4559 (2): 265 280 Article ... · types which includes; montane, sub-montane, pigmy, semi-evergreen,...

16
Accepted by T. Nguyen: 24 Dec. 2018; published: 20 Feb. 2019 ZOOTAXA ISSN 1175-5326 (print edition) ISSN 1175-5334 (online edition) Copyright © 2019 Magnolia Press Zootaxa 4559 (2): 265280 https://www.mapress.com/j/zt/ Article 265 https://doi.org/10.11646/zootaxa.4559.2.3 http://zoobank.org/urn:lsid:zoobank.org:pub:B4C51F11-7AB6-4A7F-9662-4774EA622BD2 A new species of Aspidura Wagler, 1830 (Squamata: Colubridae: Natricinae) from Knuckles, World Heritage Site, Sri Lanka L. J. MENDIS WICKRAMASINGHE 1,4 , IMESH NUWAN BANDARA 2 , DULAN RANGA VIDANAPATHIRANA 1 & NETHU WICKRAMASINGHE 1,3 1 Herpetological Foundation of Sri Lanka, 31/5, Alwis Town, Hendala, Wattala, Sri Lanka 2 Ellangàwa Unity Care for Community and Nature, Hapugoda, Ambathenna, Sri Lanka 3 Institute of Biochemistry, Molecular Biology, Biotechnology, University of Colombo, Sri Lanka 4 Corresponding author. E-mail: [email protected] or [email protected] Abstract We describe a new species, Aspidura desilvai sp. nov., closely resembling A. trachyprocta and the last addition to the ge- nus A. ravanai, from Knuckles massif, Matale District, of Sri Lanka. The species represents the ninth species of the genus known from Sri Lanka, and is readily distinguished from all other congeners by its colour pattern, the scale nature in the ischiadic region, and morphometric characteristics specially from its ratio between the snout to eye distance to its eye width. The species is currently known only from the type locality. Key words. Aspidura ravanai, A. trachyprocta, central highlands, Knuckles, Roughside snakes, South Asia, systematics Introduction The endemic snake genus Aspidura Wagler, 1830 of Sri Lanka, is currently represented by eight species, after the most recent taxonomic description of a rough-side snake Aspidura ravanai Wickramasinghe, et al. (2017). A. ravanai has hitherto been only recorded from its type locality which is the western slopes of the Sripada hills, and is sympatric with its closely resembling congener A. trachyprocta Cope, 1860, (Wickramasinghe et al. 2017). The Central Highland massif complex consists of three major massifs of significant biodiversity; Central hills, Rakwana hills, and Knuckles hills, separated by two major river basins. Knuckles massif is a part of the Central Highlands and Knuckles World Heritage Site (WHS), situated towards the north of the WHS, biogeographically separated by the Mahaweli river basin from rest of the Central Highland massifs (Fernando et al. 2007; Samarawickrama et al. 2012; Wickramasinghe et al. 2015). Its forest cover consists of a diverse range of forest types which includes; montane, sub-montane, pigmy, semi-evergreen, riverine, rock outcrop, savanna, patana grass land, and scrublands (Bambaradeniya & Ekanayake, 2003). The Knuckles range has altitudes varying from 760 - 1900 m, and the forest covers serves as an important catchment area to the longest river Mahaweli (Rajapaksha et al. 2006). The altitudinal variations and geographical location has resulted in a wide diversity of flora and fauna in this region. The biogeographical separation of the Knuckles region from a major river basin has resulted in species isolation from the rest of the region, where many of these species were separated as new from already described species spread across the Central Highlands, during the past two decades, which resulted in a rise of known species for the herpetofauna (Meegaskumbura & Manamendra-Arachchi 2005, 2011; Samarawickrama et al. 2006; Amarasinghe et al. 2014a & b). We herein describe a population from the Knuckles massif, a part of the Central Highlands and Knuckles World Heritage Site, as a new species of rough-side snake, Aspidura desilvai sp. nov., 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 ratios specially from its ratio between the snout to eye distance to its eye width, scalation, and its unique colour pattern.

Transcript of 4559 (2): 265 280 Article ... · types which includes; montane, sub-montane, pigmy, semi-evergreen,...

Page 1: 4559 (2): 265 280 Article ... · types which includes; montane, sub-montane, pigmy, semi-evergreen, riverine, rock outcrop, savanna, patana grass land, and scrublands (Bambaradeniya

Accepted by T. Nguyen: 24 Dec. 2018; published: 20 Feb. 2019

ZOOTAXA

ISSN 1175-5326 (print edition)

ISSN 1175-5334 (online edition)Copyright © 2019 Magnolia Press

Zootaxa 4559 (2): 265–280

https://www.mapress.com/j/zt/Article

265

https://doi.org/10.11646/zootaxa.4559.2.3

http://zoobank.org/urn:lsid:zoobank.org:pub:B4C51F11-7AB6-4A7F-9662-4774EA622BD2

A new species of Aspidura Wagler, 1830 (Squamata: Colubridae: Natricinae)

from Knuckles, World Heritage Site, Sri Lanka

L. J. MENDIS WICKRAMASINGHE1,4, IMESH NUWAN BANDARA2,

DULAN RANGA VIDANAPATHIRANA1 & NETHU WICKRAMASINGHE1,3

1Herpetological Foundation of Sri Lanka, 31/5, Alwis Town, Hendala, Wattala, Sri Lanka2Ellangàwa Unity Care for Community and Nature, Hapugoda, Ambathenna, Sri Lanka3Institute of Biochemistry, Molecular Biology, Biotechnology, University of Colombo, Sri Lanka4Corresponding author. E-mail: [email protected] or [email protected]

Abstract

We describe a new species, Aspidura desilvai sp. nov., closely resembling A. trachyprocta and the last addition to the ge-

nus A. ravanai, from Knuckles massif, Matale District, of Sri Lanka. The species represents the ninth species of the genus

known from Sri Lanka, and is readily distinguished from all other congeners by its colour pattern, the scale nature in the

ischiadic region, and morphometric characteristics specially from its ratio between the snout to eye distance to its eye

width. The species is currently known only from the type locality.

Key words. Aspidura ravanai, A. trachyprocta, central highlands, Knuckles, Roughside snakes, South Asia, systematics

Introduction

The endemic snake genus Aspidura Wagler, 1830 of Sri Lanka, is currently represented by eight species, after the

most recent taxonomic description of a rough-side snake Aspidura ravanai Wickramasinghe, et al. (2017). A.

ravanai has hitherto been only recorded from its type locality which is the western slopes of the Sripada hills, and

is sympatric with its closely resembling congener A. trachyprocta Cope, 1860, (Wickramasinghe et al. 2017).

The Central Highland massif complex consists of three major massifs of significant biodiversity; Central hills,

Rakwana hills, and Knuckles hills, separated by two major river basins. Knuckles massif is a part of the Central

Highlands and Knuckles World Heritage Site (WHS), situated towards the north of the WHS, biogeographically

separated by the Mahaweli river basin from rest of the Central Highland massifs (Fernando et al. 2007;

Samarawickrama et al. 2012; Wickramasinghe et al. 2015). Its forest cover consists of a diverse range of forest

types which includes; montane, sub-montane, pigmy, semi-evergreen, riverine, rock outcrop, savanna, patana grass

land, and scrublands (Bambaradeniya & Ekanayake, 2003). The Knuckles range has altitudes varying from 760 -

1900 m, and the forest covers serves as an important catchment area to the longest river Mahaweli (Rajapaksha et

al. 2006). The altitudinal variations and geographical location has resulted in a wide diversity of flora and fauna in

this region. The biogeographical separation of the Knuckles region from a major river basin has resulted in species

isolation from the rest of the region, where many of these species were separated as new from already described

species spread across the Central Highlands, during the past two decades, which resulted in a rise of known species

for the herpetofauna (Meegaskumbura & Manamendra-Arachchi 2005, 2011; Samarawickrama et al. 2006;

Amarasinghe et al. 2014a & b).

We herein describe a population from the Knuckles massif, a part of the Central Highlands and Knuckles

World Heritage Site, as a new species of rough-side snake, Aspidura desilvai sp. nov., 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 ratios specially from its ratio between the

snout to eye distance to its eye width, scalation, and its unique colour pattern.

Page 2: 4559 (2): 265 280 Article ... · types which includes; montane, sub-montane, pigmy, semi-evergreen, riverine, rock outcrop, savanna, patana grass land, and scrublands (Bambaradeniya

WICKRAMASINGHE ET AL.266 · Zootaxa 4559 (2) © 2019 Magnolia Press

Materials and methods

We sampled an unidentified Aspidura species from the Knuckles massif (Figure 1). The specimens were

photographed in life (Canon EOS 7D, Canon 100mm F/2.8L IS USM Macro Lens, and Canon MT 24+Flash dome

diffuser (1/250 Sec, f/14, ISO 100), and the entire species series was fixed and preserved in 85% ethanol and

subsequently transferred to 75% ethanol after one day. A female specimen has been selected as the holotype against a

male specimen due to the fact that this specimen has the largest SVL and shows the least amount of artifact defects.

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 (NE, 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 the nature of keels on

coastal scales of the ischiadic, anal and tail base regions which are absent in females.

Observations and drawing were made through a Leica Wild M50 microscope with drawing tube attachment.

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 (Boulenger 1890;

Wall 1921; Smith 1943; 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).

Systematics

Aspidura desilvai sp. nov.

(Figures 1–7)

Holotype. NMSL-NH 2019.01.02, adult male, 168 mm SVL (Figure 2), from Riverstone, Knuckles, Matale

District, Central Province, Sri Lanka (07°31’39” N, 80°44’01” E, elevation 1420 m). Collected by L.J.M.W and

D.R.V. on 07 July 2018.

Paratypes. NMSL-NH 2019.01.01, adult female, 208 mm SVL, from Panwila in Knuckles Mountain Range,

Kandy District, Central Province in Sri Lanka (07º22'00.36’’ N, 080º41'00.10’’ E, elevation 995 m). Collected by

L.J.M.W and I.N.B. on 13 March 2011; DWC 2019.05.01, adult female, 157 mm SVL, from Dotulugala, Knuckles

Mountain Range, Kandy District, Central Province, Sri Lanka (07º27'00.30” N, 080º45'00.20” E, elevation 1700

m). Collected by L.J.M.W and I.N.B. on 17 March 2011; DWC 2019.05.02, juvenile male, 93 mm SVL, from

Gombaniya Mountain, Knuckles Mountain Range, Matale District, Central Province, Sri Lanka (07º27'51.76’’ N,

080º45'51.79’’ E, elevation 1375 m). Collected by L.J.M.W and I.N.B. on 13 March 2011.

Diagnosis. SVL 94–216 mm; snout to eye distance 2.5 times the eye width (SE/EW); 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 124–139; subcaudals 16–29; dorsal scale rows 15–15–15;

laterally spine like tubercles present on two scale rows nearest to the subcaudals of the ischiadic, anal and tail base

regions in adult males, feeble in juvenile males, and absent in females; entire dorsum brown colour, much paler

towards anterior; three irregular dotted lines on dorsum.

Page 3: 4559 (2): 265 280 Article ... · types which includes; montane, sub-montane, pigmy, semi-evergreen, riverine, rock outcrop, savanna, patana grass land, and scrublands (Bambaradeniya

Zootaxa 4559 (2) © 2019 Magnolia Press · 267A NEW SPECIES OF ASPIDURA FROM SRI LANKA

FIGURE 1. Current distribution map of Aspidura desilvai sp. nov. (red: holotype; blue: paratypes; yellow: not collected) in the

Knuckles landscape.

Description of holotype. Adult male; SVL 168 mm; TaL 25.1 mm; TL 193.1 mm; TaL/TL 0.13; body

elongate and cylindrical; head short (SVL/HL 18.3), elliptical, indistinct from thick neck; snout long, narrowing

Page 4: 4559 (2): 265 280 Article ... · types which includes; montane, sub-montane, pigmy, semi-evergreen, riverine, rock outcrop, savanna, patana grass land, and scrublands (Bambaradeniya

WICKRAMASINGHE ET AL.268 · Zootaxa 4559 (2) © 2019 Magnolia Press

anteriorly, pointed in dorsal aspect, snout to nostril distance about 2.8 (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, distance between snout to eye about 2.6 (SE/EW) times the eye width, round pupil;

snout to eye distance 0.3 times head length (SE/HL); tail short (TaL/SVL 0.1), robust at its base, tapering

progressively to a single point.

TABLE 1. The morphometric measurements (mm) of the type series of Aspidura desilvai sp. nov.

Head scalation. Head scalation includes 1 internasal, 2 prefrontals, 2 supraoculars, 1 frontal, and 2 parietals

(Figure 3A). Rostral small, convex, wider than long and rounded in lateral, dorsal and ventral aspects. Nasal

vertically divided by a groove above nostril (Figure 3B). Internasal large, irregular hexagonal; widely in contact

with prefrontals. Two large prefrontals, longer and wider than internasals, largest distance along the longitudinal

axis of prefrontals shorter than frontal (Figure 3A) 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 (Figure 3B). 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 (Figure 3C).

Body scalation. Ventrals 124, 1st ventral longer than wide; subcaudals 24, all single; anal single and large;

dorsal scale rows 15–15–15; laterally prominent spine like tubercles present on two scale rows nearest to the

subcaudals, and its protrusion reducing towards upper scale rows in the ischiadic, anal and tail base regions (Figure

4); vertebral rows and first coastal not enlarged; no apical pit.

Hemipenis morphology. Based on Holotype specimen: right everted hemipenis extends for length of 3

subcaudals. Everted organ single subcylindrical, globular, sulcus spermaticus simple. Basal to apex region bearing

prominent spines which are evenly distributed and are in uniform length (Figure 5).

Colour in life. Supralabials and infralabials light yellow, with dark margins separating each scale (Figure 6A).

Entire dorsum reddish brown colour, much paler towards anterior and each scale having tiny dark spots (Figure 2).

Three irregular dotted lines on dorsum (Figure 6B). These are symmetrically placed and continues from neck to tail

Aspidura desilvai sp. nov. Mean SD Range

Holotype NMSL-NH

2019.01.02 (male)

Paratype DWC

2019.01.01 (female)

Paratype DWC

2019.05.01 (female)

Paratype DWC

2019.05.02 (male)

CL 4.19 3.28 2.39 1.91 2.943 0.87 1.91–4.19

DBE 2.57 2.86 2.38 1.97 2.445 0.32 1.97–2.86

DFE 2.21 2.36 2.11 1.66 2.085 0.26 1.66–2.36

EW 1.01 1.09 0.91 0.82 0.958 0.1 0.82–1.09

HD 3.96 3.74 3.33 2.72 3.438 0.47 2.72–3.96

HL 9.18 10.35 9.35 7.15 9.008 1.16 7.15–10.35

HW 3.84 3.99 3.72 3.15 3.675 0.32 3.15–3.99

IN 0.72 0.6 0.6 0.53 0.613 0.07 0.53–0.72

IO 2.69 2.93 2.65 2.08 2.588 0.31 2.08–2.93

NE 2.27 2.48 2.26 1.73 2.185 0.28 1.73–2.48

NW 0.14 0.19 0.1 0.1 0.133 0.04 0.1–0.19

SE 2.65 2.69 2.31 2.04 2.423 0.27 2.04–2.69

SG 6.68 6.81 5.97 4.63 6.023 0.87 4.63–6.81

SN 0.32 0.33 0.32 0.31 0.32 0.01 0.31–0.33

SVL 168 208 157 93 156.5 41.3 93–208

TaL 25.11 16.43 12.17 13.57 16.82 5.03 12.2–25.1

Page 5: 4559 (2): 265 280 Article ... · types which includes; montane, sub-montane, pigmy, semi-evergreen, riverine, rock outcrop, savanna, patana grass land, and scrublands (Bambaradeniya

Zootaxa 4559 (2) © 2019 Magnolia Press · 269A NEW SPECIES OF ASPIDURA FROM SRI LANKA

end. Prominent light brown stripe continues dorsolaterally from neck to tail end, marked due to much darker

regions which constitutes of dotted lines below and above this region. These lines continue from neck to tail end.

Venter primarily peach, with black blotching all over; gular region yellow.

FIGURE 2. Dorsal aspect of Aspidura desilvai sp. nov., the holotype male (NMSL-NH 2019.01.02) in life.

Colour in alcohol. Colour pattern remains unchanged. Pupil changes to off white. Darker regions fades to a

light brown.

Variations in colour. In an unpreserved male specimen (Figure 7D) except the head region and ventre the

entire body was black.

Natural History. Aspidura desilvai sp. nov. have been observed commonly in its habitat (Figure 1). The

species is confined to Knuckles conservation area, and is found in and above the lower montane forests of

Knuckles. Authors have observed the snake from 995 m up to 1700 m above sea level (Figure 8). The habitat of A.

desilvai sp. nov. is closed canopy forests dominated by Syzigium sp. (Figure 9). The moist-cooler habitat is densely

occupied with large and medium sized trees which are heavily covered with epiphytes. No direct sunlight falls to

the forest floor, and the undergrowth was not well established where the individuals were found. Relatively thin

litter cover was observed in the habitat. Commonly observed under leaf litter and loose soil while they were also

observed under rocks, boulders, and decaying logs. Individuals come out to the surface during the day time.

Reddish brown latosolic soil in the locality is more or less similar to the body colour of the snake.

Etymology. The species is named in honor of Pilippu Hewa Don Hemasiri de Silva (Dr. P. H. D. H. de Silva),

a former Director (1965-1981) of the National Museums of Sri Lanka. In recognition of his tireless services to the

country, while in service and through his many publications specially as the author of the book titled “Snake Fauna

of Sri Lanka, with special reference to skull, dentition and venom in snakes”. The species epithet desilvai is a noun in

the genitive case.

Suggested common names. desilvage madilla, and de Silva’s Rough-Side Snake in native Sinhala language

and English language respectively.

Page 6: 4559 (2): 265 280 Article ... · types which includes; montane, sub-montane, pigmy, semi-evergreen, riverine, rock outcrop, savanna, patana grass land, and scrublands (Bambaradeniya

WICKRAMASINGHE ET AL.270 · Zootaxa 4559 (2) © 2019 Magnolia Press

FIGURE 3. Aspidura desilvai sp. nov., the holotype male (NMSL-NH 2019.01.02), head; (A) dorsal, (B) lateral, and (C)

ventral, aspects. Scale bars: 1 mm.

Page 7: 4559 (2): 265 280 Article ... · types which includes; montane, sub-montane, pigmy, semi-evergreen, riverine, rock outcrop, savanna, patana grass land, and scrublands (Bambaradeniya

Zootaxa 4559 (2) © 2019 Magnolia Press · 271A NEW SPECIES OF ASPIDURA FROM SRI LANKA

FIGURE 4. Presence of spine like tubercles laterally on ischiadic, tail base and anal regions of Aspidura desilvai sp. nov., the

holotype male (NMSL-NH 2019.01.02), (A) lateral, and (B) ventral, aspects.

Page 8: 4559 (2): 265 280 Article ... · types which includes; montane, sub-montane, pigmy, semi-evergreen, riverine, rock outcrop, savanna, patana grass land, and scrublands (Bambaradeniya

WICKRAMASINGHE ET AL.272 · Zootaxa 4559 (2) © 2019 Magnolia Press

FIGURE 5. Hemipenial morphology of the right everted hemipenis of Aspidura desilvai sp. nov., the holotype male (NMSL-

NH 2019.01.02), (A) sulcate and, (B) asulcate views.

Comparison. The new species was compared with all known congeners of the genus Aspidura and the species

most closely resembles A. ravanai, and A. trachyprocta, due to the following combination of characters: one

preocular, two postoculars, 1+2 temporals, supralabials 6, 4th supralabial in contact with the eye, infralabials 6,

coastals 15, single cloacal scale, and overlapping ventral and subcaudal counts, but can easily be distinguished by

the following morphological characters: from A. ravanai: entire dorsum brown colour, much paler towards anterior

and each scale having tiny dark spots in Aspidura desilvai sp. nov. (vs. entire dorsum jet black in Aspidura

ravanai), ventrolaterally darker region which constitutes of irregular longitudinal dotted lines (vs. ventrolaterally

an irregular longitudinal yellow stripe), laterally prominent spine like tubercles present on two scale rows nearest to

the subcaudals, and its protrusion reducing towards upper scale rows (vs. entire coastal rows coarsely keeled, with

1–3 peaks on each scale) in males (Figures 4 & 10A–C), entire coastal rows of the ischiadic, anal and tail base

regions smooth (vs. feebly keeled) in females, snout to eye distance about 2.5 times its eye width (vs. 3.2 times in

A. ravanai) (Figures 3 & 10D–E); from Aspidura trachyprocta: entire dorsum brown colour, much paler towards

anterior and each scale having tiny dark spots in A. desilvai sp. nov. (vs. reddish-yellow to brown with a

longitudinal black stripe on mid dorsum in Aspidura trachyprocta), ventrolaterally darker region which constitutes

of irregular longitudinal dotted lines (vs. black stripe), laterally prominent spine like tubercles present on two scale

rows nearest to the subcaudals, and its protrusion reducing towards upper scale rows (vs. bulging spine like

tubercles prominent laterally which reduces towards dorsum) of the ischiadic, anal and tail base regions in males

(Figures 4 & 11A–C), entire coastal rows of the ischiadic, anal and tail base regions smooth (vs. feebly keeled) in

females, snout to eye distance about 2.5 times its eye width (vs. twice in A. trachyprocta) (Figures 3 & 11D–E);

from A. brachyorrhos Boie, 1827, 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 Günther, 1864 by

having coastals 15 (vs. 17), single subcaudals (vs. paired), single preocular (vs. absent); from A. deraniyagalae

Gans & Fetcho, 1982 by having 15 coastals (vs. 17), ventrals 124–139 (vs. 117–122), single subcaudals (vs.

Page 9: 4559 (2): 265 280 Article ... · types which includes; montane, sub-montane, pigmy, semi-evergreen, riverine, rock outcrop, savanna, patana grass land, and scrublands (Bambaradeniya

Zootaxa 4559 (2) © 2019 Magnolia Press · 273A NEW SPECIES OF ASPIDURA FROM SRI LANKA

paired); from A. drummondhayi Boulenger, 1904, by having single subcaudals (vs. paired), single preocular (vs.

absent); from A. guentheri Ferguson, 1876 by having 15 coastals (vs. 17), ventrals 124–139 (vs. 100–127); from A.

ceylonensis (Günther, 1858), 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).

TABLE 2. Scale count of Aspidura desilvai sp. nov.

Discussion

Aspidura desilvai sp. nov., is currently known only from the Knuckles massif. A. brachyorrhos is the only known

sympatric congener. However they can be readily distiguished from their coastal scale count, and the colour

pattern. The colouration of the novel species is distinct and unique, which also seems to ressemble the colouration

of the latosolic soil, of the habitat that the species is found in.

The novel species closely resembles Aspidura ravanai and A. trachyprocta. The Central Highlands of the

island consists of three major massifs, the Central, Knuckles and the Rakwana hills, which has been separated by

two major river basins. A. trachyprocta is distributed throughout the Central highlands (>1000 m) except the

Knuckles massif. The recently described species A. ravanai, is only found from higher altitudes of the Sripada peak

of the Central hills. However A. desilvai sp. nov. has hitherto been only observed from the Knuckles massif of the

Central Highlands from an elevation of about 995 m, and its habitat clearly does not overlap with that of A. ravanai

and A. trachyprocta, and it is therefore allopatric with both these congeners.

Aspidura desilvai sp. nov.

Holotype NMSL-NH Paratype DWC

2019.01.01

Paratype DWC

2019.05.01

Paratype DWC

2019.05.02

2019.01.02 (male) (female) (female) (male)

Internasal 1 1 1 1

Prefrontals 2 2 2 2

Frontal 1 1 1 1

Supraocular 1 1 1 1

Parietalss 2 2 2 2

Nasal 2 2 2 2

Loreals 0 0 0 0

Preocular 1 1 1 1

Postoculars 2 2 2 2

Temporals 1+2 1+2 1+2 1+2

Supralabials 6 6 6 6

Eye at 4 4 4 4

Infralabials 6 6 6 6

Anterior chin shields 2 2 2 2

Posterior chin shields 2 2 2 2

Coastals 15, 15, 15 15, 15, 15 15, 15, 15

Spine like tubercles (ischiadic region) prominent absent absent feeble

Ventrals 124 135 139 124

Preanal single/double single single single single

Anal single/double single single single single

Subcaudals 24 17 16 29

Subcaudals single/double single single single single

Page 10: 4559 (2): 265 280 Article ... · types which includes; montane, sub-montane, pigmy, semi-evergreen, riverine, rock outcrop, savanna, patana grass land, and scrublands (Bambaradeniya

WICKRAMASINGHE ET AL.274 · Zootaxa 4559 (2) © 2019 Magnolia Press

FIGURE 6. Aspidura desilvai sp. nov., the holotype (NMSL-NH 2019.01.02), head colour in life; (A) lateral and, (B) dorsal,

aspects.

Based on data gathered from the present study the area of occurrence for the sp. nov. is 120 km2, spread across

the Knuckles landscape. There has been an increase in tourism activities in the area, specially due to the

uncontrolled eco-tourism trends in the Knuckles region. The species is threatened due to habitat destruction from

expansion of existing roads (Figure 14), and an increase in the number of visitors travelling in vehicles. During the

study period we have observed as many as 7 road kills of A. desilvai, in Riverstone area alone. Apart from which

the species is threatened with habitat loss due to other anthropogenic activities, such as: forest fragmentation,

illegal cardamom plantations, uncontrolled gem mining, forest fires from human activities, introduction and spread

of invasive alien species. The recent additions to this genus is evidence that the knowledge on this genus is far from

being complete.

Page 11: 4559 (2): 265 280 Article ... · types which includes; montane, sub-montane, pigmy, semi-evergreen, riverine, rock outcrop, savanna, patana grass land, and scrublands (Bambaradeniya

Zootaxa 4559 (2) © 2019 Magnolia Press · 275A NEW SPECIES OF ASPIDURA FROM SRI LANKA

FIGURE 7. Colour variations among the population, (A) NMSL-NH 2019.01.01, adult female, (B) DWC 2019.05.02, juvenile

male, (C) unpreserved female, (D) unpreserved male.

FIGURE 8. Typical habitat of A. desilvai sp. nov., a panoramic view of the montane cloud forest covers of the Kunuckles

landscape.

Page 12: 4559 (2): 265 280 Article ... · types which includes; montane, sub-montane, pigmy, semi-evergreen, riverine, rock outcrop, savanna, patana grass land, and scrublands (Bambaradeniya

WICKRAMASINGHE ET AL.276 · Zootaxa 4559 (2) © 2019 Magnolia Press

FIGURE 9. The typical habitat of A. desilvai sp. nov., closed canopy forests dominated by Syzigium sp.

Page 13: 4559 (2): 265 280 Article ... · types which includes; montane, sub-montane, pigmy, semi-evergreen, riverine, rock outcrop, savanna, patana grass land, and scrublands (Bambaradeniya

Zootaxa 4559 (2) © 2019 Magnolia Press · 277A NEW SPECIES OF ASPIDURA FROM SRI LANKA

FIGURE 10. Ischiadic, tail base and anal regions of Aspidura ravanai, the holotype male (NMSL-NH 2017.19.01), (A) dorsal

(entire coastal rows coarsely keeled), (B) lateral (coarsely keeled, with 1–3 peaks on each scale), and (C) ventral aspects. Head,

showing snout to eye distance 3.2 times its eye width; (D) dorsal, and (E) lateral aspects. Scale bar: 1 mm.

Page 14: 4559 (2): 265 280 Article ... · types which includes; montane, sub-montane, pigmy, semi-evergreen, riverine, rock outcrop, savanna, patana grass land, and scrublands (Bambaradeniya

WICKRAMASINGHE ET AL.278 · Zootaxa 4559 (2) © 2019 Magnolia Press

FIGURE 11. Presence of bulging spine like tubercles on ischiadic, tail base and anal regions of Aspidura trachyprocta, male

(DWC 2017.05.03), (A) dorsal, (B) lateral and, (C) ventral aspects. Head showing snout to eye distance twice its eye width, (D)

dorsal and, (E) lateral aspects. Scale bar: 1 mm.

Page 15: 4559 (2): 265 280 Article ... · types which includes; montane, sub-montane, pigmy, semi-evergreen, riverine, rock outcrop, savanna, patana grass land, and scrublands (Bambaradeniya

Zootaxa 4559 (2) © 2019 Magnolia Press · 279A NEW SPECIES OF ASPIDURA FROM SRI LANKA

FIGURE 12. Threats to the species, due to habitat destruction from expansion of existing roads and an increase in the number

of visitors travelling in vehicles in Riverstone.

Acknowledgments

The Department of Wildlife Conservation for permissions granted (Permit no. WL/3/3/354), U.K.L. Peiris for all

support given. We also wish to thank the Directress of the National Museum of Sri Lanka, S. Kasthuriarachchi, and

Assistant directress L. Somarathne gratefully acknowledging their assistance in museum-reference work. A.

Medawaththa prepared the distribution map, D. Wasala, M. Chathuranga, Y. Rodrigo, Amatha and Gagana

Wickramasinghe are greatly acknowledged for their assistance in the field. We also thank our colleagues at

Herpetological Foundation of Sri Lanka (HFS), and the Institute of Biochemistry Molecular Biology and

Biotechnology (IBMBB) for various courtesies, A. Abayakoon for his encouragements. The UNDP/ GEF small

grant (No: SRL/SGP/OP6/STAR/BD/2018/09) for providing funds to carry out field research work. Nagao Natural

Environment Foundation for partial funding, K. Gunawardena and Canon Metropolitan for providing selected

camera equipment, to A. 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. Finally, we thank our Principal Sponsors

Dilmah Conservation, for funding project activities.

References

Amarasinghe, A.A.T., Karunarathna, D.M.S.S. & Fujinuma, J. (2014a) A new Calotes species from Sri Lanka with a

redescroption of Calotes liolepis Boulenger, 1885. Herpetologica, 70 (3), 323–338.

https://doi.org/10.1655/HERPETOLOGICA-D-13-00087

Amarasinghe, A.A.T., Karunarathna, D.M.S.S., Hallermann, J., Fujinuma, J., Grillitsch, H. & Campbell, P.D. (2014b) A new

species of the genus Calotes (Squamata: Agamidae) from high elevations of the Knuckles Massif of Sri Lanka. Zootaxa,

3785 (1), 59–78.

Bambaradeniya, C.N.B. & Ekanayake, S.P. (2003) A Guide to the Biodiversity of Knuckles Forest Region. IUCN, Colombo, Sri

Lanka. 68 pp.

Boie, F. (1827) Bemerkungen über Merrem’s Versuch eines Systems der Amphibien. Marburg. 1820. Erste Lieferung:

Page 16: 4559 (2): 265 280 Article ... · types which includes; montane, sub-montane, pigmy, semi-evergreen, riverine, rock outcrop, savanna, patana grass land, and scrublands (Bambaradeniya

WICKRAMASINGHE ET AL.280 · Zootaxa 4559 (2) © 2019 Magnolia Press

Ophidier. Isis von Oken, 20 (10): col. 508–566.

Boulenger, G.A. (1890) The Fauna of British India, Including Ceylon and Burma. Reptilia and Batrachia. Taylor & Francis,

London, xviii + 541 pp.

Boulenger, G.A. (1904) Description of a new snake (Aspidura drummondhayi). Spolia Zeylanica, 2 (7), 95–96.

Cope, E.D. (1860) Catalogue of Colubridae in the Museum of the Academy of Natural Sciences of Philadelphia, I.

Calamarinae. Proceedings of the Academy of Natural Sciences of Philadelphia, 74–79.

de Silva, P.H.D.H. (1980) Snake Fauna of Sri Lanka, with special reference to skull, dentition and venom in snakes. National

Museums of Sri Lanka, Colombo, 472 pp.

Ferguson, W. (1876) Description of a new snake of the genus Aspidura from Ceylon. Proceedings of the Zoological Society of

London, 819–820.

https://doi.org/10.1111/j.1096-3642.1876.tb02619.x

Fernando, S.S., Wickramasinghe, L.J.M. & Rodirigo, R.K. (2007) A New Species of Endemic Frog Belonging to Genus

Nannophrys Gunther, 1869 (Anura: Dicroglossinae) from Sri Lanka. Zootaxa, 1403 (1), 55–68.

https://doi.org/10.11646/zootaxa.1403.1.3

Gans, C. & Fetcho, J.R. (1982) The Sri Lankan genus Aspidura (Serpentes, Reptilia, Colubridae). Annals of the Carnegie

Museum, 51 (14), 271–316.

Günther, A. (1858) Catalogue of Colubrine snakes of the British Museum. London, xvi + 281 pp.

Günther, A. (1864) The Reptiles of British India. Taylor & Francis, London, xxvii + 452 pp.

Meegaskumbura, M. & Manamendra-Arachchi, K. (2005) Description of eight new species of Shrub-frogs (Ranidae:

Rhacophorinae: Philautus) from Sri Lanka. The Raffles Bulletin of Zoology, 12, 305–338.

Meegaskumbura, M. & Manamendra-Arachchi, K. (2011) Two new species of shrub frogs (Rhacophoridae: Pseudophilautus)

from Sri Lanka. Zootaxa, 2747, 1–18.

Rajapaksha, D.R.N.S., Samarawickrma, V.A.M.P.K. & Ranawana, K.B. (2006) Herpetofaunal diversity in Dothalugala Man

and Biosphere Reserve, of the Knuckles forest range, Sri Lanka. Russian Journal of Herpetology, 13 (2), 120–134.

Samarawickrama, V.A.M.P.K., Ranawana, K.B., Rajapaksha, D.R.N.S, Ananjeva, N.B., Orlov, N.L., Ranasinghe, J.M.A.S. &

Samarawickrama, V.A.P. (2006) A new species of the genus Cophotis (Squamata: Agamidae) from Sri Lanka. Russian

Journal of Herpetology, 13 (3), 207–214.

Samarawickrama, V.A.M.P.K., Samarawickrama, D.R.N.S. & Kumburegama, S. (2012) Herpetofauna in the Kaluganga upper

catchment of the Knuckles Forest Reserve, Sri Lanka. Amphibian and Reptile Conservation, 5 (2), 81–89.

Smith, M.A. (1943) The Fauna of British India, Ceylon and Burma, Including the Whole of the Indo-Chinese Sub-Region.

Reptilia and Amphibia. Vol. II. Serpentes. Taylor and Francis, London, xii + 583 pp.

Wagler, J.G. (1830) Natürliches System der Amphibien, mit vorangehender Classification der Saugthiere und Vogel. Ein

Beitrag zur vergleichenden Zoologie. J.G. Cotta’schen Buchhandlung. Munchen, Stuttgart und Tubingen, vi + 354 pp.

Wall, F. (1921) Ophidia Taprobanica or the Snakes of Ceylon. Government Printer, Colombo, 581 pp.

https://doi.org/10.1201/b16901

Wickramasinghe, L.J.M., Bandara, I.N., Vidanapathirana, D.R., Tennakoon, K.H., Samarakoon, S.R. & Wickramasinghe, N.

(2115) Pseudophilautus dilmah, a new species of shrub frog (Amphibia: Anura: Rhacophoridae) from a threatened habitat

Loolkandura in Sri Lanka. Journal of Threatened Taxa, 7 (5), 7089–7110.

Wickramasinghe, L.J.M., Vidanapathirana, D.R., Kandambi, Pyron, R.A. & Wickramasinghe, N. (2017) A new species of

Aspidura Wagler, 1830 (Squamata: Colubridae: Natricinae) from Sri Pada sanctuary (Peak Wilderness), Sri Lanka.

Zootaxa, 4347 (2), 275–292.

https://doi.org/10.11646/zootaxa.4347.2.4

APPENDIX 1. Comparative material examined.

Aspidura ravanai. Holotype NMSL-NH 2017.19.01 (male); Paratypes DWC 2017.05.01 (female), DWC 2017.05.02 (male),

Sri Pada upper region, Ratnapura district, Sabaragamuwa province, Sri Lanka.

Aspidura trachyprocta. NMSL-NH 2017.20.01 (male), DWC 2017.05.03 (male), Pidurutalagala lower region, Nuwara Eliya

district, Central province, Sri Lanka; DWC 2017.05.04 (female), Sri Pada upper region, Ratnapura district, Sabaragamuwa

province, Sri Lanka.