Aspidura Ravanai Sp

Zootaxa 4347 (2): 275–292 ISSN 1175-5326 (print edition) http://www.mapress.com/j/zt/ Article ZOOTAXA Copyright © 2017 Magnolia Press ISSN 1175-5334 (online edition) https://doi.org/10.11646/zootaxa.4347.2.4 http://zoobank.org/urn:lsid:zoobank.org:pub:1968CE67-0C8F-44C9-A12E-BD2196C9710C

A new species of Aspidura Wagler, 1830 (Squamata: Colubridae: Natricinae) from Sri Pada sanctuary (Peak Wilderness), Sri Lanka

1,4 1 L. J. MENDIS WICKRAMASINGHE , DULAN RANGA VIDANAPATHIRANA , H. K. DUSHANTHA KANDAMBI2, R. ALEXANDER PYRON3 & NETHU WICKRAMASINGHE1 1Herpetological Foundation of Sri Lanka, 31/5, Alwis Town, Hendala, Wattala, Sri Lanka 2Dangolla, Uda Rambukpitiya, Nawalapitiya, Sri Lanka 3Dept. of Biological Sciences, The George Washington University, 2029 G. St., NW, Washington, DC 20052, United States 4Corresponding author. E-mail: [email protected] or [email protected]

Abstract

We describe a new species, Aspidura ravanai sp. nov., representing the eighth species of the genus known from Sri Lanka. The new species is readily distinguished from all other congeners by its colour pattern, scalation, and genetic distinctiveness. Our molecular results indicated that Aspidura began diversifying in the Miocene approximately 18 Ma, and A. ravanai sp. nov. diverged from its likely sister lineage A. trachyprocta at least 6.2 Ma. The species is currently known only from the type locality, on the western slopes of Sri Pada Peak in the central highlands of Sri Lanka.

Key words: Aspidura trachyprocta, divergence dating, Ravana, rough-side snakes, new taxa, Sabaragamuwa province, South Asia, systematics

Introduction

The snake genus Aspidura Wagler, 1830 is endemic to Sri Lanka, and is currently represented by seven described species (Pyron et al. 2013; Wallach et al. 2014). They are commonly known as Rough-side snakes due to the presence of keeled scales on the ischiadic region. Rough-side snakes are non-venomous, burrowing species with snout-vent lengths ranging from 170 mm to 736 mm (de Silva 1980), and a subconical or semicircular head with an indistinct neck. Due to the presence of a pointed snout, they are well adapted to a burrowing and sub-terrestrial or fossorial lifestyle. They are further characterized in scalation by the absence of a loreal, single internasal, and an undivided anal (Boulenger 1890; Wall 1921; Smith 1943; Deraniyagala 1955; de Silva 1980; Gans & Fetcho 1982). The genus was erected by Wagler (1830) with Scytale brachyorrhos Boie, 1827, from Ceylon, as type species by monotypy. This species, now known as Aspidura brachyorrhos (Boie, 1827), is widely distributed in the island and is found in a variety of habitats in both the wet and the dry zones (Gans & Fetcho 1982). The species Aspidura copei Günther, 1864 is known only from seven specimens, including the holotype with missing locality data (Gans & Fetcho 1982). The taxon Aspidura guentheri Ferguson, 1876 is the smallest member of the genus, commonly found in the lowland wet zone of Sri Lanka. The species A. deraniyagalae Gans & Fetcho, 1982 is the most recently described species originally placed this genus. Another taxon, Aspidura drummondhayi Boulenger, 1904, was described from Hopewell Estate, Balangoda, from approximately 1066 to 1280 m above sea level (Boulenger 1904). The species Aspidura trachyprocta Cope, 1860 is a common snake of the Central hills distributed at ~1500 m a.s.l. Despite the genus being confined to Sri Lanka, Laidlaw (1902) erroneously reported this species from Mal (Maldives), which was subsequently repeated by several authors (Smith 1943; Deraniyagala 1955; de Silva 1972, 1980). Later, Taylor (1950) and Gans & Fetcho (1982) cast doubt on this record, since the species was otherwise undocumented in the Maldives and was unlikely to have occurred in such environmental conditions. This was later confirmed after examining the material, which was in fact a specimen of a hatchling which most likely would have been transported in trade to Mal from Ceylon (Gans 1989). Finally, the seventh member, Aspidura ceylonensis

Accepted by T. Nguyen: 31 Aug. 2017; published: 13 Nov. 2017 275 (Günther, 1858), resulted from a recent molecular analysis of Sri Lankan serpents, as a result of which the monotypic genus Haplocercus Günther, 1858 was synonymized with Aspidura Wagler, 1830 (see Pyron et al. 2013). According to the latest list of threatened taxa in Sri Lanka, five species of Aspidura are considered threatened, of which A. deraniyagalae is “Critically Endangered, A. ceylonensis, A. drummondhayi, and A. trachyprocta “Endangered, and A. brachyorrhos “Vulnerable. From the remaining two species A. guentheri is considered “Near threatened, and A. copei “Data Deficient (Wickramasinghe 2012). Highlighting the necessity of additional knowledge for these taxa, these assessments have not yet been integrated into the IUCN Red List of Threatened Species (www.iucn.org; last accessed on 31 July 2017, version 2016-3), or recent global assessments of evolutionary variation in threat risk (Tonini et al. 2016). Molecular evidence of an undescribed species was presented by Pyron et al. (2013), and the conservation status of this population is also unknown. Here, we describe this population as a new species of Rough-side snake, Aspidura ravanai sp. nov., from the Peak Wilderness (a part of the Central Highlands and Knuckles World Heritage Site) of Sri Lanka. The species was collected as a result of field surveys carried out in this region documenting the diversity of its herpetofauna. It is clearly distinct from all congeners based on multiple aspects of size and scalation, and a conspicuous yellow stripe on the ventrolateral surface of the head and tail, occasionally running the length of the body. Molecular divergence- time estimates suggest that the genus Aspidura began diversifying at least 18 Ma, and that A. ravanai diverged from its likely sister-taxon A. trachyprocta at least 6.2 Ma.

Materials and methods

We collected 3 specimens of an unidentified Aspidura species from Sri Pada Sanctuary (Figure 1). The specimens were photographed in life (Canon EOS 500D, Canon 100mm F/2.8L IS USM Macro Lens, Canon 270EX Flash), and the entire series was fixed and preserved in 85% ethyl alcohol and subsequently transferred to 75% ethyl alcohol after one day. Sixteen measurements were taken with a Mitutoyo digital vernier caliper (to the nearest 0.1 millimeter), each measurement was taken three times and their means recorded, and snout-vent length (SVL) was measured to the nearest 1 mm, using a flexible measuring tape. Nomenclature of external anatomy abbreviated in the text and 16 external measurements taken are listed herein: Cloacal width (CL, maximum width of anal scale), distance between back of eyes (DBE, maximum distance between posterior most point of eyes), distance between front of eyes (DFE, maximum distance between anterior most point of eyes), eye width (EW, measured from anterior to posterior of eye), head depth (HD, maximum height of head, across eye), head length (HL, distance between posterior edge of mandible and tip of snout), head width (HW, maximum width of head), internarial distance (IN, distance between nares), interorbital distance (IO, shortest distance between orbits), nostril to eye (distance between anterior most point of eyes and nostrils), nostril width (NW, maximum horizontal width of nostril), snout to eye distance (SE, distance between tip of snout and anterior most point of eyes), snout to gape end (SG, distance between snout tip to the posterior most point of the last supralabial), snout to nostril (SN, distance between tip of snout and anterior point of nostril), snout–vent length (SVL, from tip of snout to vent), tail length (TaL, from vent to tip of the tail), total length (TL, from snout to tip of the tail). Specimens were sexed by examining the presence of hemipenis for males, and from the nature of keels on coastal scales, which are rough in males and comparatively smooth in females. Observations were made through a Leica Wild M50 microscope. Collection localities were recorded with a Garmin E-trex venture GPS. The new material was compared with preserved specimens of other species of Sri Lankan Aspidura in the National Museum of Natural History, Sri Lanka (NMSL-NH), Natural History Museum, London (BMNH), and with data from literature (de Silva 1980; Gans & Fetcho 1982). Of the new type material discussed in this paper the holotype is deposited in the NMSL-NH, and the paratypes in the National Wildlife Research and Training Center, Department of Wildlife Conservation, Girithale, Sri Lanka (DWC). Molecular analyses. We took a comprehensive approach to estimating molecular divergence of these new populations relative to Natricinae as a whole. Figueroa et al. (2016) presented a species-level dataset consisting of all available molecular sequences for snakes from 10 genes, including 110 natricine species (of 230 total), and 6 from their sister-group Sibynophiinae (of 11). This includes the entire data matrix of Pyron et al. (2013), with all sampled Aspidura populations. We analyzed the data in MrBayes version 3.2.6 (Ronquist et al. 2012). We excised

276 · Zootaxa 4347 (2) © 2017 Magnolia Press WICKRAMASINGHE ET AL. the species from Natricinae+Sibynophiinae, and applied the same partitioning strategy by gene and codon position with the GTR+G+I model. We used the fossilized birth-death tree prior with a sampling fraction of 0.48 (116/241) and default priors on the other parameters. The April version of the Timetree databaset (http://www.timetree.org/) registers 13 studies inferring an age for this clade of 41–52Ma, with an estimate of 48Ma. Thus, we placed a uniform prior of 41–52 Ma on the root age. From the 39.2% average pairwise divergence of the matrix divided by the expected root-age of 48Ma, we derived a mean clock-rate of 0.00817 subst./site/My, and a standard deviation of e0.00817. We ran the analysis for 66 million generations, thinned every 10,000th with 25% discarded as burnin, for a total of 50 million sampled generations and 5,000 total samples. We assessed stationarity when ESS values exceeded 100 for all parameters (see Ronquist et al. 2012). This allows us to estimate the timing of divergence within Aspidura, and between A. ravanai and A. trachyprocta (Figure 2).

Systematics

Aspidura ravanai sp. nov.

Aspidura sp. Pyron et al. 2013

Holotype. NMSL-NH 2017.19.01 (Figure 3), adult male, 256 mm SVL, from Sri Pada upper region, Ratnapura district, Sabaragamuwa province, Sri Lanka, (06°48'28''N, 080°29'23''E, elevation 1,680 m a.s.l.): Coll. L.J.M. Wickramasinghe, D.R. Vidanapathirana, S.C. Ariyarathne, A.W.A. Chanaka, D. Kandambi and M.D.G. Rajeev on 05 February 2010. Paratypes. DWC 2017.05.01 (Figure 4A), adult female 414 mm SVL; DWC 2017.05.02 (Figure 4B), adult male 218 mm SVL. Collection data as for the holotype. Diagnosis. Maximum adult SVL 218–414 mm; snout to eye distance 0.3 times the head length (SE/HL); prefrontals touching eye; preocular small, does not touch supraocular; postoculars 2, lower one larger than the upper; temporal 1+2/1+2; supralabials 6/6, 4th touching eye; infralabials 6/6, first pair in contact, progressively increasing in size from 1st to 6th; anterior chin shields 2, large, touching 1–4 infralabials; posterior chin shields 2, anterior half in contact while the posterior half separated by 1st ventral; ventrals 132–139; subcaudals 20–29; dorsal scale rows 15–15–15; ischiadic, anal and tail base regions in the adults of both sexes keeled, with 1–3 peaks on each scale, more pronounced in males, spinose tubercles absent (Figures 5); entire dorsum jet black; bright yellow line from base of tail to tail end; venter black with bright yellow blotching all over. Description of holotype. Adult male; SVL 256 mm; TaL 37.2 mm; TL 293.2 mm; TaL/TL 0.13; body elongate and cylindrical; head short (SVL/HL 20), elliptical, indistinct from thick neck; snout long, narrowing anteriorly, pointed in dorsal aspect, snout to nostril distance about 2.3 (EW/SN) times as long as nostril width; nasal divided; small, triangular nostril, touching divided nasal and first supralabial, not touching rostral; eye larger than horizontal diameter of nostril, eye width about 0.3 (EW/SE) times the distance between snout to eye, round pupil; snout to eye distance 0.3 times head length (SE/HL); tail short (TaL/SVL 0.15), robust at its base, tapering progressively to a single point. Head scalation. Head scalation including 1 internasals, 2 prefrontals, 2 supraoculars, 1 frontal, and 2 parietals. Rostral small, convex, wider than long and truncate in dorsal aspect, projecting ventrally. Nasal vertically divided by a groove above nostril. Internasal single, large, irregular hexagonal; widely in contact with prefrontals. Two large prefrontals, longer and wider than internasal, largest distance along the longitudinal axis of prefrontals shorter than frontal in length, anterior-most corner of prefrontals touching nasal, bordered by 2nd and 3rd supralabial, preocular scale, eye, supraocular and frontal. Preocular small, not in contact with supraocular. Loreal and subocular scales absent. Supraocular smaller than frontal. Two postoculars, lower one larger than upper. Two parietals; largest scales on head. Temporals 1+2/1+2. Supralabials 6/6, 4th touching eye, progressively increasing in size from 1st to 6th. Mental small and triangular, wider than long. Infralabials 6/6, first pair in contact, progressively increasing in size from 1st to 6th. Anterior chin shields 2, large, touching 1–4 infralabials. Posterior chin shields 2, anterior half in contact, posterior portion separated by 1st ventral.

278 · Zootaxa 4347 (2) © 2017 Magnolia Press WICKRAMASINGHE ET AL. FIGURE 2. Timetree of Aspidura in context with other natricines. Support values are >95% Pp for all nodes unless indicated. Axis values are in millions of years before the present. We note that many genera are paraphyletic, but this may be based on erroneous or misidentified specimens from GenBank (e.g., Xenochrophis schnurrenburgeri), and should not be relied upon for higher-level taxonomy outside of Aspidura.

NEW SPECIES OF ASPIDURA FROM SRI LANKA Zootaxa 4347 (2) © 2017 Magnolia Press · 279 FIGURE 3. (A) Dorsal aspect, (B) Ventral aspect, of holotype male of Aspidura ravanai sp. nov. (NMSL-NH 2017.19.01). Small and large tick marks in the scale bar indicate mm and cm, respectively.

280 · Zootaxa 4347 (2) © 2017 Magnolia Press WICKRAMASINGHE ET AL. FIGURE 4. (A) Female paratype of Aspidura ravanai sp. nov. (DWC 2017.05.01). (B) Male paratype of Aspidura ravanai sp. nov. (DWC 2017.05.02).

NEW SPECIES OF ASPIDURA FROM SRI LANKA Zootaxa 4347 (2) © 2017 Magnolia Press · 281 FIGURE 5. (A) Coarsely keeled ischiadic region of male paratype (DWC 2017.05.02) of Aspidura ravanai sp. nov., (B) Ischiadic region of female paratype (DWC 2017.05.01) of Aspidura ravanai sp. nov., showing week keeling.

282 · Zootaxa 4347 (2) © 2017 Magnolia Press WICKRAMASINGHE ET AL. FIGURE 6. Sulcate (left), and asulcate (right) views of the right everted hemipenis of (A) Aspidura ravanai sp. nov., paratype male (DWC 2017.05.02), (B) Aspidura trachyprocta, male (NMSL-NH 2017.20.01).

NEW SPECIES OF ASPIDURA FROM SRI LANKA Zootaxa 4347 (2) © 2017 Magnolia Press · 283 FIGURE 7. (A) Aspidura trachyprocta, male (DWC 2017.05.03) in life, colour pattern-black. (B) Aspidura trachyprocta unpreserved male, colour pattern- red.

TABLE 1. The morphometric measurements (mm) of the type series of Aspidura ravanai sp. nov. Aspidura ravanai sp. nov. Mean SD Range Holotype NMSL-NH Paratype DWC Paratype DWC 2017.19.01 2017.05.01 2017.05.02 CL 3.4 6.6 3.7 4.5 1.8 3.4–6.6 DBE 3.6 5.9 3.8 4.4 1.3 3.6–5.9 DFE 2.9 4.8 3.1 3.6 1.1 2.9–4.8 EW 1.1 1.6 1.1 1.3 0.3 1.1–1.6 HD 3.2 6.1 3.3 4.2 1.7 3.2–6.1 HL 12.8 20.0 12.7 15.2 4.2 12.7–20 HW 6.4 10.3 5.8 7.5 2.4 5.8–10.3 IN 0.9 1.4 0.8 1.1 0.3 0.8–1.4 IO 3.7 5.9 3.9 4.5 1.2 3.7–5.9 NE 2.8 4.5 2.6 3.3 1.0 2.6–4.5 NW 0.4 0.4 0.3 0.4 0.0 0.3–0.4 SE 3.6 5.0 3.5 4.0 0.9 3.5–5.0 SG 8.3 11.6 8.1 9.3 1.9 8.1–11.6 SN 0.5 0.7 0.4 0.6 0.2 0.4–0.7 SVL 256.0 414.0 218.0 296.0 103.9 218–414 TaL 37.2 39.4 34.9 37.2 2.3 34.9–39.4

NEW SPECIES OF ASPIDURA FROM SRI LANKA Zootaxa 4347 (2) © 2017 Magnolia Press · 285 FIGURE 9. Lateral aspect of ischiadic, anal and tail base regions of (A) Aspidura ravanai sp. nov., holotype male (NMSL-NH 2017.19.01), (B) Aspidura trachyprocta, male (DWC 2017.05.03).

286 · Zootaxa 4347 (2) © 2017 Magnolia Press WICKRAMASINGHE ET AL. FIGURE 10. Dorsal aspect of ischiadic and tail base regions (A) Aspidura ravanai sp. nov., holotype male (NMSL-NH 2017.19.01), (B) Aspidura trachyprocta, male (DWC 2017.05.03).

NEW SPECIES OF ASPIDURA FROM SRI LANKA Zootaxa 4347 (2) © 2017 Magnolia Press · 287 FIGURE 11. Ventral aspect of anal and tail base regions of (A) Aspidura ravanai sp. nov., holotype male (NMSL-NH 2017.19.01), (B) Aspidura trachyprocta, male (DWC 2017.05.03).

288 · Zootaxa 4347 (2) © 2017 Magnolia Press WICKRAMASINGHE ET AL. TABLE 2. Scale count of Aspidura ravanai sp. nov. Aspidura ravanai sp. nov. Holotype NMSL-NH 2017.19.01 Paratype DWC 2017.05.01 Paratype DWC 2017.05.02 Internasal 1 1 1 Prefrontal 2 2 2 Frontal 1 1 1 Supraoculars 1 1 1 Parietal 2 2 2 Nasal 2 2 2 Loreals 0 0 0 Preoculars 1 1 1 Postoculars 2 2 2 Temporals 1+2 1+2 1+2 Supralabials 6 6 6 Eye at 4 4 4 Infralabials 6 6 6 Anterior chin shields 2 2 2 Posterior chin shields 2 2 2 Coastals 15, 15, 15 15, 15, 15 15, 15, 15 Keels around base of tail coarse weak coarse Ventrals 139 132 139 Preanal single/double single single single Anal single/double single single single Subcaudals 28 20 26 Subcaudals single/double single single single

Body scalation. Ventrals 139, 1st ventral longer than wide; subcaudals 29, all single; anal single and large; dorsal scale rows 15–15–15; coastal scales on posterior body from about 110th ventral feebly keeled, except the ischiadic region which is coarsely keeled, with 1–3 peaks on each scale; vertebral rows and first coastal not enlarged; no apical pit. Colour in life. Entire dorsum jet black. Fourth supralabial to neck displays bright yellow stripe ventrolaterally. Irregular bright yellow stripe continues ventrolaterally from neck to tail, becoming discontinuous midbody. Venter black with bright yellow blotching all over; gular region primarily bright yellow. Colour in alcohol. Colour pattern remains unchanged. Pupil changes to off white. Black colour fades, bright yellow changes to off white. Variation. Female paratype showed much weaker keeling on coastal scales. The first ventral wider than long. Subcaudals 20, the first subcaudal divided. Snout truncate in dorsal aspect. Male paratype showed ventrolateral bright yellow stripe only on the head and tail regions. In both paratypes, gular region black with very little bright yellow blotching. Hemipenis morphology. Based on DWC 2017.05.02 paratype specimen: right everted hemipenis extends for length of 5 subcaudals. Everted organ single subcylindrical, sulcus spermaticus simple. Basal to apex region bearing prominent spines which are evenly distributed and are in uniform length (Figure 6). Natural History. Seventeen specimens of Aspidura ravanai have been found in montane cloud forests between 1,650–2,000 m elevation. Mostly found in shady areas with dense leaf litter, hidden under loose soil or logs, and can be found up to about 30 cm below ground level. Etymology. The specific epithet ravanai is formed after Emperor Ravana, one of the greatest Emperors of all times, who is believed to have lived nearly 5000 years ago in Sri Lanka, and was said to have had immense intelligence and power. The name ravanai is a noun in the genitive case.

NEW SPECIES OF ASPIDURA FROM SRI LANKA Zootaxa 4347 (2) © 2017 Magnolia Press · 289 Suggested common names. Ravanage madilla, and Ravana’s rough-side snake in native Sinhala and English respectively. Molecular divergence. The results from our dating analysis suggest that Natricinae began diversifying approximately 37 Ma (95% Highest Posterior Density [HPD] = 29–45, Ma) in the late Eocene, consistent with most previous studies (e.g., Guo et al. 2012; Chen et al. 2013). The genus Aspidura began diversifying in the early Miocene, ~18 Ma (95% HPD = 13–24 Ma), and A. ravanai diverged from A. trachyprocta in the late Miocene, ~6.2 Ma (95% HPD = 2.8–10.0 Ma). Thus, there is substantial evolutionary and genetic divergence between these taxa, despite their general morphological similarity and ecological sympatry (Figure 2). Comparison. The new species was compared with all known congeners of the genus Aspidura and the species most closely resembles A. trachyprocta (Figure 7) due to the following combination of characters: Similar body size, one preocular, two postoculars, 1+2 temporals, supralabials 6, 4th supralabial in contact with the eye, infralabials 6 (Figure 7B), coastals 15, single cloacal scale, overlapping ventral and subcaudal counts, and similar habitat preferences, but can easily be distinguished from the following morphological characters: (1) entire dorsum jet black in Aspidura ravanai by (vs. reddish-yellow to brown with a longitudinal black stripe on mid dorsum in Aspidura trachyprocta) (Figure 7–8), (2) ventrolaterally an irregular longitudinal yellow stripe (vs. black stripe), (3) spinose tubercles absent (vs. spinose tubercles present, more pronounced in males) (Figure 9–11), (4) hemipenis extends for length of 5 subcaudals (vs. 3–4), narrow apex (vs. broader), ridges absent (vs. present) (Figure 6); from A. brachyorrhos by having 15 coastals (vs. 17), preocular not in contact with supraocular (vs. contact), prefrontal contact with eye (vs. separate), single subcaudals (vs. paired); from A. copei by having coastals 15 (vs. 17), single subcaudals (vs. paired), single preocular (vs. absent); from A. deraniyagalae by larger SVL 218– 414 mm (vs. small SVL 90–126), coastals 15 (vs. 17), ventrals 132–139 (vs. 117–122), single subcaudals (vs. paired); from A. drummondhayi by having single subcaudals (vs. paired), single preocular (vs. absent); from A. guentheri from larger SVL 218–414 mm (vs. small, SVL 170 mm), coastals 15 (vs. 17), ventrals 132–139 (vs. 100– 127); from A. ceylonensis by prefrontal touching eye (vs. not touching eye), preocular does not touch supraocular (vs. touches), lower postocular larger than the upper (vs. vise versa), mid body coastals not keeled (vs. coarsely keeled).

Discussion

Although both Aspidura ravanai and A. trachyprocta are sympatric on the western slopes of the Sri Pada peak in the Central highlands, the latter can be found across a much wide range in the Central highlands above 1,500 m elevation, whereas A. ravanai has thus far only been recorded from the western slope of Sri Pada, at ca. 1,700m. Despite similarities in morphology and overlapping distribution, the two species are widely divergent genetically (Figure 2). The genus Aspidura began diversifying at least 18 Ma, while A. ravanai diverged from A. trachyprocta roughly 6.2 Ma, during the Miocene. Without complete sampling of all species in this analysis, we can’t pinpoint the specific timing of the genus or identify the exact sister-lineage of A. ravanai. However, these dates provide minimum estimates of divergence, and show the ancient distinctiveness of A. ravanai from its sympatric congener A. trachyprocta. Ukuwela & Dawundasekara (2012) described seven colour-pattern variants of Aspidura trachyprocta from Seetha Eliya (1,800 m), Nuwara Eliya district, Sri Lanka, two of which (types 1 and 3) superficially resemble A. ravanai. Although type 1 (the most common) has the entire dorsum purplish black and the venter with orange blotches, and type 3 has a venter with yellow blotching and reddish-brown dosrum, A. ravanai is clearly distinguishable from both in having bright yellow blotching on venter, a jet-black dorsum, and the prominent bright yellow ventrolateral stripe on the neck and tail, occasionally running the length of the body discontinuously. The amount of color-pattern variation in A. trachyprocta also suggests that this taxon might represent a complex of species. Our results and others (e.g., Pyron et al. 2013) suggest that the snake diversity of Sri Lanka still contains additional species to be discovered and described.

We thank the Biodiversity Secretariat of the Ministry of Environment, Sri Lanka for funding and collaboration with this project, and thank Gamini Gamage, Dakshini Perera, and Hasula Wickramasinghe for all support rendered. We thank the Nagao Natural Environment Foundation for partial funding, Kithsiri Gunawardena and Canon Metropolitan for providing selected camera equipment, the Department of Wildlife Conservation for permissions granted (Permit no. WL/3/3/354), and the Director General, U. L Thaufeek, and U.K.L. Peeris for all support given. To Amith Perera for designing a light source “FixIt” for the microscope. LJMW acknowledges readers who purchased his self-authored coffee table book “Repertoire: A pictorial gateway to Sri Lanka’s nature, which provided part funding for purchase of necessary equipment. We are grateful for the contributions of Prasanna Samarawickrama and Mahesh Chathuranga in preparing the map and figures respectively. Kylie M. Butler, Vidya Abhayagunawardena and Toni Fernando provided constructive criticism of an earlier draft. We also wish to thank the Directress of the National Museum of Sri Lanka, Sanuja Kasthuriarachchi, and staff members Chandrika Munasinghe and Manaram de Silva, gratefully acknowledging their assistance in museum-reference work. We also thank our colleagues at Herpetological Foundation of Sri Lanka (HFS) for various courtesies, Asanka Abayakoon for his encouragements, Kanishka Ukuwela and Patrick David whose invaluable comments undoubtedly improved the quality of the paper. Finally, we thank our Principal Sponsors Dilmah Conservation, for funding the project activities.

References

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