david et al. (indian oligodon)

Accepted by D. Gower: 14 Feb. 2011; published: 24 Mar. 2011

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Zootaxa 2799: 1–14 (2011) www.mapress.com/zootaxa/ Article

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On some taxonomic and nomenclatural problems in Indian species of the genus Oligodon Fitzinger, 1826 (Squamata: Colubridae)

PATRICK DAVID1, INDRANEIL DAS2 & GERNOT VOGEL3

1Muséum national d’Histoire naturelle, Département Systématique et Évolution, Reptiles & Amphibiens, UMR 7205 OSEB, Case post-ale 30, 57 rue Cuvier, F-75231 Paris Cedex 05, France. E-mail: [email protected] of Biodiversity and Environmental Conservation, Universiti Malaysia Sarawak,94300 Kota Samarahan, Sarawak, Malaysia. E-mail: [email protected] for Southeast Asian Herpetology, Im Sand 3, D-69115 Heidelberg, Germany. E-mail: [email protected]

Abstract

This paper deals with three nomenclatural and taxonomic problems affecting two species groups of the colubrid snakegenus Oligodon Fitzinger, 1826: (i) A neotype is formally designated for Coronella cyclura Cantor, 1839, associating thisspecific nomen with populations from India, Bangladesh and Myanmar with 19 scale rows at midbody; (ii) Oligodon khe-riensis Acharji & Ray, 1936 is shown to be a valid species of the Oligodon cyclurus group occurring in northern India andNepal; (iii) The type-locality of Simotes multifasciatus Jan & Sordelli, 1865 is shown to be Sultanpur, India. This taxonis considered a synonym of Oligodon cinereus (Günther, 1864). The range of this species in India is extended. The statusof specimens of Oligodon cinereus from India and Myanmar is briefly discussed. Specimens from Thailand identified asOligodon cinereus multifasciatus and Oligodon cinereus swinhonis (Günther, 1864) are referred to Oligodon joynsoni(Smith, 1917). India is home to at least 21 species of the genus Oligodon, an updated list of which is provided.

Key words: India, neotype, Oligodon cinereus, Oligodon cyclurus, Oligodon kheriensis, Oligodon joynsoni, Simotes mul-tifasciatus, taxonomy, Thailand

Introduction

The genus Oligodon Fitzinger, 1826, widespread throughout central and tropical Asia, contains about 70 valid spe-cies (Green et al. 2010). The taxonomy of this genus, especially its contents, remains unclear, as a result of highlineage diversity, small samples for the majority of species, and continued poor sampling within the range knownto be occupied by the snakes of this genus. Several species have been described in recent years (e.g., David et al.2008a, b), whereas others have been placed in synonymy (Tillack & Günther 2010). In the present paper, weaddress one nomenclatural and two taxonomic problems affecting species of this genus present in India. These taxabelong to the Oligodon cyclurus group and Oligodon cinereus group.

Smith (1943), Wagner (1975) and Green et al. (2010) recognized several informal species groups, mostly onthe basis of hemipenial morphology. The Oligodon cyclurus group currently includes O. cyclurus (Cantor, 1839),O. fasciolatus (Günther, 1864), O. juglandifer (Wall, 1909), O. chinensis (Günther, 1888), O. formosanus (Günther,1872), O. ocellatus (Morice, 1875), O. saintgironsi David, Vogel & Pauwels, 2008, and O. macrurus (Angel,1927). This group, which contains large-growing species, is characterized by (1) long and deeply forkedhemipenes, neither spinose nor papillate (with the exception of O. formosanus, in which papillae are present), (2)17–23 dorsal scale rows, (3) 9–13 maxillary teeth, (4) complete complement of head scales, including a loreal(sometimes absent in O. macrurus) and often a presubocular, (5) anal plate entire; and (6) blotched and/or reticu-lated colour pattern, although a striped pattern may be found in some specimens of Oligodon cyclurus. These spe-cies are widespread from north-eastern India and Myanmar, to southern China and to southern Thailand.

In the present paper, we discuss the problem of the name bearing type of Coronella cyclura Cantor, 1839,which has been controversial (see Das 2004). The lack of a name-bearing type for this species has been one of thecauses for its uncertain taxonomy. A formal designation of a neotype is performed. We also address the status of

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DAVID ET AL.2 · Zootaxa 2799 © 2011 Magnolia Press

Oligodon kheriensis Acharji & Ray, 1936. This taxon from northern India and Nepal had been regarded as a syn-onym of Oligodon cyclurus by Smith (1943) but has been recognized as a valid species by Schleich & Kästle(2002) and Whitaker & Captain (2004).

Smith (1943), Wagner (1975) and Green et al. (2010) also recognized the “Oligodon cinereus group”. Thisgroup is mainly characterized by (1) short, not forked hemipenes with papillae but not spinose, (2) 15–21 dorsalscale rows (17 rows in O. cinereus), (3) 8–12 maxillary teeth, and (4) anal plate usually entire. According to Smith(1943), this group includes Oligodon cinereus (Günther, 1864), O. joynsoni (Smith, 1917), O. inornatus (Bou-lenger, 1914), O. albocinctus (Cantor, 1839), O. melanozonatus Wall, 1922, O. splendidus (Günther, 1875), and O.woodmasoni (Sclater, 1891). This group is also widespread from north-eastern India and Myanmar, to southernChina and to southern Thailand. In this paper, we discuss the distribution of Oligodon cinereus in India and the sta-tus of Simotes multifasciatus Jan & Sordelli, 1865, which Taylor (1965) considered to be a subspecies of Oligodoncinereus present in Thailand.

Material and methods

As a part of broader studies on the groups of Oligodon cyclurus and Oligodon cinereus, we investigated morpho-logical characters regarded as taxonomically significant by Smith (1943), Wagner (1975) and David et al. (2008a,b). Maxillary teeth were counted by removing the exterior gum surfaces of the right jaw in situ; tooth sockets wereincluded in the counts in cases of tooth loss. Measurements, except body and tail lengths, were taken with a slide-calliper to the nearest 0.1 mm; all measures on the body were measured to the nearest millimetre. The number ofventral scales was counted according to Dowling (1951). The numbers of dorsal scale rows are given at one headlength behind head, at midbody (i.e., at the level of the ventral plate corresponding to half of the total ventral num-ber), and at one head length before vent, respectively. Values for symmetric head characters are given in left/rightorder. Comparison with other species of the genus Oligodon was based on the literature, especially data providedby Smith (1943), Wagner (1975) and David et al. (2008a), and on examined specimens listed in Appendix I.Abbreviations of measures and other meristic characters used in the text are:

Measures and ratios. HL: head length; L-Fr: length of frontal; SVL: snout-vent length; TaL: taillength; TL: total length; TaL/TL: ratio tail length/total length; W-Fr: width of frontal. Meristic characters: ATem:anterior temporal; DSR: formula of dorsal scale rows; IL: infralabials; PreOc: preoculars; PosOc: postocularscales; SC: subcaudal scales; SL: supralabials; SubOc: suboculars; VEN: ventral scales; Temp: temporals. Museumabbreviations: BMNH: The Natural History Museum, London, UK; BNHS: Bombay Natural History Society,Mumbai, India; MNHN: Muséum National d’Histoire Naturelle, Paris, France; USNM: United States NationalMuseum, Washington, D.C., USA; ZSI: Zoological Survey of India, Kolkata, India.

Results

On the name-bearing type of Oligodon cyclurus (Cantor, 1839)

Coronella cyclura Cantor, 1839: 50. Type locality, by virtue of present neotype designation. Subdistrict of Kali-gunj, Lalmonirhat District, near Rangpur, Bangladesh (Original type locality: not given but implicitly from “Ben-gal”; see below). Neotype, by present designation. BMNH 1940.3.4.41 (adult male) (Original holotype depicted onPlate 15 of Cantor’s sketches “Drawings of animals”, deposited in the Bodleian Library, now the Radcliffe ScienceLibrary, Oxford, UK; not designated and not traced; considered lost).

There has been a substantial amount of confusion centred on the systematics of Oligodon cyclurus (Cantor,1839). Theodore Edward Cantor (1809-1860) was a Danish surgeon-naturalist with the East India Company. Hemade early collections in Peninsular Malaysia (then Malaya, described in Cantor, 1847) and most famouslydescribed the king cobra, Ophiophagus hannah, from the Sunderbans and Calcutta, in Bengal. Cantor both col-lected specimens (from Malaya) and had paintings made by local artists (for India).

For a long time, O. cyclurus was regarded as a wide-ranging, variable species (Boulenger 1894; Smith 1943).Taylor (1965), Campden-Main (1970) and especially Wagner (1975) addressed the systematics of this species. Thistaxonomic confusion arose in part because Cantor (1839) briefly described the species without indicating the num-


Zootaxa 2799 © 2011 Magnolia Press · 3TAXONOMY AND NOMENCLATURE OF INDIAN OLIGODON

ber of dorsal scale rows. Only the colour and pattern were described. Furthermore, Cantor failed to mention eithera type specimen or type locality. Consequently, the name Oligodon cyclurus has been applied to several popula-tions of Oligodon with 19, 21 or 23 dorsal scale rows at midbody and with different patterns (see, for example,Smith 1943).

A first indication on the origin of the type specimen indeed appears in the text of the original description itself.Cantor mentioned a Bengali vernacular name, Tukkr-Bora (literally, ‘sharp viper’ or ‘sharp boa’). In anotherdescription published in the same paper, Cantor associated the same Bengali vernacular name with a type localityexplicitly cited as “Bengal” (p. 51; description of Coluber curvirostris). Furthermore, a second indication appearsin the unpublished manuscript that accompanies the plates, Cantor wrote that Coronella cyclura inhabits “Bengal”.Hence, by all available evidence, the description was obviously based on a specimen from “Bengal”, which, at thetime of Cantor, was a Presidency of British India (until 1874), and included the Assam region in north-easternIndia; it also included under its administration, Penang (now Pulau Pinang, a state of the Federation of Malaysia)and Singapore, until 1867. Thereafter, Bengal included the region of eastern state of West Bengal, in addition toparts of eastern Bihar State in India (and East Pakistan, subsequently, Bangladesh, until 1947). The holotype wasdepicted on Plate 15 of Cantor’s colour sketches. These sketches and Cantor’s unpublished manuscript are depos-ited in the collection of the Radcliffe Science Library, Oxford University (Oxford, UK). References can beaccessed online at: http://www.bodleian.ox.ac.uk/rsl/ (last accessed 10 August 2010).Since Smith (1943), three authors either undertook major revisions of the group of Oligodon cyclurus and / oraddressed the status of its name-bearing type. All tried to fix the status of this specific nomen, as follows:

(1) Campden-Main (1970) addressed the taxonomy of populations of Oligodon “cyclurus” from southern Viet-nam with 19 dorsal scale rows at midbody. This author noticed that these specimens were not referable to the spe-cies with 21–23 dorsal scale rows occurring in Thailand, namely, according to Campden-Main’s taxonomicconcept, Coronella cyclura Cantor, 1839. Campden-Main showed that populations of southern Vietnam with 19dorsal scale rows were conspecific with Simotes brevicauda Steindachner, 1867. This problem has been discussedby Saint Girons (1972), Wagner (1975) and David et al. (2008a), who showed that Simotes brevicauda Steindach-ner, 1867 is a senior synonym of Oligodon ocellatus (Morice, 1875). Nevertheless, due to the absence of a type,Campden-Main (1970) considered it necessary to designate a neotype for Coronella cyclura Cantor, 1839. Thisauthor selected specimen USNM 72067 from “Bangkok, Thailand” (Campden-Main 1970: 764). Obviously, thisauthor had not investigated the original type locality of the species. The choice of “Bangkok” was merely becauseit is “in the area where most specimens that recent workers have assigned to cyclurus have been collected.” These“recent workers” were, for a large part, Taylor (1965).

(2) Subsequently, Wagner (1975) undertook a revision of Oligodon cyclurus. This author noted that two dis-tinct species were included under the specific nomen cyclurus as recognized by Campden-Main (1970). Wagnerdivided the taxon O. cyclurus into a western taxon with a constant, maximum number of 19 dorsal scale rows onthe neck and midbody, occurring in India, Bangladesh and Myanmar, and an eastern one, with a maximum numberof 21 or 23 rows (never 19) occurring east of western Myanmar (Myanmar, Thailand, Laos, Cambodia and Viet-nam). Eventually, Wagner regarded these taxa as representing three distinct species. As stated by Wagner (1975:54), the problem arose as to which of these three species the nomen cyclura Cantor, 1839 applies. It could be tospecimens with a maximum of 19 rows from India, Bangladesh and Myanmar, specimens with 19 rows from south-ern Vietnam, or eastern specimens with 21–23 dorsal scale rows.

Insightfully, Wagner (1975) noted the presence of the Bengali name Tukkr-Bora in the original description,making it obviously a species occurring, at least in part, in this region of India or Bangladesh, where this languageis spoken. On the basis of (i) this indication of an implicit type locality, and (ii) the fact that specimens from Indiaand Bangladesh have a maximum dorsal scale row number of 19 (never 21), Wagner (1975: 54–55) applied thecombination Coronella cyclura Cantor, 1839 to these more western populations with 19 dorsal scale rows, whereasSimotes fasciolatus Günther, 1864 was the next available name for eastern populations with 21−23 rows. Wagner(1975) refuted the neotype designation of Campden-Main (1970) on the basis that its locality, far from the originallocality of Cantor’s holotype, had been incorrectly selected. Nevertheless, upon the taxonomic and geographicalbases discussed above, Wagner (1975: 62) designated as neotype of Coronella cyclura Cantor, 1839 specimenBMNH 1940.3.4.41, from “Kaliganz, Rangpur, Bangladesh”, now the subdistrict of Kaligunj (or Kaligunge, Kali-gonj), Lalmonirhat District, near Rangpur (25º 44’ N, 89º 16’ E), northern Bangladesh. This latter position hasbeen accepted by Pauwels et al. (2002) and other authors.



(3) Das (2004) challenged this interpretation on the basis that Wagner (1975) is an unpublished Masters thesisand cannot be considered a published work in the sense of the International Code of Zoological Nomenclature(I.C.Z.N. 1999; referred to as the Code below). Das (2004) took this opportunity to confirm the validity of the des-ignation of the neotype from Bangkok carried out by Campden-Main (1970). The action of Das (2004) was unfor-tunate because Wagner (1975) had showed that the eastern taxon, occurring east of central Myanmar (easternMyanmar, Thailand and farther east), does not agree in body scalation with specimens of the region where Cantor(1839) obtained the holotype of Coronella cyclura.

Campden-Main’s designation created both nomenclatural and taxonomic problems, because the neotype wouldthus belong to a taxon different from Cantor’s definition. Fortunately, his designation is invalid according to twodispositions of Art. 75.3, as follows: (i) Campden-Main’s neotype designation contravenes Art. 75.3.5 of the Code,which stipulates that there should be “evidence that the neotype is consistent with what is known of the formername-bearing type from the original designation and from other sources.” Unfortunately, Campden-Main’s neotypeis a specimen with 21 dorsal scale rows, whereas Cantor had a specimen with 19 rows, as shown by the drawing ofthe specimen on Plate 15 of Cantor’s sketches. (ii) Campden-Main’s neotype designation also contravenes Art.75.3.6 of the Code, which stipulates that there should be “evidence that the neotype came as nearly as practicablefrom the original type locality (…).” Indications in Cantor’s published description, such as a vernacular name inBengali, make clear that the type was from “Bengal” (see comments above), and definitely not from Thailand.

According to Art. 75.4 of the Code, we do not consider the neotype designation of Campden Main to be validbecause it is not in accordance with two provisions of Art. 75. As a consequence, we here formally designate asneotype of Coronella cyclura Cantor, 1839, specimen BMNH 1940.3.4.41 (adult male), from “Kaliganz, RangpurDistrict, Bangladesh”, now Subdistrict of Kaligunj, Lalmonirhat District, Bangladesh. This specimen was previ-ously selected by Wagner (1975) in his unpublished thesis.

Our neotype designation complies with all provisions of Art. 75.3 as follows: this neotype is designated for thepurpose of clarifying the taxonomic status of Coronella cyclura Cantor, 1839, in associating this specific nomenwith specimens from the western part of the range of Oligodon cyclurus complex with 19 dorsal scale rows (Art.75.3.1); Oligodon cyclurus is the valid name for populations with 19 rows, distinct at specific level from popula-tions with 21 or 23 rows known as Oligodon fasciolatus (Günther, 1864) (Art. 75.3.2). The neotype BMNH1940.3.4.41 was described in detail by Wagner (1975: 62) but its main characters are: SVL 500 mm, TaL 78 mm;ratio TaL/TL 0.135; 165 VEN, 41 SC (value checked by C. J. McCarthy, pers. comm., January 2011; the valuegiven by Wagner, 18 SC, is obviously a lapsus); 19–19–15 DSR; 7/7 SL (3+4/4 entering orbit); 1/1 PreOc, 2/2PosOc; 2+2/2+2 Temp; 8/8 IL. Body tan with faint reticulations; head tan, unpatterned; venter yellow with brownspots (Art. 75.3.3); the original holotype could not be traced either in the Natural History Museum, London, in col-lections of the University of Oxford, or even in collections of the Zoological Survey of India at Kolkata, potentialcollections and/or institutions associated with Cantor and his herpetological specimens (Art. 75.3.4); the neotype isconsistent in morphology with the original holotype (see above) and comes from the same locality, namely the for-mer Bengal (Art 75.3.5 and 75.3.6); the neotype is deposited at the Natural History Museum, London (Art. 75.3.7).

As a consequence, we confirm the position of Wagner (1975) in retaining the specific epithet cyclurus for thewestern populations (India, Bangladesh and Myanmar) with 19 dorsal scale rows on the neck and at midbody.Specimens with 21 or 23 rows from Myanmar, Thailand, Cambodia, Laos and Vietnam are referred to Oligodonfasciolatus (Günther, 1864). As currently conceived, Oligodon cyclurus (Cantor, 1839) is monotypic. Oligodoncyclurus smithi (Werner, 1925) and Oligodon cyclurus superfluens Taylor, 1965 are synonyms of Oligodon fascio-latus, a species also considered monotypic (Wagner, 1975)

On the validity of Oligodon kheriensis Acharji & Ray, 1936

Oligodon kheriensis Acharji & Ray, 1936: 519. Type locality. “North Kheri Division, Eastern Circle, Kheri-Lakimpur, in the United Provinces”, currently near Kheri, 27º 57’ N, 80º 47’ E, Lakhimpur Kheri District, State ofUttar Pradesh, India. Holotype. ZSI 20391 (adult female).

This taxon was considered a synonym of Oligodon cyclurus by Smith (1943: 202) and Wagner (1975: 62).Nevertheless, an examination of the holotype of Oligodon kheriensis (Figs. 1A–C) shows that it is distinct fromOligodon cyclurus. Another specimen of O. kheriensis from Mahendranagar, in extreme southwestern Nepal, was



documented by Schleich & Kästle (2002). This specimen was deposited in the collection of the Siddarnath ScienceCampus, Mahendranagar, Nepal, and could not be examined by us. On the basis of the holotype, that we examined,and of the specimen from Nepal, this species can be defined as follows (when a single value is given, it refers to theholotype) and in Table 1.

TABLE 1. Main morphological data of Oligodon kheriensis. See Material and Methods for explanation of abbreviations. Sizesare in millimetres.

FIGURE 1. Holotype of Oligodon kheriensis Acharji & Ray, 1936, ZSI 20391. A: General view of body dorsum. B: Head indorsal view. C: Body venter.

Description and variation. Body robust, subcylindrical; head ovoid, short, thick, barely distinct from the thickneck; snout projecting over the lower jaw, long, rounded, amounting to 40.7% of HL, or 6.1 times as long as diam-eter of eye; eye rather small, with a round pupil; tail thick, short and tapering. Maximal total length: 1,010 mm(specimen from Nepal, presumed male). Dentition: typical of the Oligodon cyclurus group: 8 maxillary teeth,namely 6 small + the last two being strongly enlarged and blade-like. Body scalation: DSR 19–19–17 rows; allscales smooth, small and ovoid. Ventral plates slightly angulated; subcaudal all paired; anal plate entire. Head sca-lation: rostral enlarged, thick, curved onto the upper snout surface, visible from above, not separating internasals;nasals slightly “butterfly-shaped”, ca. 0.6 times as long as high, vertically divided, with the posterior part distinctlysmaller than anterior one; nostril crescentic, piercing middle of nasal just in front of the division; internasals sub-rectangular, in broad contact, shorter than prefrontals; prefrontals subrectangular, distinctly wider than long; frontalpentagonal, trapezoid, as long as wide; a supraocular on each side, distinctly longer than wide, wider than prefron-tals; two very large subtriangular parietals, much longer than the frontal, in broad contact; loreal scales, in contactwith the nasal; 8/8 supralabials, 1st SL small, 2nd and 3rd in contact with the loreal, 4th and 5th entering orbit, 6thand 7th largest; preoculars high and narrow; no presubocular; small postoculars; 2+2 or 1+1 temporals, anteriorones elongated; 8/8 infralabials, first pair in contact, IL 1–5 in contact with anterior chin shields, 5th IL the largest.Colour and pattern in alcohol: the holotype is bleached presumably through long-term exposure to daylight. Thedorsum is currently unpatterned ivory yellow; venter paler, without dark blotches. The original description givesthe following description “… coral red above, yellowish below”. The specimen from Nepal described by Schleich& Kästle (2002: 901; pls. 96–97), although also preserved, is in better condition. These authors described it as

Number Sex SVL TaL TaL/TL VEN SC DSR SL PreOc PosOc SubOc Lor Temp

ZSI 20391 & 840 102 0.108 196 38 19–19–17 8/8 1/1 1/1 0/0 1/1 2+2 / 2+2

“Nepal” % 870 140 0.139 196 43 19–19–17 8/8 1/1 2/2 0/0 1/1 2+2 / 1+1



almost uniform coral above, with some faint brownish spots on dorsal scales of the anterior body. No dark headpattern such as chevrons typical of the Oligodon cyclurus group. The venter is pink.

Distribution (Fig. 2). India. State of Uttar Pradesh. Near Kheri, in Lakhimpur Kheri District. – Nepal. Farwestern Region. Mahendranagar, in Kanchanpur District.

Both localities are separated by only 120 km. Lakhimpur Kheri District borders Nepal.

FIGURE 2. Ranges of Oligodon kheriensis and Oligodon cyclurus in India and adjacent countries. Oligodon kheriensis. Star =type locality; triangle = specimen cited by Schleich & Kästle (2002), Mahendranagar, southwestern Nepal. Oligodon cyclurus.Pentagon = locality of the neotype; solid circles = examined specimens; open circles = literature records.

Comparison with Oligodon cyclurus. We refer Oligodon kheriensis to the Oligodon cyclurus group on thebasis of its external morphology, especially the number of dorsal scale rows and the presence of two anterior tem-porals. Its affinities will be confirmed when hemipenes have been described. Oligodon kheriensis is thus the west-ernmost representative of the Oligodon cyclurus group. It can be separated from O. cyclurus by the followingcharacters: (1) higher number of ventral scales, 196 vs. 159–178 VEN (latter values based on examined specimens,plus a few literature records dealing only with specimens from India) in O. cyclurus; (2) uniform dorsum vs. retic-ulated or blotched (or blotched and striped) in O. cyclurus, and (3) the absence of the large, arrow-like cephalic andnape marking in O. kheriensis.

In India, Oligodon cyclurus has been recorded from a few localities in north-eastern India (see Fig. 2, based onWagner 1975; Ahmed & Dasgupta 1992; Mathew 1995; and our material). Acharji (1958: 389) recorded a speci-men from “Inspection Bungalow, Inanpur, Manbhum District, Bihar”, now Ramkanali (23° 35’ N, 86° 76’ E),Puruliya district, State of West Bengal. The morphological data provided by the author do not allow a formal iden-tification of this specimen but its colour and pattern are consistent with those of Oligodon cyclurus.



Further evidence in favour of the distinct specific status of O. kheriensis is its geographical isolation from Oli-godon cyclurus. Its type locality is about 840 km from the westernmost confirmed locality of Oligodon cyclurus(Jalpaiguri, State of West Bengal), or ca. 750 km from the locality of the specimen recorded by Acharji (1958).

Note on Oligodon cinereus (Günther, 1864) in India

Simotes cinereus Günther, 1864: 215. Type locality. “Gamboja”, now Cambodia. Holotype. BMNH RR1946.1.1.25 (adult male).

This species is widespread from extreme north-eastern India and eastern Bangladesh to south-eastern Chinaand Taiwan, and southwards up to Peninsular Thailand across the whole of Myanmar, Laos, Cambodia and Viet-nam (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 scalesincluding a loreal, (5) 8 (rarely 7) SL, SL 4–5 (rarely 3–4) entering orbit, (5) and 1 anterior temporal. Specimenswith 15 DSR are referable to Oligodon inornatus (Boulenger, 1914); specimens with 17–17–15 DSR and morethan 185 VEN belong to Oligodon joynsoni (Smith, 1917). This latter species is quite similar to reticulate patternedspecimens 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.

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

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 ofthis species remains unresolved. Although Smith (1943) did not recognize any subspecies, other authors such asTaylor (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 wereunclear. 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 isunclear, and would appear to have been collected in the first half of the 1800s, a period when only two notable col-lections 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 threeSchlagintweit brothers, Hermann (1826–1882), Adolphe (1829–1857) and Robert (1833–1885), who originatefrom 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 anumber 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. toP. David, January 2009), the zoological museum in Bonn at that time was the “Naturhistorisches Museum Poppels-dorf” (situated in what is now a suburb of the city of Bonn), but was destroyed during World War II (see Friedrich2006), 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). Thedepicted specimen shows a dorsal pattern made of irregular, dark crossbars, a dark nuchal marking pointing for-wards, some blotches on the head, and a venter and under tail surface strongly marked with dark, square blotches.

TABLE 2. Main morphological data of Oligodon cinereus in India. “Type J&S” = holotype of type of Simotes multifasciatusJan & Sordelli, 1865. See Material and Methods for explanation of other abbreviations. Lengths in millimetres.

Taylor (1965) recognized O. cinereus swinhonis (Günther, 1864) and O. cinereus multifasciatus (Jan & Sor-delli, 1865) for populations from Thailand. However, Taylor (1965) pointed out that the type locality of Simotesmultifasciatus Jan & Sordelli, 1865 was “Sultanepore”, “a locality unknown to me”. Although there is no suchlocality in Thailand, there are indeed several cities known as Sultanpoor, now Sultanpur in India. All of them arelocated 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, themost 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 specimendescribed 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 scalerows allow us to agree with previous authors, such as Smith (1943) and Wagner (1975), in referring this taxon tothe Oligodon cinereus complex, namely O. cinereus and its current numerous synonyms. However, this locality isout 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 previouslywesternmost 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 thisspecimen (Figs. 3A–C), which shares with the holotype of Simotes multifasciatus the number of 17 DSR, the dorsalpattern with distinct black crossbands and the blotched venter. We here refer it to Oligodon cinereus, as currentlydefined. 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 ofOligodon cinereus in northern India (see Fig. 4). In the north-western part of its range, this species is now knownfrom 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 Dis-trict. 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 toits scalation data, especially the number of dorsal scale rows, the specimen recorded from Changlang District, Stateof 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 severalmorphs within this species, based on the dorsal, head and ventral patterns. Smith’s “Var. I” is nearly unpatternedabove and with a powdered venter. This morph, to which the holotype of Simotes cinereus belongs, is widespreadin Myanmar, Thailand and the Indochinese Peninsula. Smith (1943) also recognized a “Var. II” with strong, dark

Number Sex SVL TaL TaL/TL VEN SC DSR SL PreOc PosOc SubOc Temp

Type J&S % ---- ---- ---- ---- 32 ?–17–? 8/8 ?/1 ?/2 ?/1 ? / 1+2

ZSI 12356 & 150 17 0.102 169 30 17–17–15 8/7 1/1 2/2 1/1 1+2 / 1+2



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 spec-imens with distinct dorsal black crossbars and strong reticulations between the crossbars, head with at least anuchal 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, northernThailand, and Vietnam. Smith also recognized a “Var. IV”, which is not relevant to the present discussion becauseit 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 therange of species in correlation with the dorsal pattern of specimens examined by him. The range of the “black-barred” 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 andconfirm that they are identical in pattern and scalation with those from northern Vietnam.

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

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 twomorphs. In contrast, specimens with dark, solid dorsal crossbars (Form III of Smith; black-barred morph of Wag-ner) 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 twomorphs 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 darknuchal chevron and a heavily blotched venter. These characters are similar to those of Smith’s Form III. We hererefer the holotype of Simotes multifasciatus to this morph. A discussion on the intraspecific taxonomy of Oligodoncinereus will be addressed elsewhere (Vogel et al., unpublished data) but black barred specimens of India, Myan-



mar and Thailand and those of northern Vietnam and Laos might prove to be distinct at the species level from othermorphs of Oligodon cinereus.

TABLE 3. Comparison between morphs of Oligodon cinereus. See Material and Methods for explanation of abbreviations.

The precision of the type locality of Simotes multifasciatus Jan & Sordelli, 1865 has implications for the taxon-omy of the genus Oligodon in Thailand. Taylor (1965) referred a total of four specimens of Oligodon from northernThailand to the subspecies O. cinereus swinhonis (Günther, 1864) (one specimen; from “12 km north of ChiangDao, Chiang Mai Province) and O. cinereus multifasciatus (Jan & Sordelli, 1865) (three; from “Chang Dow [12km, 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 ofirregular dark crossbars with either faint (“swinhonis”) or strong (“multifasciatus”) reticulations.

TABLE 4. Comparison between specimens of Oligodon joynsoni and Thai specimens referred to Oligodon cinereus by Taylor(1965). See Material and Methods for explanation of abbreviations. Lengths in millimetres. * The value of the total lengthgiven by Taylor (90 mm) is obviously a lapsus for 190 mm.

We have discussed above the status of Simotes multifasciatus Jan & Sordelli, 1865, showing that its type local-ity is indeed ‘India’. The type locality of Simotes swinhonis Günther, 1864 is “Amoy”, now Xiamen, Fujian Prov-ince, People’s Republic of China. The number of ventrals in specimens examined by Taylor is much higher thanthose of the syntypes of Simotes swinhonis (156–168) and of the Indian specimen identified by us above as similarto 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 pat-tern with the description of Simotes longicauda joynsoni Smith, 1917 (p. 276. Type locality: “Maa Yome, MuangNgow, N. Thailand”, now Muang Ngao or Muang Ngau, 18° 45' 27” N, 99° 58' 42” E, Lampang Province). Oligo-don joynsoni (Smith, 1917) is now considered a valid species of the O. cinereus group based on a high number ofventrals (> 185) and subcaudals (> 36). Furthermore, Taylor’s specimens were obtained from the provinces of Chi-ang 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 swinhonisand O. cinereus multifasciatus. The current range of O. joynsoni, a relatively rarely encountered species, knownfrom northern Thailand and Laos, is given in Fig. 5 (after Smith 1943; Taylor 1965; Nabhitabhata et al. 2004; ourmaterial). This species is now known from the following localities: Thailand. Chiang Mai Province: North of

Morph Sex TaL/TL VEN Sc L-Fr / W-Fr

Smith’s I + II“cinereus”(n = 18)

%

&

0.129–0.149 ( = 0.140)

0.123–0.131 ( = 0.127)

156–168 ( = 162.9)

167–178 ( = 173.5)

35–42 ( = 37.2)

33–37 ( = 35.3)

1.20–1.37( = 1.28)

Smith’s III“tamdaoensis”(n = 19)

%

&

0.122–0.140 ( = 0.128)

0.101–0.120 ( = 0.112)

169–180 ( = 173.3)

173–184 ( = 176.4)

30–42 ( = 36.6)

30–36 ( = 32.9)

1.11–1.30( = 1.20)

Specimen Sex SVL TaL TaL/TL VEN SC DSR SL ATem

O. joynsoni

BMNH 1946.1.4.23 % 655 105 0.138 190 47 17–17–15 8 / 8 1 / 1

BMNH 1969.1809 % 570 100 0.149 186 50 17–17–15 7 / 7 2 / 2

MNHN 1896.0633 & 572 80 0.123 194 39 17–17–15 8 / 8 2 / 2

Taylor’s specimens

3464 & 548 84 0.133 196 42 17–17–15 7 / 7 2 / 2

EHT-HMS 3623 & 606 77 0.113 196 38 17–17–15 7 / 7 2/1 / 2/1

31699 % 529 71 0.119 190 32 17–17–15 8 / 7 -----

28 % 169* 21 0.111 197 42 17–17–15 7 / 7 -----

xx

xx

xx

x

xx

xx

xx

x



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 (KhlongLan) and Nakhon Ratchasima (Khao Yai). However, no information on their scalation was provided and given thatconfusion 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 tothe fact that the holotype of S. multifasciatus agrees in pattern with O. joynsoni (Smith, 1917).

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

Conclusion

Whitaker & Captain (2004) recognized 22 species of the genus Oligodon Fitzinger, 1826 in India, whereas Sharma(2007) listed only 19. According to our data, 21 species have been recorded within the political limits of India, asfollows: Oligodon affinis Günther, 1862; Oligodon albocinctus (Cantor, 1839); Oligodon arnensis (Shaw, 1802);Oligodon brevicauda Günther, 1862; Oligodon calamarius (Linnaeus, 1758); Oligodon catenatus (Blyth, 1854);Oligodon cinereus (Günther, 1864); Oligodon cyclurus (Cantor, 1839); Oligodon dorsalis (Gray, 1835); Oligodonerythrogaster Boulenger, 1907; Oligodon erythrorhachis (Wall, 1910); Oligodon juglandifer (Wall, 1909); Oli-godon kheriensis Acharji & Ray, 1936; Oligodon melaneus (Wall, 1909); Oligodon melanozonatus Wall, 1922;Oligodon nikhili Whitaker & Dattatri, 1982; Oligodon taeniolatus (Jerdon, 1854); Oligodon theobaldi (Günther,1868); Oligodon travancoricus Beddome, 1877; Oligodon venustus (Jerdon, 1854); Oligodon woodmasoni(Sclater, 1891).



The occurrence of O. calamarius is based on Whitaker & Captain (2004). Oligodon dorsolateralis (Wall,1910), recognized as valid by Whitaker & Captain (2004), is currently regarded as a synonym of O. cyclurus fol-lowing Wagner (1975).

The revalidation of Oligodon kheriensis and the intricate intraspecific taxonomy of O. cinereus suggest thatrevisions of the two main groups of Oligodon defined by Smith (1943) are necessary in order to fully understandthe diversity within the genus. This is especially true for Oligodon cinereus. Our material suggests notable differ-ences in scalation correlated with the dorsal pattern which may warrant the recognition of distinct species.

Acknowledgements

We are grateful to Olivier S.G. Pauwels (Institut Royal des Sciences Naturelles de Belgique, Brussels, Belgium) forhis constructive comments on the draft of this paper. We thank Colin McCarthy (BMNH) and G. Ramakrishna,B.H.C.H. Murthy and S. Raha (ZSI) for their kind permission to examine specimens in their collections. We alsothank Theo Dunnet, former librarian (The Radcliffe Science Library, Oxford University, Oxford, UK), for his assis-tance with T. Cantor’s unpublished sketches.

References

Acharji, M.N. (1958) On a collection of chelonians and snakes from Chota Nagpur, Bihar. Records of the Indian Museum,53[1955] (3–4), 383−392.

Acharji, M.N. & Ray, H.C. (1936) A new species of Oligodon from the United Provinces (India). Records of the IndianMuseum, 38, 519−520.

Agarwal, I., Mistry, V.K. & Athreya, R. (2010) A preliminary checklist of the reptiles of Eaglenest Wildlife Sanctuary, WestKameng District, Arunachal Pradesh, India. Russian Journal of Herpetology, 17 (2), 81–93.

Ahmed, S. & Dasgupta, G. (1992) Reptilia. In: Anonymous, Fauna of West Bengal. Part 2. Calcutta, Zoological Survey ofIndia, State Fauna Series, 3, 1–65.

Angel, F. (1927) Liste des reptiles et des batraciens rapportés de l’Indo-Chine par M. P. Chevey. Description d’une variété nou-velle de Simotes violaceus Cantor. Bulletin du Muséum d’Histoire naturelle, Paris, 33 (6), 496–498.

Armitage, G. (1989) The Schlagintweit collections. Indian Journal of History of Science, 24, 67–83.Beddome, R.H. (1877) Descriptions of new reptiles from the Madras Presidency. Proceedings of the Zoological Society of Lon-

don, 1877 (4), 685–686.Blyth, E. (1854) Notices and descriptions of various reptiles, new or little known. Journal of the Asiatic Society of Bengal, 23

(3), 287–302.Boulenger, G.A. (1894) Catalogue of the Snakes in the British Museum (Natural History). Volume II., containing the conclusion

of the Colubridae Aglyphae. British Museum (Natural History), London, xi + 382 pp., Pls. 1–20.Boulenger, G.A. (1907) Description of a new snake from Nepal. Records of the Indian Museum, 1 (3), 217.Boulenger, G.A. (1914) Descriptions of new reptiles from Siam. With notes by Malcolm Smith, M.R.C.S., L.R.C.P. The Jour-

nal of the Natural History Society of Siam, 1(2), 67–70, Plate.Bourret, R. (1934) Notes herpétologiques sur l’Indochine française. IV. Sur une collection d’Ophidiens de Cochinchine et du

Cambodge. Bulletin général de l’Instruction Publique, 13e Année (1, septembre), 13–30. [Separate reprint: p. 3–20].Bourret, R. (1935) Notes herpétologiques sur l’Indochine française. X. Les serpents de la station d’altitude du Tam-dao. Bulle-

tin Général de l’Instruction Publique, 14e Année (8, avril), 259–271 [Separate reprint: p. 1–13].Campden-Main, S.M. (1970) The identity of Oligodon cyclurus (Cantor, 1839) and revalidation of Oligodon brevicauda (Stein-

dachner, 1867) (Serpentes: Colubridae). Proceedings of the Biological Society of Washington, 82 (58), 763–766.Cantor, T.E. (1839) Spicilegium serpentium indicorum. Proceedings of the Zoological Society of London, 1839, 31–34 & 49–

55.Cantor, T.E. (1847) Catalogue of reptiles inhabiting the Malay Peninsula and islands, collected or observed by Theodore Can-

tor, Esq., M.D., Bengal Medical Service. Journal of the Asiatic Society of Bengal, 16, 607–656, 897–952, 1026–1078Das, I. (2004) On neotype designations for Coronella cyclura Cantor, 1839 (Serpentes: Colubridae). Hamadryad, 29 (1), 133–

134.Dasgupta, G. & Raha, S. (2004) Reptilia. In: Anonymous, Fauna of Bihar (including Jharkhand). Part-1. Kolkata, Zoological

Survey of India, State Fauna Series, 11, 143–179.Dasgupta, G. & Raha, S. (2006) Reptilia. In: Anonymous, Fauna of Nagaland. Kolkata, Zoological Survey of India, State

Fauna Series, 12, 433–460, Figs. 1–21.David, P., Vogel, G. & Pauwels, O.S.G. (2008a) A new species of the genus Oligodon Fitzinger, 1826 (Squamata: Colubridae)

from southern Vietnam and Cambodia. Zootaxa, 1939, 19–37.David, P., Vogel, G. & van Rooijen, J. (2008b) A revision of the Oligodon taeniatus (Günther, 1861) (Squamata: Colubridae)



group, with the description of three new species from the Indochinese Region. Zootaxa, 1965, 1–45.Dowling, H.G. (1951) A proposed standard system of counting ventrals in snakes. British Journal of Herpetology, 1 (1), 97−99.Fitzinger, L.J. (1826) Neue classification der Reptilien nach ihren natürlichen Verwandtschaften. Nebst einer Verwandtschafts-

tafel und einem Verzeichnisse der Reptilien-sammlung des K. K. Zoologischen Museum’s zu Wien. J. G. Heubner, Wien, (6unnumbered pages) + 66 pp., Table.

Friedrich, J. (2006) The fire: the bombing of Germany, 1940-1945. Columbia University Press, New York, 532 pp.Gans, C. (1955) Localities of the herpetological collections made during the “Novara Reise”. Annals of the Carnegie Museum,

33, 275–285.Gray, J.E. (1832–1835) Illustrations of Indian zoology; chiefly selected from the collection of Major-General Hardwicke, F.R.S.,

L.S., M.R.A.S., M.R.I.A., & C. Volume II (parts XI-XX). London, Adolphus Richter and Co & Parbury, Allen, and Co, (3unnumbered pages), Pls. 1–102.

Green, M.D., Orlov, N.L. & Murphy, R.W. (2010) Toward a phylogeny of the Kukri snakes, genus Oligodon. Asian Herpetolo-gical Research, 1 (1), 1–21.

Günther, A. (1862) On new species of snakes in the collection of the British Museum. Annals and Magazine of Natural History,(3), 9 (49), 52–67, Pls. 9–10.

Günther, A. (1864) The reptiles of British India. Ray Society, London, xxvii + 452 pp., Pls. 1–26.Günther, A. (1868) Sixth account of new species of snakes in the collection of the British Museum. Annals and Magazine of

Natural History, (4), 1 (6), 413–429, Pls. 17–19.Günther, A. (1872) Seventh account of new species of snakes in the collection of the British Museum. Annals and Magazine of

Natural History, (4), 9 (49), 13–37, Pls. 3–6.Günther, A. (1875) Second report on collections of Indian Reptiles obtained by the British Museum. Proceedings of Zoological

Society of London, 1875, 224–234, Pls. 30–34.Günther, A. (1888) On a collection of reptiles from China. Annals and Magazine of Natural History, (6), 1 (3), 165–172, Pl. 12.I.C.Z.N. (International Commission on Zoological Nomenclature) (1999) International Code of Zoological Nomenclature.

Fourth Edition adopted by the International Union of Biological Sciences. International Trust of Zoological Nomencla-ture, London, xxix + 306 pp.

Jan, G. & Sordelli, F. (1860–1881) Iconographie générale des ophidiens. Atlas. Milano, F. Sordelli, London, Baillière Tindal &Co, Paris, J.-B. Baillière & Fils; Madrid, C. Bailly-Baillière, Vol. I [1860–1866], Livraisons 1–17, Pls. 1–102.

Jerdon, T.C. (1854) Catalogue of the Reptiles inhabiting the Peninsula of India. Journal of the Asiatic Society of Bengal, 22[1853] (6), 522–534.

Linnaeus, C. (1758) Systema Naturae per regna tria naturae, secundum classes, ordines, genera, species, cum characteribus,differentiis, synonymis, locis. Tomus I. Editio decima, reformata. Holmiae 5stockholm), Laurentius Salvius, 824 pp.

Mathew, R. (1995) Reptilia. In: Anonymous, Fauna of Meghalaya. Part 1. Vertebrates. Calcutta, Zoological Survey of India,State Fauna Series, 4, 379–454, Figs. 1–4, Pls. 1–38.

Morice, A. (1875) Coup d’oeil sur la faune de la Cochinchine française. Archives de Médecine navale, 24, 5–101.Nabhitabhata, J., Chan-ard, T. & Chuaynkern, Y. (2004 [dated “2000”]) Checklist of amphibians and reptiles in Thailand.

Office of Environmental Policy and Planning (OEPP), Bangkok, Biodiversity Series, Vol. 9, 152 pp.Pauwels, O.S.G., Wallach, V., David, P. & Chanhome, L. (2002) A new species of Oligodon Fitzinger, 1826 (Serpentes, Colu-

bridae) from southern Peninsular Thailand. Natural History Journal of the Chulalongkorn University, Bangkok, 2 (2), 7–18.

Praschag, P., Sommer, R.S., McCarthy, C., Gemel, R. & Fritz, U. (2008) Naming one of the world’s rarest chelonians, thesouthern Batagur. Zootaxa, 1758, 61–68.

Saint Girons, H. (1972) Les serpents du Cambodge. Mémoires du Muséum National d’Histoire Naturelle, Paris, (Nouvellesérie), (Série A, Zool.), 74, 1−170, Pls. 1−42.

Sanyal, D.P. & Gayen, N.C. (2006) Reptilia. In: Anonymous, Fauna of Arunachal Pradesh (Part-1). Kolkata, Zoological Sur-vey of India, State Fauna Series, 13, 247–284.

Schleich, H.H. & Kästle, W. (Eds.) (2002) Amphibians and Reptiles of Nepal. Ruggell (Germany), A. R. G. Gantner Verlag K.G., x + 1,201 pp.

Sclater, W.L. (1891) Notes on a collection of snakes in the Indian Museum, with descriptions of several new species. Journal ofthe Asiatic Society of Bengal, 60 (2), 230–250, Pl. 6.

Shaw, G. (1802) General zoology, or systematic natural history. Vol. III. Part II. Amphibia. G. Kearsley, London, i–vii + 313–615, Pls. 87–140. [Reprinted in 1999 by the Society for the Study of Amphibians and Reptiles, Ithaca, New York].

Sharma, R.C. (2007) Fauna of India and adjacent countries. Reptilia. Volume-III (Serpentes). Kolkata, Zoological Survey ofIndia, xxvi + 410 pp.

Smith, M.A. (1917) Description of a new snake and a new frog from Siam. The Journal of the Natural History Society of Siam,2 (4), 276–278

Smith, M.A. (1943) The Fauna of British India, Ceylon and Burma, including the whole of the Indo-chinese subregion. Reptiliaand Amphibia. Vol. III, Serpentes. Taylor & Francis, London, xii + 583 pp.

Steindachner, F. (1867) Reptilien. In: Reise der Österreichischen Fregatte Novara um die Erde in den Jahren 1857, 1858, 1859,unter dem Befehln des Commodore B. von Wüllerstorf-Urbair. Zoologischer Theil. Erster Band (Wirbelthiere). Vienna,Kaiser-König. Hof-u. Staatsdrückerei, 1–98, Pls. 1–3.

Sthanapati, J. (1996) History of magnetic studies in India 1850–1980: a bibliography. Indian Journal of History of Science, 31,



205–302.Taylor, E.H. (1965) The serpents of Thailand and adjacent waters. University of Kansas Science Bulletin, 45 (9), 609−1096.Tillack, F. & Günther, R. (2010) Revision of the species of Oligodon from Sumatra and adjacent islands, with comments on the

taxonomic status of Oligodon subcarinatus (Günther, 1872) and Oligodon annulifer (Boulenger, 1893) from Borneo (Rep-tilia, Squamata, Colubridae). Russian Journal of Herpetology, 16[2009] (4), 265–294.

Wagner, F.W. (1975) A revision of the Asian colubrid snakes Oligodon cinereus (Günther), Oligodon joynsoni (Smith), and Oli-godon cyclurus (Cantor). Unpublished MS thesis, Baton Rouge, Louisiana State University, 97 pp.

Wall, F. (1909) Notes on snakes from the neighbourhood of Darjeeling. Journal of the Bombay Natural History Society, 19 (2),337–357.

Wall, F. (1910) A new snake from Assam. Oligodon erythrorachis. Journal of the Bombay Natural History Society, 19 [1909](4), 923–924.

Wall, F. (1922) A new snake from the Northern Frontier of Assam. Records of the Indian Museum, 24, 29–30.Werner, F. (1925) Neue oder wenig bekannte Schlangen aus dem Wiener naturhistorischen Staatsmuseum (2. Teil). Sitzungs-

berichte der Kaiserlichen Akademie der Wissenschaften. Mathematisch-Naturwissenschaftliche Classe. (Abt. 1. Mineralo-gie, Botanik, Zoologie, Geologie und Paläontologie), 134, 45–66.

Whitaker, R. & Captain, A. (2004) Snakes of India. The Field Guide. Draco Books, Chennai, xiv + 481 pp., map.Whitaker R. & Dattatri, S. (1982) A new species of Oligodon from the Palni Hills, south India (Serpentes: Colubridae). Journal

of the Bombay Natural History Society, 79 (3), 630–631.

Appendix. Specimens examined

Oligodon cyclurus (21 specimens). Bangladesh. ZSI 7087, “Kaligunz, Rangpore”, now Subdistrict of Kaligunj, LalmonirhatDistrict, near Rangpur. India. State of Meghalaya: BNHS 804, “Khassia Hills, Assam”, now Khasi Hills. BNHS 806,Tura, Garo Hills. State of West Bengal: BMNH 1936.7.2.6–7, “Mal District, Dooars, 400’, N. Bengal”, now Mal, Jal-paiguri District; ZSI 8611–12, Darjeeling; BNHS 812–813, Tinelhari (= Tindharia?), Darjeeling District; BMNH1969.1802, BNHS 808–811; ZSI 12878, Jalpaiguri, Jalpaiguri District; ZSI 18674, Sevoke, DarjeelingDistrict. Myanmar. Ayeyarwady Division: BNHS 802, “Bassein”, now Pathein. Bago Division. ZSI 7091, ZSI 12390,“Pegu”, now Bago. Kachin State: MNHN 1893.0389, Bhamo. Yangon Division: BNHS 799, ZSI 11571, “Rangoon”, nowYangon. No locality: MNHN 4241, “Birmanie”, Burma, now Myanmar.

Oligodon kheriensis (1 specimen). India. State of Uttar Pradesh: ZSI 20391 (holotype), “North Kheri Division, Eastern Circle,Kheri-Lakimpur, in the United Provinces”, currently near Kheri, 27º57’N, 80º47’E, Lakhimpur Kheri District.

Oligodon joynsoni (5 specimens). Thailand. Lampang Province: BMNH 1946.1.4.23 (Holotype), BMNH 1969.1809, MuangNgao or Muang Ngau, 18°45'27N, 99°58'42E. Chiang Mai Province: BMNH 1938.8.7.40, BMNH 1969.1808, “Me Wang,N. Siam”, now Mae Wang. Laos. MNHN 1896.0633, “Laos”, without locality.

Oligodon cinereus (39 specimens). Cambodia. BMNH RR 1946.1.1.25 (BM 61.8.12.8), “Camboja” (holotype of Simotes cin-ereus Günther, 1864). MNHN 1970.0437–0440, Trapeang–Chan, Kompong Chhnang Province. India. State of Assam. S.Sengupta’s private collection SS057, Tezpur, Sonitpur District; S. Sengupta’s private collection SS067, Jaipur, TinsukiaDistrict. State of Bihar: ZSI 12356, “Cargola, Purnea”, Kargola, District of Purnia. Laos. MNHN 1928.0070, MNHN1999.8150–8151, “Xieng-Khouang”, Xiengkhuang, Xiengkhuang Province. Myanmar. Kayah State: MNHN 1893.0387–0388, “Monts Carin, alt. 1200–1300 m”, now Mts. Karen. Kayin State: BMNH 91.11.26.32–33, “Toungyi, S. Shan States,5,000 ft”, now Taunggyi; BNHS 845, Taunggyi. Sagaing Division: BNHS 848, Mansi. No locality: BNHS 846–847,“Burma”. – Thailand. Lampang Province: BMNH 1921.4.1.19, “Me Wang, Siam”, now Mae Wang. Kanchanaburi Prov-ince: BMNH 1921.4.1.20, Sai Yok. Vietnam. Bac Thai Province: MNHN 1897.0407–0408, “Bac Kan (Tonkin)”, now BacKan. Cao Bang Province: MNHN 1900.0330, “Région de Cao Bang (Tonkin)”, now Cao Bang. Ha Giang Province:MNHN 1997.4359–5360, Takke Sector, Bana Nature Reserve, Na Hang. Ha Noi Province: MNHN 1911.0040, Hanoi. HaTinh Province: ZFMK 81475–476, Ky Anh - Ke Go, Ky Thuong Commune. Lai Chau Province: MNHN 1892.0264, BanNam. Lao Cai Province: MNHN 1901.0373, “Bao-Ha, Tonkin”, now Bao Hà; MNHN 1948.0087, “Coc-xan, région deLao Kay”. Tuyên Quang Province: MNHN 1901.0507, “Minh Luong (Tonkin)”, now Minh Lang. Vinh Phù Province:MNHN 1938.0136, Tam Dao (syntype of Holarchus violaceus tamdaoensis Bourret, 1935); MNHN 1996.7436, Tam Dao;PSGV 487–1, Tam Dao. North Vietnam: MNHN 1908.0203, “Haut Tonkin”, Upper Tonkin; MNHN 1958.0465–0466,“Bavi et Nghan Son (Tonkin)”.


david et al. (indian oligodon)

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