On the Taxonomic Status of Uropeltis Bicatenata (Günther) (Reptilia: Serpentes: Uropeltidae)

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ON THE TAXONOMIC STATUS OF UROPELTIS BICATENATA (GÜNTHER) (REPTILIA: SERPENTES: UROPELTIDAE)

David J. Gower1, Ashok Captain2 and Sanjay S. Thakur3

1Department of Zoology, The Natural History Museum, London SW7 5BD, United Kingdom. Email: [email protected] 23/1 Boat Club Road, Pune 411001, Maharashtra, India 3666/1 Bhoi-Ali, Raviwar Peth, Talegaon Dabhade 410506, Maharashtra, India (with six text-ﬁgures) ABSTRACT.– The Indian uropeltid snake Uropeltis bicatenata (Günther, 1864) has been considered a junior synonym of Uropeltis ceylanica Cuvier, 1829, implicitly or explicitly, since Beddome (1886). Re-examination of the holotype and historical and newly collected material of U. bicatenata conﬁrms Günther’s (1864) assessment that this form represents a distinct species of Uropeltis. We resurrect U. bicatenata from the synonymy of Uropeltis ceylanica, provide a new diagnosis of the species, redescribe the holotype, document variation among all known material, and discuss aspects of the state of the taxonomy of Uropeltis.

KEY WORDS.– India, shieldtail, snakes, systematics, taxonomy, Western Ghats.

INTRODUCTION the 1800s, a flurry of taxonomic action saw sev- Uropeltidae (sensu McDiarmid et al., 1999) is a eral genera (Siluboura Gray, 1845; Coloburus family of charismatic, burrowing alethinophid- Duméril in Duméril and Duméril, 1851; Crealia ian snakes endemic to peninsular India and Sri Gray, 1858) erected and subsequently relegated Lanka (Gans, 1973, 1976, 1979; Cadle et al., to the junior synonymy of Uropeltis, and many 1990; Bossuyt et al., 2004). At first glance, the species synonymised within the genus, but lit- taxonomy of Uropeltidae appears stable, the tle of this has been reassessed in any detail in vast majority of taxonomic actions having been the intervening period. This early work was of- executed in the 1800s. For example, only seven ten conducted in a more casual framework than of the 47 currently recognised species were de- would occur today, where type specimens were scribed after 1896 (McDiarmid et al., 1999), and not designated and synonymies often listed only one of these in the last 50 years (Deraniya- without any discussion. This has resulted in an gala, 1975). However, this lack of recent taxo- intricate and often confusing taxonomic history nomic activity creates the false impression of a (see Gans, 1966; McDiarmid et al., 1999). This well-established systematic framework. In real- is exemplified by the type species of Uropeltis, ity, most species are poorly characterised on the U. ceylanica Cuvier, 1829, for which McDiar- basis of few character systems for which varia- mid et al. (1999) list many unjustified emendation has been studied across only small samples. tions, junior synonyms and varieties, as well as In addition, much of the type and important documenting that the type locality “Ceylan” is historical material has poor locality data, and is both imprecise and presumably incorrect. housed in London and Paris, with limited acces- During examination of new Uropeltis materi- sibility to modern Indian and Sri Lankan work- al from the Western Ghats of Maharashtra, India, ers. The robustness of the current taxonomy is we recognised an apparently distinct form that uneven across the family, and it is our impression keys out (using the most recent keys of Smith, that the most unsatisfactory situation relates to 1943; Rajendran, 1985; Sharma, 2003) as U. cey- the most speciose (c. 23 species) of the currently lanica but which has a distinctive colour pattern recognised genera, Uropeltis Cuvier, 1829. In and differs in other characters from the lecto- October, 2008] Taxonomic status of Uropeltis bicatenata 65 type of U. ceylanica and many other specimens anteriorly and posteriorly. Fourth upper previously referred to that species. Furthermore, labial as high as long. Caudal disk flat, the new material closely resembles two his- well defined, not much shorter than tail, torical specimens in separate collections—the terminating in a broad, horny, bicuspid type and previously only reported specimen of scale which is slightly turned upwards; U. bicatenata (Günther, 1864) in the Natural each scale composing the caudal disc is History Museum, London, UK (BMNH), and provided with one or two or three keels. a specimen wrongly identified by M. A. Smith The body is surrounded by seventeen se- (Ali, 1949) as U. rubrolineatus (Günther) in the ries of scales on the neck as well as in collections of the Bombay Natural History So- its middle; ventral shields 135; twelve ciety, Mumbai, India (BNHS). Uropeltis bicate- pairs of subcaudals. The circumference nata was described by Günther (1864) but has of the thickest (anterior) part of the body been subsequently considered a junior synonym is one-eleventh of the total length. Black of U. ceylanica (see below). Here we reassess above and below, each scale on the back U. bicatenata, resurrect it from the synonymy with a yellowish margin. A yellow band of U. ceylanica, rediagnose the species (based runs along each side of the body; it corre- on historical and new material), and redescribe sponds to the joining edges of the fourth and figure the holotype. The Muséum National and fifth outer series of scales; anteriorly d’Histoire Naturelle, Paris, France is abbrevi- it is broken up into a series of large spots, ated as MNHN. posteriorly it flanks the lower part of the tail. Lower parts entirely black. TAXONOMIC HISTORY A single example of this beautiful Günther (1863: 350) included “Silybura bi- species, 9½ inches long, was brought catenata. Dekkan. East India Company.” in a by Colonel Sykes from the Deccan. The brief report listing new species to be described specimen is a male, with the tail 8 lines in a subsequent monograph. The latter work long; it is figured on Plate XVII. Of its (Günther, 1864) presented a formal description natural size; figure H’ represents the up- of S. bicatenata (p. 191), a figure of the whole per side of the head.” body (plate XVII H) and a line drawing of head Eleven years later, Günther (1875) still recog- scalation in dorsal view (plate XVII H’). These nised S. bicatenata as a distinct species (with are reproduced here, below and in Fig. 1. no indication that any more specimens had been “SILYBURA BICATENATA. (Plate XVII. Figs. found), and included it in a key to the species of H, H’.) Silybura. Theobald (1868, 1876) also listed bi- Snout obtusely conical; rostral round- catenata as a valid species. Beddome (1886) and ed, very short, shorter than the nasals; Boulenger (1890, 1893) listed, without discus- vertical square, its front part, which sion, Silybura bicatenata under the synonymy extends between the frontals being as of their preferred names for S. ceylanica, namely large as its hind part; it is rectangular S. nilgherriensis Beddome, 1863 and S. brevis

Figure 1. Reproduction of Günther’s (1864: plate XVII. ﬁgs. H, H’) original ﬁgures of Uropeltis bicatenata (Günther). 66 Hamadryad [Vol. 33, No. 1

Figure 2. Holotype (BMNH 1946.1.16.8) of Uropeltis bicatenata (Günther). Scale in millimetres. October, 2008] Taxonomic status of Uropeltis bicatenata 67

Günther, 1862, respectively. Silybura Peters, Silybura brevis Günther, 1862: Boulenger 1861 is an unjustified emendation of Siluboura (1890: 269; 1893: 158) Gray, 1845 which is a junior synonym of Uro- Uropeltis ceylanicus Cuvier, 1829: Smith peltis Cuvier, and treatment of U. bicatenata as (1943: 80) a junior synonym of U. ceylanica has been fol- Uropeltis rubrolineatus (Günther, 1875): Ali lowed, without further comment, by all four of (1949: 376) the main subsequent comprehensive taxonomic Uropeltis (Siluboura) ceylanicus Cuvier, treatments of Uropeltidae: Smith (1943: 80), 1829: Mahendra (1984: 85–86) Gans (1966: 18—the type locality of Silybura U[ropeltis]. ceylanicus bicatenata (Günther, bicatenata is incorrectly given as “Wynad, Mal- 1864): Murthy (1990: 15) abar, 3500 feet elevation”), Mahendra (1984: Uropeltis ceylanica Cuvier, 1829: McDiar- 85–86) and McDiarmid et al. (1999). This tax- mid et al. (1999: 144) onomy has also been followed implicitly or ex- Holotype.– BMNH 1946.1.16.8 (formerly plicitly by authors of post-Smith (1943) faunal BMNH 60.3.19.1277), male, from “Dekkan” lists (e.g., Das, 1997, 2003; Murthy, 1982, 1990; or “the Deccan” according to Günther (1863) Sharma, 2003; Whitaker, 1978). Despite inter- and Günther (1864) respectively. The BMNH preting U. bicatenata as a junior synonym of catalogue and the jar label gives “Deccan (?)”, U. ceylanica, several authors continued to list which McDiarmid et al. (1999: 144) interpret as “bicatenata” as a colour variant of the senior a questioning of the locality. Other than that the synonym. This appears to have been initiated by specimen was presented by Colonel Sykes, there Smith’s (1943: 80) diagnosis of U. ceylanica, are no further collection data, although the first which includes: “with a lateral yellow stripe (bi- part of the original BMNH specimen number catenata)”. Rajendran (1985: 65) imprecisely indicates that the specimen was catalogued in quoted Smith by reporting a “bicarinate” variant 1860. The BMNH accessions register entry for of U. ceylanica. Murthy (1990: 15) upgraded 60.3.19.1277 states only “Typhlops”, presented Smith’s variant to U. ceylanicus bicatenata but by East India House. confused things further by attributing various Referred material.– BNHS S225 (female, ventral colour patterns to the form that are not collected by S. Ali, September 1948), Bhi- present in Günther’s material, and which were mashankar, Pune District, Maharashtra, India; not ascribed by Smith to any particular one of BNHS 3251 and 3252 (male and female respec- his listed varieties. tively, I. Agarwal and S. Kehimkar, 2004), close Ali (1949) reported a uropeltid specimen to Bhimashankar Wildlife Sanctuary, Pune Dis- from Bhimashankar, Maharashtra, India, that trict, Maharashtra, India; BNHS 3265 (male), M. A. Smith had identified for him as Uro- 3266 (male) and 3267 (female) Fangul Gawhan, peltis rubrolineatus (Günther, 1875). We refer Pune District, Maharashtra, India (all three col- this specimen (BNHS S225) to U. bicatenata, lected by S. Thakur, October 2003). See Table 1 which we consider a valid species. We follow for details and morphometric and meristic data, the taxonomic nomenclature of McDiarmid et and Fig. 6 for distribution of localities. al. (1999). Diagnosis.– A Uropeltis with 17 dorsal scale rows at midbody and a notably flat-to-mildly- TAXONOMY concave tail shield (distinctly not convex). Uropeltis bicatenata (Günther, 1864) Ranges of variation of seven known speci- (Figs. 1–6, Table 1) mens: total length 155–264 mm; ventral scales Silybura bicatenata Günther, 1863: Nomen nu- 130–141; subcaudal scales 8–9 (three females) dum. Günther (1863: 350) or 10–12 (four males); tail shield with 34–43 Silybura bicatenata Günther 1864: Günther keeled scales; typically 7 (uniquely 6, on one (1864: 191, Pl. XVII H, H”; 1875: 229); Theo- side only) maxillary and dentary teeth per row. bald (1868: 43; 1876: 134); Gans (1966: 18) Uropeltis bicatenata differs from type speci- Silybura nilgherriensis Beddome, 1863: Bed- mens of all other similarly scaled and shielded, dome (1886: 15) nominate species of the genus, namely Smith’s (1943: 74) group IIA and IIB species (see Table 68 Hamadryad [Vol. 33, No. 1

2) in the following ways: Uropeltis arcticeps In addition to the combinations of charac- (Günther, 1875) has fewer teeth (4–5 per row), ters listed above, U. bicatenata has a distinctive a shorter ocular and proportionately smaller colour pattern that serves to separate it from all eye, and fewer ventrals (< 130). Uropeltis cey- similarly scaled species. For example, U. rubro- lanica has fewer teeth (4 maxillary, 5 dentary), lineata and U. rubromaculata have vivid red (in a proportionately larger 3rd supralabial, a longer life) and not yellowish markings, and the lat- midline suture between prefrontals than nasals, eral stripes in the types of U. rubrolineata are a rostral that extends posterior to the nares, few- broader, occupying dorsal scale rows 1–3 or 1–4 er subcaudals (6), a proportionately broader tail (versus rows 4–5 in U. bicatenata). Unlike the shield with fewer keeled scales (28), smaller oc- unblemished belly of all known U. bicatenata, ular and eye, narrower frontal, shorter parietal, the ventrals of the type specimens of U. ceylani- and narrower ventrals. Uropeltis rubrolineata ca, U. arcticeps, U. myhendrae, U. rubrolineata (Günther, 1875) has more ventrals (>164), few- and U. rubromaculata have pale specks, blotch- er subcaudals (6–7 female, 9 male), fewer max- es and/or bands. The types of U. ceylanica, U. illary teeth (typically 5 per row), fewer keeled phipsonii, and U. rubrolineata also differ from shield scales (27–30), a proportionately shorter all known material of U. bicatenata in having a tail, and smaller eye relative to ocular scale. broad transverse ventral band in the region of Uropeltis rubromaculata (Beddome, 1867) has the anus, linking the left and right lateral stripes fewer maxillary teeth (5 per row), a longer head on the tail (although this character is known to relative to snout-vent length, and more keeled vary in some other Uropeltis, e.g., U. macrolepis scales on the tail shield (45–52). Uropeltis my- macrolepis, AC, pers. obs.). Finally, although hendrae (Beddome, 1886) and Uropeltis phip- varying in their clarity, the speckled yellow sonii (Mason, 1888) have more dentary teeth chevron markings on the dorsum of U. bicate- (typically 9 per row), and proportionately longer nata are not seen in the other species. Günther’s rostrals—so that the portion visible from above (1875) key separated bicatenata from other is clearly longer than its distance from the fron- Uropeltis species having 17 scale rows, <160 tal. Uropeltis myhendrae has more keeled scales ventrals, flat tail shield, and lacking a sharply on the tail shield (47). Uropeltis phipsonii has pointed snout, on the basis of its regular, nar- more ventrals (>143) and more supralabials (5; row, lateral yellow stripes, and this serves still one type has 4 on one side), although four BNHS to identify all the known material of this species non-type specimens (BNHS S231–234) that we except for one heavily blotched individual (see are confident can be referred to Uropeltis phip- below). sonii all have 4 (AC, pers. obs). Uropeltis bicatenata is distinct also (DJG, Remarks.– We consider many if not most spe- pers. obs.) in colour pattern and meristic and cies of Uropeltis to be poorly characterised, and morphometric characters from all types of all have therefore restricted ourselves here to com- other species (brevis Günther, 1862; short- parisons of all (arcticeps, ceylanica, phipsonii, tii Beddome, 1863; nilgherriensis Beddome, rubrolineata, rubromaculata) or all BMNH- 1863; annulata Beddome, 1886) recognised as housed (myhendrae—one, possibly two MNHN junior synonyms of U. ceylanica in the most types not examined, see McDiarmid et al., 1999) recent comprehensive treatments (Smith, 1943; type material of Smith’s Group IIA and IIB (Ta- Gans, 1966; McDiarmid et al., 1999). Detailed ble 2). Although we are confident that U. bicate- data for these types are not presented here be- nata is a distinct, clearly diagnosable, valid spe- cause a much-needed, full re-evaluation of the cies, we anticipate that the ranges of variation of taxonomy of U. ceylanica is beyond the scope individual characters will increase when larger of the present study. Some of our new observa- samples are considered. That the material newly tions lie outside Smith’s (1943: 61) diagnosis of referred to U. bicatenata encompasses the size Uropeltidae—the five (versus a constant four) range of at least one of the types of each of the supralabials in at least some U. phipsonii, and other species (compare Tables 1 and 2) lends the several instances of tooth counts beyond the some confidence to distinguishing the species reported range of 6–8 per maxilla and 8–10 per based on small samples. mandible (Tables 1, 2). October, 2008] Taxonomic status of Uropeltis bicatenata 69

Redescription of holotype.– Some morphomet- parietals not notably longer than frontal, with ric and meristic data are given in Table 1. New broadly rounded posterior margins. Two small photographs and drawings of the holotype are scales in temporal region between and in contact presented in Fig. 2 and 3, respectively. The with fourth supralabial and posterior of parietal. specimen is an adult male in fair condition, pre- Three elongate infralabials: second and third served in a single loose, flat coil. Some parts of subequal in length, notably longer than first. the body are soft, especially in the anterior half First infralabials make minimal midline contact of the specimen. In particular, the head is soft, immediately behind small, slightly protuberant and here the outermost layer of keratin of the mental. Beyond first infralabials, single pair of scales has been lost so that determining exact scales (left substantially overlapping right, ante- squamation patterns is difficult in some places. riorly) lies between mental and first single mid- The colour is somewhat faded, with the black ventral scale (latter = first ventral sensu Gower and yellow described by Günther (1864) now and Ablett, 2006). First ventral longer than wide, dark brown and pale, golden yellow. There are these proportions reversed by third ventral. no incisions into the specimen, its sex has been Inside of mouth pale, without notable pigmen- inferred here from the relatively long tail and tation. Tongue deeply forked, dorsal surfaces of high number of subcaudal scales via compari- pointed tips with some midline pigmentation. son with the dissected, referred, sexually dimor- Seven teeth in each maxillary row and seven phic BNHS material. It is unclear how Günther (left) and eight (right) in dentary rows. No signs (1864) sexed the holotype as a male. of palatal teeth. All teeth simple, pointed, back- Snout tip a little squashed, but capped by ward pointing, rather straight. Spacing of teeth short, rounded (dorsal and lateral views) ros- even in all rows. No great variation in tooth size, tral shorter (dorsal view) than gap between it but largest maxillary teeth towards middle of and anterior tip of frontal scale (= “vertical” of row, anterior teeth largest in dentary row. Den- Günther, 1864). Rostral extends back dorsally tary teeth hidden deeper in gingivae and less no further than level of nares. Ventral surface of prominent than maxillary rows. Anteriormost rostral gently notched at margin of mouth. Un- maxillary teeth approximately aligned with su- paired hexagonal frontal distinctively shaped, ture between first and second supralabials, pos- being marginally longer than broad, with short teriormost tooth just behind posterior margin of lateral (ocular) margins that are not parallel (di- third supralabial. Dentary row of similar length vergent anteriorly), and slightly concave pos- and alignment. terolateral margins. Anterolateral margins also Body subcylindrical to slightly dorsoventral- slightly concave posteriorly, and subequal in ly compressed. All head and body scales lack length to posterolateral margins. Paired nasals keels, macroscopically smooth, with iridescent (there are no separate internasals) not greatly outer keratin layer. Dorsal body scales evenly outsized by prefrontals (= “frontals” of Günther, sized around and along body. Midline ventral 1864), with subequal midline contacts between scales between mental and anal 134 (versus two pairs both being asymmetric. Small (c. 0.3 Günther’s count of 135), generally evenly sized mm diameter) subcircular external naris slightly except for gradually narrowing anterior- and countersunk within small depression, lying in posteriormost members. At midbody, ventrals anteroventral corner of undivided nasal. Four approximately 1.5 times as broad as exposed supralabials: first smallest, making shortest part of adjacent, first row of dorsals. At level of contribution to margin of mouth. Second a lit- fifth ventral, 19 dorsal scale rows, reducing to tle longer, much larger. Third (low posteriorly) 17 rows soon thereafter, maintained until at least and especially fourth much the largest. Nasal up to tenth ventral anterior to anals. At one ven- contacts supralabials 1 and 2; ocular contacts tral anterior to anals, 15 dorsal scale rows. Im- supralabials 3 and 4. Ocular large, conspicuous mediately anterior to tail shield, 12 dorsal scale (but slightly less than half ocular length), cir- rows. Paired anal scales (right overlying left) cular eye in anteroventral corner. Eye bulges in considerably larger than posteriormost ventrals dorsal view, shrivelled pupil appears subcircu- and all subcaudals. Distal margin of each anal lar. Pre-, supra- and postoculars absent. Paired overlaps two other scales in addition to anteri- 70 Hamadryad [Vol. 33, No. 1 covered 1 f 6 9 6 measured 1 31 42 20 25 31 3.2 1.2 7.3 2.8 5.9 6.3 6.1 4.5 7.8 239 10.7 22.3 BNHS 3267 region Fangul Gawhan by = width. W circumferences defined 9 7 m 26 41 18 21 27 3.1 1.1 6.8 2.3 5.3 5.7 6.5 4.6 5.4 7.1 192 10.1 13.6 14.1 here ruler; BNHS 3266 is Fangul Gawhan with shield mm = snout-vent length; ail 1 T 9 6 6 m 27 43 20 22 29 2.9 1.3 7.2 2.6 5.4 5.7 5.3 6.3 7.1 204 to 10.5 12.9 15.8 BNHS 3265 Fangul Gawhan those (Günther).

callipers, f 6 7 27 38 16 18 34 2.9 1.1 5.9 2.3 4.9 5.3 7.5 3.5 4.3 7.4 4.5 4.8 165 22.3 dial bicatenata BNHS 3252 Bhimashankar with opeltis Ur taken = length; m male; r right; SVL of 6 7 5 m 24 41 14 16 31 2.5 1.2 2.2 3.9 5.2 8.5 3.9 7.1 4.2 8.4 5.4 155 18.5 mm BNHS 3251 0.1 Bhimashankar to specimens f 7 6 32 34 10 25 27 12 22 30 3.4 1.6 7.9 3.2 6.1 6.9 6.9 7.4 4.5 8.7 264 referred BNHS S225 Measurements Bhimashankar and (*) surface. 3 9 6 6 m 31 41 22 26 15 34 3.1 1.7 7.5 6.1 6.7 4.9 6.7 7.3 248 10.7 16.5 holotype their BMNH “Deccan” the 1946.1.16.8* half for least data at over (mm) W matt are that Abbreviations: C = circumference; D distance; f female; l left; L morphometric . of parietal scale TL and W scales

(TL) W (tl) at anus keeled frontal scale ail L otal L Specimen Locality Sex L Midline D between rostral and frontal scales D between snout tip and posterior of last supralabial scale (= HL) SVL/HL Maximum L D between rostral and posterior of midline suture between parietals D between snout tip and posterior of midline suture between parietals No. keeled scales on shield Max. no. scales across shield L Min. no. scales across shield Shield Shield L W C at anus Midbody C T TL/tl tl as % of Midbody TL/W T Meristic

1. those 1 1 2 3 24 21 22 23 18 19 20 17 16 15 13 14 1 12 10 5 6 7 8 9 4 able T by with string and ruler October, 2008] Taxonomic status of Uropeltis bicatenata 71 15 17 17 17 19 1.7 3.2 2.1 3.3 4.5 3.6 2.2 7,7 7,7 4,4 3,4 2.8 1.8 2.5 1.2 137 8, 8 2 r/l 2.08 1.64 2 4 2 15 17 17 18 19 1.6 3.1 1.9 3.1 2.8 1.4 7,7 7,7 4,4 3,4 2.5 1.8 2.2 1.1 131 2 r/l 12, 12 2 15 16 17 18 19 1.9 3.2 1.9 3.3 4.2 3.4 2.3 7,7 7,7 4,4 3,3 2.5 1.9 2.2 1.1 130 2 r/l 1.48 12, 10 2 15 15 17 17 19 1.6 2.7 1.7 2.7 3.8 2.5 1.6 7,6 7,7 4,4 3,3 2.2 1.5 1.8 0.9 141 9, 8 2 r/l 1.56 1 1 15 17 17 19 19 1.5 2.6 1.7 2.7 3.7 2.2 1.4 7,7 7,6 4,4 3,3 2.1 1.4 1.9 0.9 135 1, 1 2 r/l 2.1 1.57 1 4 3 15 17 17 18 19 1.9 3.7 2.3 3.4 4.9 2.6 7,7 7,7 4,4 3,3 1.7 2.5 1.1 135 8, 8 2 r/l 1.54 2.27 15 17 17 17 19 1.8 3.5 2.3 3.4 4.8 3.5 2.3 7,7 7,8 4,4 3,3 2.8 2.2 2.3 1.1 134 2 r/l 1.52 2.09 12, 12 - - at corner of mouth of ventral scales at midbody of 1st dorsal scale row at mid prefrontal ocular scale (LO) W W W frontal scale entral scales Anal scales and nature of overlap V No. dorsal scale rows at level of one ventral scale anterior to anal No. dorsal scale rows at level of ten ven tral scales anterior to anal scale No. dorsal scale rows at midbody No. dorsal scale rows at level of 10th ventral D between nares D between eyes D between eye and naris D between snout tip and eye Head Max. (WV) Max. body (WD) WV/WD Maxillary teeth (left, right) Dentary teeth (left, right) Supralabials (left, right) Infralabials (left, right) No. dorsal scale rows at level of 5th ventral Subcaudals (left, right) W Max. L Max L Eye diameter (ED) LO/ED 48 47 46 45 44 43 30 31 32 33 34 35 36 37 38 39 40 41 42 49 25 26 27 28 29 72 Hamadryad [Vol. 33, No. 1 s Smith’ 5 27 29 37 5.3 8.6 2.2 4.2 3.6 2.7 3.6 1.3 7.5 3.2 8.7 7.2 in 398 10.9 13.2 2.77 12.9 30.9 10.7 12.7

BMNH abbreviations 1946.1.15.53 U. rubrolineata opeltis and Ur of 6 Methods 30 7.5 . BMNH jar species 1946.1.16.26† U. rubrolineata of same in 1 2 6 27 29 33 examined) 2.6 4.5 5.5 6.3 2.5 1.5 1.6 0.6 7.5 4.6 3.6 4.2 6.8 6.4 5.4 177 2.67 27.7 BMNH not 1946.1.15.63 U. rubrolineata specimens was 1 untagged 8 6 30 34 28 1.9 3.8 4.4 1.2 4.7 2.4 1.5 1.3 1.4 0.6 4.6 5.8 4.4 7.4 8.6 5.3 148 2.33 17.2 material BMNH two U. arcticeps 1946.1.16.1 ‡ type taken. MNHN 6 8 6 1.1 30 32 30 2.7 4.9 5.4 1.4 2.9 1.9 1.8 1.7 0.6 5.3 5.7 5.2 9.5 6.4 not 194 1 2.67 17.5 BMNH U. arcticeps 1946.1.16.12 which counts for , some 1 1 8 1.7 1 29 47 29 14 4.7 7.4 9.5 1.8 3.7 3.3 2.3 2.3 2.3 7.5 9.8 4.1 9.4 334 1 13.7 24.4 BMNH and 1946.1.16.9 ts U. myhendrae myhendrae U. 7 29 28 28 1.9 4.2 4.9 1.3 5.3 2.4 1.7 1.6 1.5 0.7 6.5 5.1 6.8 4.2 5.7 4.8 7.3 161 2.14 23.7 measuremen (except MNHN 39 U. ceylanica so types - all for shrivelled, data badly (mm) W specimen † IIB. morphometric of parietal scale and

TL and W IIA prefrontal (TL) W ocular scale (LO) (tl) frontal scale at anus frontal scale otal L ail L able 1. Maximum L D between rostral and posterior of midline suture between parietals D between snout tip and posterior of midline suture between parietals Midline D between rostral and frontal scales D between snout tip and posterior of last suprala bial scale (= HL) SVL/HL L W Max. L Max L Eye diameter (ED) LO/ED No. keeled scales on shield Max. no. scales across shield L W T T TL/tl tl as % of Midbody TL/W Shield Shield L Min. no. scales across shield Specimen Meristic Group

T 2. 1 4 5 6 7 8 9 1 1 18 19 20 21 22 23 24 25 26 27 28 29 17 16 13 14 15 able (1942) as for T October, 2008] Taxonomic status of Uropeltis bicatenata 73

ormost subcaudals. Twelve pairs of macroscopically smooth subcaudal scales between anus and single terminal tail scute. 17 2.9 4.8 3.4 5.7 8.7 5.7 3.4 2r/l 6, 6 3, 3 165 4, 4 9, 9 6, 5 1.68 Tail shield (= “caudal disk” of Günther, 1864) conspicuous, well defined. Flat to gently concave, oval, longer than head. Shield scales matt, minutely pitted. Some dorsal body scales anterior and anterolateral to shield (as defined here) 2 9, 9 3, 3 4, 4 8, 8 5, 5 bear low carinae but distinct from shield scales by being mostly or entirely shiny. There are 41 matt, keeled scales lying entirely or mostly within the shield. Transversely, shield is maximally six keeled scales wide; longitudinally minimal- 3 17 1.6 2.7 1.8 4.1 2.3 1.5 2r/l 167 7, 6 3, 3 4, 4 7, 8 5, 5 1.53 ly nine keeled scales long (excluding terminal scute). Anteriormost shield scale bears four sub- parallel, low carinae or keels, all other shield scales bear one, two (mostly) or three, generally more prominent carinae. Shield carinae straight, 17 1.1 2.1 1.6 2.4 3.2 2.1 1.4 1.5 2r/l 128 9, 9 3, 3 4, 4 5, 5 5, 4 longitudinal, hardened keels, with perpendicular to mildly concave posterior margin, so that hardened posterodorsal tips are square to pos- terodorsally-pointed in lateral view. Terminal scute mildly transversely convex, dorsally and 17 1.3 2.6 2.2 3.1 3.9 2.7 1.9 2r/l 127 8, 8 3, 3 4, 4 4, 5 4, 4 1.42 ventrally. Terminally it bears pair of parame- dian, posteriorly directed short spines. Upper surface of terminal scute bears few irregularly scattered, small, hardened pointed tubercles. 7 17 2.7 4.5 3.1 5.4 5.5 3.3 2r/l 140 7, 8 3, 3 4, 4 9, 8 7, 8 Background colour an even chocolate-brown 1.67 across dorsal and ventral surfaces of body, head and tail. Body scales slightly paler distally, with yellowish halo immediately inside transparent outer rim. Some notable pale golden-yellow 17 1.1 2.3 1.8 2.5 3.4 2.1 1.6 2r/l 130 6, 6 3, 3 4, 4 5, 5 4, 4 1.31 markings stand out against background. Lateral stripe begins narrowly on margin of mouth, on second supra- and infralabials. Stripe remains narrow on upper jaw until broadening behind eye, passes over most of large fourth supralabial; broadens at posterior of third infralabial. Behind corner of mouth, lateral stripe two to three scales wide. Stripe becomes broken on right (level with ninth ventral) and left (20th ventral) to form three blotches on right and two on left. Four dorsal crossbars (widely incomplete middorsally) arise from blotched region, anteriormost lies anterior to first lateral blotch. Backwards from level of 32nd ventral at corner of mouth of ventral scales at midbody (WV) of 1st dorsal scale row at midbody (WD) lateral stripe again complete along most of body W W W as regular, narrow, zigzag line (with rounded entral scales Dorsal scale rows at midbody V Anal scales and nature of overlap Subcaudal scales (left, right) Infralabials (left, right) Supralabials (left, right) Dentary teeth (left, right) D between nares D between eyes D between eye and naris D between snout tip and eye Head Max. Max. WV/WD Maxillary teeth (left, right) edges). Zigzag formed by yellowish markings 44 47 48 49 41 40 39 30 31 32 33 34 35 36 37 38 on posterodorsal edge of each scale in fourth dorsal row and posteroventral edge of each fifth 74 Hamadryad [Vol. 33, No. 1 9 36 49 13 27 9.9 9.2 6.5 5.4 8.2 2.2 4.9 4.2 3.3 3.4 1.5 2.3 2.9 5.2 4.5 368 10.2 17.1 10.1 16.5 22.3 BMNH 1946.1.15.52 U. rubromaculata 7 3 1.4 32 52 13 27 24 9.8 8.5 5.8 8.4 2.3 4.7 3.8 3.3 1.5 2.2 3.1 5.1 4.2 365 1 10.8 16.9 10.5 15.2 BMNH 1946.1.15.51 U. rubromaculata 3 30 10 52 26 15 7.7 7.2 6.5 4.1 6.5 7.7 1.8 9.4 3.5 2.3 2.8 1.3 2.2 2.3 3.8 5.9 8.6 256 14.2 17.1 BMNH 1946.1.15.84 U. rubromaculata 6 32 50 27 9.3 8.1 9.5 5.3 8.2 9.9 2.2 4.2 3.8 3.1 3.3 1.5 2.2 2.7 4.9 5.4 349 17.5 12.2 18.7 18.7 10.8 BMNH 1946.1.15.83 U. rubromaculata 9 6 9 5 2 35 45 12 27 8.6 8.1 9.7 4.7 3.6 2.9 3.3 1.5 2.2 2.7 4.6 4.3 9.9 343 15.1 14.6 23.5 BMNH 1946.1.15.82 U. rubromaculata 5 5 26 30 30 6.8 4.8 8.5 2.4 4.6 0.9 6.8 2.9 2.2 1.2 1.9 0.8 1.5 2.9 5.6 8.3 218 10.7 2.38 12.3 17.7 BMNH U. phipsonii 1946.1.16.33-34‡ 8 6 1 33 34 31 7.1 6.3 5.5 2.8 5.3 1.3 8.5 3.5 2.7 1.7 2.2 2.2 1.7 3.3 5.4 8.5 277 12.3 14.9 18.6 BMNH U. phipsonii 1946.1.16.33-34‡ W of parietal scale TL W prefrontal (TL) W ocular scale (LO) (tl) at anus frontal scale frontal scale otal L ail L W Specimen TL/W Shield Shield L Min. no. scales across shield Max. no. scales across shield L No. keeled scales on shield Maximum L D between rostral and posterior of midline suture between parietals D between snout tip and posterior of midline suture between parietals Midline D between rostral and frontal scales D between snout tip and posterior of last supralabial scale (= HL) SVL/HL L W Max. L Max L Eye diameter (ED) LO/ED D between nares D between eyes T T TL/tl tl as % of Midbody contd. 1 1 9 4 5 6 7 8 1 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 able 2. T October, 2008] Taxonomic status of Uropeltis bicatenata 75

row scale. Lateral stripes again broken brieﬂy about ten ventral scales in front of anus. Stripes remain narrow zigzags until three ventrals in 17 3.3 5.5 8.2 5.6 3.6 4,4 5, 5 7, 7 3, 3 133 8, 8 2 r/l 1.56 front of anus, here extending onto tail as broad continuous stripes (about two scales wide). Left and right stripes converge a little toward end of slightly tapered tail, barely crossing lateral- most margins of penultimate subcaudal scales 17 3.3 5.7 9.4 5.7 3.4 4,4 5, 5 6, 6 3, 3 135 9, 8 2 r/l 1.68 where stripes terminate one scale prior to terminal scute. Anal scales with off-white posterior margin, just inside transparent outer edge. No transverse bands extending onto ventral surface of tail from lateral stripes.

4 6 Posterior to irregular and incomplete anterior 17 2.4 3.9 2.7 4,4 5, 5 6, 7 3, 3 127 2 r/l 1.44 10, 10 cross-bars, majority of dorsal surface of body marked with delicate, largely regular pattern of forward-pointing V-shapes (chevrons) spaced one dorsal scale row apart. Each V formed by yellowish blotches on distal tips of midline 17 3.1 5.1 7.6 5.2 3.1 4,4 5, 5 7, 7 3, 3 131 2 r/l 1.68 10, 9 (ninth) dorsal scale row and posteromedial margin of next two (seventh and eight) scale rows, although even here, scales have transparent distalmost edge. Dorsal V pattern continues up to level of anus with varying completeness. 3 5 5 17 7.3 3.4 4,4 5, 5 7, 7 3, 3 133 9, 8 2 r/l 1.47 Between anus and tail shield dorsal surface un- patterned, uniform brown. Dorsal pattern Vs often incompletely formed, in particular the pale spot on distal tip of scale row nine not always contacting the generally more continuous

3 5 patches on rows seven and eight, particularly 17 2.1 3.6 2.5 5,4 7, 7 9, 9 3, 3 144 1, 12 2 r/l 1.44 1 further posteriorly, so that pattern (\ /) can also be described as herringbone- or tyre-tread-like. Tail-shield scales uniform pale brown except for translucent tips of carinae. Terminal scute with

1 midline whitish stripe on posterior half, two 17 3.6 5.2 4.1 2.7 5,5 2.3 8, 8 9, 9 4, 4 147 1, 1 2 r/l 1.52 pointed tips also pale. 1 Additional information from referred specimens.– Some meristic and morphometric data are presented in Table 1. Line drawings of head scalation and photographs of some of the referred material are shown in Fig. 4 and 5, respectively. The referred material comprises six additional specimens (three males, three females) rang- ing from 155 to 264 mm total length (TL), thus encompassing the holotype (male, 248 mm). Sexual dimorphism in tail length (4.5% of TL at corner of mouth of ventral scales at midbody of 1st dorsal scale row at in females, 5.4–7.1% in males) and number of W W W subcaudal scales (females 8–9, males 10–12) is entral scales D between snout tip and eye Head Max. (WV) Max. midbody (WD) WV/WD Maxillary teeth (left, right) Dentary teeth (left, right) Supralabials (left, right) Infralabials (left, right) Dorsal scale rows at midbody V Anal scales and nature of overlap Subcaudal scales (left, right) D between eye and naris pronounced and non-overlapping. No other no- 33 34 35 36 37 38 39 40 41 44 47 48 49 32 table dimorphism was observed. 76 Hamadryad [Vol. 33, No. 1

Head scalation patterns in referred material S225, but black (as originally described for hol- generally match holotype. Portion of rostral vis- otype by Günther, 1864) to dark blue-black in all ible dorsally always shorter than its distance more recently preserved specimens. Paler mark- from frontal, only in BNHS 3251 does it extend ings always shades of yellow (never red), more as far back as level with posterior margin of lemon-yellow anteriorly and more orange-yel- nares (Fig. 4B). Frontal generally with concave low posteriorly in larger Fangul Gawhan indi- antero- and posterolateral margins, but extent viduals. Body scales resemble those of holotype varies—anterolateral margins strongly concave in consistently having transparent distal margins in BNHS 3265 and 3267, posterolateral margins lying beyond thin, translucent, yellowish halo. straight to mildly convex in smallest specimen Apart from translucent distal margins, ventrals (BNHS 3251). Midline nasal and prefrontal uniformly darkly coloured in all specimens. sutures straight only in BNHS 3251 and 3267. Scales under lower jaw uniform in all speci- Supra- and infralabials constant in number and mens except BNHS 3265 and 3266, which have relative sizes. Under front of lower jaw, first pair small, pale-yellowish spots on each of parame- of infralabials make broad midline contact be- dian scales contacting first and second ventrals. hind mental only in BNHS 3267. Only in BNHS BNHS 3265 has an additional small spot nearby 3251 do first pair of chin scales behind first in- on second ventral. fralabials not make broad, overlapping (some Lateral body stripes and dorsal chevron left over right, some vice versa) contact, so that markings constant and distinctive in all referred first ventral contacts mental (Fig. 4C). Ocular specimens except for notable variant BNHS and eye consistently large and maintain fairly 3267 (Fig. 5e, f), which has extensive lateral constant relative proportions. Parietals generally blotches and faint dorsal speckles only occasion- short, rounded, longest in BNHS 3267. BNHS ally coming close to forming Vs. BNHS 3267 3251 has asymmetric pair of small scales im- is interpreted as a rare exception—it is the only mediately between back of irregularly sutured unusually marked individual seen among tens of parietals. Left side of BNHS S225 has three (not uncollected live animals at Bhimishankar (AC, two) small scales between and contacting fourth SST, pers. obs.) and c. 10 animals seen at Fangul supralabial and parietal (Fig. 4A). Teeth almost Gawhan (SST, pers. obs.). Indeed, it was col- constant in number. Pupil in preserved speci- lected especially because of its unusual colour mens generally an irregular blob, most circular pattern. in BNHS 3267. Anal scales always paired, right Dorsal chevrons vary in completeness, best overlying left. Subcaudals always macroscopi- defined in holotype, in referred specimens spots cally smooth. Tail shield similarly proportioned on distal ends of midline (ninth) dorsal scale in all specimens, with 34 to 43 keeled, matt row small or absent, so that pattern is more her- scales, most of which are bicarinate, a few uni- ringbone (\ /) or tyre-tread than chevron-like, carinate, fewer tricarinate, and none tetracari- and arms of Vs or /s are sometimes incomplete. nate. A single tetracarinate, glossy scale lies just Midline dorsal scales often with yellow marks anterior to shield of BNHS 3251. Largest keels on posterolateral margins instead of posterior resemble closely those of holotype in being tip, so that V or \ / pattern is more U-U like. sharply pointed with perpendicular to concave Dorsal markings extend onto tail but fade before posterior margins, so that shield as a whole is shield, this varying from two (BNHS 3265) to rough. Terminal scute of BNHS S225 broken; seven (BNHS 3266) scales anterior to first shield that of 3251 lacks right posterior spine. Length scale. Lateral stripe extends forwards generally of spines varies (long in e.g., BNHS 3265, onto second supralabial and posterior of third 3266). Most specimens have small additional, infralabial, but may continue further forwards less acutely pointed lateral and/or posteromedi- as a thin line on lips, most notably in BNHS al spines. Terminal scute spines of BNHS 3265 3252 where it extends onto first supra- and in- have small additional lateral cusps, BNHS 3265 fralabials. First break in stripes behind head has a small posteromedial cusp. ranges from level with ninth (BNHS 3267) to Background body colour not chocolate brown fifteenth (BNHS 3266) ventral, though in BNHS in any referred specimen. Grey-brown in BNHS 3251 it remains complete. Anteriorly, stripe nar- October, 2008] Taxonomic status of Uropeltis bicatenata 77

Figure 3. Outline scale drawings of head of holo- Figure 4. Outline scale drawings of heads of referred type (BMNH 1946.1.16.8) of Uropeltis bicatenata specimens of Uropeltis bicatenata (Günther). Up- (Günther) in lateral, ventral and dorsal views. For per figure: dorsal view of BNHS S255 (female, TL = scale see Fig. 2. 264); Middle, lower figures: anterodorsolateral view of head and ventral view of lower jaw of BNHS 3251 (male, TL = 155 mm). rows after final main break between points level with ventrals 29 to 39. Anterior to anus, stripe broadens at a point between one and four ventral 3251, BNHS 3252) in the smallest specimens to scales further forwards than in holotype (i.e., a thin faint arc (BNHS 3267) or clearer but still three to six scales anterior to anal). Stripe on tail narrow (BNHS 3265) band in larger animals. As generally two scales wide, three in BNHS 3265 in the holotype, hardened spines on distal edge and BNHS 3267, never encroaching onto sub- of terminal scute, and short midline stripe are caudals. In all referred specimens except BNHS yellow (larger animals) to off-white in all re- 3267, lateral stripes generally thin, complete, ferred specimens. In life, the recently collected regular, and zigzagged. Except in BNHS 3267, referred specimens were blackish with vibrant posteriorly the stripes are never broken for the golden/orange yellow markings. From photo- entire length of one scale, in BNHS 3252 they graphs taken in life, the pupil is circular. are unbroken. Distribution, ecology and conservation.– Uro- Whitish line toward distal margins of anal peltis bicatenata is known with certainty from scales of holotype is yellow in referred speci- only two localities, Bhimashankar and Fangul mens, varying from faint and diffuse (BNHS Gawhan (locally known as Fangli) both in Pune 78 Hamadryad [Vol. 33, No. 1

Figure 5. Habitat at Fangul Gawhan, and variation in colour pattern in referred specimens of Uropeltis bicatenata (Günther) from this locality: a), b) typical colour pattern for species, as seen in BNHS 3265; c) forest ﬂoor in summer (May, dry season); d) hill seen in May; forest in which U. bicatenata have been found is seen as a thin green horizon towards lower part of hill; e), f) exceptional colour variant BNHS 3267. October, 2008] Taxonomic status of Uropeltis bicatenata 79

Figure 6. Map showing known (Bhimashankar, Fangul Gawhan) and possible (Torna Fort) localities for Uro- peltis bicatenata (Günther).

District, Maharashtra (Fig. 6), separated by c. The specimens from Fangul Gawhan (Fig. 30 km. Further fieldwork is required to ascer- 5c, d) were found under a log in secondary tain whether the species occurs at intervening semi-evergreen forest: 10–15 m tall with 70% and surrounding localities, and at other altitudes canopy (19°15’11”N, 73°42’27”E, 803 m asl), and habitats. The type locality of “Deccan” is a short distance from the village (19°15’55”N, imprecise but can be considered to include the 73°43’02”E, 740 m asl). Vegetation in the im- two known localities, which lie in the higher al- mediate vicinity of the collection site included titudes of the Ghats at this part of their range, at Mallotus philippensis, Albizia amara, Ficus the western edge of the Deccan plateau. In addi- racemosa, Atalantia racemosa, Carvia callosa, tion to the two known localities, a superficially Olea dioica, Mangifera indica, Pittosporum similar (in colour and pholidosis), potentially dasycaulon; Piper sp. and Memycylon umbel- conspecific form has been seen (but not col- latum. lected) at the more southerly locality of Torna In addition to the recently collected material, (c. 40 km south-west of Pune), Pune District, several other sightings of Uropeltis bicatenata Maharashtra (Fig. 6). have been made inside the protected area (130 km2) of Bhimashankar Wildlife Sanctuary (cen- 80 Hamadryad [Vol. 33, No. 1 trally 19°14’N, 73°35’E; 650–1,140 m asl) (AC, need of substantial revision. This should ideally SST, pers. obs.; I. Agarwal, S. Kehimkar, pers. be based on investigation of a wider range of comm.). The species can be seen occasionally characters for type, historical, and newly col- on roads (including roadkills), and in and near lected material. The latter is needed in many waste heaps, but little is known about toler- cases to establish distributions because locality ance to habitat disturbance. The site at Fangul data of type and referred material is often im- Gawhan is not officially protected, but the For- precise. Newly collected material would also est Department has attempted to get local peo- enable taxonomic hypotheses to be more readily ple to prevent further degradation of the forest. tested with DNA sequence data. Although there is no indication that the species Previously, the taxonomy of uropeltids has is currently threatened, we suggest that it is rec- been founded on a small set of characters, most- ognised as data deficient based on IUCN criteria ly colour, size, number of ventral and subcaudal pending further, especially distributional data. scales, the relative size of the eye and ventrals, Uropeltis bicatenata is closer in appearance the form of the tail tip, size of the rostral scale, and ground colour to U. m. macrolepis, which and snout shape. Some of these have been dealt has 15 midbody scale rows, than it is to U. phip- with in a confusing manner. For example, terms sonii, which like U. bicatenata, has 17 scale previously used to describe snout shape (some rows. All three species as (as presently under- of which are used in diagnoses and keys) in- stood) are found in the Bombay Ghats/ Hills. clude obtusely pointed, acutely pointed, pointed More work is required to determine if any of and rounded. Sometimes different terms have these species are sympatric. been applied to the same species—the snout Suggested common name.– We prefer “Bicate- of U. macrolepis has been described as both nate Uropeltis” or “Two-chained Uropeltis”. “rounded” (Smith, 1943) and obtusely conical” We assume bicatenata to stem from the Latin (Günther, 1864). Characters describing the size catena, meaning chain—this perhaps in refer- of the eye relative to the ocular, and the ventral ence to the superficially chain-link-like lateral scale width relative to adjacent dorsal rows have stripes that are formed by rounded-zigzag lines, been imprecise, with little or no raw data or ex- or alternatively to the arms of the Vs on the dor- act proportions presented. Our study has high- sal surface of the body, although this seems less lighted the potential utility of several previously likely given that these markings are more com- un- or underexploited character systems for plete Vs and less herringbone (\ /) like in the Uropeltis systematics. Although further work only specimen available to Günther. Uropeltis is required to further test this potential, these translates as shield-tail, from the Greek and characters include tooth counts, the number of Latin pelte for small shield, and the Greek oura keeled shield scales, and morphometrics (Tables for tail. However, “shieldtail” is widely used to 1, 2). Investigating new characters as part of fu- refer to uropeltids as a whole, rather than Uro- ture work will be an important component of the peltis (e.g., Whitaker and Captain, 2004; Das much needed revision of uropeltid taxonomy. and de Silva, 2005), and we suggest it is best avoided as a common name for members of the ACKNOWLEDGEMENTS genus. For assistance with access to material, advice, critical discussion, information and encourage- DISCUSSION ment we are grateful to Ishan Agarwal, Varad Uropeltis bicatenata is a valid species. That it re- Giri, Sameer Kehimkar, Colin McCarthy, mained hidden in the synonymy of U. ceylanica Romulus Whitaker and Mark Wilkinson. We for more than 100 years, with the second known also thank the Director of BNHS (Asad Rah- specimen being referred to a third species (U. mani) for support and access to collections and rubrolineata) by one of the foremost workers in facilities. Ivan Ineich (MNHN) kindly loaned the field (M. A. Smith) illustrates the inadequate the holotype of U. ceylanica to London. Harry state of the taxonomy of uropeltids, especially Taylor (BMNH) photographed the holotype Uropeltis. It is our belief that Uropeltis is taxo- of Uropeltis bicatenata. Sujoy Chaudhuri of nomically extremely poorly understood, and in ECOLLAGE made the excellent map- thanks October, 2008] Taxonomic status of Uropeltis bicatenata 81

Suj. Reviews by Patrick David and Van Wallach 22:32–45. improved an earlier draft of this paper. DJG was _____. 2003. Growth of knowledge on the reptiles able to visit BNHS and Pune thanks to the NHM of India, with an introduction to systematics, Zoology Research Fund, and an India-UK Sci- taxonomy and nomenclature. Journal of the ence Networks award from the Indian Depart- Bombay Natural History Society 100:446– ment of Science and Technology and The Royal 501. Society, the latter for which he acknowledges _____ & A. DE SILVA. 2005. A photographic guide to also his host at the National Centre for Cell Sci- snakes and other reptiles of Sri Lanka. New ence (Pune), Yogesh Shouche. AC & SST thank Holland Publishers (UK) Ltd., London. 144 Baburao Kashinath Korde of Fangul Gawhan pp. for local support. We thank Aparna Watve for DERENIYAGALA, P. E. P. 1975. A new fossorial snake identifying plant species around the site of col- of the genus Rhinophis Hemprich. Spolia lection in Fangul Gawhan. We applaud Prakash Zeylanica 33:535–536. Thosre (Director, Social Forestry, Pune divi- GANS, C. 1966. Liste der rezenten Amphibien und sion, Maharashtra) for his efforts in protecting Reptilien. Uropeltidae. Das Tierreich 84:1– the habitat around Fangul Gawhan. 29. _____. 1973. Uropeltid snakes—survivors in a LITERATURE CITED changing world. Endeavour 32:60–65. ALI, S. 1949. Extension of range of the earth snake _____. 1976. Aspects of the biology of uropeltid Uropeltis rubrolineatus (Günther). Journal of snakes. In: Morphology and biology of rep- the Bombay Natural History Society 48:376. tiles. pp:191–204, pl.:1–4. A. d’A. Bellairs BEDDOME, R. H. 1886. An account of the earth & C. B. Cox (Eds). Linnean Society Sympo- snakes of the peninsula of India and Ceylon. sium Series Number 3. Academic Press, New Annals and Magazine of Natural History, Se- York. ries 5, 17:3–33. _____. 1979. A subterranean snake with a funny BOSSUYT, F., N. BEENAERTS, M. MEEGASKUMBURA, tail. Natural History 88:70–75. D. J. GOWER, R. PETHIYAGODA, K. ROELANTS, GOWER, D. J. & J. D. ABLETT. 2006. Counting ven- A. MANNAERT, M. WILKINSON, M. M. BAHIR, K. tral scales in Asian anilioid snakes. Herpeto- MANAMENDRA-ARACHCHI, O. V. OOMMEN, P. K. logical Journal 16:259–263. L. NG, C. J. SCHNEIDER & M. C. MILKINKOVITCH. GÜNTHER, A. C. L. 1863. Third account of new spe- 2004. Local endemism within the Western cies of snakes in the collection of the British Ghats–Sri Lanka biodiversity hotspot. Sci- Museum. Annals and Magazine of Natural ence 306:479–481. History, Series 3, 12:348–365. BOULENGER, G. A. 1890. The fauna of British In- _____. 1864. The reptiles of British India. Ray So- dia, including Ceylon and Burma. Reptilia ciety, London. xxvii + 433 pp. and Batrachia. Taylor & Francis, London. _____. 1875. Second report on collections of In- 541 pp. dian reptiles obtained by the British Museum. _____. 1893. Catalogue of the snakes in the British Proceedings of the Zoological Society of Museum (Natural History). Volume I: Con- London 1875:224–234. taining the families Typhlopidae, Glauconi- MAHENDRA, B. C. 1984. Handbook of the snakes of idae, Boidae, Ilysiidae, Uropeltidae, Xeno- India, Ceylon, Burma, Bangladesh, and Paki- peltidae, and Colubridae Aglyphae, Taylor & stan. Annals of Zoology 22B:1–412. Francis, London. xiii + 448 pp. McDIARMID, R. W., J. A. CAMPBELL & T. A. TOURÉ. 1999. CADLE, J. E., H. C. DESSAUER, C. GANS & D. F. GART- Snake species of the World, Volume 1. The SIDE. 1990. Phylogenetic relationships and Herpetologists’ League, Washington, D.C. molecular evolution in uropeltid snakes (Ser- MURTHY, T. S. N. 1982. An illustrated ﬁeld guide to pentes: Uropeltidae): allozymes and albumin the rough tailed snakes of India. The Snake immunology. Biological Journal of the Lin- 14:119–135. nean Society 40:293–320. _____. 1990. Illustrated guide to the snakes of the DAS, I. 1997. Checklist of the reptiles of India Western Ghats, India. Records of the Zoo- with English common names. Hamadryad logical Survey of India, Occasional Paper 82 Hamadryad [Vol. 33, No. 1

114:1–69. Baptist Mission Press, Calcutta. 91 pp. RAJENDRAN, M. V. 1985. Studies in uropeltid _____. 1876. Descriptive catalogue of the reptiles snakes. Madurai Kamaraj University, Ma- of British India. Thacker, Spink & Co., Cal- durai. v + 132 pp. cutta. x + 238 + xxxviii + xiii pp; 5 pl. SHARMA, R. C. 2003. Handbook Indian snakes. WHITAKER, R. 1978. Common Indian snakes. A Zoological Survey of India, Kolkata. xx + ﬁeld guide. Macmillan India Ltd., New Del- 292 pp + 69 plates. hi. xiv + 154 pp. SMITH, M. A. 1943. The fauna of British India. Rep- _____ & A. CAPTAIN. 2004. Snakes of India, the ﬁeld tiles and Amphibia Vol. III: Serpentes. Taylor guide. Draco Books, Chennai. xiv + 481 pp. and Francis, London. 583 pp. THEOBALD, W. 1868. Catalogue of reptiles in the Received: 23 May 2007. Museum of the Asiatic Society of Bengal. Accepted: 12 November 2007.