Cnemaspis Revision- Sri Lanka

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Kelum Manamendra-Arachchi * , Sudesh Batuwita * & Rohan Pethiyagoda *,**

* Wildlife Heritage Trust of Sri Lanka, 95 Cotta Road, Colombo 8, Sri Lanka.** Author for correspondence.

Abstract

All nominal species assigned to the diurnal geckos of the genusCnemaspis Strauch, 1887 from SriLanka are reviewed and re-described on the basis of their name-bearing types. Based on a recentsurvey, eleven new species are described from Sri Lanka:C. clivicola, C. kallima, C. latha, C. amith,C. menikay, C. pava, C. phillipsi, C. pulchra, C. punctata, C. silvula and C. upendrai. A total of 21 validspecies of Cnemaspisare now recognized from the island, all of them endemic. Two speciesC. amith and C. tropidogasterare not presently known from extant populations. A neotype is designated forGymnodactylus scalpensisFerguson, 1877 (type locality Gannoruwa Forest, near Kandy) and lectotypesselected forGymnodactylus kandiana Kelaart, 1852, andGonatodes kandianus var. tropidogaster Boulenger,1885 (the type locality of the latter, which is not yet known from an extant population, is restrictedto Sri Lanka).Cnemaspis ranwellai Wickramasinghe, 2006, is shown to be a junior objective synonymof C. scalpensis.An examination of type material showsGymnodactylus humei Theobald, 1876, to be a junior subjective synonym of G. kandianus Kelaart, 1852.

The name-bearing types of all nominal peninsular Indian species described prior to 1984 are re-described, and lectotypes selected forGoniodactylus boieiGray, 1842,Gymnodactylus beddomeiTheobald,

1876,G. gracilis Beddome, 1870,G. indicus Gray, 1846,G. ornatus Beddome, 1870,G. wicksiiStoliczka,1873, andG. wynadensis Beddome, 1870. Three new South Indian species discovered among thesehistorical specimens are described:C. monticola(type locality Wynaad, Kerala State);C. australis (typelocality Tirunelveli, Tamil Nadu State); andC. nilagirica (type locality: Nilgiri Hills, Tamil Nadu;formerly a syntype of C. tropidogaster).Cnemaspis wicksii(Stoliczka, 1873) andC. andersonii (Annandale,1904), hitherto buried in the synonymy, are shown to be valid species. Examination of their name- bearing types showsGonatodes bireticulatus Annandale, 1915, to be a junior subjective synonym of Gymnodactylus sisparensis Theobald, 1876. The identities of Gymnodactylus mysoriensis Jerdon, 1853 andG. littoralis Jerdon, 1853, two valid species, remain to be fixed through the designation of neotypesselected from topotypical specimens so as to facilitate future description of new taxa from southernIndia.

Zeylanica, ISSN 1391-6270. October, 2007. Vol. 7, No. 1, pp. 9122, incl. 6 tabs., 46 figs. and 12 pls. 2007, Wildlife Heritage Trust of Sri Lanka.


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MANAMENDRA-ARACHCHI ET AL.

geographical specific name), have historically beenconsidered to have extremely wide ranges, having been recorded from as far afield as the Andamans. Asconcluded by Baueret al. (2007), these were probablyfor the most part misidentified (undescribed) species,

and we hope future workers will describe them.Given the clarity of the present taxonomy, werefrain from providing a discussion of the historicalsynonymy because this is unlikely to be useful tofuture workers.

Materials and methods

Material for this study were collected in Sri Lanka aspart of a survey of the islands Sauria commissioned by the Department of Wildlife Conservation in 1998.Specimens were fixed in 10% formalin and preservedin 70% ethanol; samples of tissue for DNA analysiswere stored in 95% ethanol. The new materialcollected in this survey is deposited in the collectionof the Wildlife Heritage Trust of Sri Lanka (WHT).All available types of Sri Lankan and peninsularIndian Cnemaspis, deposited in The Natural HistoryMuseum, London (BMNH), the National Museum of Sri Lanka, Colombo (NMSL), and Zoological Surveyof India, Calcutta (ZSI), were also examined.

The following measurements were taken withdial Vernier callipers (to the nearest 0.1 mm): AG,distance between axilla and groin; ED, eye diameter(horizontal diameter of orbit); EN, distance betweenanteriormost point of orbit and middle of nostril; ES,snout length (distance between anteriormost point of orbit and tip of snout); FEL, femur length (distance between groin and knee); FL, finger length (distance between tip of claw and the nearest fork); FOL, footlength (distance between heel and tip of longest toe,with both foot and tibia flexed); HL, head length(distance between posterior edge of mandible andtip of snout); HW, head width (maximum width of head); IN, internarial distance (least distance betweeninner margins of nares); IO, interorbital width (leastdistance between upper margins of orbits); LAL,lower-arm length (distance from elbow to wrist with

both upper arm and palm flexed); MBE, distance between angle of gape and posterior-most pointof orbit; MFE, distance between angle of gape andanteriormost point of orbit; MN, mandible to nostrildistance (distance between angle of gape and middleof nostril); PAL, palm length (distance betweenposterior-most margin of palm and tip of longestfinger); SVL, snoutvent length (distance betweentip of snout and anterior margin of vent); TAL, taillength (distance between anterior margin of vent totail tip); TBL, tibia length (distance between knee

and heel, with both tibia and tarsus flexed); TBW,tail base width; TL, toe length (distance betweentip of claw and nearest fork); TYD, tympanum-eyedistance (distance between anterior-most marginof tympanum and posterior-most margin of eye);

TYE, tympanum diameter; UAL, upper-arm length(distance between axilla and angle of elbow); UEW,supercilium width (distance between supraciliaryscale row and lateral bony margin of frontal).

Scale counts and morphological observationswere made using an Olympus SZ40 dissectingmicroscope. Meristic characters were taken asfollows: supralabials and infralabials, first labial scaleto last labial scale towards gape, which is distinctlylarger than the granular scales at gape; subdigitallamellae, from first proximal enlarged scansor greaterthan twice width of the largest palm scale, to distal-most lamella at tip of digit; ventrals, from mental toa scale on preanal scale row, separately for each sex.Sex was determined by the presence of hemipenial bulges and preanal and/or femoral pores. Caudal-scale characters are given only for specimens withunregenerated tails (or unregenerated portions of tails), except in the case of type specimens in whichthe tails have been lost or regenerated.

Photographs were taken with a Cannon IXUS50 digital camera. Colour photographs are of livingspecimens unless otherwise stated. Altitudes aregiven in metres above mean sea level; geographicco-ordinates were taken using a Magellan 12-channel GPS (geodetic datum WGS-84: WorldGeodetic System of 1984), and in some instances,using topographic maps (1 inch: 1 mile, SurveyDepartment, Colombo).

The identification of sympatric species (except forCnemaspis), is based on Smith (1935), Taylor (1950,1953), Deraniyagala (1953), Greer (1991), Batuwita &Bahir (2005), Bahir & Maduwage (2005), Bahir & Silva(2005); nomenclature follows Kluge (2001).

This work necessitated examination of the typespecimens in the Zoological Survey of India, Kolkata(ZSI). This was done during a visit to that institution by K.M.-A. in February, 2007. The ZSIs collection is

of immense historical and scientific interest, and itis regrettable that the custodians of this collectionpermitted access to the specimens only after repeatedrequests, and even then for less than three hours perday. Most frustratingly, it was prohibited to photo-graph specimens. Such arbitrary restrictions sub-vert sound science are a sad indictment on a oncefamous institution; they also call into question theresponsiveness of ZSI to Recommendation 72F of the International Code of Zoological Nomenclature(1999).


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REVISION OF CNEMASPIS

TAXONOMY

Gekkonidae: Cnemaspis

PENINSULAR INDIAN SPECIES

Cnemaspis boiei (Gray, 1842)Goniodactylus boieiGray, 1842

(Figures 1a h; Table 2; Plate 1a, b)

Lectotype (here designated). BMNH 1962.181A(male), 35.3 mm SVL, India, Presented by GeneralT. Hardwicke.

Paralectotypes. BMNH 1962.181B (female), 38.4mm SVL; BMNH 1962.181C (female), 38.8 mm SVL;BMNH 1962.181D (male), 30.7 mm SVL, India,presented by General T. Hardwicke.

DiagnosisCnemaspis boiei differs from its peninsular Indianand Sri Lankan congeners by a combination of the following characters. Maximum SVL 38.8 mm;paired postmentals separated by a medial scale; eachpostmental bounded by 4 scales; ventrals, 117; dorsalscales homogeneous; spine-like tubercles absent onflank; gular, pectoral and abdominal scales smooth;pectoral and abdominal scales juxtaposed; no femoralor preanal pores in either sex; subcaudals on medianrow enlarged, subequal; enlarged tubercles absent on base of tail; supralabials to angle of jaws, 6; subdigitallamellae on digit IV of pes, 17. Description of lectotypeAn adult male, 35.3 mm SVL (for measurements, seeTable 2). Head relatively short (HL 25.2% of SVL),moderately broad (HW 15.2% of SVL, HW 60.6% of HL), distinct from neck. Snout length greater thaneye diameter (ED 76.7% of ES); interorbital broad(IO 15.7% of HL); pupil rounded. Granules on snoutsmooth, equal in size, larger than those of occipitalregion; scales on interorbital and superciliumsmooth; scales on ventral surface of head conical, not

keeled. Rostral scale in contact with first supralabial,partially divided by a medial groove: a narrowtransverse suture begins from medial groove (Fig.1a). Nares separated medially by two enlargedsupranasals and 2 small internasals (Fig. 1a). Naresrounded, dorsally orientated, not in contact withfirst supralabial. Three postnasals, the lower onein broad contact with first supralabial on left side(Fig. 1a). Mental subpentagonal, wider than long,posteriolaterally in contact with a single elongate,enlarged, postmental and a single subpentagonal

medial scale (on left side; right side damaged);postmentals widely separated; postmental onleft side bordered posteriorly by 4 smooth scales,including a medial scale (probably 8 scales borderingpostmentals) (Fig. 1b). Supralabials 6, infralabials

6, decreasing in size towards gape (Fig. 1c). Earopening oval, higher than wide, oblique (TW 50.0%of TH). Scales on ventral surface of neck smooth.Granules between anterior margin of ear openingand posterior margin of eye, 24, larger than those of occipital region.

Body slender, elongate. Mid-dorsal granulesisolated, scattered, homogeneous, keeled (Fig. 1d), alittle larger than scales of occipital region; spine-liketubercles absent on flank; abdominal scales smooth(Fig. 1e). Ventrals, 117. Pectoral and abdominal scales juxtaposed (Fig. 1e), those around vent and base of tail smooth, subimbricate to imbricate (Fig. 1 f ). Nopreanal or femoral pores (Fig. 1 f ).Forelimbs moderately long, slender (LAL 13.5%SVL, UAL 13.0% SVL); hind limbs long (TBL 16.9%SVL, FEL 18.6% SVL); tibia short. Dorsal scales on both fore and hind limbs smooth; ventral scales onupper arm narrow, pointed, smooth; on lower armsmooth. Scales on anterior surface of thigh small,granular, subequal to those on ventral surface (Fig.1 f ). Scales on dorsal and ventral surface of shanksmooth. Scales on manus and pes smooth, rounded.Digits elongate, slender, all bearing slightly recurvedclaws; subdigital lamellae entire; subdigital lamellaeon finger I, 9, finger II, 12, finger III, 14, finger IV,14, finger V, 12; toe I, 8, toe II, 10, toe III, 15, toe IV,17, toe V, 13; interdigital webbing absent; relativelength of digits (fingers): IV (3.6 mm) > III (3.2 mm)> V (2.7 mm) > II (2.5 mm) > I (1.8 mm); (toes) IV(4.0 mm) > III (3.9 mm) > II (3.4 mm) > V (3.0 mm)> I (2.2 mm).

Tail subcylindrical, tapering, its length greaterthan SVL (TAL 129.7% of SVL); tail base distinctlyswollen. Enlarged tubercles absent on base of tail.Dorsal scales on mid-tail smooth, imbricate directed backwards (Fig. 1 g); post-cloacal spurs absent;subcaudals smooth, hexagonal (some scales divided),

in a regular enlarged median series (Fig. 1h).Coloration. In preservative: (see Plate 1a, b) dorsallyand ventrally olive; no blotches or markings.

Variation. Supralabials 7 in paralectotype BMNH1962.181B. Pores absent in both sexes. SVL range30.7 38.8 mm (4 ex.).

ComparisonsCnemaspis boiei differs from each of its peninsular


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Figure 1. Cnemaspis boiei: BMNH 1962.181A, lectotype, male,a, dorsal aspect of snout (right side damaged);b, ventralaspect of chin (right side damaged);c, lateral aspect of gape, showing arrangement of supralabials and infralabials;d, e,arrangement of scales on dorsal and ventral regions, respectively, of mid-body; f , preanal and right hind-limb regions,showing absence of pores; g, arrangement of scales on dorsal aspect of tail;h, arrangement of subcaudal scales. Scale bars: 1 mm.

a b

c d

e f

g h


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Indian and Sri Lankan congeners, respectively, bythe following opposing suites of characters:

Cnemaspis beddomei: preanal pores 6 9. Cnemaspis goaensis: presence of pores.Cnemaspis littoralis :a black marking on nape.Cnemaspis mysoriensis:

spine-like tubercles present on flank, femoral pores2 or 3.Cnemaspis indraneildasii and C. heteropholis:dorsal scalation heterogeneous.Cnemaspis jerdonii:spine-like tubercles present on flank, presence of femoral pores.Cnemaspis nairi: presence of preanalpores. Cnemaspis otai: 4 preanal and 3 femoral pores.Cnemaspis sisparensis: postmentals in contact, spine-like tubercles absent on flank, 7 or 8 femoral pores. Cnemaspis wynadensis: 5 femoral pores.Cnemaspis

yercaudensis: 2 preanal and 3 femoral pores.Cnemaspis gracilis: pores present, postmentals in contact,subcaudals in median series not enlarged.Cnemaspisindica: postmentals in contact, presence of femoralpores. Cnemaspis ornata: dorsal tubercles arrangedin 15 longitudinal rows, a dark-brown collar presenton nape, preanal pores 68.Cnemaspis wicksiiand C.andersonii: presence of pores.Cnemaspis monticola:spine-like tubercles on flank; pectoral and abdominalscales imbricate.Cnemaspis australis : subcaudals onmedian row pointed, strongly keeled.Cnemaspisnilagirica: pectoral and abdominal scales weaklykeeled; subcaudals pentagonal or quadrate.

Cnemaspis tropidogaster: gular, pectoral andabdominal scales keeled, spine-like tubercles presenton flank.Cnemaspis kandiana,C. podihuna,C. scalpensis, C. gemunu and C. phillipsi: spine-like tubercles presenton flank.Cnemaspis alwisi: 9 femoral pores, spine-liketubercles present on flank.Cnemaspis molligodai: 5preanal and 8 or 9 femoral pores, spine-like tuberclespresent on flank. Cnemaspis samanalensis: gular,pectoral and abdominal scales tricarinate, subcaudalskeeled, spine-like tubercles present on flank.Cnemaspis retigalensis: dorsal scales heterogeneous,gulars carinate, a single preanal pore and 3 or 4femoral pores, spine-like tubercles present on flank;subcaudals in an irregular enlarged median series.Cnemaspis punctata : 5 7 femoral pores, spine-liketubercles present on flank, distinctive spotted dorsal

coloration.Cnemaspis kumarasinghei, C. menikay andC. latha: presence of pores; presence of spine-liketubercles on flank.Cnemaspis silvula, C. pulchra and C. pava: pectoral and abdominal scales keeled, spine-like tubercles present on flank.Cnemaspis clivicola:pectoral and abdominal scales keeled, spine-liketubercles present on flank, subcaudals in medianseries not enlarged.Cnemaspis upendrai: some gularscales tricarinate, pectoral and abdominal scaleskeeled, spine-like tubercles on flank.Cnemaspis kallima: spine-like tubercles present on flank,

subcaudals on distal half of tail carinate.Cnemaspisamith: gular scales carinate; pectoral and abdominalscales imbricate; dorsal granules heterogeneous;subcaudals on median row enlarged, irregular.

RemarksThe precise type locality of Cnemaspis boiei withinIndia is not known. However, it is immediatelyidentifiable in that both males and females lack bothfemoral and preanal pores, a character otherwiseshared only with the northeast-Indian species,C.assamensis Das and Sengupta, 2000.

Cnemaspis indica (Gray, 1846)Gymnodactylus indicus Gray, 1846(Figures 2a h; Table 2; Plate 1c, d)

Lectotype (here designated). BMNH 46.11.22.22b(male), 29.3 mm SVL, Madras.

Paralectotypes. BMNH 46.11.22.22a (male), 30.4 mmSVL; BMNH 46.11.22.22c (female), 30.0 mm SVL(deformed by dehydration), Madras.

DiagnosisCnemaspis indica differs from its peninsular Indianand Sri Lankan congeners by a combination of the following characters. Maximum SVL 30.4 mm;paired postmentals separated by a medial scale; eachpostmental bounded by 3 scales including medialscale; ventrals, 94; dorsal scales homogeneous,imbricate, carinate; spine-like tubercles absent onflank; ventrals smooth, imbricate; no preanal pores;5 femoral pores on each side; subcaudals smooth,median series slightly enlarged; supralabials toangle of jaws, 7; subdigital lamellae on digit IV of pes, 14. Description of lectotypeAn adult male, 29.3 mm SVL (for measurements,see Table 2). Head relatively short (HL 24.9% of SVL), broad (HW 20.8% of SVL, HW 83.5% of HL),distinct from neck. Snout length somewhat greater

than eye diameter (ED 66.6% of ES); interorbital broad (IO 16.4% of HL); pupil rounded. Granuleson snout intermixed with both smooth and keeledscales, larger than those of occipital region; scaleson interorbital and supercilium granular, keeled,those on ventral surface of head weakly carinate.Rostral scale partially divided by a medial groove,in contact with first supralabial. Nares separated by two supranasals and a single internasal scale;internasal as large as supranasals (Fig. 2a). Naresrounded, dorsally orientated, not in contact with


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Figure 2. Cnemaspis indica: BMNH 46.11.22.22b, lectotype, male,a, dorsal aspect of snout;b, ventral aspects of chin;c,lateral aspect of gape, showing arrangement of supralabials and infralabials;d, e, arrangement of scales on dorsal andventral regions, respectively, of mid-body; f , preanal and left hind limb regions, showing arrangement of femoral pores;

g, arrangement of scales on dorsal aspect of tail;h, arrangement of subcaudal scales. Scale bars: 1 mm.

a b

c d

e f

g h


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first supralabial. Two postnasals, the lower one in broad contact with first supralabial (Fig. 2a). Mentalsubpentagonal, as wide as long, posteriolaterallyin contact with two enlarged postmentals andmedially with a single hexagonal postmental scale;

postmentals bordered posteriorly by 5 smooth scalesincluding medial scale (Fig. 2b). Supralabials 7,infralabials 7, decreasing in size towards gape (Fig.2c). Ear opening circular (TW 100.0% of TH). Scaleson ventral surface of neck smooth. Nineteen granules between anterior margin of ear opening and posteriormargin of eye.

Body robust, short. Mid-dorsal granuleshomogeneous, imbricate and carinate (Fig. 2d),intermixed with a few smooth scales; tuberclesabsent on flank; abdominal scales smooth, imbricate,obtusely pointed and wider than long (Fig. 2e).Ventrals, 94; scales around vent and base of tailsmooth, imbricate. No preanal pores; 5 femoralpores (Fig. 2 f ).

Forelimbs relatively short, slender (LAL 12.2%SVL, UAL 9.8% SVL); hind limbs moderately long(TBL 18.4% SVL, FEL 18.7% SVL); tibia short. Dorsalscales on both fore and hind limbs with 13 keel-like ridges; ventral scales on upper and lower armsmooth, a few of them carinate. Scales on anteriorsurface of thigh carinate, those on posterior surfacesmooth (Fig. 2 f ). Scales on dorsal and ventral surfaceof shank mostly smooth, a few carinate. Dorsalscales on manus carinate, ventrals smooth. Digits

elongate, slender, all bearing slightly recurved claws;subdigital lamellae unnotched, entire, except for 2or 3 fragmented scales at base; subdigital lamellaeon finger I, 8, finger II, 11, finger III, 11, finger IV, 11,finger V, 9; toe I, 9, toe II, 12, toe III, 11, toe IV, 14,toe V, 9; interdigital webbing absent; relative lengthof digits (fingers): III (2.9 mm) > IV (2.5 mm) > II(2.4 mm) > V (2.1 mm) > I (1.8 mm); (toes) > III (3.8mm) > IV (3.7 mm) > II (3.3 mm) > V (3.0 mm) > I(2.1 mm).

Tail tapering (damaged); tail base distinctlyswollen. Enlarged tubercles absent on base of tail.Dorsal scales on mid-tail imbricate, carinate, directed

backwards (Fig. 2 g); a conical post-cloacal spurpresent on each side; subcaudals smooth, those onmedian row slightly enlarged (Fig. 2h).

Variation. Five infralabials and 2 internasals inparalectotype BMNH 46.11.22.22a; 35 femoral pores;pores absent in females; female paralectotype BMNH46.11.22.22c with enlarged endolymphatic sacs,suggestive of the specimen having been collectedduring the breeding season; SVL range 29.330.4mm (3 ex.).

Coloration. In preservative: (see Plate 1c, d) dorsallylight brown with indistinct dark marking on dorsum;tail with a longitudinal mid-dorsal stripe; venterpinkish-white.

Comparisons. Cnemaspis indica differs from each of its peninsular Indian and Sri Lankan congeners bythe following opposing suites of characters:

Cnemaspis beddomei: no interanasals, preanal pores6 9, no femoral pores.Cnemaspis goaensis: preanalpores present. Cnemaspis littoralis : body slender,dorsal scales granular, median series of subcaudalsenlarged, hexagonal.Cnemaspis mysoriensis: spine-like tubercles present on flank, femoral pores 2 or3. Cnemaspis indraneildasii, C. wicksii, C. monticola and C. andersonii: spine-like tubercles present onflank. Cnemaspis heteropholis: heterogeneous dorsalscalation. Cnemaspis jerdonii: spine-like tuberclespresent on flank.Cnemaspis nairi: no femoral pores.Cnemaspis otai and C. sisparensis: paired postmentalsin contact. Cnemaspis wynadensis: subcaudals ina regular enlarged (widened) series.Cnemaspis

yercaudensis: 2 preanal pores.Cnemaspis boiei: no preanal or femoral pores.Cnemaspis gracilis:postmentals in contact.Cnemaspis ornata: dorsaltubercles arranged in 15 longitudinal rows, dark- brown collar on nape, no femoral pores, preanalpores 6 8. Cnemaspis australis : subcaudals pointed,strongly keeled.Cnemaspis nilagirica: dorsal scalesisolated, scattered; subcaudals on median rowenlarged, subequal.

Cnemaspis tropidogaster: gular, pectoral andabdominal scales keeled, spine-like tuberclespresent on flank.Cnemaspis kandiana, C. podihuna,C. scalpensis, C. gemunu and C. phillipsi: spine-liketubercles present on flank.Cnemaspis alwisi: 9femoral pores, spine-like tubercles present on flank.Cnemaspis molligodai: 5 preanal and 8 or 9 femoralpores, spine-like tubercles present on flank.Cnemaspissamanalensis: gular, pectoral and abdominal scalestricarinate, subcaudals keeled, spine-like tuberclespresent on flank. Cnemaspis retigalensis: dorsalscales heterogeneous, gulars carinate, pectoral and

abdominal scales smooth, a single preanal pore and3 or 4 femoral pores, spine-like tubercles presenton flank. Cnemaspis punctata : 5 7 femoral pores,spine-like tubercles present on flank, distinctivespotted dorsal Coloration.Cn emaspis kumarasinghei, C. menikay and C. latha: presence of preanal pores,tubercles present.Cnemaspis silvula,C. pulchra,C. pava and C. clivicola: pectoral and abdominal scales keeled,spine-like tubercles present on flank.Cnemaspisupendrai: some gular scales tricarinate, pectoral andabdominal scales keeled, spine-like tubercles present


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on flank. Cnemaspis kallima: gular scales smooth,spine-like tubercles present on flank; subcaudalson distal half of tail carinate.Cnemaspis amith: gularscales carinate; pectoral and abdominal scalessmooth; dorsal granules heterogeneous; subcaudals

on median row enlarged, irregular.RemarksCnemaspis indica is distinguished from all its penin-sular Indian and Sri Lankan congeners by possessingimbricate dorsal scales.

Cnemaspis jerdonii (Theobald, 1868)Gymnodactylus jerdonii Theobald, 1868

Syntypes. ZSI 6179 (adult), 9.6 mm HL; ZSI 6180(juvenile), 3.4 mm HL, no specific locality; bothspecimens too badly damaged and fragmented tofacilitate measurements.

Note. The syntypes of this species are in too poora state to facilitate detailed description, for whichreason we also refrain from selecting a lectotype.Fortunately, Theobalds original description isremarkably detailed for its time: the followingdiagnosis and description, based in part on thesyntypes, is supplemented by data provided in theoriginal description.

DiagnosisCnemaspis jerdonii differs from its peninsular Indianand Sri Lankan congeners by a combination of the following characters. Maximum SVL 38.1 mm;paired postmentals separated by a medial scale;each postmental bounded by 2 scales includingmedial scale; dorsal scales homogeneous; spine-liketubercles present on flank; ventrals rhomboid; noerect spines laterally on tail; tail laterally keeled; scales on dorsal and ventral surface of neck imbricate,smooth; dorsal scales on both fore and hind limbsimbricate. Preanal pores absent; 8 femoral pores oneach side; median series of subcaudals enlarged;supralabials and infralabials to angle of jaws, 9 and

7 respectively.Redescription of syntype ZSI 6179Pupil rounded. Granules on snout smooth; scaleson interorbital, supercilium and ventral surface of head, smooth. Mental subpentagonal, wider thanlong, posteriolaterally in contact with two enlargedpostmentals, medially with a single hexagonalpostmental scale; postmentals bordered posteriorly by 5 smooth scales including medial scale. Earopening elongate, oblique. Scales on dorsal and

ventral surface of neck smooth, imbricate. Dorsalsurfaces of both fore and hind limbs covered byimbricate scales.

This is consistent with the original description of C. jerdonii (Theobald, 1868), which was as follows:

Male. Back uniformly granular. Tail with a singlerow of large subcaudal plates or scuta, with a fewlarge scales along their edges. A strong double foldon the throat. Scales of the belly small, rhomboid.Along the sides two rows of distant erect spines, become obsolete on the tail. Sides keeled. Pubicregion covered with same scales as the belly. No largepranals. Femoral pores eight on each thigh; large,cup-shaped, placed along the hind edge of limb, bordered in font by rather largish scales, and behind by the granular scales of the back. Length, headand body 1.50, tail 1.50==3.00. Colour (in spirits)above, greenish grey mottled with brown, beneath,greenish white. Nine upper and seven lower labials.Nostrils close behind the rostral. Lower rostral large, just separates a pair of small triangular chin plates.Allied toG. Mysoriensis by its spines, but not to beidentified with any species described by Jerdon(Theobald, 1868).

Comparisons. Cnemaspis jerdonii differs from eachof its peninsular Indian and Sri Lankan congeners,respectively, by the following opposing suites of characters:

Cnemaspis beddomei: spine-like tubercles absenton flank, preanal pores 69, femoral pores absent.Cnemaspis goaensis: postmentals widely separated bya medial scale.Cnemaspis littoralis: a black markingon nape. Cnemaspis mysoriensis: femoral pores 2or 3. Cnemaspis indraneildasii: paired postmentalsseparated by three small scales.Cnemaspis indica,C. nairi , C. otai, C. sisparensis and C. yercaudensis:spine-like tubercles absent on flank.Cnemaspis boiei: no preanal or femoral pores.Cnemaspis gracilis:postmentals in contact.Cnemaspis ornata: dorsaltubercles arranged in 15 longitudinal rows, dark- brown collar on nape, femoral pores absent, preanalpores 6 8. Cnemaspis wicksii, C. monticola and C.

andersonii: gular scales carinate.Cnemaspis australis :subcaudals pointed, strongly keeled, in a regularenlarged median series.Cnemaspis nilagirica: gularscales carinate; enlarged caudal tubercles present.

Cnemaspis tropidogaster: gular, pectoral andabdominal scales keeled.Cnemaspis kandiana and C.menikay: gular scales carinate.Cnemaspis podihuna:preanal pores present. Cnemaspis scalpensis, C.

gemunu and C. phillipsi: 12 16 femoral pores.Cnemaspis alwisi: caudal tubercles present.Cnemaspismolligodai: 5 preanal pores.Cnemaspis samanalensis:


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gular, pectoral and abdominal scales tricarinate,subcaudals keeled. Cnemaspis retigalensis: dorsalscales heterogeneous, gulars carinate.Cnemaspis

punctata : 5 7 femoral pores, distinctive spotted dorsalcoloration. Cnemaspis kumarasinghei and C. latha:

subcaudal scales in an enlarged irregular medianseries. Cnemaspis silvula, C. pulchra, C. pava andC. clivicola: pectoral and abdominal scales keeled.Cnemaspis upendrai: some gular scales tricarinate;pectoral and abdominal scales keeled.Cnemaspis kallima: subcaudals on distal half of tail carinate. Cnemaspis amith: gular scales carinate; dorsalgranules heterogeneous.

Cnemaspis gracilis (Beddome, 1870)Gymnodactylus gracilis Beddome, 1870

(Figures 3a g; Table 2; Plate 1e, f )

Lectotype (here designated). BMNH 74.4.29.393(male), 29.7 mm SVL, Palghat Hills (Kerala State,India), coll. Colonel R. H. Beddome.

Paralectotypes. BMNH 74.4.29.394 (female), 32.9mm SVL; BMNH 74.4.29.395 (female), 27.9 mm SVL;BMNH 74.4.29.396 (female), 25.4 mm SVL; BMNH74.4.29.397 (male), 30.6 mm SVL; BMNH 74.4.29.398(male), 27.6 mm SVL, Palghat hills (Kerala State,India), coll. Colonel R. H. Beddome.

DiagnosisCnemaspis gracilis differs from its peninsular Indianand Sri Lankan congeners by a combination of thefollowing characters. Maximum SVL 32.9 mm; pairedpostmentals broadly in contact, each postmental bounded by 2 scales; ventrals, 111; dorsal scalesheterogeneous with large, keeled tubercles in 11 or12 rows on dorsum; spine-like tubercles present onflanks; ventrals smooth, imbricate; 2 preanal pores,separated by 2 unpored scales; 35 femoral poreson each side; subcaudals smooth, imbricate, notenlarged; supralabials to angle of jaws, 7; subdigitallamellae on digit IV of pes, 22.

Description of lectotypeAn adult male, 29.7 mm SVL (for measurements, seeTable 2). Head long (HL 27.9% of SVL), broad (HW20.8% of SVL, HW 74.6% of HL), distinct from neck.Snout length somewhat greater than eye diameter(ED 64.1% of ES); interorbital broad (IO 12.0% of HL);pupil rounded. Granules on snout smooth, largerthan those of occipital region; scales on interorbitaland supercilium conical, granular; scales on ventralsurface of head smooth. Rostral scale partially

divided by a medial groove, in contact with firstsupralabial (Fig. 3a). Nares separated by two slightlyenlarged supranasals and a single internasal scale;internasal extends towards snout tip (Fig. 3a). Naresrounded, dorsally orientated, not in contact with

first supralabial. Two postnasals, the lower one in broad contact with first supralabial (Fig. 3a). Mentalsubtriangular, wider than long, posteriolaterally incontact with two enlarged postmentals; postmentals broadly in contact with each other, borderedposteriorly by 5 smooth scales (Fig. 3b). Supralabials7, decreasing in size towards gape; infralabials 6,decreasing in size towards gape (Fig. 3c). Ear openingoval, higher than wide, oblique (TW 66.6% of TH).Scales on ventral surface of neck smooth. Seventeenscales between anterior margin of ear opening andposterior margin of eye. Conical granules laterally onneck, larger than those of interorbital region.

Body slender, short. Mid-dorsal granulesheterogeneous, keeled or smooth, as large as scalesof occipital region; large, keeled tubercles (Fig. 3d) in11 or 12 rows on dorsum; spine-like tubercles presenton flank; ventrals, 111, imbricate, smooth (Fig. 3e);2 preanal pores separated by 2 unpored scales; 4femoral pores on each side (Fig. 3 f ).

Forelimbs moderately long, slender (LAL 14.4%SVL, UAL 14.8% SVL); hind limbs long (TBL 18.1%SVL, FEL 21.2% SVL); tibia short. Dorsal scales on both fore and hind limbs smooth, a few of themcarinate; ventral scales on upper and lower armsmooth. Dorsal scales on thigh carinate, a few of them smooth. Scales on dorsal and ventral surface of thigh (Fig. 3 f ), shank, manus and pes, smooth. Digitselongate, slender, all bearing slightly recurved claws;subdigital lamellae entire, unnotched; subdigitallamellae on finger I, 10, finger II, 14, finger III, 16,finger IV, 17, finger V, 14; toe I, 10, toe II, 16, toe III,20, toe IV, 22, toe V, 17; interdigital webbing absent;relative length of digits (fingers): IV (2.9 mm) > III(2.7 mm) > V (2.6 mm) > II (2.2 mm) > I (2.0 mm);(toes) IV (3.9 mm) > III (3.5 mm) ~ V (3.5 mm) > II(3.1 mm) > I (2.1 mm).

Tail tapering, with 6 large, conical tubercles at

its base, about as large as tubercles on mid-dorsum.Dorsal scales on mid-tail similar to scales on mid- body, directed backwards (Fig. 3 g). Tail with about6 oval, keeled tubercles separated by 26 scales;a conical post-cloacal spur present on each side;subcaudals smooth, not enlarged, imbricate (Fig.3 f ).

Coloration. In preservative: (see Plate 1e, f ) dorsallypinkish brown; dark spots along mid-dorsum, fromnape to sacrum; two black streaks behind eye; mid-


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Figure 3. Cnemaspis gracilis: BMNH 74.4.29.393, lectotype, male,a, dorsal aspect of snout;b, ventral aspect of chin;c,lateral aspect of gape, showing arrangement of supralabials and infralabials;d, arrangement of scales on dorsal regionof mid-body, showing enlarged tubercles;e, preanal and hind-limb regions, showing arrangement of femoral andpreanal pores; f, arrangement of scales on dorsal aspect of tail, showing whorls of enlarged tubercles; g, arrangement of subcaudal scales of BMNH 74.4.29.394, paralectotype, female. Scale bars: 1 mm.

a b

c d

e

f

g


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dorsal tubercles pale; limb with pale tubercles; digits banded; venter whitish, head with dark streaks.

Variation. Supralabials 6 or 7; infralabials 6 or 7; 35femoral pores; pores absent in females. SVL range

25.4 32.9 mm (7 ex.).ComparisonsCnemaspis gracilis differs from each of its peninsularIndian and Sri Lankan congeners, respectively, by thefollowing opposing suites of characters:

Cnemaspis beddomei: spine-like tubercles absenton flank, no interanasals, subcaudals on median rowenlarged, in a series of alternating wider and narrowerscales, preanal pores 69, no femoral pores.Cnemaspis

goaensis: postmentals widely separate by a medialscale.Cnemaspis littoralis: dorsal scales homogeneous,median series of subcaudals enlarged, hexagonal.Cnemaspis mysoriensis: femoral pores 2 or 3.Cnemaspisindraneildasii: paired postmentals separated by smallscale. Cnemaspis heteropholis: paired postmentalsseparated by three small scales, no post-cloacal spurs.Cnemaspis jerdonii: no preanal pores.Cnemaspis nairi:no femoral pores.Cnemaspis otai: 4 preanal pores.Cnemaspis sisparensis: 7 or 8 femoral pores.Cnemaspiswynadensis: subcaudals enlarged, subequal; no preanalpores, 5 femoral pores, postmentals widely separated.Cnemaspis yercaudensis: postmentals separated by asingle scale.Cnemaspis boiei: no preanal or femoralpores, postmentals widely separated by 2 medialscales.Cnemaspis indica: no preanal pores.Cnemaspisornata: dorsal tubercles arranged in 15 longitudinalrows, a dark-brown collar on nape, no femoral pores,preanal pores 6 8. Cnemaspis wicksii, C. andersoniiand C. monticola: gulars carinate.Cnemaspis australis :subcaudals pointed, strongly keeled, those on medianrow of slightly enlarged.Cnemaspis nilagirica: dorsalscales homogeneous; pectoral and abdominal scalesweakly keeled; subcaudals on median row enlarged,subequal.

Cnemaspis tropidogaster: gular, pectoral andabdominal scales keeled.Cnemaspis kandiana andC. menikay: gulars carinate.Cnemaspis podihuna, C.

scalpensis, C. gemunu and C. phillipsi, homogeneousdorsal scalation.Cnemaspis alwisi, 9 femoral pores.Cnemaspis molligodai: 5 preanal and 8 or 9 femoralpores, dorsal scales homogeneous; subcaudalson median row enlarged, subequal.Cnemaspissamanalensis: gular, pectoral and abdominal scalestricarinate, subcaudals keeled.Cnemaspis retigalensis:gulars carinate, a single preanal pore and 3 or 4 femoralpores; subcaudals on median row enlarged, irregular.Cnemaspis punctata : 5 7 femoral pores; distinctivespotted dorsal coloration.Cnemaspis kumarasinghei

and C. latha: postmentals separated medially by asmall scale.Cnemaspis silvula, C. pulchra, C. pava andC. clivicola: pectoral and abdominal scales keeled.Cnemaspis upendrai: some gular scales tricarinate,pectoral and abdominal scales keeled.Cnemaspis

kallima: subcaudals on distal half of tail carinate. Cnemaspis amith: gular scales carinate; subcaudalson median row enlarged, irregular.

RemarksKluge (2001) listedCnemaspis gracilis (Beddome, 1870)(=Gymnodactylus gracilis) in synonymy of C. kandiana(Kelaart, 1852). Examination of the respective typesshow that they are clearly two distinct taxa (seerespective diagnoses and comparisons).

Cnemaspis ornata (Beddome, 1870)Gymnodactylus ornatus Beddome, 1870

(Figures 4a j; Table 2; Plate 2c f )

Lectotype (here designated). BMNH 74.4.29.400(male), 42.0 mm SVL, South Tinnevelly Hills(Tirunelveli, Tamil Nadu State, India).

Paralectotypes. BMNH 74.4.29.401 (male), 41.7mm SVL; BMNH 74.4.29.402 (male), 37.6 mmSVL; BMNH 74.4.29.403 (juvenile), 25.6 mm SVL;BMNH 74.4.29.404 (juvenile), 26.6 mm SVL; BMNH74.4.29.405 (female), 37.2 mm SVL; BMNH 74.4.29.406(juvenile), 25.6 mm SVL; BMNH 74.4.29.407 (juvenile),26.7 mm SVL; BMNH 74.4.29.408 (juvenile), 27.4 mmSVL; BMNH 74.4.29.409 (juvenile), 22.6 mm SVL,South Tinnevelly Hills (Tirunelveli, Tamil NaduState, south-western India).

DiagnosisCnemaspis ornata differs from its peninsular Indianand Sri Lankan congeners by a combination of thefollowing characters. Maximum SVL 42.0 mm; pairedpostmentals widely separated by a medial scale; eachpostmental bounded by 3 or 4 scales including medialscale; ventrals, 161; dorsal scales heterogeneous withenlarged tubercles; spine-like tubercles absent on

flank; ventrals smooth, subimbricate; 68 preanalpores; no femoral pores; subcaudals smooth,median series with individual large, divided scalesalternating with pairs of smaller undivided ones;supralabials to angle of jaws, 79; subdigital lamellaeon digit IV of pes, 30.

Description of lectotypeAn adult male, 42.0 mm SVL (for measurements, seeTable 2). Head long (HL 29.0% of SVL), broad (HW20.0% of SVL, HW 68.8% of HL), distinct from neck.


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Snout length greater than eye diameter (ED 59.0% of ES); interorbital broad (IO 11.4% of HL); pupil rounded.Granules on snout smooth, larger than those of occipitalregion; scales on interorbital and supercilium granular;scales on ventral surface of head smooth. Rostral

scale partially divided by a medial groove, in contactwith first supralabial (Fig. 4a). Nares separated bytwo enlarged scales, in contact with supranasals; nointernasals (Fig. 4a). Nares rounded, dorsally orientated,not in contact with first supralabial. Two post-nasals,the upper one enlarged, the lower one in contact withfirst supralabial (Fig. 4a). Mental subpentagonal, widerthan long, truncate posteriorly; mental posteriolaterallyin contact with two enlarged postmentals, mediallywith a single hexagonal postmental scale; postmentalswidely separate, bordered posteriorly by 6 smoothscales including a medial scale (Fig. 4b). Supralabials8, infralabials 8, decreasing in size towards gape (Fig.4c). Ear opening elliptical, much higher than wide,oblique (TW 12.5% of TH). Dorsal and lateral scaleson neck granular, intermixed with enlarged tubercles.Twenty-three granules between anterior margin of earopening and posterior margin of eye. A large conicalscale laterally on mid-neck.Body robust, short. Mid-dorsal granules isolated,scattered, not keeled, intermixed with large tubercles(Fig. 4d); spine-like tubercles on flank; ventrals, 161,subimbricate, longer than wide, smooth (Fig. 4e); 6preanal pores; no femoral pores (Fig. 4 f ).

Forelimbs long, slender (LAL 16.9% SVL, UAL13.0% SVL); hind limbs moderately long (TBL 18.3%SVL, FEL 18.3% SVL); tibia short. Dorsal scales on both fore and hind limbs smooth; ventral scales onupper arm and lower arm smooth. Scales on thighdorsally and ventrally smooth (Fig. 4 f ). Scales ondorsal and ventral surface of shank smooth. Scaleson manus and pes smooth, rounded. Digits elongate,slender, all bearing slightly recurved claws (Fig. 4 j);subdigital lamellae fragmented on proximal portion;an enlarged scansor near middle of digits, more thantwice width of other lamellae; 2 or 3 fragmentedscales behind each enlarged scansor; distal lamellaeentire (Fig. 4i); subdigital lamellae on finger I, 21,

finger II, 21, finger III, 28, finger IV, 29, finger V, 22;toe I, 14, toe II, 23, toe III, 28, toe IV, 30, toe V, 27;interdigital webbing absent; relative length of digits(fingers): IV (5.0 mm) III (4.6 mm) > V (4.2 mm) ~ II(4.2 mm) > I (2.7 mm); (toes) IV (6.0 mm) > III (5.1mm) ~ V (5.1 mm) > II (4.5 mm) > I (2.8 mm).

Tail (damaged), tapering; tail base distinctlyswollen. Enlarged tubercles absent at base of tail.Dorsal scales on mid-tail imbricate, smooth, directed backwards, with 26 rows of enlarged tubercles(Fig. 4 g); one conical post-cloacal spur present

on each side (Fig. 4 f ). Subcaudals on median rowenlarged, smooth, with individual large, dividedscales alternating with pairs of smaller undividedones (Fig. 4h).

Variation. Supralabials 7 9; infralabials 7 or 8;postmentals bordered posteriorly by 5 smooth scalesin BMNH 74.4.29.401 and in BMNH 74.4.29.402; 68preanal pores; pores absent in females. SVL range26.6 42.0 mm (9 ex.).

Coloration. In preservative: colour variable (see Plate2c f ), dorsally pinkish brown, with a pair of darkspots along mid-dorsal line; several pale spots forman uninterrupted mid-dorsal line behind and betweendark spots in juveniles. A pale chevron on head, witha dark spot at its angle; another pale semicircularmarking originates behind each eye and extendsto angle of chevron. Neck with a pale band (whichprecedes a darker band in some adult specimens).Venter dusky white with dark markings.

Comparisons. Cnemaspis ornata differs from eachof its peninsular Indian and Sri Lankan congeners,respectively, by the following opposing suites of characters:

Cnemaspis beddomei: subdigital lamellae onproximal series entire.Cnemaspis goaensis: presenceof femoral pores.Cnemaspis littoralis: body slender,dorsal scales homogeneous, subcaudals in medianseries enlarged, hexagonal.Cnemaspis mysoriensis:femoral pores 2 or 3.Cnemaspis indraneildasii: irregularrows of dorsal tubercles.Cnemaspis heteropholis:enlarged conical tubercles absent on neck, no post-cloacal spurs.Cnemaspis jerdonii: no preanal pores.Cnemaspis nairi: subdigital lamellae on proximal seriesentire. Cnemaspis otai: paired postmentals in broadcontact, 4 preanal and 3 femoral pores.Cnemaspissisparensis: postmentals in contact, no preanal pores, 7or 8 femoral pores.Cnemaspis wynadensis: subcaudalsenlarged, subequal; no preanal pores, 5 femoralpores.Cnemaspis yercaudensis: 2 preanal and 3 femoralpores. Cnemaspis boiei: no preanal or femoral pores.

Cnemaspis gracilis: postmentals in contact.Cnemaspisindica: postmentals in contact.Cnemaspis wicksii, C.andersonii and C. monticola: gulars carinate, spine-like tubercles present on flank.Cnemaspis australis :subcaudals pointed, strongly keeled scales.Cnemaspisnilagirica: dorsal scales homogeneous; pectoral andabdominal scales weakly keeled; subcaudals onmedian row enlarged, subequal.

Cnemaspis tropidogaster: gular, pectoral andabdominal scales keeled.Cnemaspis kandiana, C.

podihuna, C. scalpensis, C. gemunu, C. phillipsi, C. men-


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Figure 4. Cnemaspis ornata: BMNH 74.4.29.400, lectotype, male,a, dorsal aspect of snout;b, ventral aspect of chin;c,lateral aspect of gape, showing arrangement of supralabials and infralabials;d, e, arrangement of scales on dorsal andventral regions, respectively, of mid-body; f , preanal and left hind-limb regions, showing arrangement of preanal pores;

g, arrangement of scales on dorsal aspect of tail;h, arrangement of subcaudal scales;i, ventral aspect of digit IV of leftpes; j, lateral aspect of claw of digit IV of left pes. Scale bars: 1 mm.

a b

c d

e f g

h i j


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ikay, C. alwisi and C. molligodai: presence of femoralpores. Cnemaspis samanalensis: gular, pectoral andabdominal scales tricarinate, subcaudals keeled.Cnemaspis retigalensis: gulars carinate, a singlepreanal pore and 3 or 4 femoral pores.Cnemaspis

punctata: 5 7 femoral pores, distinctive spotted dorsalcoloration.Cnemaspis upendrai, C. kumarasinghei, C.latha, C. pulchra, C. pava, C. kallima, C. silvula and C.clivicola: presence of femoral pores.Cnemaspis amith:gular scales carinate.

Cnemaspis wynadensis (Beddome, 1870)Gymnodactylus wynadensis Beddome, 1870

(Figures 5a g; Table 2)

Lectotype (here designated). BMNH 74.4.29.355(male), 33.4 mm SVL, Wynaad (Kerala, India), coll.Colonel Beddome.

Paralectotypes. BMNH 74.4.29.356 (male), 36.6 mmSVL; BMNH 74.4.29.357 (male), 34.2 mm SVL; BMNH74.4.29.358 (male), 39.4 mm SVL; BMNH 74.4.29.359(male), 40.3 mm SVL; BMNH 74.4.29.360 (male), 30.7mm SVL; BMNH 74.4.29.361 (male), 31.3 mm SVL;BMNH 74.4.29.362 (male), 40.6 mm SVL; BMNH74.4.29.363 (male), 38.9 mm SVL; BMNH 74.4.29.364(male), 34.3 mm SVL; BMNH 74.4.29.365 (male),33.7 mm SVL; BMNH 74.4.29.366 (female), 42.1 mmSVL; BMNH 74.4.29.367 (female), 41.2 mm SVL;BMNH 74.4.29.368 (female), 40.8 mm SVL; BMNH74.4.29.369 (subadult male), 25.3 mm SVL; BMNH74.4.29.370 (subadult female), 25.5 mm SVL; BMNH74.4.29.371 (juvenile), 18.4 mm SVL, Wynaad, coll.Colonel Beddome.

DiagnosisCnemaspis wynadensis differs from its peninsularIndian and Sri Lankan congeners by a combinationof the following characters. Maximum SVL 40.6 mm;paired postmentals separated by 3 medial scales; eachpostmental bounded by 4 scales including medialscale; dorsal scales homogeneous, smooth, isolatedand scattered; spine-like tubercles absent on flank;

abdominal scales smooth, subimbricate; no preanalpores; 4 6 femoral pores on each side; subcaudalson median series enlarged, subequal; supralabialsto angle of jaws, 5; subdigital lamellae on digit IVof pes, 18. Description of lectotypeAn adult male, 33.4 mm SVL (for measurements,see Table 2). Head moderately long (HL 27.2% of SVL), broad (HW 19.1% of SVL, HW 70.3% of HL),distinct from neck. Snout length a little greater

than eye diameter (ED 72.2% of ES); interorbitalnarrow (IO 13.1% of HL); pupil rounded. Granuleson snout smooth; scales on ventral surface of headsmooth. Rostral scale partially divided by a medialgroove, in contact with first supralabial (Fig. 5a).

Nares separated by two enlarged supranasals; smallinternasal scales separate supranasals posteriorly(Fig. 5a). Nares rounded, dorsally orientated, notin contact with first supralabial (Fig. 5a). Aboutfour postnasals, the uppermost one enlarged(Fig. 5a). Mental subpentagonal, wider than long,posteriolaterally in contact with two widelyseparated, enlarged postmentals and medially withthree small postmental scales; postmentals borderedposteriorly by 9 smooth (2 enlarged lateral and 7 smallmedial) scales (Fig. 5b). Supralabials 5, infralabials 6,decreasing in size towards gape (Fig. 5c). Ear openingoval, higher than wide, oblique (TW 40.0% of TH).Scales on ventral surface of neck smooth.Body robust. Mid-dorsal granules homogeneous,isolated and scattered, smooth (Fig. 5d); spine-liketubercles absent on flank; abdominal scales smooth,subimbricate, longer than wide, obtuse (Fig. 5e).Scales around vent and base of tail smooth, imbricate(Fig. 5 f ). No preanal pores, 5 femoral pores (Fig.5 f ).

Forelimbs relatively short, slender (LAL 14.0%SVL, UAL 10.1% SVL); hind limbs long (TBL 18.2%SVL, FEL 20.6% SVL); tibia short. Dorsal scales on both fore and hind limbs smooth; ventral scales onupper arm and lower arm smooth. Scales on anteriorsurface of thigh and ventral side smooth (Fig. 5 f );those on ventral surface of shank, and on manusand pes, smooth. Digits elongate, slender, all bearingslightly recurved claws; subdigital lamellae entire,unnotched; subdigital lamellae on finger I, 9, fingerII, 12, finger III, 16, finger IV, 15, finger V, 13; toe I, 10,toe II, 13, toe III, 18, toe IV, 18, toe V, 17; interdigitalwebbing absent; relative length of digits (fingers): III(3.0 mm) > IV (2.8 mm) > II (2.7 mm) > V (2.6 mm) >I (1.8 mm); (toes) IV (3.5 mm) > III (3.4 mm) ~ V (3.4mm) > II (2.8 mm) > I (2.1 mm).

Tail tapering, its length greater than SVL (TAL

112.8% of SVL); tail base slightly swollen, lackingenlarged tubercles. Dorsal scales on tail smooth, imbricate, directed backwards (Fig. 5 g); no post-cloacalspurs; subcaudals smooth, median series regular,enlarged, widened, circular or hexagonal (Fig. 5 f ).

Variation. Femoral pores, 46; pores absent infemales; SVL range from 18.4 to 42.1 mm (16 ex.).

ComparisonsCnemaspis wynadensis differs from each of its


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Figure 5. Cnemaspis wynadensis: BMNH 74.4.29.355, lectotype, male,a, dorsal aspect of snout;b, ventral aspect of chin;c,lateral aspect of gape, showing arrangement of supralabials and infralabials;d, e, arrangement of scales on dorsal andventral regions, respectively, of mid-body; f , preanal, hind limb and tail regions, showing arrangement of femoral poresand arrangement of subcaudal scales; g, arrangement of scales on dorsal aspect of tail. Scale bars: 1 mm.

a b

c d

e f

g


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peninsular Indian and Sri Lankan congeners,respectively, by the following opposing suites of characters:

Cnemaspis beddomei: preanal pores 69, no femoralpores.Cnemaspis goaensis: presence of preanal pores.

Cnemaspis littoralis : body slender, dorsal scaleshomogeneous. Cnemaspis mysoriensis: spine-liketubercles present on flank, femoral pores 2 or 3.Cnemaspis indraneildasii: spine-like tubercles presenton flank. Cnemaspis heteropholis and C. sisparensis:subcaudals on median series divided.Cnemaspis

jerdonii: spine-like tubercles present on flank.Cnemaspis nairi: no femoral pores.Cnemaspis otai:paired postmentals in broad contact, 4 preanal and3 femoral pores.Cnemaspis yercaudensis: 2 preanaland 3 femoral pores.Cnemaspis boiei: no preanal orfemoral pores. Cnemaspis gracilis: postmentals incontact, subcaudals in median series not enlarged.Cnemaspis indica: postmentals in contact.Cnemaspisornata: dorsal tubercles arranged in 15 longitudinalrows, dark-brown collar on nape, no femoral pores,preanal pores 6 8. Cnemaspis wicksii, C. andersoniiand C. monticola: gulars carinate.Cnemaspis australis :subcaudals pointed, strongly keeled.Cnemaspisnilagirica: pectoral and abdominal scales weaklykeeled.

Cnemaspis tropidogaster: gular, pectoral andabdominal scales keeled, spine-like tuberclespresent on flank.Cnemaspis kandiana, C. podihuna,C. scalpensis, C. gemunu and C. phillipsi: spine-like tubercles present on flank.Cnemaspis alwisi:9 femoral pores, spine-like tubercles present onflank. Cnemaspis molligodai: 5 preanal and 8 or9 femoral pores, spine-like tubercles present onflank. Cnemaspis samanalensis: gular, pectoral andabdominal scales tricarinate, subcaudals keeled,spine-like tubercles present on flank.Cnemaspisretigalensis : a single preanal pore and 3 or 4 femoralpores; spine-like tubercles present on flank;subcaudals of median row enlarged, irregular.Cnemaspis punctata : spine-like tubercles presenton flank. Cnemaspis kumarasinghei, C. menikay, C.

pulchra, C. pava, C. upendrai, C. latha, C. kallima, C.

silvula and C. clivicola: presence of preanal pores. Cnemaspis amith: gular scales carinate; subcaudalson median row enlarged, irregular.

Cnemaspis wicksii (Stoliczka, 1873)Gymnodactylus wicksii Stoliczka, 1873

(Figures 6a c)

Lectotype (here designated). ZSI 5868 (male),28.6 mm SVL, from Preparis Island (AndamansArchipelago, Bay of Bengal).

Paralectotypes. ZSI 5866 (male), 27.9 mm SVL; ZSI5867 (female), 26.6 mm SVL, from Preparis Island,in the Andaman Archipelago, Bay of Bengal.

Diagnosis

Cnemaspis wicksii differs from its peninsular Indianand insular congeners by a combination of thefollowing characters. Maximum SVL 28.6 mm;paired postmentals separated by a medial scale,each bounded by 3 scales including medial scale;dorsal scales heterogeneous; gulars weakly carinate;other ventrals smooth, imbricate; dorsal scales on both fore and hind limbs smooth; ventral scales onarm smooth; scales on anterior and ventral surfacesof thigh smooth; supralabials to angle of jaws, 5; 4preanal pores; 4 or 5 femoral pores on each side;subcaudals smooth, 2 smaller obtuse scales betweensingle larger rounded ones in median series. Description of lectotypeAn adult female, 28.6 mm SVL (for measurements,see Table 2). Head moderately long (HL 26.9% of SVL), moderately broad (HW 15.3% of SVL, HW57.1% of HL), distinct from neck. Pupil rounded.Granules on snout smooth; scales on ventralsurface of head feebly keeled. Mental subtriangular,wider than long, posteriolaterally in contact withtwo enlarged postmentals, medially with a singlehexagonal postmental scale; postmentals borderedposteriorly by 5 smooth scales including medial scale(Fig. 6a). Supralabials 5, infralabials 7, decreasing insize towards gape. Scales on ventral surface of neckfeebly carinate.

Body slender. Mid-dorsal scales granular;abdominal scales smooth, those around vent and base of tail smooth, imbricate; 4 preanal and 4 (leftside) and 5 (right side) femoral pores (Fig. 6b).

Dorsal scales on both fore and hind limbs smooth;ventral scales on upper arm and lower arm smooth.Scales on both anterior and ventral surfaces of thighsmooth (Fig. 6b). Scales on ventral surface of shankfeebly carinate, unnotched; subdigital lamellae onfinger I, 10, finger II, 12, finger III, 16, finger IV, 14,

finger V, 11.Tail broken (a broken tail remains in the specimen jar). Enlarged tubercles absent on base of tail. Dorsalscales on tail oval, directed backwards; subcaudalssmooth, 2 smaller obtuse scales between single largerrounded ones in median series (Fig. 6c).

Measurements of lectotype (those of paralectotypesZSI 5866 and ZSI 5867 in parentheses; measurementsin mm): HL, 7.7 (7.6, 6.9); HW, 4.4 (4.4, 4.1); SVL, 28.6(27.9, 26.6).


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Coloration. In preservative: general body colourgreyish-dark brown. No prominent markings on body.

Variation. Preanal and femoral pores absent infemales. SVL ranges from 26.6 to 28.6 mm (3 ex.).

ComparisonsCnemaspis wicksiidiffers from each of its peninsularIndian and Sri Lankan congeners, respectively, by thefollowing opposing suites of characters:

Cnemaspis beddomei: spine-like tubercles absent onflank, preanal pores 69, no femoral pores.Cnemaspis

goaensis: subcaudals in median series not enlarged.Cnemaspis littoralis: dorsal scales homogeneous,median series of subcaudals enlarged, hexagonal.

Cnemaspis mysoriensis: femoral pores 2 or 3.Cnemaspisindraneildasii and C. heteropholis: gulars smooth.Cnemaspis jerdonii: no preanal pores.Cnemaspis nairi:spine-like tubercles absent on flank, no femoral pores.Cnemaspis otai: paired postmentals in broad contact.Cnemaspis sisparensis: postmentals in contact, spine-like tubercles absent on flank, no preanal pores, 7 or8 femoral pores.Cnemaspis wynadensis: subcaudalsenlarged, subequal; no preanal pores, 5 femoralpores, spine-like tubercles absent on flank.Cnemaspis

yercaudensis: 2 preanal and 3 femoral pores, spine-like

tubercles absent on flank, gulars smooth.Cnemaspisboiei: no preanal or femoral pores, spine-like tuberclesabsent on flank.Cnemaspis gracilis: postmentals incontact, subcaudals in median series not enlarged.Cnemaspis indica: postmentals in contact, spine-liketubercles absent on flank.Cnemaspis ornata: dorsaltubercles arranged in 15 longitudinal rows, dark- brown collar on nape, no femoral pores, preanalpores 6 8. Cnemaspis andersonii: 7 supralabials,scales on ventro-lateral region carinate, subcaudalsin median series not enlarged.Cnemaspis monticola:subcaudals carinate.Cnemaspis australis : subcaudalspointed, strongly keeled.Cnemaspis nilagirica: dorsalscales homogeneous; pectoral and abdominal scalesweakly keeled; subcaudals on median row enlarged,subequal.

Cnemaspis tropidogaster: gular, pectoral and ab-dominal scales keeled.Cnemaspis kandiana: exceptin the median series, subcaudals carinate.Cnemaspis

podihuna, C. scalpensis, C. gemunu and C. phillipsi: 12 16 femoral pores.Cnemaspis alwisi: no preanalpores. Cnemaspis molligodai: 5 preanal and 8 or 9femoral pores, gular, pectoral and abdominal scalessmooth; subcaudals on median row enlarged,subequal. Cnemaspis samanalensis: gular, pectoraland abdominal scales tricarinate, subcaudals keeled.Cnemaspis retigalensis: a single preanal pore and 3 or 4

Figure 6. Cnemaspis wicksii: ZSI 5868, lectotype, male,a, ventral aspect chin;b, preanal and left hind-limb regions, showingarrangement of femoral and preanal pores;c, arrangement of subcaudal scales. Scale bars: 1 mm.

a

b

c


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femoral pores.Cnemaspis punctata : no preanal pores,distinctive spotted dorsal coloration.Cnemaspiskumarasinghei and C. latha: gular scales smooth.Cnemaspis silvula, C. pulchra, C. pava, C. clivicolaandC. upendrai: pectoral and abdominal scales keeled.

Cnemaspis kallima: gular scales smooth, subcaudals ondistal half of tail carinate.Cnemaspis menikay: dorsalgranules conical, 1 or 2 preanal pores.Cnemaspisamith: dorsal granules unkeeled.

RemarksWe have included a description of Cnemaspis wicksii here, even though its range is outside our region of interest (peninsular India and Sri Lanka), because itwas previously listed in the synonymy of C. kandaiana by Smith (1935), where it has hitherto remained.Based on an examination of their name-bearingtypes, however, Cnemaspis wicksii is immediatelydistinguishable fromC. kandaiana by having thescales of the dorsal surface of both limbs smooth(vs carinate inC. kandaiana); all subcaudals smooth(vs subcaudals carinate except for those on medianrow, which are smooth); 5 supralabial scales (vs 8 or9); and no tubercles on tail base (vs about 6 enlargedtubercles on tail base). We considerCnemaspis wicksii to be a valid species, clearly distinguishable from allits Sri Lankan and peninsular Indian congeners.

Cnemaspis beddomei (Theobald, 1876)Gymnodactylus beddomeiTheobald, 1876

Gymnodactylus marmoratus Beddome, 1870(Figures 7a g; Table 2; Plate 1 g, h)

Lectotype (here designated). BMNH 1946.9.4.83(male), 46.6 mm SVL, South Tinnevelly (Tirunelveli,southern Tamil Nadu State, India) and TravankorHills 30005000 feet (Travancore, Kerala State,India), coll. Colonel R. H. Beddome.

Paralectotypes. BMNH 1946.9.4.82 (female), 48.2 mmSVL; BMNH 1946.9.4.84 (male), 42.2 mm SVL; BMNH1946.9.4.85 (male), 43.9 mm SVL; South Tinnevelly[Tirunelveli, southern Tamil Nadu State, India] and

Travankor Hills 30005000 feet [Travancore, KeralaState, India], coll. R. H. Colonel Beddome.

DiagnosisCnemaspis beddomeidiffers from its peninsular Indianand Sri Lankan congeners by a combination of thefollowing characters. Maximum SVL 50.6 mm;paired postmentals separated by a medial scale; eachpostmental bounded by 4 or 5 scales including medialscale; ventrals, 154; dorsal scales heterogeneous;spine-like tubercles absent on flank; ventrals smooth,

isolated and scattered; 7 preanal pores; no femoralpores; subcaudals on median row widened, irregular,some scales divided; supralabials to angle of jaws, 6;subdigital lamellae on digit IV of pes, 16.

Description of lectotypeAdult male, 46.6 mm SVL (for measurements, seeTable 2). Head long (HL 28.1% of SVL), broad (HW20.3% of SVL, HW 72.5% of HL), distinct from neck.Snout length greater than eye diameter (ED 57.3%of ES); interorbital broad (IO 16.7% of HL); pupilrounded. Granules on snout smooth, larger thanthose of occipital region; scales of occipital regiongranular, those on interorbital and superciliumconical, and those on ventral surface of headgranular, smaller than those on abdominal. Rostralscale partially divided by a medial groove, in contactwith first supralabial (Fig. 7a). Nares separated bytwo enlarged supranasals; no internasals (Fig. 7a).Nares rounded, dorsally orientated, not in contactwith first supralabial. Two enlarged postnasalspresent, the lower one in broad contact with firstsupralabial (Fig. 7a). Mental subpentagonal, truncateposteriorly, as wide as long, posteriolaterallyin contact with two widely separated quadratepostmentals and medially in contact with a singleelongate postmental scale; postmentals borderedposteriorly by 8 smooth scales including medial scale(Fig. 7b). Supralabials 6, infralabials 7, decreasingin size towards gape (Fig. 7c). Ear opening oval,higher than wide, oblique. Scales on ventral surfaceof neck granular. Granules between anterior marginof ear opening and posterior margin of eye 29.Conical granules on sides of neck larger than thoseof occipital region.

Body robust, short. Mid-dorsal scales granular,isolated and scattered, conical, intermixed withlarge scales (Fig. 7d); spine-like tubercles absent onflank; abdominal scales smooth (Fig. 7e). Ventrals154, isolated and scattered (Fig. 7e); scales aroundpreanal area enlarged; 7 preanal pores, no femoralpores (Fig. 7 f ).

Forelimbs moderately long, slender (LAL 13.5%

SVL, UAL 13.9% SVL); hind limbs long (TBL 17.3%SVL, FEL 24.4% SVL); tibia short. Dorsal scales on both fore and hind limbs granular, conical; ventralscales on upper arm and lower arm granular andsmaller than dorsals. Dorsal scales of thigh andshank granular (Fig. 7 f ); ventral scales on shankimbricate, smooth, those on thigh enlarged, conical.Dorsal sales on manus and pes granular, ventralsimbricate, smooth. Digits elongate, slender, all bearing slightly recurved claws; subdigital lamellaeentire, unnotched; subdigital lamellae on finger I,


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Figure 7. Cnemaspis beddomei: BMNH 1946.9.4.83, lectotype, male,a, dorsal aspect of snout;b, ventral aspect of chin;c,lateral aspect of gape, showing arrangement of supralabials and infralabials;d, e, arrangement of scales on dorsal andventral regions, respectively, of mid-body; f , preanal and hind-limb regions, showing arrangement of preanal pores; g,arrangement of scales on ventral side of tail-base. Scale bars: 1 mm.

a b

c d

e f

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11, finger II, 14, finger III, 18, finger IV, 19, finger V,15; toe I, 17, toe II, 19, toe III, 18, toe IV, 16, toe V, 10;interdigital webbing absent; relative length of digits(fingers): V (5.6 mm) > IV (5.2 mm) > III (5.0 mm) >II (4.7 mm) > I (3.6 mm); (toes) IV (6.7 mm) > III (6.1

mm) > V (5.8 mm) > II (5.6 mm) > I (3.4 mm).Regenerated tail tapering, its length less thanSVL (TAL 74.6% of SVL); tail base distinctlyswollen. Scales on tail base conical to imbricate;enlarged tubercles absent on tail. Dorsal scales ontail imbricate; a single enlarged, oval-shaped, post-cloacal spur present on each side; subcaudal scaleson tail base imbricate to subimbricate, increasing insize towards tail tip (Fig. 7 g).

Variation. Supralabials, 6 or 7; infralabials, 6 or 7;preanal pores 6 8; femoral pores absent; pores absentin females; SVL range 42.246.6 mm (4 ex.).

Unregenerated tail of paralectotype BMNH1946.9.4.86 with a median series of enlarged, dividedsubcaudals, with individual large scales betweenpairs of smaller ones (see Fig. 4h of C. ornata), mostsubcaudal scales divided on distal portion of tail.

Coloration. In preservative, dorsally light brownwith dark mottling in region of head, pale mottlingon body; tail with pale bands; venter brownish white;two dark, V-shaped streaks on chin and gular region(see Plate 1h).

Comparisons. Cnemaspis beddomeidiffers from eachof its peninsular Indian and Sri Lankan congeners,respectively, by the following opposing suites of characters:

Cnemaspis goaensis: presence of femoral pores.Cnemaspis littoralis: body slender, subcaudals of median series enlarged, hexagonal.Cnemaspismysoriensis: spine-like tubercles present on flank,femoral pores 2 or 3.Cnemaspis indraneildasii and C. heteropholis: dorsal scales heterogeneous. Cnemaspis jerdonii: no preanal pores.Cnemaspisnairi: a dark gular band present.Cnemaspis otai:paired postmentals in broad contact, 4 preanal and

3 femoral pores.Cnemaspis sisparensis: postmentalsin contact, no preanal pores, 7 or 8 femoral pores. Cnemaspis wynadensis: subcaudals enlarged, subequal;no preanal pores, 5 femoral pores.Cnemaspis

yercaudensis: 2 preanal and 3 femoral pores.Cnemaspisboiei: no preanal or femoral pores.Cnemaspis gracilis:postmentals in contact, subcaudals in median seriesnot enlarged.Cnemaspis indica: postmentals in contact,no preanal pores.Cnemaspis ornata: dorsal tuberclesarranged in 15 longitudinal rows, a dark-browncollar on nape.Cnemaspis wicksii and C. andersonii:

presence of femoral pores.Cnemaspis monticola: spine-like tubercles present on flank.Cnemaspis australis :subcaudals pointed, strongly keeled.Cnemaspisnilagirica: dorsal scales homogeneous; pectoral andabdominal scales weakly keeled; subcaudals on

median row enlarged, subequal.Cnemaspis tropidogaster: gular, pectoral andabdominal scales keeled, spine-like tuberclespresent on flank.Cnemaspis kandiana, C. podihuna,C. scalpensis, C. menikay, C. gemunu and C. phillipsi:spine-like tubercles present on flank.Cnemaspisalwisi: no preanal pores, spine-like tubercles presenton flank.Cnemaspis molligodai: 8 femoral pores, spine-like tubercles present on flank; subcaudals in medianrow enlarged, subequal.Cnemaspis samanalensis:gular, pectoral and abdominal scales tricarinate,subcaudals keeled, spine-like tubercles present onflank.Cnemaspis retigalensis: gulars carinate, a singlepreanal pore and 3 or 4 femoral pores, spine-liketubercles present on flank.Cnemaspis punctata : nopreanal pores, spine-like tubercles present on flank,distinctive spotted dorsal coloration.Cnemaspiskumarasinghei: spine-like tubercles present on flank. Cnemaspis latha: ventrals 111; gular scales smooth,abdominal scales and subcaudal scales smooth,spine-like tubercles present on flank.Cnemaspissilvula,C. pulchra and C. pava: pectoral and abdominalscales keeled, spine-like tubercles present on flank.Cnemaspis clivicola pectoral and abdominal scaleskeeled, spine-like tubercles present on flank,subcaudals in median series not enlarged.Cnemaspisupendrai: some gular scales tricarinate, pectoral andabdominal scales keeled, spine-like tubercles presenton flank. Cnemaspis kallima: gular scales smooth,spine-like tubercles present on flank, subcaudalson distal half of tail carinate.Cnemaspis amith: gularscales carinate; pectoral and abdominal scalessmooth; dorsal granules heterogeneous; subcaudalson median row enlarged, irregular.

Cnemaspis sisparensis (Theobald, 1876)Gymnodactylus sisparensis Theobald, 1876Gymnodactylus maculatus Beddome, 1870

Gonatodes bireticulatus Annandale, 1915Cnemaspis anaikattiensis Mukherjee, Bhupathy &Nixon, 2005

(Figures 8a g, 9a f ; Table 2; Plate 2a, b)

Holotype. BMNH 74.4.29.383 (male) (fragmented),Sholakal, the foot of Sispara Ghat, coll. ColonelBeddome.

Other material. ZSI 17970 (male), 42.7 mm SVL,holotype of Gonatodes bireticulatus Annandale,


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1915, Karalai 13003000 ft, Cochin State, coll. F.H. Gravely (see Mukherjeeet al., 2005); ZSI 25601,(male), 61.0 mm SVL, ZSI 25602, (female), 58.0 mm(respectively, holotype and paratype of Cnemaspisanaikattiensis Mukherjee, Bhupathy & Nixon, 2005),

Anaikatti Hills, Western Ghats, Coimbatore district,Tamil Nadu, India, 110530.9N, 764736.2E, coll.A. M. A. Nixon & D. Mukherjee, 17 September, 2003(not examined).

DiagnosisCnemaspis sisparensis differs from its peninsularIndian and Sri Lankan congeners by a combinationof the following characters. Maximum SVL 42.7mm; paired postmentals widely separated by twomedial scales; each postmental bounded by 4 scalesincluding medial scale; dorsal scales homogeneous,isolated and scattered; spine-like tubercles absenton flank; ventrals smooth, imbricate; supralabials toangle of jaws 7 or 8; no preanal pores, 7 or 8 femoralpores; subcaudals smooth; those on median seriesenlarged, with individual large scales alternating between pairs of smaller ones.

Description of holotypeAn adult male. Head broad (HW 65.6% of HL),pupil rounded. Granules on snout smooth, largerthan those of occipital region; scales on ventralsurface of head with smooth tubercles. Rostral scalepartially divided by a medial groove, in contactwith first supralabial (Fig. 8a). Nares separated bytwo enlarged supranasals and three internasals,the median one of which extends towards snout tip(Fig. 8a). Nares rounded, dorsally orientated, notin contact or in contact with first supralabial (Fig.8a). Three postnasals, the upper one enlarged, thelower one in contact with first supralabial (Fig. 8a).Mental subtriangular, truncate posteriorly, as wideas long, posteriolaterally in contact with two widelyseparate, enlarged postmentals (damaged) andmedially with a small, hexagonal postmental scale;postmentals bordered posteriorly by several smallscales (damaged) (Fig. 8b). Supralabials 7, infralabials

8, decreasing in size towards gape (Fig. 8c).Mid-dorsal granules homogeneous, isolatedand scattered; spine-like tubercles absent on flank;abdominal scales smooth, imbricate; scales aroundvent and base of tail smooth, imbricate (Fig. 8d). Nopreanal pores; 8 femoral pores (right side); left sidedamaged (Fig. 8e).

Dorsal scales on both fore and hind limbs smooth;ventral scales on upper arm and lower arm smooth.Scales on anterior surface of thigh (Fig. 8e) andventral surface of shank smooth.

Tail tapering, enlarged tubercles absent at its base.Dorsal scales on tail imbricate, directed backwards(Fig. 8 f ). No post-cloacal spurs. Subcaudals smooth;scales on median series enlarged with single largescales alternating between pairs of smaller ones

(Fig. 8 g).Measurements of holotype (in mm): HL, 13.1;HW, 8.6; TAL, 55.2 (other measurements could not be taken because the specimen is fragmented: seePlate 2e, f ).

Variation (based on ZSI 17970): an adult male, 42.7mm SVL. Head long (HL 28.1% of SVL), moderately broad (HW 20.3% of SVL, HW 72.5% of HL), distinctfrom neck. Snout length greater than eye diameter(ED 69.5% of ES). No internasals; lower postnasals in broad contact with first supralabial (Fig. 9a). Mentalsubtriangular, as wide as long, posteriolaterallyin contact with two widely separated, enlargedpostmentals and medially with 2 small medial scales;postmentals bordered posteriorly by 8 smooth scales(2 enlarged lateral scales and 6 small medial scales)(Fig. 9b). Supralabials 8, infralabials, 10, decreasing insize towards gape (Fig. 9c). Ear opening oval, higherthan wide, oblique (TW 70.0% of TH) (Fig. 9c). Scaleson ventral surface of neck smooth, tuberculate. Bodyrobust. Femoral pores on each side, 7 (Fig. 9d). Scaleson manus and pes smooth, rounded. Tail tapering,its length greater than SVL (TAL 129.7% of SVL);enlarged tubercles present on base of tail. Dorsalscales on tail imbricate, directed backwards (Fig. 9e).Subcaudals smooth; scales of median series enlarged,with individual large scales alternating between pairsof smaller ones (Fig. 9 f ).

Measurements of ZSI 17970 (in mm): ED, 3.2; ES4.6; HL, 12.0; HW, 8.7; IN, 2.5; SVL, 42.7; TAL, 55.4;TYD, 3.6; TYE, 0.71.0. Coloration. In preservative: dorsally light to dark brown with dark-brown mottling on dorsum (seePlate 2a, b); nape with a dark-brown marking; tailwith pale bands; digits banded; venter brownishwhite with indistinct light-brown patches.

Comparisons. Cnemaspis sisparensis differs from eachof its peninsular Indian and Sri Lankan congeners,respectively, by the following opposing suites of characters:

Cnemaspis beddomei: preanal pores 69, no femoralpores.Cnemaspis goaensis: presence of preanal pores.Cnemaspis littoralis: subcaudals of median seriesenlarged, hexagonal.Cnemaspis mysoriensis: spine-like tubercles present on flank, femoral pores 2 or 3.Cnemaspis indraneildasii: spine-like tubercles present


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Figure 8. Cnemaspis sisparensis: BMNH 74.4.29.383, holotype, male,a, dorsal aspect of snout;b,ventral aspect of chin (dottedline indicates damaged area);c, lateral aspect of gape, showing arrangement of supralabials and infralabials;d,arrangementof scales on dorsal scales of mid-body;e, preanal and right hind-limb regions, showing arrangement of femoral pores; f ,arrangement of dorsal scales of mid-tail; g, arrangement of subcaudal scales. Scale bars: 1 mm.

a b

c

d

e

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ercles present on flank.Cnemaspis retigalensis: singlepreanal pore and 3 or 4 femoral pores, spine-liketubercles present on flank.Cnemaspis punctata : 5 7femoral pores, spine-like tubercles present on flank,distinctive spotted dorsal coloration.Cnemaspis

upendrai , C. kumarasinghei, C. pulchra, C. pava, C. latha, C.kallima, C. silvula and C. clivicola: presence of preanal pores.Cnemaspis amith: gular scales carinate;dorsal granules unkeeled.

RemarksMukherjeeet al. (2005) describedCnemaspis anai-kattiensis from the Anaikatti Hills (110530.9Nand 764736.2E), Western Ghats, a part of theNilgiri Biosphere Reserve in Tamil Nadu. Whileacknowledging that of its Western-Ghats congenersthe new species most closely resembledC. sisparensis, Mukherjeeet al. (2005) compared it with what theyconsidered to be The holotype of C. sisparensis(Theobald) [=Gonatodes bireticulatus] Annd. collected by F. H. Gravely from Kavalai, 13003000 ft, Co-chin State ZSI 17970, Zoological Survey of India,Kolkata. The holotype of C. sisparensis is in factBMNH 74.4.29.383, a male specimen collected fromSholakal, the foot of Sispara Ghat.

We examined the types of C. bireticulatus at ZSIand C. sisparensis at the BMNH. There is no doubt thatthese two species are conspecific, the distinguishingcharacters used by Mukherjeeet al. (2005)nointernasal scale inC. sisparensis (vs presence inthe new species), nostril not in contact with firstsupralabial (vs in contact inC. anaikattiensis)beingextremely variable inCnemaspis. The internasal scaleis often present or absent in different individualswithin a single population, and the nostril in contactor not in contact on either side of the same individual(e.g., see Remarks underC. podihuna, below). Thetype localities of C. anaikattiensis (Anaikatti Hills nearCoimbatore) andC. sisparensis (Sholakal, at the foot of Sispara Ghat, 1112N, 7628E, at the northeast endof Silent Valley National Park), are also relativelyclose to each other. In any event, the descriptionand diagnosis of C. anaikattiensis provided by

Mukherjeeet al. (2005) exactly match the holotypeof C. sisparensis, and we therefore here consider theformer to be a junior synonym of the latter.

Cnemaspis andersonii (Annandale, 1904)Gonatodes andersonii Annandale, 1904

(Figures 10a d)

Holotype. ZSI 15012 (male), 31.5 mm SVL; Nar-condum, Andaman Archipelago (India), Bay of Bengal.

DiagnosisCnemaspis andersonii differs from its peninsularIndian and Sri Lankan congeners by a combinationof the following characters. Maximum SVL 31.5 mm;paired postmentals separated by a medial scale; each

postmental bounded by 3 scales including medialscale; dorsal scales heterogeneous; gulars carinate;other ventrals smooth, imbricate; supralabials toangle of jaws, 7; preanal pores 3, femoral pores 4 oneach side; no median series of enlarged subcaudals. Redescription of holotypeAn adult male, 31.5 mm SVL (for measurements, seeTable 2). Head relatively short (HL 25.3% of SVL),moderately broad (HW 14.5% of SVL, HW 57.5% of HL), distinct from neck. Snout length greater thaneye diameter (ED 63.8% of ES), pupil rounded. Scaleson ventral surface of head and neck weakly carinate.Rostral scale partially divided by a medial groove,in contact with first supralabial (Fig. 10a). Naresseparated by two enlarged supranasals (damaged onleft side) and a single internasal, as large as supranasal(Fig. 10a). Nares rounded, dorsally orientated, incontact with first supralabial (Fig. 10a). Mentalsubpentagonal, wider than long, posteriolaterally incontact with two enlarged postmentals and mediallywith a single small hexagonal postmental scale;postmentals bordered posteriorly by 5 smooth scalesincluding medial scale (Fig. 10b). Supralabials 7,infralabials 7, both series decreasing in size towardsgape; ear opening oval, higher than wide, oblique(TW 60.0% of TH) (Fig. 10c).

Body slender. Mid-dorsal granules heterogeneous;pectoral and abdominal scales smooth; scales onventro-lateral region weakly carinate; 3 preanal and4 femoral pores (Fig. 10d). Tail tapering; a singleenlarged, conical, post-cloacal spur present on eachside; subcaudals in median series not enlarged.

Measurements of holotype (in mm): ED, 2.3; ES,3.6; HL, 8.0; HW, 4.6; SVL, 31.5; TAL, 40.2; TYD, 2.4;TYE, 0.50.3.

Coloration. In preservative: overall body colourdark-greyish brown.

Comparisons. Cnemaspis andersonii differs from eachof its peninsular Indian and Sri Lankan congeners,respectively, by the following opposing suites of characters:

Cnemaspis beddomei: preanal pores 6 9, nofemoral pores.Cnemaspis goaensis: subcaudals onmedian row enlarged. Cnemaspis littoralis: dorsalscales homogeneous, median series of subcaudals


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enlarged, hexagonal.Cnemaspis mysoriensis: femoralpores 2 or 3.Cnemaspis indraneildasii: dorsal scales of both limbs smooth; ventral, gular and caudal scalessmooth. Cnemaspis heteropholis: gulars smooth, nopost-cloacal spurs.Cnemaspis jerdoniiand C. nairi : nofemoral pores.Cnemaspis indica, C. otai, C. sisparensis,C. wynadensis and C. yercaudensis: spine-like tuberclesabsent on flank.Cnemaspis boiei: no preanal or femoralpores. Cnemaspis gracilis: 2 preanal pores separated by 2 unpored scales, postmentals broadly in contact.Cnemaspis ornata: no femoral pores, preanal pores6 8, dark brown collar on nape.Cnemaspis wicksii: 5supralabials, scales on ventro-lateral region smooth,subcaudals in median series enlarged.Cnemaspis

monticola: no large, conical tubercles at base of tail, spine-like tubercles present on flank, pectoraland abdominal scales smooth.Cnemaspis australis :subcaudals pointed, strongly keeled.Cnemaspisnilagirica: dorsal scales homogeneous; subcaudalson median row enlarged, subequal.

Cnemaspis tropidogaster: gular, pectoral andabdominal scales keeled.Cnemaspis kandiana:subcaudals on median row enlarged, irregular.Cnemas pis podihuna: subcaudals on median rowenlarged, subequal.Cnemaspis scalpensis, C. gemunu

and C. phillipsi: 12 16 femoral pores.Cnemaspisalwisi: 7 9 femoral pores.Cnemaspis molligodai: 5preanal and 8 or 9 femoral pores, dorsal scaleshomogeneous; subcaudals on median row enlarged,subequal.Cnemaspis samanalensis: gular, pectoral andabdominal scales tricarinate, subcaudals keeled.Cnemaspis retigalensis: single preanal pore and 3 or 4femoral pores; subcaudals on median row enlarged,irregular. Cnemaspis punctata : 5 7 femoral pores,distinctive spotted dorsal coloration.Cnemaspiskumarasinghei: gular scales smooth.Cnemaspis latha:gular scales smooth.Cnemaspis silvula, C. pulchraand C. pava: pectoral and abdominal scales keeled.Cnemaspis clivicola: pectoral and abdominal scales

keeled. Cnemaspis upendrai: some gular scalestricarinate, pectoral and abdominal scales keeled.Cnemaspis kallima: gular scales smooth.Cnemaspismenikay: dorsal granules conical, 1 or 2 preanalpores. Cnemaspis amith: dorsal granules unkeeled;subcaudals on median row enlarged, irregular.

RemarksCnemaspis andersonii was listed in the synonymyof C. kandiana by Smith (1935), where it has sinceremained. Examination of its holotype (the only

Figure 10. Cnemaspis andersonii: ZSI 15012, holotype, male,a, dorsal aspect of snout (left side damaged);b,ventral aspectof chin;c, lateral aspect of head, showing arrangement of supralabials and infralabials;d, preanal and right hind-limbregions, showing arrangement of femoral and preanal pores. Scale bars: 1 mm.

a b

cd


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known specimen representing this taxon) shows,however, that it is distinguished fromC. kandiana by having the subcaudals of the median row notenlarged (vs enlarged in C. kandiana); and scales of the ventro-lateral region of body carinate (vs smooth).

We therefore considerCnemaspis andersonii to be avalid species.

Cnemaspis goaensis Sharma, 1976Cnemaspis goaensis Sharma, 1976

(Figures 11a e; Table 2)

Holotype. ZSI 22110 (male), 27.5 mm SVL; ca. 3 kmS. of Forest Rest House, Canacona (Poinguinim),Goa (south-western India).

Paratypes. ZSI 22215 (male), 28.3 mm SVL; samelocality data as holotype.

DiagnosisCnemaspis goaensisdiffers from its peninsular Indianand Sri Lankan congeners by a combination of thefollowing characters. Maximum SVL 28.3 mm;paired postmentals separated by a medial scale;each postmental bounded by 3 scales includingmedial scale; dorsal scales heterogeneous mixed withlarge keeled tubercles; gulars weakly carinate; otherventral scales smooth, imbricate; 3 preanal pores and2 4 femoral pores on each side; subcaudals slightlyenlarged; supralabials to angle of jaws, 8; subdigitallamellae on digit IV of pes, 16.

Redescription of holotypeAn adult male, 27.5 mm SVL (for measurements,see Table 2). Head long (HL 30.1% of SVL), broad(HW 19.2% of SVL, HW 63.8% of HL), distinct fromneck. Snout length about equal to eye diameter (ED96.5% of ES); interorbital narrow (IO 6.0% of HL);pupil rounded. Granules on snout keeled; scaleson interorbital and supercilium granular; scales onventral surface of head weakly carinate. Rostral scalepartially divided by a medial groove, in contact withfirst supralabial (Fig. 11a). Nares separated by two

enlarged supranasals, pentagonal internasal andthree small scales; internasal extends towards snouttip (Fig. 11a). Nares rounded, dorsally orientated,not in contact or in contact with first supralabial.Two postnasals, the lower one in broad, in contactwith first supralabial (Fig. 11a). Mental subtriangular,wider than long, posteriolaterally in contact withtwo enlarged postmentals and medially with asingle hexagonal postmental scale; postmentals bordered posteriorly by 5 smooth scales includingmedial scale (Fig. 11b). Supralabials and infralabials

8 each, decreasing in size towards gape (Fig. 11c).Ear opening oval, higher than wide, oblique (TW66.6% of TH) (Fig. 11c). Scales on ventral surface of neck feebly carinate.

Body robust. Mid-dorsal granules heterogeneous,

mixed with large keeled tubercles from head tosacrum; ventral abdominal scales smooth; scalesaround vent and base of tail smooth, imbricate; 3preanal and 4 femoral pores (Fig. 11d).

Forelimbs moderately long, slender (LAL 13.4%SVL, UAL 12.3% SVL); hind limbs long (TBL 18.1%SVL, FEL 29.0% SVL); tibia short. Dorsal scales on both fore and hind limbs weakly carinate; ventralscales on upper arm and lower arm smooth. Scales onthigh and shank smooth. Digits elongate, slender, all bearing slightly recurved claws; subdigital lamellaeentire, unnotched; subdigital lamellae on finger I,9, finger II, 12, finger III, 13, finger IV, 15, finger V,11; toe I, 9, toe II, 12, toe III, 16, toe IV, 16, toe V, 16;interdigital webbing absent; relative length of digits(fingers): IV (2.5 mm) ~ II (2.5 mm) > III (2.3 mm) > V(2.0 mm) > I (1.4 mm); (toes) II (3.5 mm) > I (2.8 mm)> III (2.7 mm) > IV (2.5 mm) > V (1.8 mm).

Tail tapering (its length less than SVL in paratype,ZSI 22215: TAL 85.0% of SVL); tail base distinctlyswollen; 4 6 enlarged, conical, keeled tubercles on base of tail, about as large as granules on mid-body.Dorsal scales on mid-tail imbricate, carinate, directed backwards; a conical post-cloacal spur presenton each side; median row of subcaudals slightlyenlarged on proximal half of tail, subquadrangular,smooth (Fig. 11e); scales on distal half carinate.

Coloration. In preservative: in general body colourgreyish dark brown.

Variation. Seven supralabials and 7 infralabials inparatype ZSI 22215. Femoral pores, 24 on each side(2 ex.); pores absent in females.

Comparisons. Cnemaspis goaensis differs from eachof its peninsular Indian and Sri Lankan congeners,respectively, by the following opposing suites of

characters:Cnemaspis beddomei: no interanasals, subcaudals ina series of alternating wide and narrow enlarged scales,no femoral pores.Cnemaspis littoralis: body slender,dorsal scales homogeneous, subcaudals on medianseries enlarged, hexagonal.Cnemaspis mysoriensis:femoral pores 2 or 3.Cnemaspis indraneildasii and C.heteropholis: gulars smooth.Cnemaspis jerdonii: nopreanal pores. Cnemaspis nairi: no femoral pores.Cnemaspis otai: paired postmentals in broad contact,gulars smooth. Cnemaspis sisparensis: postmentals


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in contact, no preanal pores, 7 or 8 femoral pores. Cnemaspis wynadensis: subcaudals enlarged, subequal;no preanal pores, 5 temporal pores, postmentalswidely separated. Cnemaspis yercaudensis: gulars

smooth.Cnemaspis boiei: no preanal or femoral pores,postmentals widely separated by 2 medial scales.Cnemaspis gracilis: postmentals in contact, subcaudalsin median series not enlarged.Cnemaspis indica:postmentals in contact, no preanal pores.Cnemaspisornata: dorsal tubercles arranged in 15 longitudinalrows, dark-brown collar on nape, no femoral pores,preanal pores 6 8.Cnemaspis wicksii: 5 supralabials,subcaudals enlarged.Cnemaspis andersonii: ventro-lateral region carinate, subcaudals in median seriesnot enlarged, internasal as large as supranasal.

Cnemaspis monticola: Dorsal scales on mid-tail keeled,isolated and granular.Cnemaspis australis : subcaudalspointed,

Cnemaspis Revision- Sri Lanka

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