of water-colour sketches deposited in the Zoological Library of the Natural History Museum, London, where he determined that four snakes depicted in sketch 55 and 61-63 were examples of Coluber fasciolatus ( Smith 1943, p. 529). In contrast, our re-examination of the illustrations revealed that sketch no. 55 depicts a wolfsnake, Lycodon sp., and no. 61 a Common ratsnake, Ptyas mucosa and only sketch 62 and 63 illustrates a Banded racer (see Fig. 8 ).

It is also to be noted that from the initial confusion caused by Günther (1864), the original scalation data of Shaw’s Coluber fasciolatus (192 ventrals and 62 subcaudals) were no longer considered representative of the scalation variation in the Banded racer. Not a single subsequent author, even when citing the original source, has included the values of the "Nooni Paragoodoo" in their variation of Coluber fasciolatus ; not even Wilson (1967) in his revision discusses the very low number of 62 subcaudals recorded for the type specimen.

Bauer et al. (2015) investigated two collections of dried snake skins housed in the Natural History Museum, London. The first collection arrived in 1837, the second in 1904 and are preliminarily attributed to Patrick Russell (see also Campbell 2015). The skins are attached to paper and in addition to their inventory numbers, for some specimens, handwritten notes are added which refer to Russell’s books of 'Indian Serpents’ (see Fig. 4 ). It is remarkable that all specimens from the 1904 collection labelled with the vernacular name "Nooni Paragoodoo" have been identified as Lycodon aulicus by Bauer et al. (2015). Our reexamination of the Lycodon specimens amongst Russell’s dried skin collections (including BMNH 1837.9.26.50-52; BMNH 1904.7.27.34, 43, 48, 52, 55, 72, 89 and 93) revealed that none of these specimens qualifies as the holotype of Coluber fasciolatus , as the measured lengths of these skins (ranging from 37 to 55 cm) are well below the 2 feet [60.96 cm] length reported by Russell for his "Nooni Paragoodoo".

In this work, we clarify the identity of C. fasciolatus , by analysing Russell’s (1796) original data and comparing it with specimens from various museums, data and photographs from the authors and literature concerning Indian snakes (e.g. Günther 1864; Boulenger 1890; Smith 1943; Whitaker and Captain 2004). We conclude that, in contrast to earlier determinations ( Günther 1864; Boulenger 1890, 1893; Smith 1943, to name a few authorities only), Russell’s "Nooni Paragoodoo" and therefore Coluber fasciolatus Shaw (incl. C. hebe Daudin) is not a racer but a wolfsnake of the genus Lycodon .

Wall (1914, p. 34) explained the meaning of the vernacular name "Nooni Paragoodoo" as: ‘Nooni’ = oil in Telugu language and ‘Paragoodoo’ stands for a runner or glistening, concerning the smooth and polished character of the scales in the snake. The extraordinary glossy character of dorsal scalation is well known for Lycodon species.

To determine the species affiliation we carefully compared Russell’s description and the sketch on plate 21 with published data of Lycodon species from southern and eastern India (Table 1 ). In terms of the combination of pholidosis, colouration and pattern, Russell’s specimens are generally comparable to a short banded morphotype of the Common Indian wolfsnake, Lycodon aulicus , a widespread species in the Indian subcontinent.

In a recent study on population systematics of the L. aulicus complex Ganesh and Vogel (2018) identified two different species-groups. Based on head dimensions, collar band pattern, hemipenal morphology, preocular-frontal and supraocular-prefrontal scale contact conditions, they reassessed the systematics of Lycodon aulicus , provided a redefined diagnosis for Coluber aulicus (sensu stricto) and revalidated Lycodon anamallensis Günther, 1864, a taxon so far regarded as conspecific with L. aulicus . Another result of Ganesh and Vogel’s (l.c.) analysis is the synonymization of Lycodon osmanhilli Taylor, 1950 with L. anamallensis .

Our comparison of the original information of Coluber fasciolatus Shaw, with data from the two morphotypes identified by Ganesh and Vogel (l.c.), showed the closest match with L. anamallensis (see Table 1 ). Although ventral and subcaudal counts of female Coluber fasciolatus lie within the ranges of females from both Lycodon anamallensis and Coluber aulicus , it is clearly distinguished from the latter by the combination of following diagnostic characters: (1) preoculars not in contact with frontal vs. in contact, (2) supraoculars in contact with prefrontals vs. not in contact, (3) collar absent vs. collar present and in contact with parietals, (4) upper labials yellow with dark brown center vs. upper labials whitish, (5) short whitish bands stippled with black, not extending into lateral edges of ventrals vs. elongated cream-white bands, extending onto lateral edges of ventrals.

Based on these consistencies of its morphological characters, colour and pattern, we consider Coluber fasciolatus Shaw, 1802 and Coluber hebe Daudin, 1803 as conspecific with Lycodon anamallensis Günther, 1864, including its synonym Lycodon osmanhilli Taylor, 1950.

With the reallocation of the taxon Coluber fasciolatus Shaw to the genus Lycodon we follow the lead of H. Boie (1826) and regard Lycodon fasciolatus (Shaw, 1802) as a valid combination with priority over its younger synonyms L. hebe , L. anamallensis and L. osmanhilli .

Description.

According to Taylor (1950, as Lycodon osmanhilli ), Ganesh and Vogel (2018, as Lycodon anamallensis ) and our own observations, Lycodon fasciolatus can be diagnosed as follows: (1) dorsal scales smooth in 17 rows at fore- and midbody and 15 before vent, (2) one to three preventrals followed by 174-204 laterally angulate ventrals (max. 197 for Indian populations), (3) 59-74 paired subcaudals, (4) a divided anal scale, (5) nasals in contact with first supralabial, naris large (6) internasals much smaller than prefrontals, (7) frontal shorter than its distance to tip of snout, shorter than parietals, (8) loreal elongated, nearly twice as long as high, (9) one preocular, usually not in contact with frontal (10) two postoculars, (11) supraocular usually in contact with prefrontal, (12) 2+3+4 temporals, (13) nine supralabials, third to fifth in contact with eye, (14) 10-11 sublabials, first five in contact with anterior submaxillars, (15) a thin, small head, reddish brown coloured, slightly darker than body, (16) iris black, pupil scarcely visible, (17) dorsal colour brown or reddish brown, (18) collar absent, first band just behind neck, curved backwards, (19) whole body with short cream white or yellow, irregularly shaped bands or blotches strippled with black, scarcely extending to the flanks, not reaching the lateral edges of ventrals, (adults sometimes patternless), (20) body lateral with four to five thin longitudinal interrupted whitish lines (21) supralabials creme white or yellow, anterior ones with a distinct dark brown spot in its center, (22) short hemipenis reaching to eighth subcaudal, covered with numerous long flounces and spines, (23) total length up to 52 cm, (24) relative tail length 0.14-0.20.

Distribution.

Russell (1796) does not provide specific information for the origin of his "Nooni Paragoodoo", Shaw (1802) only states “India” as the type locality for C. fasciolatus and Daudin (1803), referring to the title of Russell’s publication, i.e. “Coromandel” as the origin of his C. hebe . As already noted by Bauer (2015, p. 35) the description of C. fasciolatus is based on a single specimen and the snake depicted on plate 21 in Russell (1796) is the holotype. This makes Wallach et al’s (2014, p. 54) lectotype designation unnecessary and their type locality restriction "Vizagapatam, coast of Coromandel, India" [= Vishakhapatnam, Andhra Pradesh state, SE India, 17°41'N, 83°13'E, elevation 25 m] via lectotype selection" is unjustified because no such information was provided by Russell for the holotype. Even considering that Russell spent most of the time of his service for the East India Company in the Visakhapatnam district (Andhra Pradesh state of India), it is not clear whether he captured the individual specimen himself. Russell (1796) mentioned 11 persons by name who sent him material but does not name anybody in relation to the "Nooni Paragoodoo". Other specimens described by him without a named donor and therefore possibly captured by himself personally came from Lake Ankapilly, Bimblipatam, Boni, Masulupatam and Nerva in Andhra Pradesh, from Tanjore in Tamil Nadu, from Ganjam in Odisha and Hyderabad in Telangana. Strictly speaking, besides Vizagapatnam any of these places could be considered as the possible type locality of L. fasciolatus .

Lycodon fasciolatus seems to be widespread in southern peninsular India and recent records from the Coromandel region support the possible demarcation of the type locality in this area. According to Das and de Silva (2005, as L. osmanhilli ), Ganesh and Chandramouli (2011, as L. aulicus morph 2), Ganesh and Vogel (2018, as L. anamallensis ), and Madawala et al. (2019, as oanamallensis [sic] or anamalensis [sic]), L. fasciolatus is known from the Indian states of Kerala (Kannur, Mundakayam), Tamil Nadu (Anaimalai Hills, Chennai, Mannampandal), Andhra Pradesh (Nallamala Hills) and from Sri Lanka (Ampara, Andigama, Chilaw, Colombo, the Jaffna Peninsula, Kandy, Matara, Peradeniya, Tabbowa, Yala). In the present study, the species is found to be common along the Ganjam coast up to Chhatrapur and we also report the northernmost record from Ganjam, Odisha state of India (see Appendix 10). This species was found inhabiting Pandanus odorifer ( Forsskål), Kuntze bushes, other scrub vegetation, agricultural fields and occasionally entering human habitation.

Some old records mention a specimen under the name Coluber or Coryphodon fasciolatus ( Blyth 1855a, p. 291; Günther 1860, p. 163) collected by Major W.S. Sherwill at Darjeeling (West Bengal, India) but this refers to another taxon, probably also a Lycodon species.

Günther’s (1861, p. 218) record of C. fasciolatus from the central hilly region of Nepal, based on a coloured sketch in the B. H. Hodgson’s collection, shows a colubrid snake different from both wolfsnakes and racers and will be discussed below.

Recently, O’Shea et al. (2018) reviewed the complex taxonomic history and confusion surrounding L. capucinus and L. aulicus and finally considered Lycodon capucinus , also based on molecular data published by Siler et al. (2013), as a valid species. They discussed the distribution of both species and presented a map showing the actual known range of L. capucinus with its westernmost mainland border running between northeastern India and Bangladesh. At the same time, O’Shea et al. (l.c., p. 71) stated that "[...] L. hebe ( Daudin 1803), [is] now a synonym of L. capucinus ; [...]" but neglected the fact that the origin of Coluber hebe Daudin, with its type locality on the Coromandel Coast of India, contradicts their presented distribution for L. capucinus .

Remarks.

Blyth (1855b, p. 740, smallprint) described Coluber vittacaudatus (collector unknown, holotype not located) from the vicinity of Darjeeling (West Bengal state, India) and considered it as allied to Coluber fasciolatus Shaw. The taxon fell into oblivion until Das et al. (1998, p. 157) listed it again as a valid species based on a personal communication by Van Wallach. Whitaker and Captain (2004, footnote 6) were unsure where C. vittacaudatus should be placed and Schätti et al. (2014, p. 384, smallprint) regarded it as "possibly a senior synonym of the Oriental ratsnake Orthriophis taeniurus yunnanensis (Anderson, 1879), and of O. taeniurus (Cope, 1861) as well." The alleged relationship to C. fasciolata Shaw, as originally suggested by Blyth (1855b), prompted Wallach et al. (2014) and later Das and Das (2017) to place Coluber vittacaudatus into the previously monotypic genus Argyrogena Werner, unaware that Coluber fasciolatus Shaw is not a racer.

It is undisputed that Blyth (l.c.) considered the resemblance of C. vittacaudatus to C. fasciolatus in the sense of Shaw, so he clearly did not mean what is now commonly referred to as the Banded racer. Because of our clarification that C. fasciolatus is a wolfsnake, all previous assumptions concerning the identity of Coluber vittacaudatus Blyth are questionable.

Figures

Figure 3. Lycodon fasciolatus comb. nov. A. Holotype of Coluber fasciolatus Shaw, 1802, as well as holotype of Coluber hebe Daudin, 1803, " Nooni Paragoodoo " reproduced from Russell (1796, pl. 21), B. uncollected live specimen from Rushikulya, Ganjam, Odisha state, India, and C. " Nuhni Paragudih oder Russelsche Hofnatter ", rearranged sketch from Russell (1796, pl. 21) reproduced by Bechstein (1802, pl. 9, fig. 3).

Figure 4. Lycodon fasciolatus comb. nov. Previously unpublished extract from a paper sheet with glued specimens from the 1904 collection of dried snake skins preliminarily attributed to Patrick Russell showing a specimen of the " Nooni Paragoodoo " kept in the herpetological collection of the Natural History Museum, London, U. K. (BMNH 1904.7.27.93).

Figure 8. Platyceps plinii comb. nov. Previously unpublished original colour-painting from the Thomas Hardwicke collection of sketches (Volume II, sketch 63) kept in the Zoological Library of the Natural History Museum, London, U. K. (see also Smith 1943, p. 529). Reproduced with permission from the Natural History Museum, London, U. K.