Platyceps plinii (Merrem, 1820) comb. nov. - Banded racer

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

Platyceps plinii (Merrem, 1820) comb. nov. - Banded racer

Coluber . - Russell 1796, p. 34-35, pl. 29 [reproduced here in Fig. 5 A ], "Patza Tutta", from “Casemcottah” [=Kasimkota village Visakhapatnam district, Andhra Pradesh state, India; see Note 1], two specimens coll. Captain Gowdie and Lieutenant Whyte; Anonymous 1796, p. 1667, "Patza Tutta"; Gmelin 1798, p. 26, "Patza Tutta"; Bechstein 1802, p. 273 f., pl. 44, fig. 2, "Die halbbandierte Natter" or "Patza Tutta" [reproduced here in Fig. 5 B ]; Gmelin 1809, p. 685 f. "Patza Tutta"; Wilkes 1810, p. 813 "Patza tutta"; Buhle et al. 1835, p. 97, "Die halbbandierte Natter, (Patza Tutta)".

Coluber pictus Daudin, 1803, p. 347-349, "La Coleuvre Peinte", type locality "Casem-Cottah, au Bengale", based on Russell’s (1796, p. 34-35, pl. 29) account of "Patza Tutta", holotype not located; Cuvier and Duméril 1818, p. 190, “patza-tutta”; Boie 1827, col. 524; Schlegel 1837, pt. Descriptive, p. 172; Traill 1843, p. 148; Cantor 1847a, p. 67; Cantor 1847b, p. 913; Jerdon 1853, p. 529; Blyth 1854, p. 416; Blyth 1855a, p. 292 [nec Coluber pictus Gmelin, 1789, p. 1116; nec Coluber pictus Georgi, 1801, p. 1883; nec Coluber pictus Pallas, 1814, p. 54 [sic], 45; nec Coluber pictus Theobald, 1868a, p. 45; all primary homonyms].

Natrix [ Natrix ] Plinii Merrem, 1820, p. 101 f., "halbbandierte [Natter]", type locality “Bengalen”, restricted here to “Casemcottah” (= Kasimkota village, Andhra Pradesh state, India; see Appendix Note 1), based on Russell’s (1796, p. 34-35 pl. 29) account of "Patza Tutta", type material not located, lectotype by present designation: specimen depicted on plate 29 in Russell (1796); Anonymous 1822, col. 693.

Coluber plinii . - Boie 1827, col. 524; Schlegel 1837, pt. Descriptive, p. 172; Traill 1843, p. 148; Cantor 1847a, p. 67; Cantor 1847b, p. 913; Blyth 1855a, p. 292; Wallach et al. 2014, p. 219 [wrongly attributed as a synonym of D. pictus Gmelin].

Coluber curvirostris Cantor, 1839: 51, type locality “Bengal”, holotype not located, an unpublished original sketch depicting the iconotype is kept in the Bodleian Library (Oxford, U.K.), [reproduced here in Fig. 7 ]; Cantor 1841, p. 82.

Coluber fasciolatus . - Mason 1861, p. 708; Smith 1928, p. 495; Smith 1943, p. 170, [partim], p. 529 [partim, Hardwicke sketches vol. II, plate 62, 63 only, the latter reproduced here in Fig. 8 ]; Constable 1949, p. 127; Taylor 1950, p. 546; Deranyiagala 1955, p. 25; Minton 1962, p. 17; Swan and Leviton 1962, p. 140; Anderson 1964, p. 163; Minton 1966, p. 123; Underwood 1967, p. 134; Mertens 1969, p. 54; Singh 1972, p. 190 [as Coluber fasciolata [sic]; Waltner 1973, p. 15; Sharma and Diksit 1976, p. 171; Khan 1977, p. 147; de Silva, P.H.D.H. 1980, p. 256 f; Mahendra 1984, p. 289; Shrestha and Shah 1985, p. 51; Amr and Disi 2011, p. 191.

Coronella baliodeira . - Cantor 1847a, p. 67 [partim, "Patza Tutta"]; Cantor 1847b, p. 913 [partim, "Patza Tutta"], [non Coronella baliodeira Boie, 1827].

Coryphodon fasciolatus . - Günther 1858, p. 109 [partim]; Beddome 1862, p. 16 f. [partim]; Cope 1863, p. 338; Gray 1863, p. 83; Jan 1863, p. 64 [partim].

Tyria [ Coluber fasciolata ]. - Cope 1863, p. 338.

Zamenis fasciolatus . - Günther 1864, p. 254 f., [partim]. pl. 21, fig. F; Theobald 1868a, p. 47; Anonymous 1870, p. 46; Anderson 1871, p. 34; Stoliczka 1871a, p. 191; Stoliczka 1871b, p. 431; Nicholson 1874, p. 87, pl. 13, fig. 1; Theobald 1876, p. 170 f. [partim]; Anonymous 1877, p. 44; Ferguson 1877, p. 19; Blanford 1879, p. 114; Müller 1884, p. 285; Phipson 1886, p. 84; Haly 1886, p. 10; Boulenger 1890, p. 327 [partim]; Haly 1891, p. 3, 21; Sclater 1891, p. 28; Boulenger 1893, p. 404 [partim]; Bethencourt Ferreira 1897, p. 225; Cardew 1897, p. 593; Willey 1906, p. 233; Wall 1907, p. 115; Wall, 1908, p. 21, footnote; Abercromby 1910, p. 6; Abercomby 1913, p. 304; Wall 1913, p. 20, footnote; Cazaly 1914, p. 26 f.; Wall 1914, p. 34, pl. 22 [partim]; Wall 1921, p. 191 ff.; Prater 1924, p. 157, 169 f; Wall 1924, p. 618; Mullan 1927, p. 380; Amaral 1929, p. 11; Nicholls 1929, p. 116, 128; Werner 1929, p. 65, 71 [partim]; Gharpurey 1931, p.1084; Gharpurey 1932a, p. 465; Gharpurey 1932b, p. 906; Gharpurey 1932c, p. 273 f.; Lindberg 1932, p. 691 ff.; Gharpurey 1935a, p. 943; Gharpurey 1935b, p. 198; Bourret 1936, p. 175; Fraser 1936, p. 58 ff.; Lindberg 1939, p. 333; Themido 1941, p. 20.

A. [rgyrogena] Argyrogena rostrata Werner, 1924, p. 51, fig, 4a-c, type locality “Argentinien” [in error], holotype NMW 18160; Dunn 1928 p. 23 [ Argyrogena ]; Amaral 1929, p. 11.

Argyrogena fasciolata . - Wilson 1967, p. 261 [new comb.], [partim]; Kumar et al. 1976, p. 92; Kramer 1977, p. 735; Khan 1980, p. 135, 144; Tiedemann and Häupl 1980, p. 50; Khan 1982, p. 4; Murthy and Acharjyo 1987, p. 140; Welch 1988, p. 36; Williams and Wallach 1989, p. 12; de Silva 1990, p. 77 f.; Schleich 1993, p. 146; de Silva 1994, p. 15; Shah 1995, p. 38; Anders et al. 1997, p. 53; Das and Andrews 1997, p. 4; Shah 1997, p. 27; Das 1998, p. 44; Das and Ota 1998, p. 184; Shah 1998b, p. 27; Shrestha 1998, p. 39; Gayen 1999, p. 15; Smith and David 1999, p. 46; Daniels 2000, p. 8; Vyas 2000a, p. 20 [as Argyrogena stolatum ]; Shrestha 2001, p. 171; Gruber 2002, p. 820 ff.; Khan 2002, p. 79 ff.; Das 2003, p. 474; Shrestha 2003, p. 634; Khan 2004, p. 196; Shah and Tiwari 2004, 154; Whitaker and Captain 2004, p. 138; Somaweera 2006, p. 50: Khan 2006, p. 192 ff.; map and p. 275; Srinivasulu et al. 2006, p. 113; Whitaker 2006, p. 31 ff.; Hutton and David 2008, p. 314; Khaire 2008, p.68; Rameshwaran 2008, p. 22, back cover [partim]; Satish 2008, p. 88; Srinivasulu and Das 2008, p. 119; Dutta et al. 2009, p. 108; Aengals 2009, p. 193; de Silva and Jinasena 2009, p. 6; Kabir et al. 2009, p. 118 f.; Prakash and Raziuddin 2009, p. 12; Ingle 2011, p. 83; Joshi 2011; p.1 f.; Sharma and Warman 2011, p. 62; Thakur 2011, p. 3; Vijayaraghavan and Ganesh 2011, p. 40 [partim]; Amol et al. 2012, p. 364; Bhandarkar et al. 2012, p. 45; Chikane and Bhosale 2012, p. 7 image 5D, p. 14 table; Das 2012, p. 73; Khan et al. 2012, p. 166; Kumbhar et al. 2012, p. 364; Walmiki et al. 2012a, p. 96; Walmiki et al. 2012b, p. 9; Kästle et al. 2013 p. 197; Khaire 2010, p. 36; Pande et al. 2013, p. 29 f.; Vyas 2013, p. 38; Fellows 2014, p. 2; Hasan et al. 2014, p. 183; Ingle et al. 2014, p. 136; Joshi et al. 2014, p. 168; Khaire 2014, p. 44; Khedkar et al. 2014, p. 8; Porob et al. 2014, p. 179, image; Pradhan et al. 2014, p. 244; Schätti et al. 2014, p. 384, smallprint; Seetharamaraju 2014, p. 95 ff.; Wallach et al. 2014, p. 54; Ahmed et al. 2015, p. 56; Bauer 2015, p. 35, 37; Bauer et al. 2015, p. 76; Charjan and Joshi 2015, p. 2; Deshmukh et al. 2015, p. 38 f.; Joshi et al. 2015, p. 24; Koushik, 2015, p. 91; Manhas et al. 2015a, p. 1870; Palot 2015, p. 8016; Patel et al. 2015, p. 122, fig. 3; Pradhan and Talmale 2015, p. 117; Sayeswara et al. 2015, p. 498; Bansode et al. 2016, p. 105 f.; Dileepkumar et al. 2016, p. 53 f.; Khan 2016, p. 23; Pasar and Paul 2016, p. 343; Tambre and Chavan 2016, p. 108 f.; Das and Das 2017, p, 165; de Silva and Ukuwela 2017, p. 171; Gayen et al. 2017, p. 19; Joshi et al. 2017, p. 584; Kanaujia et al. 2017, p. 121; Paliwal and Bhandarkar 2017, p. 3440; Sheikh 2017, p. 56; Aengals et al. 2018, p. 22; Bansode and More 2018, p. 377; Ganesh et al. 2018, p. 42; Jadhav et al. 2018, p. 1858, 1860; Khan et al. 2018, p. 1731; Manhas et al. 2018, p. 109; Pasar 2018, p. 5; Patil 2018, p. 6; Das and Pramanick 2019, p. 94; Das et al. 2019, p. 311 ff.; Gemel et al. 2019, p. 171; Ingle et al. 2019, p. 81; Madawala et al. 2019, p. 251; Mohalik et al. 2019, p. 150; Patel and Vyas 2019, p. 771; Patel et al. 2019a, p. 23; Patel et al. 2019b, p. 148 f.; Sagadevan et al. 2019, p. 13567 [partim], Rajabizadeh et al. 2020, p. 6.

Argyrogena fasciolatus [sic]. - Whitaker 1978, p. 31. [partim]; Koul and Murphy 1979, p. 48; Kutty et al. 1981, p. 167; Sharma 1982, p. 123 [partim]; Whitaker 1982, p. 31. [partim]; Murthy 1985, p. 64; Vyas 1987, p. 228; Green 1988, p. 144; Khan 1988, p. 13 [as Argyrogene [sic] Coluber fasciolatus ]; Daniel 1989, p. 78; Dutta and Acharjyo 1990, p. 41; Murthy 1990, p. 27 f.; Ahmed and Dasgupta 1992, p.42; Murthy 1992, p.173; Negi 1992, p. 129; Khan 1993, p. 89, fig. 8; Das 1994, p. 31; Sanyal and Sur 1995, p. 60; Das 1996, p. 54; Khaire 1996, p. 69; Das 1997, p. 40; de Silva 1998a: 64; de Silva 1998b, p. 7; Shah 1998a, p. 29; Captain and Thakur 1999, p. 147 ff.; Khan 1999, p. 286; Sharma 1999a, p. 42; Sharma 1999b, p. 95; Sharma 1999c, p. 112; Vyas 1999, p. 5; Islam et al. 2000, p. 77; Sharma 2000, p. 277; Vyas 2000b, p. 388; Daniels 2001a, p. 7; Daniels 2001b, p. 16; de Silva 2001, p. 59; Ingle 2001a, p. 14; Ingle 2001b, p. 23; Vyas 2001a, 204; Vyas 2001b, p. 23; Vyas 2001c, p. 18; Daniel 2002, p. 94 f.; Das 2002, p. 20; Ingle 2002, p. 9; Khan 2002, p. 79 f.; Lobo 2002, p. 6; Sharma 2002a, p. 51; Sharma 2002b, p. 63; Bhuvaneswari 2003, p. 169; Ingle 2003, p. 4; Palot and Radhakrishnan 2003, p. 4; Shah et al. 2003, p.13; Sharma 2003, p. 82 f.; Dasgupta and Raha 2004, p. 167; Khan, M.A.R. 2004, p. 18; Pradhan 2004, p. 111; Somaweera 2004: 37; Vyas 2004, p. 1514; Chandra and Gajbe 2005, p. 83; Das and de Silva 2005, p. 24; Gajbe and Gupta 2005, p. 10; Rao et al. 2005, p. 1739; Sharma 2007, p. 132 f.; Vyas 2007, p. 13; Chandra et al. 2008, p. 52, 54, 71; Ingle 2008, p. 15; Pradhan 2008, p. 327; Anonymous 2010, p. 56; Bahuguna 2010, p. 501; Singh and Ngullie 2010, p. 56; Thakur 2010, p. 15; Patankar 2011, p. 9; Sawant 2011, appendix 2, p. 2; Vyas 2011, p. 11; Chauhan and Shingadia 2012, p. 64; Dasgupta et al. 2012, p. 217; Upadhye et al. 2012, p. 16; Aengals and Pradhan 2013, p. 454; Reddy et al. 2013, p. 519; Raut et al. 2014, p. 374, 376; Manhas et al. 2015b, p. 1550 [as Argyrogene [sic] Coluber fasciolatus ]; Anonymous 2017, Annex 5, p. 5; Vibhute 2018, p. 79; Anonymous n. d., p. 13.

Platyceps plinii . - this work.

The preceding list of synonyms and chresonyms contains quotations which to our best knowledge refer to the snake species so far known as Banded racer ( Argyrogena fasciolata auctt.) in the sense of Günther (1864). Because of our clarification that Coluber fasciolatus Shaw represents a wolfsnake of the genus Lycodon (see above), it is necessary to assign an adequate scientific name to the Banded racer. In this context, two well-known names are available from the list of synonyms, i.e. Coluber curvirostris Cantor, 1839, and Argyrogena rostrata Werner, 1924. The first name was synonymized with C. fasciolatus by Günther (1864) and the latter by Smith (1928).

Coluber curvirostris was based on a single specimen from “Bengal” described by Theodore Cantor (1839) in his "Specilegium Serpentium Indicorum". A coloured sketch of the holotype is deposited in the Bodleian Library (Oxford, U.K.) and reproduced here for the first time (Fig. 7 ).

Franz Werner (1924) described Argyrogena rostrata (genus et species nova) based on a specimen stored in the Natural History Museum, Vienna, which was originally listed as having come from “Argentinien” and for a few years it was considered a valid neotropical taxon ( Dunn 1928; Amaral 1929). Malcolm A. Smith, who visited the herpetological collection of the Natural History Museum, Vienna in 1927, examined several of Werner’s type specimens, including the holotype A. rostrata , recognized its true identity, synonymized it with Coluber fasciolatus and relegated Argyrogena to the synonymy of Coluber . This classification was largely maintained until Wilson (1967) dedicated a study to the Banded racer and in this context revalidated Werner’s name Argyrogena . Subsequently, the combination Argyrogena fasciolata (Shaw, 1802) was established and has been used for the Banded racer until today (see list of synonyms and chresonyms and e.g. Williams and Wallach 1989; Smith and David 1999; Whitaker and Captain 2004; Khan 2006; Somaweera 2006; Wallach et al. 2014; Koushik, 2015; Das and Das 2017; Uetz et al. 2019, to name a few general publications). In a short overview of the taxonomy of Coluber , Amr and Disi (2011, p. 191) erroneously stated that Wilson (1967) reallocated the taxon Coluber fasciolatus to the genus Platyceps .

We also tracked down two older and previously overlooked or wrongly assigned names that clearly refer to what is currently understood to be the Banded racer, viz. Coluber pictus Daudin, 1803 and Natrix plinii Merrem, 1820. Again both names are entirely based on the same description in Russell (1796, p. 34-35, pl. 29: " Coluber " named "Patza Tutta" by natives).

Boie (1827) considered Coluber plinii and Coluber pictus as synonymous and a Coluber close to Coelognathus helena (= Coelognathus helena (Daudin, 1803)). Cantor (1847a, b) on the other hand regarded Daudin’s Coluber pictus and Merrem’s Coluber plinii to be identical to Coronella baliodeira Boie, 1827, an assessment that Blyth (1854, p. 416, 1855a, p. 292) contradicts by addressing both taxa as true Coluber based on a juvenile specimen procured in South India by Jerdon (1853, p. 529; ZSI K 7333), which was later listed by Theobald (1868a, p. 47) under Zamenis fasciolatus .

From the mid of the 19th century onwards both names fell into oblivion as potential synonyms of the Banded racer. This was probably also due to Schlegel (1837, pt. Descriptive p. 172) who regarded both Coluber pictus Daudin and Coluber plinii Merrem as possibly identical with Coluber triscalis Linnaeus, 1758 and Coluber corallinus Linnaeus, 1758. The latter two are currently regarded as synonyms of the Caribbean colubrid snake Liophis (now Erythrolamprus ) Coluber triscalis (see Dundee 1994).

Coluber pictus Daudin, 1803 however, is threatened for reasons of homonymy by an older name, Coluber pictus Gmelin, 1789, the accepted original combination for the Painted bronzeback (now Dendrelaphis pictus , see ICZN 1958). Therefore, Coluber pictus Daudin, 1803 represents a younger primary homonym of Coluber pictus Gmelin, 1789 and is unavailable (ICZN 1999, Art. 57.2; see also above the list of synonyms and chresonyms for further homonymous names).

Merrem (1820) used Russell’s (1796) account of the "Patza Tutta" and formally named it Natrix Plinii . His diagnosis was very short and besides some general features concerning habitus and characteristics of dorsal scales, he mentioned only the number of ventral (202) and subcaudal (91) scales. Additional information can be found in Russell’s (1796, p. 34-35, pl. 29) original text and plate. He described the "Patza Tutta" with the following characters: 1) head small, very little broader than neck, ovate, depressed, 2) two internasals, 3) two prefrontals, larger than internasals but similar in form, 4) two supraoculars, purse-shaped, 5) frontal bell-shaped, 6) parietals semi heart-shaped, truncate, 7) mouth wide jaws unequal, lower considerably shorter, 8) teeth small sharp, reflex, two palatal rows, and one marginal [maxillar], in the upper jaw, 9) eyes distant, lateral, large, orbicular, 10) body round, covered with very small smooth, ovate scales, outer row orbicular, 11) length one foot eleven inches [58.4 cm], tail five inches [12.7 cm], very tapered and sharply pointed, 12) dorsal colour brown, forebody with narrow crossbands, composed of short, black and white lines, behind these, a few obscure reddish-brown bands interspaced by white colour of interstitial skin; bands fading at posterior part of body, 13) outermost dorsal scale row yellow, 14) ventral with pale-yellowish cast. The description resembles the specimen depicted on plate 29 and in comparison with our own and published data (see below) it corresponds to male specimens of the Banded racer.

To our best knowledge, Natrix plinii Merrem, 1820 is the oldest available name for the snake species which is commonly now regarded as Banded racer. Merrem (l.c.) simplified the type locality to “Bengal”, but according to Russell (1796, p. 35) two specimens of the "Patza Tutta" were sent to him from “Casemcottah” by Captain Gowdie in 1788 and by Lieutenant Whyte without year specification (see Appendix Note 1).

Type material.

We could not locate the two type specimens of Natrix plinii Merrem, 1820 among Russell’s dry and wet preserved material stored in the collections of the Natural History Museum, London, U.K. A wet preserved topotypic male ( BMNH 37a) donated by P. Russell according to Günther (1858, p. 110) from "East Indies", restricted to "Vizagapatam District" by Boulenger (1893, p. 405), does not correspond with the original data for the "Patza Tutta" published by Russell (1796 p. 34-35, pl. 29). Because of its lower ventral scale count (199 vs. 202), an incomplete tail (with 51 vs. 91 subcaudals) and a shorter snout-vent length (43.2 vs. 58.5 cm [1 ft 11 inch]), specimen BMNH 37a can be excluded as one of the syntypes. Also among the dried skin collections attributed to Russell (including BMNH 1837.9.26.48-49 and BMNH 1904.7.27.81) no specimen corresponds to his original data of the "Patza Tutta".

The same applies to the type specimen of Coluber curvirostris Cantor, 1839 from “Bengal” which is not yet located. Although some of Cantor’s original specimens are known to be stored in the Natural History Museum, London ( Adler 2007, 2016), we were unable to identify the holotype of C. curvirostris among the preserved specimens in the collection. A possible candidate donated by the Zoological Society of London to the British Museum (Nat. Hist.) ( BMNH 1851.9.13.260 from “India”), can be excluded as the holotype because it does not correspond to Cantor’s original data and plate, in respect of the number of ventrals (226 vs. 220) and subcaudals (88 vs. 85). A transcript of Cantor’s unpublished original manuscript for C. curvirostris , with an expanded description, is presented here in the Appendix (Note 2).

Description.

Based on our own data and summarized information from literature ( Russell 1796; Smith 1943; Wilson 1967; Whitaker and Captain 2004), Platyceps plinii is characterized as follows: (1) dorsal scales smooth in 23-25 (exceptionally 21 or 22) rows at forebody, 23 (exceptionally 21 or 25) at midbody and 17 (exceptionally 16 or 15) before the vent, (2) one to three preventrals followed by 190-234 ventrals, (3) 73-101 paired subcaudals, (4) a divided cloacal plate, (5) nasals divided, in contact with first and second supralabial, naris medium sized (6) internasals smaller than prefrontals, (7) frontal shorter than its distance to tip of snout, as long as or little shorter than parietals, (8) loreal usually slightly longer than high, rarely as long as high or higher than long, exceptionally fused with prefrontal, (9) usually two (80% of examined specimens) exceptionally three preoculars, one larger upper preocular usually in contact with frontal and one (exceptionally two) very small lower presuboculars wedged between third and fourth supralabial, (10) two (exceptionally three) postoculars, (11) supraocular not in contact with prefrontal, (12) temporals variable, in anterior two (rarely three or one) and secondary three (rarely two) rows, (13) eight (exceptionally seven, nine or ten) supralabials, fourth and fifth (rarely fifth and sixth or sixth and seventh) in contact with the eye, (14) usually ten, sometimes nine or eleven sublabials, first four to five in contact with anterior submaxillars, (15) head wide and moderately pointed, barely distinct from neck, body round, olive brown to reddish brown coloured, patternless in adults or with light vermicular pattern in juveniles, (16) iris dark grey or blackish, pupil black with light brown corona, (17) dorsal colour greyish or reddish brown, (18) collar absent, (19) whole body patternless or with indistinct small light bands on forebody in adults; in juveniles with distinct black and white strippled bands reaching lateral edges of ventrals, extending to near vent, (20) supralabials creme brown, patternless, or with small light spots in juveniles (21) throat, ventral body and tail yellowish cream without pattern, (22) hemipenis simple; proximal fourth smooth; distal three fourths covered with small spines gradually decreasing in size distally; distal area with irregular shaped calyces, denser towards the apex (23) total length up to 148 cm, (24) relative tail length 0.19-0.31.

Variation.

Pholidosis: Head 2.17-2.14 times longer than broad (males 1.17-2.14, females 1.22-2.05), canthus rostralis moderate developed. Rostral nearly twice as broad as high, clearly visible from above. Internasals usually shorter than prefrontals, sometimes equal in length in, e.g. BNHS 646 and BNHS 319, or slightly longer in, e.g. BMNH 62.8.14.29, MCZ R-28645, NMW 25465:3 and SMF 50409. Frontal 1.01-1.46 times longer than maximum width (males 1.01-1.46, females 1.02-1.32), 1.05-1.63 times longer than internasals and prefrontals (males 1.05-1.63, females 1.16-1.42), in Pakistani population from southern Sind equal in length in, e.g. SMF 50410 or slightly shorter incl. SMF 50406-407, SMF 50409, SMF 50446, SMF 62922 and in a specimen from ‘Bombay’ BNHS 646. Posterior border of parietals usually more or less straight or slightly indented (< shaped) at the median suture or, sometimes, forming an obtuse angle in, e.g. MNHN RA-0.6230. Posterior edge of parietals less than half of the maximum width. Loreal usually longer than high, equal in BNHS 3193, shorter in, e.g. BNHS 2857, BNHS 3107, ZMB 8053 (on right side) and SMF 50407 (on right side) or fused with prefrontal in SMF 62921 (on right side) and SMF 50446 (on both sides). Preocular single or with an additional small pre-subocular below it or rarely with two small pre-suboculars in SMF 62922. Preocular in contact with frontal or rarely separated in, e.g. MNHN RA-0.6230.

Predominantly eight supralabials, last two or three being larger (longer), fifth highest; nine in BMNH 1921.6.15.14-15, SMF 57311 (on left side), BNHS 3175 (on both sides), or seven in MCZ R-28645 (on right side). Usually fourth and fifth supralabials in contact with eye, fifth and sixth sometimes in specimens with nine supralabials, e.g. SMF 57311. Two postoculars except in a specimen from “India” without exact locality ( BMNH 62.8.14.29) with three postoculars on left side; upper scale somewhat wider, lower usually higher. Normally two anterior temporals, sometimes three including BMNH 62.8.14.29 and NMW 18160 (on left side) and SMF 57311 (on right side), exceptionally one in NMW 25465:3 (on both sides). Secondary temporals variable, mostly three, sometimes two e.g. SMF 57312, SMF 50407 and SMF 50446 (on both sides), BNHS 642, SMF 62921, SMF 50406, and SMF 50410 (on left side) and BNHS 3107, SMF 50409 and SMF 62922 (on right side), or exceptionally four in e.g. ZMA RENA 12554 (left side). Mostly ten sublabials, sometimes eleven, including BMNH 1921.6.15.14 and SMF 57311 (on both sides), SMF 50407 and BMNH 37A (on left side), BNHS 646, BNHS 3175, SMF 50406 and SMF 50410 (on right side), or rarely nine, e.g. SMF 62922 (both sides); the anterior four to five scales in contact with first inframaxillary, the sixth or rarely seventh largest. Anterior inframaxillars normally longer and wider than, or about equal to, posterior pair, except in BMNH 62.8.14.29, BNHS 657, BNHS 3093, BNHS 3107, MNHN RA-0.6230, SMF 50407, SMF 62921-62922 and ZMB 8053. The posterior pair of inframaxillars usually separated by two or three rows of gular scales of variable shape and size. Gulars in three to four oblique rows between the apical edge of the posterior inframaxillars and first ventral. Ventrals in examined material 190-231 (males 190-220, females 215-231), last plate divided in SMF 50407. Usually two, sometimes one or three preventrals. Cloacal plate divided, right part overlapping left. Subcaudals in 73-101 pairs (males 81-101, females 73-89). Total body scales, including preventrals and terminal scale 273-321 (males 273-315, females 296-321). Dorsal scales, usually arranged in 23-25/23/17 rows along the trunk. Three specimens from Maharashtra ( BMNH 3093, BMNH 642 and BMNH 646) show 27 anterior DSR; one specimen each from “Calcutta” (NMW 25463: 3), “India” ( BMNH 37 A), and “Bangalore” ( BMNH 1920.7.7.5) shows 21, 22 and 24 anterior DSR respectively. One specimen from Sind ( SMF 50409) with 25 and one from “India” ( BMNH 37 A) with only 21 midbody DSR and two specimens from Bangalore ( BMNH 1921.615.14, BMNH 1920.7.7.5) with 16 DSR at posterior part of body. In contrast to Constable (1949) we counted 23 midbody DSR instead of 21 for MCZ R-28645 from “Bangalore”. Dorsal scale with paired apical pits. Supracaudal scales usually with two apical pits, up to three pits on the first three scales followed the supracaudal reduction.

Dorsal scale reduction formula summarized from 31 examined specimens (see Appendix 10; divergent data from literature [ Wilson 1967] in square brackets). Only main reductions are given.

2+3, 3+4, or 10+11 (6-11) 2+3, 3+4, 7+8, 8+9, 9+10, or 10+11 (65-131) [79-133]

25 -------------------------------- 23 -------------------------------------------------------------------

3+4, or 10+11 (5-9) 2+3, 3+4, 7+8, 8+9, 9+10, or 10+11 (63-131) [82-129]

2+3, 3+4, 4+5, 8+9, 9+10, or 10+11 (68-145) [105-137]

21 ------------------------------------------------------------------------

1+2, 2+3, 3+4, 4+5, 7+8, 8+9, or 9+10 (68-142) [107-138]

4+5, 6+7, 7+8, 8+9, or 9+10 (111-192) [157-174] 7+8 (188-194)

19 ------------------------------------------------------------- 17 --------------------------- 15.

3+4, 4+5, 7+8, 8+9, or 9+10 (112-191) [156-178] 7+8, or 8+9 (190-195)

Six examined specimens ( BMNH 1851.9.13.260, BMNH 1920.7.7.5, BMNH 1921.6.15.14, BMNH 1940.3.4.45, NMW 18160, and SMF 50409, all females) show an additional lateral or bilateral increase to 18 or 19 dorsal scale rows involving rows one to three between 89 and 99% of ventrals.

Hemipenis (based on ZSI-CZRC-V-6416, Pune, left organ, Fig. 11 ).

The hemipenis reaches up to 13th subcaudals and is 36.9 mm in length and 8.2 mm maximum width at midbody and apex. It is divided into three distinct areas; the proximal area (nearly 30% of the total hemipenial length) is smooth without any ornamentation, middle zone (about 25% of the total hemipenial length) is with evenly scattered spinules and the apical calyculated portion (45% of the total hemipenial length). No enlarged spines are present, but the size of spinules at the proximal end are slightly larger than the ones at the distal end. The calyculated area can be further divided almost equally into proximal half with large calyces and distal half with smaller and denser calyces. The sulcus spermaticus is single, bounded with thick walls, runs straight across its length and opens into a delta (3 mm wide at apex, 1.9 mm in length), which is nude. The delta is subapical in position and the apex is calyculated, edged with fewer papillae. At the sulcate side, the calyces are larger proximally and gradually smaller and denser towards the distal end. The distal calyces are scalloped and edged with papillae. Along the asulcate side the spinous area starts more distally than the sulcate and lateral side. After the spinous zone, there are 2-3 rows of large calyces followed by densely packed calyces towards the apex. The proximal large calyces (9-10 in number) are of uneven size (mostly rounded or pentagonal) and among them the distal ones are scalloped. The large scalloped calyces are of 2.2-2.7 mm in length and 1.9-2.5 mm in width.

Wilson (1967) mentioned that the hemipenis is clavate, but in a fully inflated organ it starts widening from the middle and from the asulcate side there is a slight constriction between the midbody and apex. The ornamentation in ZSI-CZRC-6416 from Pune and ZSI-CZRC-6284 matches with that of LSUMZ 9425 as provided by Wilson (1967). However, the only available drawing of the organ as provided by the author is of the sulcate side, but apparently, differentiations in calyces are seen at the asulcate side.

Dentition (based on ZMB 8053, male, Bengalen).

15/15 maxillary teeth, the anterior 13/13 precranterian teeth are increasing in size posteriorly and are followed by a small diastema and 2/2 enlarged roundish cranterian teeth without groove. All maxillary teeth are slightly curved posteriorly, without significant interspaces. Medial to the precranterian teeth are single replacement tooth at different growth stages. One to two replacement teeth per cranterian tooth are found posteromedially to each tooth, showing different growth stages. 11/11 palatine teeth, decreasing in size posteriorly. All are curved posteriorly shortly above the base. Lateral to the palatine teeth there is a single replacement tooth at different growth stages. No significant interspace exists between the different palatine teeth. Posteromedial process of palatine large, expanding two pterygoid teeth. 13/12 pterygoid teeth, decreasing in size posteriorly. All are curved posteriorly shortly above the base. Lateral to the pterygoid teeth there is a single replacement tooth at different growth stages. No significant interspace exists between the different palatine teeth. The posterior 57% of the pterygoid are without teeth. 17/17 mandibular teeth, increasing in size up to tooth 5/5 and from there decreasing in size posteriorly. All are slightly curved posteriorly. Medial to the mandibular teeth there are 1-2 replacement teeth at different growth stages.

Variation in dentition.

Wall (1914) based on one skull from his collection described 13 subequal maxillary teeth which are followed by a diastema which is as large as one tooth and followed by two little larger posterior teeth. He described 11 palatine, 15 to 16 pterygoid teeth all decreasing in length posteriorly and 18 mandibular teeth which increasing in length up to the fifth tooth and from there decrease in length posteriorly. Smith (1943, p. 159, fig. 48 D) depicted an upper jaw with 13 precranterian teeth which are increasing very slightly in size posteriorly and are followed by a distinct diastema and a pair of enlarged cranterian teeth. Wilson (1967, p. 271) mentioned like Smith (1943, p. 170), 12 -14 maxillary teeth followed by a distinct diastema but in contrast to the latter, Wilson (l. c.) identified the following two cranterian teeth as unenlarged and counted 9-11 palatine teeth, 14-17 pterygoid teeth, and 14-18 mandibular teeth. The dentition of four Indian specimens ( BMNH 37a, see Fig. 12 ; MCZ R-28645; NMW 25465:3 and ZMB 4786a) is characterized by 12-14 maxillary teeth followed by a diastema (except MCZ R-28645, without diastema) and two enlarged cranterian teeth, 9-11 palatine teeth with a posteriomedial process of palatine expanding two or three pterygoid teeth, 13-17 pterygoid teeth with 48-52% of the posterior pterygoid without teeth, and 16-19 mandibular teeth, increasing in size up to tooth 4-6 and from there decreasing in size posteriorly.

Four specimens from Pakistan ( SMF 50410, SMF 57312 and SMF 62921-22) show 12-14 maxillary teeth followed by a diastema (except SMF 50410, without diastema) and two enlarged cranterian teeth, 9-11 palatine teeth with a posteriomedial process of palatine expanding two pterygoid teeth, 14-17 pterygoid teeth with 48-58% of the posterior pterygoid without teeth, and 17-20 mandibular teeth, increasing in size up to tooth 4-7 and from there decreasing in size posteriorly.

Dimensions and proportions.

The body is robust, moderately stout, roundish in cross-section at midbody. Head moderately pointed, barely distinct from neck. Eye large, 16-24% of head length. Male specimens grow slightly larger than females; the longest examined specimens are from Sind, Pakistan including a male ( SMF 50446) with 1482 mm and a female ( SMF 50409) with 1361 mm total length. The longest Indian specimen was reported by Gharpurey (1931) from Ahmednagar, Maharashtra with 1422 mm total length. The smallest male (NMW 25465: 3) with 336 mm and the smallest female ( SMF 57311) with 553 mm total length came from “Calcutta” and Sind respectively. The tail versus body length ratio ranges from 0.21 to 0.31 (males 0.21-0.31, females 0.23-0.28). For ten specimens from Telangana state, India measured by Seetharamaraju (2014, p. 95 ff.) the tail versus body length ratio ranges from 0.19 to 0.23 (males 0.19, females 0.19-0.23).

Colour and pattern (Fig. 9 A-K and Fig. 10 A-J ).

Dorsal colour and pattern show a significant ontogenetic change. In juvenile and subadult specimens the dorsal ground colour is olive-brown to light brown, the head dorsally slightly darker with narrow elongated irregular shaped whitish markings. Along the body up to 70 narrow whitish/black or dark brown stippled crossbars, usually two scales wide and in contact with lateral edges of ventrals. The first band is separated from the posterior border of parietals, the interspace between bands is three to four dorsal scales wide. The intensity of bands decreases posteriorly and the contours become blurred near to the vent. Dorsocaudally the bands are dissolved in an irregular fine pattern of light and dark points. Temporals, preoculars and supralabials can show small whitish spots. From about 600 mm total length the colour and pattern of young specimens begin to fade and disappear gradually. Adult specimens are dorsally dark brown to dark reddish-brown without any markings on head, body or tail. The ventral side of head, body and tail in all age classes is patternless yellowish cream.

Distribution.

Platyceps plinii is commonly believed to occur from Pakistan in the West to Bangladesh in the East with a North to South distribution from Nepal to Sri Lanka (see e.g. Smith 1943; Mahendra 1984; Welch 1988; Daniel 2002; Gruber 2002; Whitaker and Captain 2004; Sharma 2007; Wallach et al. 2014; Das et al. 2019; Uetz et al. 2019). We analyzed pertinent literature and locality information from museum specimens to present a list of localities corrected and updated for each country (see below and Appendix 9 Gazetteer).

According to our current knowledge, Platyceps plinii is only known with certainty from India and Pakistan. In India it is recorded from the states of Andhra Pradesh (Kasimkota [Casemcottah, type locality], Eluru, Nallamala Hills, Pulicat Lake, Rajahmundry, Rollapenta, Sundipentha/Sikharam, Thummalapalle mine and Visakhapatnam), Bihar (Munger, Patna, Tinpahar and Rajmahal), Chhattisgarh (Pharsabhar), Goa (Bondla Wildlife Sanctuary and Panaji), Gujarat (Baroda, Bharuch, Bhavnagar, Dahod, Dangs, Gandhinagar, Girnar Hill, Gir Forest, Kala Gadba, nr. Mahal, Mehsana, Panchmahal, Sabarkantha, Samot, Surat, Valsad, Vansda National Park and Vijapur), Haryana (Ambala), Jharkhand (Hazaribag District), Karnataka (Balekola, Chincholi Forest, Belgaum, Coimbatore, Coorg, Hosar, Mysuru and Shivamogga), Kerala (Muthanga in Wayanad Wildlife Sanctuary), Madhya Pradesh (Bhopal district, Barkatullah University Campus, Dewas, Dumna Nature Park, Gwalior, Indore, Jabalpur, Kanha National Park, Katra Hills, Mandla, Pachmarhi Biosphere Reserve, Sagar, Satpura Tiger Reserve, Shahdol, Sita Hill and Ujjain), Maharashtra (Akola district, Ahmednagar, Ale nr. Narayangon, Atpadi, Aurangabad, Bassein Fort, Bohali, Borivali (Gorai) mangroves, Buldhana district, Chink Hill, Dahanu Forest Division, Deolali, Devagad, Ghorawadi nr. Talegaon, Jalgaon, Jawhar, Juvem, Kaas Plateau, Khandala, Kolhapur district, Konkan region, Kurduvadi, Malegaon, Marol, Mokhada, Mumbai, Nagar, Nagpur, Nanded, Navegaon National Park, Navi, Panchgani, Parel, Parvati-Pachgaon Hills, Powai, Pune, Ranidoh region, Thana, Vidarbha region, Vidyanagari, Visapur and Yavatmal), Odisha (Gandhamardan Hills, Nandankanan, Choudwar, Similipal Biosphere Reserve, Baripada and Bhadrak), Puducherry, Rajasthan (near Hemavas Dam, Pratapgarh district and Ramri), Tamil Nadu (Chennai and Hosur Hills), Telangana (Farahabad, Hyderabad region, Ippalapally, Jannaram, Kerimeri, Khanapur, Kinnerasani, Old Bowenpally, Rushulacheruvu, Tarnaka and Vijayapuri), Uttar Pradesh (Mathura region, Faizabad, Prayagraj and Varanasi), West Bengal (Dum Dum, Durgapur, Kolkata and Panchet Hill). The westernmost published record of the Banded racer comes from the Girnat Hill or Girnagar in the Junagadh district of Gujarat, approximately 70°31' East ( Patel et al. 2019a). According to Anonymous (1870), Anderson (1871), Stoliczka (1871b), Sclater (1891), Smith (1943) and Ahmed and Dasgupta (1992) the actual westernmost distributional border for P. plinii lies in West Bengal state of India, near Kolkata approximately 88°25' East (incl. ZSI-K 7332, don. Joseph Fayrer 1870, ZSI-K 7334 don. John F. Galiffe, and the specimen from Dum Dum illustrated on sketch 63 of the Hardwicke collection in the Natural History Museum, London; see Fig. 8 ).

In Pakistan confirmed records are known from the Indus plain in Sindh province (nr. Badin, Hala, Jati, Lakarna, Makli Hills, Mohenjo-daro, Pir Patho, Raj Malk, Sonda, and Tatta) and some authors mentioned southern and central Punjab without specific locality (e.g. Khan 1980, 1982, 2002, 2006). We found only one record with more precise information for Punjab from the Lal Suhanra National Park in Bahawalpur published by Khan et al. (2018, p. 1731).

Unconfirmed or erroneous distribution records.

Sporadically the Banded racer is mentioned for the Himalayan region in northwestern and northeastern India. Mahajan and Agrawal (1976) and Agrawal (1979) published records (as Coluber fasciolatus ) from Saproon village and the Gambhar bridge from an elevation of 1513 m and 1485 m a.s.l. respectively. Both localities are situated in the Solan district of Himachal Pradesh state of India. From Uttarakhand state the Banded racer was listed by Sharma (2004) from Sokharak, Mandal Rest House and Mandal-Chopta road in Kedarnath Wildlife Sanctuary located between 1500 and 3300 m a.s.l. in the Chamoli and Rudraprayag districts, and by Bahuguna (2010) for the Corbett Tiger Reserve (now Jim Corbett National Park). Since these reports, no further specimens from the Himalayan range of Himachal Pradesh and Uttarakhand have become known and because of the fact that these localities are considerably above the average altitudinal distribution of this species (see below), we suspect that these records are based on incorrect determinations (Abhijit Das, pers. comm. 2020) and consider the records to be very doubtful. The Banded racer is also listed from the Eastern Himalayas by e.g. Jha and Thapa (2002), Chettri and Bhupathy (2007) and Chettri et al. (2011). Whereas the first authors consider it “rare”, Chettri and Bhupathy (l.c.) regarded it surprisingly as “common” in Sikkim. Since none of the above-mentioned authors provided a single voucher specimen or a specified locality in that East Himalayan state, and no preserved material is known from any collection, so we consider its distribution in Sikkim to be unproven.

Based on water-colour sketch no. 61 from the Hardwicke collection, Smith (1943, p. 529) mentioned “Cawnpore” [Kanpur, Uttar Pradesh state of India] for Coluber fasciolatus . Although the Banded Racer is known to occur in this state (see above), the snake depicted on this drawing is in fact, a Common ratsnake, Ptyas mucosa .

Wallach et al. (2014) added the Pakistani province of Balochistan to the distribution range, but the literature cited provided no such indications and we could not identify a single credible record for this state.

When it comes to the distribution of the Banded racer in Sri Lanka, there is a consistent reference to Haly (1886, 1891), who allegedly provided the first evidence. But, as already noted by Somaweera (2006), Ferguson (1877) was the first who listed this species for “Ceylon”. Ferguson (l.c.) stated that the specimen “[…] was sent to the Museum [Colombo] lately by Mr. Deputy Queen’s Advocate Thwaites, and supposed to have been found by Mr. Twynam, the Government Agent of the Northern Province, about Aripo" (see Appendix Note 3). Since Haly (1886) the Banded racer is consistently listed as part of the Sri Lankan snake fauna (e.g. Wall 1921; Deranyiagala 1955; P. de Silva 1980; de Silva 1990, 2001; Das and de Silva 2005; Somaweera 2006; de Silva and Jinasena 2009; Wallach et al. 2014; de Silva and Ukuwela 2017) although no additional specimen has been collected, no preserved voucher specimens are known from any collection, and recent fieldwork on Jaffna Peninsula revealed no further material ( Abyerami and Sivashanthini 2008; Madawala et al. 2019). We follow Taylor (1950) and regard this species as doubtful for Sri Lanka.

In several publications, especially in overview works and faunal lists, the Banded racer is mentioned for Bangladesh. This is probably because in older publications (e.g. Cantor 1839; Günther 1864; Theobald 1876; Boulenger 1890) “Bengal” is listed as a known distribution, a region currently divided between Bangladesh and the West Bengal state of India. After the separation of the central and eastern regions of Bengal from India in 1947 and the foundation of the state of Bangladesh in 1971, the majority of (Indian) authors changed the original information “Bengal” to "West Bengal"(India), but others still maintained the distribution for Bangladesh (e.g. Welch 1988; Daniel 1989, 2002; Das 2002; Whitaker and Captain 2004; Wallach et al. 2014; Uetz et al. 2019), although Wall (1914) already clarified that the species is not found further east than Calcutta in West Bengal state of India. Authors who refer exclusively to the herpetofauna of Bangaladesh (e.g. Islam et al. 2000; Khan 1988, 2004; Kabir et al. 2009) do not present more detailed localities and while Hasan et al. (2014) listed the Banded racer as expected to occur in the Northeast and Southeast of the country, he did not provide any supporting evidence. We could not find any historical or current evidence that this snake species occurs in Bangladesh.

From Nepal, the Banded racer was first mentioned by Günther (1861) in a list of “[…] Cold-Blooded Vertebrata [ …]” collected in that country. Günther’s source was a water-colour painting of a specimen from the "Central hilly region" (reproduced here in Fig. 6 ) probably made by Raj Man Sinh Chitrakar, a Nepalese draftsman employed by Brian H. Hodgson ( McClelland and Griffith 1844; Cocker and Inskipp 1988). The locality most probably refers to the complex of hills and valleys surrounding the Kathmandu valley where the majority of specimens in the Hodgson Nepalese collection originated. Our examination of Hodgson’s original drawing, in the Zoological Library of Natural History Museum, London, revealed that the specimen depicted is not a Banded racer but a natricid snake, Rhabdophis himalayanus ( Günther, 1864), a species reported from that region by e.g. Gruber in Schleich et al. (2002). Shrestha (2001) stated that the Banded racer has been reported from the Royal Chitwan National Park, and “[…] lives in dense Sal forest as well as grassland of Chitwan valley in Nepal [ …]”. We regard this statement as based on a misidentification because no voucher is known. Similar to the aforementioned record for Sri Lanka, the occurrence of the Banded racer in Nepal relies solely on an old and unreviewed original source, so the presence of this species in Nepal could not be confirmed. Although Kramer (1977) already considered the distribution in Nepal as questionable, and no further material has been collected (Karan B. Shah in litt. 2019), most subsequent authors continue to list the Banded racer in the Nepalese herpetofauna (e.g. Swan and Leviton 1962; Welch 1988; Schleich 1993; Shrestha 1998, 2001, 2003; Gruber 2002; Shah and Tiwari 2004; Wallach et al. 2014; Uetz et al. 2019).

Based on records of the Banded racer in Bengal, Schleich and Kästle (2000) and Lenz (2012) suspected its possible occurrence in the Himalayan Kingdom of Bhutan which was adopted as questionable by Uetz et al. (2019). As explained above for the Himalayan regions of India and Nepal such a presumption lacks any basis and we refer to the list of verified records above (see also Appendix 9 Gazetteer).

Cantor (1847a, b) mentioned a specimen from the Province Wellesley (Seberang Perai, Penang state, West Malaysia) which corresponds with Russell’s "Nooni Paragoodoo", and determined it as Coluber fasciolatus . Based on dentitional characters, which he regarded as similar to Russell’s (1796) description, he concluded that C. fasciolatus “[…] cannot be placed in the genus Lycodon , to which it has been referred by M.M. H. Boie, -Wagler and Schlegel". Wall (1908, p. 21 footnote, 1913, p. 20 footnote, 1921, p. 195 footnote, 1924, p. 619) and Wilson (1967, p. 266) already doubted the authenticity of Cantor’s record. Our re-examination of this juvenile specimen ( BMNH 60.3.19.1121) confirmed its identity as P. plinii , thus the locality given by Cantor must be considered as an error.

Mason (1861) listed " Coluber fasciolatus " for “Burmah” and Theobald (1868b) and Nicholson (1874) mentioned, " Zamenis fasciolatus Shaw" from “Mergui” [Mergui or Myeik Archipelago, Myanmar]. The latter quoted “Blyth” but gives no specific reference, that makes a reassessment impossible. Theobald (1880) added "Tenasserim, &c." to the distribution of the Banded racer, without reference to specific specimens or exact localities. As indicated above, the easternmost records are from West Bengal and we, therefore, consider the records for Myanmar to be based on misidentification.

Habitat and natural history.

Platyceps plinii is widespread over the Indian Subcontinent (see above, Map 1 and Appendix 9 Gazetteer). It is reported as regionally common in e.g. Maharashtra and Karnataka ( Murthy and Acharjyo 1987; Daniel 1989, 2002; Deshmukh et al. 2015; Ingle et al. 2014) and abundant in parts of Goa ( Murthy and Acharjyo 1987). It inhabits plains, plateaus and mid hills covered with mixed and dry deciduous forest, scrub forest as well as semi-arid and rocky areas (for habitat examples see Fig. 13 A and B ).

The vertical distribution ranges from sea level to over 1000 m altitudes, with about 85% of all finds below 600 m a.s.l. A few unusually high sites are known from the states of Madhya Pradesh (Pachmarhi Biosphere Reserve, 1053 m a.s.l.) and Maharashtra (Kaas Plateau, 1191 m a.s.l., and Panchgani, 1258 m a.s.l.) (see Appendix 9 Gazetteer).

It is a mainly terrestrial species but also observed to be a good climber ( Ingle 2008) and found in open places, tall grass, dense bushes, under roots of old trees, in spiles of stones and bricks, and rodent holes. The Banded racer is also reported from urban and cultivated areas where it was found in houses, village environs, agricultural open fields, gardens, parks and farms ( Stoliczka 1871a; Wall 1914; Lindberg 1932; Khaire 1996, 2014; Ingle 2002; Whitaker and Captain 2004; Rao et al. 2005; Srinivasulu et al. 2006; Chandra et al. 2008; Chikane and Bhosale 2012; Seetharamaraju 2014; Charjan and Joshi 2015; Sayeswara et al. 2015; Dileepkumar et al. 2016; Ingle et al. 2019). In some places, e.g. Bhadrak, Odisha, 30-40 individuals were found hibernating together in the cob walls in human habitations (Mohalik pers. comm.).

It is a diurnal species that is only exceptionally observed active during the night and known for its defensive and pugnacious behaviour. If threatened or captured it erects itself, flattens the forebody and furiously attacks when further provoked which one can confuse with a cobra ( Wall 1907, 1914, 1921; Whitaker and Captain 2004). Its antipredator mechanism was summarized by Green (1988) as follows: (1) dorsoventral body compression, (2) dorsoventral neck compression, (3) lifting of head and neck, (4) S-coil striking posture, (5) lunging and striking, and (6) bite. One female caught while crossing a road immediately defecated and started attempting to bite. In other instances, individuals were observed climbing electric cables and visiting switch boxes, possibly looking for hiding geckos ( Hemidactylus spp.).

The Banded racer is an active hunter that usually kills its prey by constriction or body pressure. Its prey consists of small mammals like rats, mice, shrews and bats, but also frogs, lizards, birds, worms, and for juveniles insects are part of the food spectrum ( Wall 1914; Daniel 1989, 2002; Sharma 1999a; Das and de Silva 2005; Satish 2008; Whitaker and Captain 2004).

Mating of the Banded racer was observed between late winter and mid-monsoon ( Dileepkumar et al. 2016). The reproduction is oviparous with clutch sizes between 2 and 23 eggs and egg sizes between 26.8-34.7 mm length and 16.5-22.4 mm width, laid between January and September ( Vyas 1987; Khaire 1996, 2008, 2010, 2014; Das 2002; Das and de Silva 2005; Satish 2008; Ingle et al. 2014; Tambre and Chavan 2016; Parsar 2018; Patel et al. 2019b). Hatchlings have been observed in May ( D’Abreu in Wall 1914) and July ( Wall 1914; Daniel 1989, 2002).

Manhas (2015a) observed the Banded racer as roadkill during monsoon in Bhopal district of Madhya Pradesh state and Mohalik et al. (2019) reported the predation of this species by a leucistic Krait, Bungarus caeruleus , from the Patharakali area of the Bhadrak district in Odisha state of India.

For Pakistani specimens, Minton (1966) and Wilson (1967) observed semifossorial behaviour and Khan (2002, 2006) reported that reproduction in Banded racer occurs twice a year, during the months April to May and August to September, with clutch sizes from 23 to 30 eggs.

Figures

Figure 5. Platyceps plinii comb. nov. A. Lectotype of Natrix plinii Merrem, 1820, " Patza Tutta " from " Casemcottah " reproduced from Russell (1796, pl. 29), and B. " Die halbbandierte Natte Bechstein (1802, pl. 44, fig. 2).

Figure 6. Extract from a previously unpublished original water-colour painting from Brian H. Hodgson's collection of drawings kept in the Zoological Library of the Natural History Museum, London, U. K. The depicted specimen from the " Central hilly region " of Nepal, determined by Guenther (1861) as "? Coryphodon fasciolatus " (= Platyceps plinii comb. nov.), is the original and sole source for the distribution of the Banded Racer in Nepal. In fact, the specimen shown is an adult Rhabdophis himalayanus (Guenther, 1864) (see chapter " Unconfirmed or erroneous distribution "). The inscription in Devanagari script on the lower right side of the sheet states that this specimen was preserved in alcohol on 27 September 1857. Reproduced with permission from the Natural History Museum, London, U. K.

Figure 7. Platyceps plinii comb. nov. Previously unpublished original painting (no. 7) of the holotype of Coluber curvirostris Cantor, 1839 from Cantor's original manuscript entitled " Indian Serpents-Innocuous-Collected, figured & described (1831 - 1837) ", kept in the Bodleian Library, Oxford, U. K. (see Appendix, Note 2). Reproduced with permission from the Bodleian Library, Oxford, U. K.

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.

Figure 9. Platyceps plinii comb. nov. in life from various parts of India. Adults: A. ZSI-CZRC- 6384 (male, SVL: 491) from Baripada, Odisha state, B. Uncollected specimen from Pune, Maharashtra state, C. ZSI-CZRC-V- 6416 (male, SVL: 815) from Pune, Maharashtra state, and D. Uncollected female from Amaravati, Maharashtra state. Hatchlings / juveniles: E. ZSI-CZRC-V- 6329 (subadult male, SVL: 355) from Choudwar, Odisha state, F. Uncollected specimen from Pune, Maharashtra state, G. Uncollected specimen from Amboli, Maharashtra state, and H. Uncollected specimen from Yavatmal, Maharashtra state. SVL in mm.

Figure 10. Platyceps plinii comb nov. Variation of neck pattern in specimens from various parts of India. Adults: A. NCBS-AQ 492 (female, SVL: 820) from Bangalore, Karnataka state B. BMNH 62.8.14.29 (female, SVL: 854 mm) from Bengal, C. ZSI-CZRC- 7111 (female, SVL: 965) Kabeerdham, Chhattisgarh state and D. BNHS 2857 (female, SVL: 746) Pune, Maharashtra state, E. ZSI-CZRC 6331 (male, SVL: 713) from Bhandara, Maharashtra state. Hatchlings / juveniles: F. BNHS 657 (male, SVL: 331) from Satna, Madhya Pradesh state, G. BNHS 639 (sex unknown, SVL: 212) from Saugor, Madhya Pradesh state, H. BNHS 637 (sex unknown, SVL: 222) from Ambala, Punjab state, I. BMNH 37 A (male, SVL: 432) from Visakhapatnam, Andhra Pradesh state, and J. ZSI-CZRC 6330 (male, SVL: 350) from Cuttack, Odisha state. Scale bar 10 mm; SVL in mm.

Figure 11. Hemipenis of Platyceps plinii comb. nov. (ZSI-CZRC-V- 6416, left organ) from Pune, Maharashtra state, India: A. lateral, B. asulcal, and C. sulcal view. Scale bar 10 mm.

Figure 12. Skull of Platyceps plinii comb. nov. (BMNH 37 a) from Visakhapatnam, Andhra Pradesh state, India: A. lateral, B. ventral (tooth bearing and connecting bones virtually extracted), C. dorsal and D. ventral view (lower jaw virtually extracted). Scale bar 1.5 mm.

Figure 13. Habitats of Platyceps plinii comb. nov. in India: A. near type locality Kasimkota village, Andhra Pradesh state, and B. from Pune, Maharashtra state.