Oligodon cinereus ( Günther, 1864 ), Gunther, 1864

David, Patrick, Das, Indraneil & Vogel, Gernot, 2011, On some taxonomic and nomenclatural problems in Indian species of the genus Oligodon Fitzinger, 1826 (Squamata: Colubridae), Zootaxa 2799, pp. 1-14 : 7-10

Note on Oligodon cinereus ( Günther, 1864) in India

Simotes cinereus Günther, 1864: 215 . Type locality. “Gamboja”, now Cambodia. Holotype. BMNH RR 1946.1.1.25 (adult male).

This species is widespread from extreme north-eastern India and eastern Bangladesh to south-eastern China and Taiwan, and southwards up to Peninsular Thailand across the whole of Myanmar, Laos, Cambodia and Vietnam ( Wagner 1975; our material). Oligodon cinereus is mostly characterized by (1) 17–17–15 DSR, (2) anal entire, (3) no more than 182 VEN (no more than 176 in the nominative subspecies), (4) a full complement of head scales including a loreal, (5) 8 (rarely 7) SL, SL 4–5 (rarely 3–4) entering orbit, (5) and 1 anterior temporal. Specimens with 15 DSR are referable to Oligodon inornatus ( Boulenger, 1914) ; specimens with 17–17–15 DSR and more than 185 VEN belong to Oligodon joynsoni ( Smith, 1917) . This latter species is quite similar to reticulate patterned specimens of O. cinereus cinereus but the number of ventrals easily separate these two distinct species, with 156– 178 in O. c. cinereus vs. 186–197 in O. joynsoni .

A morphologically variable species, Oligodon cinereus shows a variety of dorsal patterns (see Smith 1943), which have been used as a basis for describing several subspecies. However, the current intraspecific taxonomy of this species remains unresolved. Although Smith (1943) did not recognize any subspecies, other authors such as Taylor (1965) accepted the validity of several subspecies, including O. cinereus tamdaoensis ( Bourret, 1935) , O. cinereus pallidocinctus Bourret, 1934 , O. cinereus swinhonis ( Günther, 1864) and O. cinereus multifasciatus (Jan & Sordelli, 1865) . More recently, Wagner (1975) showed that the differences between the various subspecies were unclear. Wagner refrained from accepting subspecies, a position that we follow here.

Among these taxa, we here address the status of Simotes multifasciatus Jan & Sordelli, 1865 (in 1860–1881: Livraison 12, pl. IV: fig. 2. Type locality: “Sultanepore”, now Sultanpur, State of Uttar Pradesh, northern India. Holotype: not given, stated as deposited in the “ Musée de Bonn ”, presumably lost). The origin of the specimen is unclear, and would appear to have been collected in the first half of the 1800s, a period when only two notable collections were made by European naturalists outside England. Perhaps the most well-known is that of the Austro- Hungarian Novara Expedition, whose collections are in Vienna ( Gans 1955). The second was made by the three Schlagintweit brothers, Hermann (1826–1882), Adolphe (1829–1857) and Robert (1833–1885), who originate from Germany (see Armitage 1989). Made incidental to geomagnetic studies, on behalf of the East India Company ( Sthanapati 1996), this large collection of botanical, zoological as well as geological material was deposited in a number of museums in Europe, including Bonn ( Armitage 1989). Significantly, the Schlagintweits followed “the course of the Ganges River upstream over Varanasi, Allahabad and Fatehgarh to the Himalaya” ( Praschag et al. 2008), clearly approaching Sultanpur in modern day Uttar Pradesh.

This taxon was depicted on a plate of Jan & Sordelli (1865 in 1860–1881). On page 8 of the Index to the plates, the holotype was said to have been deposited in the Museum of Bonn. According to Dr. W. Böhme (pers. comm. to P. David, January 2009), the zoological museum in Bonn at that time was the “Naturhistorisches Museum Poppelsdorf” (situated in what is now a suburb of the city of Bonn), but was destroyed during World War II (see Friedrich 2006), together with nearly all its holdings. The type of Simotes multifasciatus is therefore considered lost.

Jan & Sordelli (1865 in 1860–1881) merely stated that the specimen had 17 dorsal scale rows. Nevertheless, although incomplete, the drawing is detailed enough to derive some data of the head scalation (see Table 2). The depicted specimen shows a dorsal pattern made of irregular, dark crossbars, a dark nuchal marking pointing forwards, some blotches on the head, and a venter and under tail surface strongly marked with dark, square blotches.

Number Sex SVL TaL TaL/TL VEN SC DSR SL PreOc PosOc SubOc Temp Type J&S ɗ ---- ---- ---- ---- 32?–17–? 8/8?/1?/2?/1? / 1+2 ZSI 12356 Ψ 150 17 0.102 169 30 17–17–15 8 /7 1/1 2/2 1/1 1+2 / 1+2 Taylor (1965) recognized O. cinereus swinhonis ( Günther, 1864) and O. cinereus multifasciatus (Jan & Sordelli, 1865) for populations from Thailand. However, Taylor (1965) pointed out that the type locality of Simotes multifasciatus Jan & Sordelli, 1865 was “Sultanepore”, “a locality unknown to me”. Although there is no such locality in Thailand, there are indeed several cities known as Sultanpoor, now Sultanpur in India. All of them are located in the northern part of India, in the states of Gujarat, Jammu and Kashmir, Punjab, Haryana, Rajasthan, Maharashtra, Madhya Pradesh, Andhra Pradesh, Himachal Pradesh, Uttaranchal, and Uttar Pradesh. However, the most famous of them is the legendary capital of Ram, a locality in the ancient Kingdom of Oudh in northern India. It is now the Indian city of Sultanpur, south-east of Lucknow, State of Uttar Pradesh. We suggest that the specimen described by Jan & Sordelli (1865 in 1860–1881) originated from this once famous city.

The general habitus of the type of Simotes multifasciatus Jan & Sordelli, 1865 and the number of dorsal scale rows allow us to agree with previous authors, such as Smith (1943) and Wagner (1975), in referring this taxon to the Oligodon cinereus complex, namely O. cinereus and its current numerous synonyms. However, this locality is out of the known range of O. cinereus in India, from where it was previously known only from the state of Assam (Tinsukia, Tinsukia District); see Wagner (1975). Sultanpur in Uttar Pradesh is ca. 1,200 km from the previously westernmost known locality of O. cinereus .

Dasgupta & Raha (2004) mentioned, as Oligodon cyclurus , a specimen (ZSI 12356), from “Cargola, Purnea”, the latter, an old spelling for (the District of) Purnia, in the north-eastern Indian State of Bihar. We examined this specimen ( Figs. 3 A–C), which shares with the holotype of Simotes multifasciatus the number of 17 DSR, the dorsal pattern with distinct black crossbands and the blotched venter. We here refer it to Oligodon cinereus , as currently defined. Its main morphological characters are given in Table 2.

Distribution of Oligodon cinereus in India and Bangladesh ( Fig. 4 ). On this basis, we extend the range of Oligodon cinereus in northern India (see Fig. 4 ). In the north-western part of its range, this species is now known from the following localities: India (based on Agarwal et al. 2010; Dasgupta & Raha 2004, 2006; Mathew 1995; Wagner 1975; examined material). State of Arunachal Pradesh: Eaglenest Wildlife Sanctuary. State of Assam: Jaipur, Tinsukia District; Tezpur, Sonitpur District; Tinsukia, Tinsukia District. State of Bihar: Cargola, Purnia District. State of Meghalaya: West Garo Hills; East Khasi Hills. State of Nagaland: Naga Hills. State of Uttar Pradesh: Sultanpur, Sultanpur District. Bangladesh (based on Smith 1943; Wagner 1975). “ Chittagong Hills”. According to its scalation data, especially the number of dorsal scale rows, the specimen recorded from Changlang District, State of Arunachal Pradesh, and cited by Sanyal & Gayen (2006), is not conspecific with O. cinereus .

Comparison with other populations of Oligodon cinereus . Smith (1943) and Wagner (1975) defined several morphs within this species, based on the dorsal, head and ventral patterns. Smith’s “Var. I” is nearly unpatterned above and with a powdered venter. This morph, to which the holotype of Simotes cinereus belongs, is widespread in Myanmar, Thailand and the Indochinese Peninsula. Smith (1943) also recognized a “Var. II” with strong, dark reticulations or irregular crossbars, and a uniform head; this author referred Simotes multifasciatus to this morph, which is considered to be widespread from southern Myanmar to southern China. Smith’s “Var. III” includes specimens with distinct dorsal black crossbars and strong reticulations between the crossbars, head with at least a nuchal chevron, and a venter strongly patterned with rectangular blotches. According to Smith (1943), this morph, described as Holarchus violaceus tamdaoensis Bourret, 1935 , occurs in India, Bangladesh, Myanmar, northern Thailand, and Vietnam. Smith also recognized a “Var. IV”, which is not relevant to the present discussion because it occurs in Vietnam and Laos and presents a dorsal pattern made of white, dark-edged crossbars.

Wagner (1975) recognized the same patterns as Smith, and he (p. 80) published an interesting map showing the range of species in correlation with the dorsal pattern of specimens examined by him. The range of the “blackbarred” morph is especially noteworthy and confirms Smith’s (1943) suggestions. It occurs in Assam (Tinsukia), Myanmar, northern and central Thailand and northern Vietnam. We examined four specimens from Myanmar and confirm that they are identical in pattern and scalation with those from northern Vietnam.

According to our material, we do not separate Smith’s forms I and II. Some “immaculate” specimens (Form I) show some reticulations and, furthermore, there is no difference in morphometry and scalation between these two morphs. In contrast, specimens with dark, solid dorsal crossbars (Form III of Smith; black-barred morph of Wagner) are distinct not only in both their pattern on dorsum and belly, but also in morphometry (relative tail of length, and body scalation, see Table 3). There is no difference in various characters of head scalation between these two morphs but the length:width ratio of the frontal is greater in Morphs I+II.

The drawing of the holotype of Simotes multifasciatus Jan & Sordelli, 1865 shows irregular crossbars, a dark nuchal chevron and a heavily blotched venter. These characters are similar to those of Smith’s Form III. We here refer the holotype of Simotes multifasciatus to this morph. A discussion on the intraspecific taxonomy of Oligodon cinereus will be addressed elsewhere (Vogel et al., unpublished data) but black barred specimens of India, Myanmar and Thailand and those of northern Vietnam and Laos might prove to be distinct at the species level from other morphs of Oligodon cinereus .

TABLE 3. Comparison between morphs of Oligodon cinereus . See Material and Methods for explanation of abbreviations. Morph Sex TaL/TL VEN Sc L-Fr / W-Fr Smith’s I + II ɗ 0.129–0.149 (x = 0.140) 156–168 (x = 162.9) 35–42 (x = 37.2) 1.20–1.37(x = 1.28) “ cinereus ” Ψ 0.123–0.131 (x = 0.127) 167–178 (x = 173.5) 33–37 (x = 35.3) (n = 18)

Smith’s III ɗ 0.122–0.140 (x = 0.128) 169–180 (x = 173.3) 30–42 (x = 36.6) 1.11–1.30(x = 1.20) “ tamdaoensis ” Ψ 0.101–0.120 (x = 0.112) 173–184 (x = 176.4) 30–36 (x = 32.9) (n = 19)

The precision of the type locality of Simotes multifasciatus Jan & Sordelli, 1865 has implications for the taxonomy of the genus Oligodon in Thailand. Taylor (1965) referred a total of four specimens of Oligodon from northern Thailand to the subspecies O. cinereus swinhonis ( Günther, 1864) (one specimen; from “ 12 km north of Chiang Dao, Chiang Mai Province) and O. cinereus multifasciatus (Jan & Sordelli, 1865) (three; from “Chang Dow [12 km, north], Chiang Mai Province; Loei Province), respectively. According to the descriptions given by Taylor, these specimens have 17–17–15 DSR, 190–197 VEN and 32–42 SC (see Table 4). Their dorsal pattern is made of irregular dark crossbars with either faint (“ swinhonis ”) or strong (“ multifasciatus ”) reticulations.

Specimen Sex SVL TaL TaL/TL VEN SC DSR SL ATem O. joynsoni

BMNH 1946.1.4.23 ɗ 655 105 0.138 190 47 17–17–15 8 / 8 1 / 1 BMNH 1969.1809 ɗ 570 100 0.149 186 50 17–17–15 7 / 7 2 / 2 MNHN 1896.0633 Ψ 572 80 0.123 194 39 17–17–15 8 / 8 2 / 2 Taylor’s specimens

We have discussed above the status of Simotes multifasciatus Jan & Sordelli, 1865 , showing that its type locality is indeed ‘India’. The type locality of Simotes swinhonis Günther, 1864 is “Amoy”, now Xiamen, Fujian Province, People’s Republic of China. The number of ventrals in specimens examined by Taylor is much higher than those of the syntypes of Simotes swinhonis (156–168) and of the Indian specimen identified by us above as similar to the type of Simotes multifasciatus . We will not discuss here the status of Simotes swinhonis Günther, 1864 , regarded by Smith (1943) and Wagner (1975) as a synonym of Oligodon cinereus (reticulated morph).

The four Thai specimens of the Oligodon cinereus group examined by Taylor (1965) agree in scalation and pattern with the description of Simotes longicauda joynsoni Smith, 1917 (p. 276. Type locality: “Maa Yome, Muang Ngow, N. Thailand ”, now Muang Ngao or Muang Ngau, 18° 45' 27” N, 99° 58' 42” E, Lampang Province). Oligodon joynsoni ( Smith, 1917) is now considered a valid species of the O. cinereus group based on a high number of ventrals (> 185) and subcaudals (> 36). Furthermore, Taylor’s specimens were obtained from the provinces of Chiang Mai and Loei, in localities quite close to the type locality of O. joynsoni .

We identify as Oligodon joynsoni the four specimens described by Taylor and referred to O. cinereus swinhonis and O. cinereus multifasciatus . The current range of O. joynsoni , a relatively rarely encountered species, known from northern Thailand and Laos, is given in Fig. 5 (after Smith 1943; Taylor 1965; Nabhitabhata et al. 2004; our material). This species is now known from the following localities: Thailand. Chiang Mai Province: North of Chiang Dao. Lampang Province: Muang Ngao; Mae Wang. Loei Province: No locality. Laos. No precise locality. Nabhitabhata et al. (2004:120) also recorded this species from the Thai provinces of Kamphaeng Phet (Khlong Lan) and Nakhon Ratchasima (Khao Yai). However, no information on their scalation was provided and given that confusion with Oligodon cinereus is easy, we regard these localities as in need of confirmation. Taylor (1965:781) mentioned O. joynsoni from Thailand but stated not to have seen specimens. This confusion probably arose due to the fact that the holotype of S. multifasciatus agrees in pattern with O. joynsoni ( Smith, 1917) .

Figures

FIGURE 3. Oligodon cinereus, ZSI 12356. A: General view. B: Head in dorsal view. C: Body venter.

FIGURE 4. Range of Oligodon cinereus in India and Bangladesh. Solid circles = examined specimens; open circles = literature records; star = type locality of Simotes multifasciatus Jan & Sordelli, 1865. Specimens from Myanmar are not included.

FIGURE 5. Range of Oligodon joynsoni. Solid circles = examined specimens; star = type locality of Simotes longicauda joynsoni Smith, 1917.