Zeylanica, ISSN 1391-6270. October, 2007. Vol. 7, No. 1, pp. 9–122, incl. 6 tabs., 46 figs. and 12 pls. © 2007, Wildlife Heritage Trust of Sri Lanka.

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 genus Cnemaspis Strauch, 1887 from Sri Lanka are reviewed and re-described on the basis of their name-bearing types. Based on a recent survey, 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 valid species of Cnemaspis are now recognized from the island, all of them endemic. Two species—C. amith and C. tropidogaster—are not presently known from extant populations. A neotype is designated for Gymnodactylus scalpensis Ferguson, 1877 (type locality Gannoruwa Forest, near Kandy) and lectotypes selected for Gymnodactylus kandiana Kelaart, 1852, and Gonatodes kandianus var. tropidogaster Boulenger, 1885 (the type locality of the latter, which is not yet known from an extant population, is restricted to Sri Lanka). Cnemaspis ranwellai Wickramasinghe, 2006, is shown to be a junior objective synonym of C. scalpensis. An examination of type material shows Gymnodactylus 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 for Goniodactylus boiei Gray, 1842, Gymnodactylus beddomei Theobald, 1876, G. gracilis Beddome, 1870, G. indicus Gray, 1846, G. ornatus Beddome, 1870, G. wicksii Stoliczka, 1873, and G. wynadensis Beddome, 1870. Three new South Indian species discovered among these historical specimens are described: C. monticola (type locality Wynaad, Kerala State); C. australis (type locality Tirunelveli, Tamil Nadu State); and C. nilagirica (type locality: Nilgiri Hills, Tamil Nadu; formerly a syntype of C. tropidogaster). Cnemaspis wicksii (Stoliczka, 1873) and C. andersonii (Annandale, 1904), hitherto buried in the synonymy, are shown to be valid species. Examination of their name- bearing types shows Gonatodes bireticulatus Annandale, 1915, to be a junior subjective synonym of Gymnodactylus sisparensis Theobald, 1876. The identities of Gymnodactylus mysoriensis Jerdon, 1853 and G. littoralis Jerdon, 1853, two valid species, remain to be fixed through the designation of neotypes selected from topotypical specimens so as to facilitate future description of new taxa from southern India. Ma n a m e n d r a -Ar a c h c h i e t a l .

Table 1. Checklist of nominal species of Cnemaspis from peninsular India and Sri Lanka and their type localities (cross (†) = subjective synonym; double-cross (‡) = objective synonym; asterisk (*) = junior primary homonym; all other species are considered valid).

Indian species Locality

† Cnemaspis anaikattiensis Mukherjee et al., 2005 Anaikatti Hills, Western Ghats, Tamil Nadu State

Cnemaspis australis new species Tirunelveli, Tamil Nadu State Gymnodactylus beddomei Theobald, 1876 S Tirunelveli, Tamil Nadu State † Gonatodes bireticulatus Annandale, 1915 Sholakal, Sispara Ghat (Nilgiris), Tamil Nadu State Goniodactylus boiei Gray, 1842 India Cnemaspis goaensis Sharma, 1976 3 km S of Forest Rest House, Canacona, Goa Gymnodactylus gracilis Beddome, 1870 Palghat [Palakkad], Kerala State Cnemaspis heteropholis Bauer, 2002 North Kanara, Karnataka State Goniodactylus indica Gray, 1846 Madras (Chennai, or ‘Madras Presidency’=Tamil Nadu) Cnemaspis indraneildasii Bauer, 2002 North Kanara, Karnataka State Gymnodactylus jerdonii Theobald, 1868 No specific locality: India Gymnodactylus littoralis Jerdon, 1853 Sea Coast in Malabar [Kerala State] * Gymnodactylus maculatus Beddome, 1870 Replacement name: C. sisparensis (Theobald, 1876) Gymnodactylus malabarica Jerdon, 1853 Forests of Malabar [Kerala State] Cnemaspis monticola new species Wynaad, Kerala State * Gymnodactylus marmoratus Beddome, 1870 Replacement name: C. beddomei (Theobald, 1876) Gymnodactylus mysoriensis Jerdon, 1853 Bangalore, Karnataka State Cnemaspis nairi Inger, Marx & Koshy, 1984 Ponmudi, Trivandrum District, Kerala State Cnemaspis nilagirica new species Nilgherries, Nilgiri District, Tamil Nadu State Gymnodactylus ornata Beddome, 1870 Tirunelveli, Tamil Nadu State Cnemaspis otai Das and Bauer, 2000 Vellore, North Arcot District, Tamil Nadu State Gymnodactylus planipes Beddome, 1870 Nellicootah, below the Nilgiris Gymnodactylus sisparensis Theobald, 1876 Sholakal, Sispara Ghat (Nilgiris), Tamil Nadu State Gymnodactylus wynadensis Beddome, 1870 Wynaad, Kerala State Cnemaspis yercaudensis Das and Bauer, 2000 Yercaud, Shevaroyan (Shevroy) Range, Tamil Nadu State Sri Lankan species

Cnemaspis alwisi Wickramasinghe & Munindradasa, 2007 Dolukanda, Kurunegala District Cnemaspis amith new species Ceylon (= Sri Lanka) Cnemaspis clivicola new species Nanu Oya, Nuwara Eliya District Cnemaspis gemunu Baueret al., 2007 Hakgala, Nuwaraeliya District † Gymnodactylus humei Theobald, 1876 Kandy Cnemaspis kallima new species Gammaduwa, Matale District Gymnodactylus kandianus Kelaart, 1852 Ceylon (= Sri Lanka) Cnemaspis kumarasinghei Wickramasinghe & Munindradasa, 2007 Maragala, Monaragala District Cnemaspis latha new species Bandarawela, Nuwara Eliya District Cnemaspis menikay new species Ihala-Kalugala, Allauwa, Kegalle District Cnemaspis molligodai Wickramasinghe & Munindradasa, 2007 Waratalgoda, Rathnapura District Cnemaspis pava new species Nawalapitiya, Kandy District Cnemaspis phillipsi new species Gammaduwa, Matale District Cnemaspis podihuna Deraniyagala, 1944 Lahugala, in the Eastern Province Cnemaspis pulchra new species Rakwana, Ratnapura District Cnemaspis punctata new species Gammaduwa, Matale District ‡ Cnemaspis ranwellai Wickramasinghe, 2006 Gannoruwa, Kandy District Cnemaspis retigalensis Wickramasinghe & Munindradasa, 2007 Weweltenna, Ritigala, Anuradhapura District Cnemaspis samanalensis Wickramasinghe & Munindradasa, 2007 Adam’s Peak, Ratnapura District Gymnodactylus scalpensis Ferguson, 1877 Gannoruwa, Kandy District Cnemaspis silvula new species Kombala-Kottawa Forest Reserve, Galle District Gonatodes kandianus tropidogaster Boulenger, 1885 Ceylon (= Sri Lanka) Cnemaspis upendrai new species Pussellawa, Kandy District

10 Zeylanica Re v i s i o n o f Cn e m a s p i s

Introduction As a result, three new Indian species are described (all with well-defined type localities) based on The diminutive day-geckos of the tropical Asian and historical museum material, for which we hope African genus Cnemaspis Strauch, 1887, are known extant populations will be discovered in the course from 34 nominal species in peninsular India and Sri of future exploration. Lanka (Table 1). Accretion of species in the genus in In consequence of this paper, all but two of the this region, having begun with C. boiei (Gray, 1842), species of Cnemaspis from peninsular India and has been irregular: 18 species-group names became Sri Lanka (viz., Gymnodactylus mysoriensis Jerdon, available in the 19th century, four in the 20th century 1853, and G. littoralis Jerdon, 1853) now possess and 12 since 2000. The relatively large number of recently-described name-bearing types, facilitating names created in the past few years is suggestive of the reliable description of newly-discovered species an increased pace of biodiversity exploration in this by other workers in the future. We hope that this region, and hints at the possibility of many more will enable future workers discovering putative species of these geckos being discovered. Indeed, new species from this region to avoid creating based on our experience in Sri Lanka, we predict unnecessary new names (e.g., see Remarks under that if present trends continue, species-richness C. sisparensis, below). in the genus could double in the coming decade, In addition to a new species discovered among especially as more exploration takes place in the the syntype series of C. kandiana, we also describe Western Ghats. here 11 new species of Cnemaspis from Sri Lanka, This being so, a stable nomenclature for the which were discovered in the course of a saurian geckos in this group is clearly desirable. Given the survey we conducted in the island during the past wide distribution of the genus, the large number of decade. While this project was nearing completion, species involved (~51; see Bauer, 2000; Bauer et. al., Wick­rama­singhe & Munin­dra­dasa (2007), despite 2007; Das, 1993; Das & Bauer, 2000; Das & Bauer, 2000; being aware of our work and disregarding the Code Das & Sengupta. 2000; Das & Grismer. 2003; Das & of Ethics of the International Code of Zoological Leong, 2004; Das, 2005; Kluge, 2001; Mukherjee et. Nomen­clature (1999), published five new species and al, 2005; Wickramasinghe, 2006; Wickramasinghe in­validly designated a number of neotypes and new & Munindradasa, 2007), and the scattering of types ‘syntypes’ (sic). The consequential nomenclatural and among repositories in several continents, a revision ethical issues have been dealt with by Pethiyagoda of the entire genus is unlikely to take place in the (2007). We examined the type material of Wickrama­ near term. Nevertheless, the identities of the ‘old’ singhe & Munindra­dasa (2007) in the National Mus­ species need to be clarified in order to facilitate the eum of Sri Lanka and found their descriptions to be reliable description of new ones. This is particularly in several respects at variance with their types. Given necessary given that the descriptions of the most that their species were only very recently published, recent 12 species were made largely without refer­ and in a journal to which there is ready electronic ence to the older type material. access, we here refrain from re-describing them, In this paper we re-describe (based on an but instead provide diagnoses that we hope will aid examination of all extant type material) the species identification of these species by future workers; we published prior to the modern period of Indian also provide brief remarks by way of corrigenda. and Sri Lankan Cnemaspis taxonomy (in which new The subjective synonymy of Cnemaspis in our names were accompanied by detailed descriptions region of interest is not rich, and prior to this work and diagnoses), which we consider as having contained only a single name, Gymnodactylus humei commenced with Inger et al. (1984). So as to stabilize Theobald, 1876. We have strived to maintain this the nomenclature, we select lectotypes for species happy situation, and chosen (in the interests of that were based on series of syntypes. Bearing in ‘nomenclatural parsimony’: see Bossuyt & Dubois, mind the need to preserve stability, in some cases 2001) to minimize the synonymy by stabilizing old we designate neotypes for species whose types we names through the designation of name-bearing are confident are lost; in others, we refrain from types consistent with their original descriptions and doing so in the hope that a future revision of the type localities. By these means we hope we have left Indian members of the genus (by other authors) will the taxonomy of this group in a better state than that provide for this. in which we found it. The examination of old museum material brought A few Cnemaspis, most notoriously the catchall to light cases among the Indian taxa in which the species C. kandiana (type locality ‘Ceylon’ but syntype series comprised more than a single species. restricted to the vicinity of Kandy by virtue of the

Vol. 7, No. 1 11 Ma n a m e n d r a -Ar a c h c h i e t a l .

geographical specific name), have historically been and heel, with both tibia and tarsus flexed); TBW, considered to have extremely wide ranges, having tail base width; TL, toe length (distance between been recorded from as far afield as the Andamans. As tip of claw and nearest fork); TYD, tympanum-eye concluded by Bauer et al. (2007), these were probably distance (distance between anterior-most margin for the most part misidentified (undescribed) species, of tympanum and posterior-most margin of eye); and we hope future workers will describe them. TYE, tympanum diameter; UAL, upper-arm length Given the clarity of the present taxonomy, we (distance between axilla and angle of elbow); UEW, refrain from providing a discussion of the historical supercilium width (distance between supraciliary synonymy because this is unlikely to be useful to scale row and lateral bony margin of frontal). future workers. Scale counts and morphological observations were made using an Olympus SZ40 dissecting Materials and methods microscope. Meristic characters were taken as follows: supralabials and infralabials, first labial scale Material for this study were collected in Sri Lanka as to last labial scale towards gape, which is distinctly part of a survey of the island’s Sauria commissioned larger than the granular scales at gape; subdigital by the Department of Wildlife Conservation in 1998. lamellae, from first proximal enlarged scansor greater Specimens were fixed in 10% formalin and preserved than twice width of the largest palm scale, to distal- in 70% ethanol; samples of tissue for DNA analysis most lamella at tip of digit; ventrals, from mental to were stored in 95% ethanol. The new material a scale on preanal scale row, separately for each sex. collected in this survey is deposited in the collection Sex was determined by the presence of hemipenial of the Wildlife Heritage Trust of Sri Lanka (WHT). bulges and preanal and/or femoral pores. Caudal- All available types of Sri Lankan and peninsular scale characters are given only for specimens with Indian Cnemaspis, deposited in The Natural History unregenerated tails (or unregenerated portions of Museum, London (BMNH), the National Museum of tails), except in the case of type specimens in which Sri Lanka, Colombo (NMSL), and Zoological Survey the tails have been lost or regenerated. of India, Calcutta (ZSI), were also examined. Photographs were taken with a Cannon IXUS The following measurements were taken with 50 digital camera. Colour photographs are of living dial Vernier callipers (to the nearest 0.1 mm): AG, specimens unless otherwise stated. Altitudes are dis­tance between axilla and groin; ED, eye diameter given in metres above mean sea level; geographic (horizontal diameter of orbit); EN, distance between co-ordinates were taken using a Magellan 12- anteriormost point of orbit and middle of nostril; ES, channel GPS (geodetic datum WGS-84: World snout length (distance between anteriormost point of Geodetic System of 1984), and in some instances, orbit and tip of snout); FEL, femur length (distance using topographic maps (1 inch: 1 mile, Survey between groin and knee); FL, finger length (distance Department, Colombo). between tip of claw and the nearest fork); FOL, foot The identification of sympatric species (except for length (distance between heel and tip of longest toe, Cnemaspis), is based on Smith (1935), Taylor (1950, with both foot and tibia flexed); HL, head length 1953), Deraniyagala (1953), Greer (1991), Batuwita & (distance between posterior edge of mandible and Bahir (2005), Bahir & Maduwage (2005), Bahir & Silva tip of snout); HW, head width (maximum width of (2005); nomenclature follows Kluge (2001). head); IN, internarial distance (least distance between This work necessitated examination of the type inner margins of nares); IO, interorbital width (least specimens in the Zoological Survey of India, Kolkata distance between upper margins of orbits); LAL, (ZSI). This was done during a visit to that institution lower-arm length (distance from elbow to wrist with by K.M.-A. in February, 2007. The ZSI’s collection is both upper arm and palm flexed); MBE, distance of immense historical and scientific interest, and it between angle of gape and posterior-most point is regrettable that the custodians of this collection of orbit; MFE, distance between angle of gape and permitted access to the specimens only after repeated ant­erior­most point of orbit; MN, mandible to nostril requests, and even then for less than three hours per distance (distance between angle of gape and middle day. Most frustratingly, it was prohibited to photo­ of nostril); PAL, palm length (distance between graph specimens. Such arbitrary restrictions sub­ posterior-most margin of palm and tip of longest vert sound science are a sad indictment on a once finger); SVL, snout–vent length (distance between famous institution; they also call into question the tip of snout and anterior margin of vent); TAL, tail res­ponsiveness of ZSI to Recommendation 72F of length (distance between anterior margin of vent to the International Code of Zoological Nomenclature tail tip); TBL, tibia length (distance between knee (1999).

12 Zeylanica Re v i s i o n o f Cn e m a s p i s

TAXONOMY medial scale (on left side; right side damaged); postmentals widely separated; postmental on Gekkonidae: Cnemaspis left side bordered posteriorly by 4 smooth scales, including a medial scale (probably 8 scales bordering PENINSULAR INDIAN SPECIES postmentals) (Fig. 1b). Supralabials 6, infralabials 6, decreasing in size towards gape (Fig. 1c). Ear Cnemaspis boiei (Gray, 1842) opening oval, higher than wide, oblique (TW 50.0% Goniodactylus boiei Gray, 1842 of TH). Scales on ventral surface of neck smooth. (Figures 1a–h; Table 2; Plate 1a, b) Granules between anterior margin of ear opening and posterior margin of eye, 24, larger than those of Lectotype (here designated). BMNH 1962.181A occipital region. (male), 35.3 mm SVL, “India”, Presented by General Body slender, elongate. Mid-dorsal granules T. Hardwicke. isolated, scattered, homogeneous, keeled (Fig. 1d), a little larger than scales of occipital region; spine-like Paralectotypes. BMNH 1962.181B (female), 38.4 tubercles absent on flank; abdominal scales smooth mm SVL; BMNH 1962.181C (female), 38.8 mm SVL; (Fig. 1e). Ventrals, 117. Pectoral and abdominal scales BMNH 1962.181D (male), 30.7 mm SVL, “India”, juxtaposed (Fig. 1e), those around vent and base of presented by General T. Hardwicke. tail smooth, subimbricate to imbricate (Fig. 1f). No preanal or femoral pores (Fig. 1f). Diagnosis Forelimbs moderately long, slender (LAL 13.5% Cnemaspis boiei differs from its peninsular Indian SVL, UAL 13.0% SVL); hind limbs long (TBL 16.9% and Sri Lankan congeners by a combination of SVL, FEL 18.6% SVL); tibia short. Dorsal scales on the following characters. Maximum SVL 38.8 mm; both fore and hind limbs smooth; ventral scales on paired postmentals separated by a medial scale; each upper arm narrow, pointed, smooth; on lower arm postmental bounded by 4 scales; ventrals, 117; dorsal smooth. Scales on anterior surface of thigh small, scales homogeneous; spine-like tubercles absent on granular, subequal to those on ventral surface (Fig. flank; gular, pectoral and abdominal scales smooth; 1f). Scales on dorsal and ventral surface of shank pectoral and abdominal scales juxtaposed; no femoral smooth. Scales on manus and pes smooth, rounded. or preanal pores in either sex; subcaudals on median Digits elongate, slender, all bearing slightly recurved row enlarged, subequal; enlarged tubercles absent on claws; subdigital lamellae entire; subdigital lamellae base of tail; supralabials to angle of jaws, 6; subdigital on finger I, 9, finger II, 12, finger III, 14, finger IV, lamellae on digit IV of pes, 17. 14, finger V, 12; toe I, 8, toe II, 10, toe III, 15, toe IV, 17, toe V, 13; interdigital webbing absent; relative Description of lectotype length of digits (fingers): IV (3.6 mm) > III (3.2 mm) An adult male, 35.3 mm SVL (for measurements, see > V (2.7 mm) > II (2.5 mm) > I (1.8 mm); (toes) IV Table 2). Head relatively short (HL 25.2% of SVL), (4.0 mm) > III (3.9 mm) > II (3.4 mm) > V (3.0 mm) moderately broad (HW 15.2% of SVL, HW 60.6% of > I (2.2 mm). HL), distinct from neck. Snout length greater than Tail subcylindrical, tapering, its length greater eye diameter (ED 76.7% of ES); interorbital broad than SVL (TAL 129.7% of SVL); tail base distinctly (IO 15.7% of HL); pupil rounded. Granules on snout swollen. Enlarged tubercles absent on base of tail. smooth, equal in size, larger than those of occipital Dorsal scales on mid-tail smooth, imbricate directed region; scales on interorbital and supercilium backwards (Fig. 1g); post-cloacal spurs absent; smooth; scales on ventral surface of head conical, not subcaudals smooth, hexagonal (some scales divided), keeled. Rostral scale in contact with first supralabial, in a regular enlarged median series (Fig. 1h). partially divided by a medial groove: a narrow transverse suture begins from medial groove (Fig. Coloration. In preservative: (see Plate 1a, b) dorsally 1a). Nares separated medially by two enlarged and ventrally olive; no blotches or markings. supranasals and 2 small internasals (Fig. 1a). Nares rounded, dorsally orientated, not in contact with Variation. Supralabials 7 in paralectotype BMNH first supralabial. Three postnasals, the lower one 1962.181B. Pores absent in both sexes. SVL range in broad contact with first supralabial on left side 30.7–38.8 mm (4 ex.). (Fig. 1a). Mental subpentagonal, wider than long, posteriolaterally in contact with a single elongate, Comparisons enlarged, postmental and a single subpentagonal Cnemaspis boiei differs from each of its peninsular

Vol. 7, No. 1 13 Ma n a m e n d r a -Ar a c h c h i e t a l . R 8×2 7.2 1.3 1.2 5.6 4.9 6.7 9.8 1.3 5.7 5.0 4.6 3.1 4.0 5.2 8.0 3.0 3.7 4.2 4.2 4.0 8.9 7.0 2.7 4.5 5.5 5.5 4.7 5.6 4.0 5.8 2.4 42.3 10.6 32.7 BMNH 74.4.29.729 C. nilagirica holotype (ZSI holotype C. ornata ; lectotype C. boiei; C. 6×3 5.3 1.0 1.0 4.2 3.7 5.5 7.4 0.7 3.3 4.5 3.4 2.3 3.1 4.0 6.2 2.0 2.5 2.5 2.7 2.4 4.9 8.6 5.4 1.9 3.0 3.1 3.5 3.2 3.7 2.9 1.6 3.3 30.4 35.0 BMNH 82.5.22.67 C. australis holotype (BMNH 74.4.29.729)

R 74.4.29.400) of ) of 1962.181A ) 1.7 4.6 1.0 0.8 3.7 3.3 4.9 7.2 0.6 3.6 3.6 2.5 2.1 2.9 3.6 5.0 1.4 2.1 2.4 2.5 2.2 4.5 7.7 4.4 3.0 – – 3.0 3.5 2.9 2.7 3.1 1.4 17.5 26.0 BMNH 74.4.29.372 C. monticola C. australis ; and 5×2 – 6.4 1.2 1.2 4.7 4.7 6.3 8.4 0.9 4.3 4.7 2.8 2.6 3.1 3.6 6.9 1.8 2.7 3.0 2.8 2.6 5.5 9.1 6.1 3.8 2.1 2.8 3.4 3.5 3.4 3.6 3.4 1.8. lectotype (BMNH 33.4 BMNH 74.4.29.355 C. wynadensis 82.5.22.67) of C. indica; ; lectotype (BMNH lectotype beddomii ; 8×1 C. – 8.4 1.4 1.4 7.1 5.4 7.9 1.4 6.3 5.7 7.7 4.4 3.6 4.5 6.1 7.7 2.7 4.2 4.6 5.0 4.2 8.1 5.5 2.8 4.5 5.1 6.0 5.1 4.0 5.5 1.6 11.3 42.0 12.2 BMNH C. ornata 74.4.29.400 holotype (BMNH 5×5 – 6.1 1.1 1.2 3.6 3.6 5.3 7.1 1.1 3.9 3.8 5.4 3.0 2.4 2.7 3.6 5.5 1.8 2.4 2.9 2.5 2.1 5.3 7.3 4.1 2.1 3.3 3.8 3.7 3.0 2.9 2.9 1.2 29.3 BMNH C. indica 46.11.22.22b 46.11.22.22b C. monticola; 4×6 – 6.2 0.9 1.0 4.3 3.5 5.3 7.4 0.7 3.5 4.0 5.4 3.7 2.5 3.0 3.9 6.3 2.0 2.2 2.7 2.9 2.6 5.5 8.3 4.0 2.1 3.1 3.5 3.9 3.5 2.5 4.4 1.4 29.7 BMNH 74.4.29.393 C. gracilis C. gracilis; lectotype (BMNH of 46.11.22.22b) 3×2 – 5.3 0.9 0.5 3.7 3.0 5.3 7.6 0.7 3.4 3.1 5.0 2.0 2.8 2.5 2.9 8.0 1.4 2.5 2.3 2.5 2.0 4.9 8.3 2.7 2.8 3.5 2.7 2.5 1.8 2.3 3.4 1.2 ) and paralectotype (ZSI 5859) of 5859) (ZSI paralectotype and 1946.9.4.83 ) ZSI 27.5 22110 22110 C. goaensis 8×4 4.8 4.4 5.4 1.3 1.4 4.0 4.7 7.9 0.9 4.1 6.0 6.6 2.8 3.3 3.5 4.3 1.8 2.5 3.2 3.6 2.7 6.2 8.9 4.0 2.2 3.4 3.9 4.0 3.0 3.1 4.6 0.6 holotype (BMNH 74.4.29.372) of 45.8 35.3 BMNH C. boiei 1962.181A 1962.181A 8×7 – 7.2 – 9.1 1.5 2.0 5.6 8.9 1.5 7.3 8.8 8.7 4.1 4.3 4.4 5.3 – – – – – – 5.4 – – – – – 3.8 5.2 1.5 ZSI 12.5 46.3 13.7 5859 C. wynadensis; lectotype (BMNH 74.4.29.393) of C.beddomii 1.1 6.3 6.7 9.5 1.6 2.2 6.2 8.1 1.2 6.1 8.1 3.2 3.5 3.8 6.1 3.6 4.7 5.0 5.2 5.6 5.9 3.4 5.6 6.1 6.7 5.8 6.2 6.0 2.5 11.4 34.8 12.1 46.6 10.3 13.1 BMNH 1946.9.4.83 C. goaensis;

TAL HW IN IO LAL MBE MFE MN NS PAL SVL DBE TBL DFE ED EN ES FEL I FL II FL III FL IV FL V FL FOL HL TBW I TL II TL III TL IV TL V TL TYD TYE (×10) Measurements (in mm) of lectotype (BMNH lectotype of mm) (in Measurements 2. Table of 22110) UAL UEW (BMNH 74.4.29.355) of of C. nilagirica .

14 Zeylanica Re v i s i o n o f Cn e m a s p i s

a b

c d

e f

g h

Figure 1. Cnemaspis boiei: BMNH 1962.181A, lectotype, male, a, dorsal aspect of snout (right side damaged); b, ventral aspect 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.

Vol. 7, No. 1 15 Ma n a m e n d r a -Ar a c h c h i e t a l .

Indian and Sri Lankan congeners, respectively, by subcaudals on distal half of tail carinate. Cnemaspis the following opposing suites of characters: amith: gular scales carinate; pectoral and abdominal Cnemaspis beddomei: preanal pores 6–9. Cnemaspis scales imbricate; dorsal granules heterogeneous; goaensis: presence of pores. Cnemaspis littoralis: subcaudals on median row enlarged, irregular. a black marking on nape. Cnemaspis mysoriensis: spine-like tubercles present on flank, femoral pores Remarks 2 or 3. Cnemaspis indraneildasii and C. heteropholis: The precise type locality of Cnemaspis boiei within dorsal scalation heterogeneous. Cnemaspis jerdonii: India is not known. However, it is immediately spine-like tubercles present on flank, presence of identifiable in that both males and females lack both femoral pores. Cnemaspis nairi: presence of preanal femoral and preanal pores, a character otherwise pores. Cnemaspis otai: 4 preanal and 3 femoral pores. shared only with the northeast-Indian species, C. Cnemaspis sisparensis: postmentals in contact, spine- assamensis Das and Sengupta, 2000. like tubercles absent on flank, 7 or 8 femoral pores. Cnemaspis wynadensis: 5 femoral pores. Cnemaspis Cnemaspis indica (Gray, 1846) yercaudensis: 2 preanal and 3 femoral pores. Cnemaspis Gymnodactylus indicus Gray, 1846 gracilis: pores present, postmentals in contact, (Figures 2a–h; Table 2; Plate 1c, d) subcaudals in median series not enlarged. Cnemaspis indica: postmentals in contact, presence of femoral Lectotype (here designated). BMNH 46.11.22.22b pores. Cnemaspis ornata: dorsal tubercles arranged (male), 29.3 mm SVL, “Madras”. in 15 longitudinal rows, a dark-brown collar present on nape, preanal pores 6–8. Cnemaspis wicksii and C. Paralectotypes. BMNH 46.11.22.22a (male), 30.4 mm andersonii: presence of pores. Cnemaspis monticola: SVL; BMNH 46.11.22.22c (female), 30.0 mm SVL spine-like tubercles on flank; pectoral and abdominal (deformed by dehydration), “Madras”. scales imbricate. Cnemaspis australis: subcaudals on median row pointed, strongly keeled. Cnemaspis Diagnosis nilagirica: pectoral and abdominal scales weakly Cnemaspis indica differs from its peninsular Indian keeled; subcaudals pentagonal or quadrate. and Sri Lankan congeners by a combination of Cnemaspis tropidogaster: gular, pectoral and the following characters. Maximum SVL 30.4 mm; abdominal scales keeled, spine-like tubercles present paired postmentals separated by a medial scale; each on flank. Cnemaspis kandiana, C. podihuna, C. scalpensis, postmental bounded by 3 scales including medial C. gemunu and C. phillipsi: spine-like tubercles present scale; ventrals, 94; dorsal scales homogeneous, on flank. Cnemaspis alwisi: 9 femoral pores, spine-like imbricate, carinate; spine-like tubercles absent on tubercles present on flank. Cnemaspis molligodai: 5 flank; ventrals smooth, imbricate; no preanal pores; preanal and 8 or 9 femoral pores, spine-like tubercles 5 femoral pores on each side; subcaudals smooth, present on flank. Cnemaspis samanalensis: gular, median series slightly enlarged; supralabials to pectoral and abdominal scales tricarinate, subcaudals angle of jaws, 7; subdigital lamellae on digit IV of keeled, spine-like tubercles present on flank. pes, 14. Cnemaspis retigalensis: dorsal scales heterogeneous, gulars carinate, a single preanal pore and 3 or 4 Description of lectotype femoral pores, spine-like tubercles present on flank; An adult male, 29.3 mm SVL (for measurements, subcaudals in an irregular enlarged median series. see Table 2). Head relatively short (HL 24.9% of Cnemaspis punctata: 5–7 femoral pores, spine-like SVL), broad (HW 20.8% of SVL, HW 83.5% of HL), tubercles present on flank, distinctive spotted dorsal distinct from neck. Snout length somewhat greater coloration. Cnemaspis kumarasinghei, C. menikay and than eye diameter (ED 66.6% of ES); interorbital C. latha: presence of pores; presence of spine-like broad (IO 16.4% of HL); pupil rounded. Granules tubercles on flank. Cnemaspis silvula, C. pulchra and on snout intermixed with both smooth and keeled C. pava: pectoral and abdominal scales keeled, spine- scales, larger than those of occipital region; scales like tubercles present on flank. Cnemaspis clivicola: on interorbital and supercilium granular, keeled, pectoral and abdominal scales keeled, spine-like those on ventral surface of head weakly carinate. tubercles present on flank, subcaudals in median Rostral scale partially divided by a medial groove, series not enlarged. Cnemaspis upendrai: some gular in contact with first supralabial. Nares separated scales tricarinate, pectoral and abdominal scales by two supranasals and a single internasal scale; keeled, spine-like tubercles on flank. Cnemaspis internasal as large as supranasals (Fig. 2a). Nares kallima: spine-like tubercles present on flank, rounded, dorsally orientated, not in contact with

16 Zeylanica Re v i s i o n o f Cn e m a s p i s

a b

c d

e f

g h

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 and ventral 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.

Vol. 7, No. 1 17 Ma n a m e n d r a -Ar a c h c h i e t a l .

first supralabial. Two postnasals, the lower one in Coloration. In preservative: (see Plate 1c, d) dorsally broad contact with first supralabial (Fig. 2a). Mental light brown with indistinct dark marking on dorsum; subpentagonal, as wide as long, posteriolaterally tail with a longitudinal mid-dorsal stripe; venter in contact with two enlarged postmentals and pinkish-white. medially with a single hexagonal postmental scale; postmentals bordered posteriorly by 5 smooth scales Comparisons. Cnemaspis indica differs from each of including medial scale (Fig. 2b). Supralabials 7, its peninsular Indian and Sri Lankan congeners by infralabials 7, decreasing in size towards gape (Fig. the following opposing suites of characters: 2c). Ear opening circular (TW 100.0% of TH). Scales Cnemaspis beddomei: no interanasals, preanal pores on ventral surface of neck smooth. Nineteen granules 6–9, no femoral pores. Cnemaspis goaensis: preanal between anterior margin of ear opening and posterior pores present. Cnemaspis littoralis: body slender, margin of eye. dorsal scales granular, median series of subcaudals Body robust, short. Mid-dorsal granules enlarged, hexagonal. Cnemaspis mysoriensis: spine- homogeneous, imbricate and carinate (Fig. 2d), like tubercles present on flank, femoral pores 2 or intermixed with a few smooth scales; tubercles 3. Cnemaspis indraneildasii, C. wicksii, C. monticola absent on flank; abdominal scales smooth, imbricate, and C. andersonii: spine-like tubercles present on obtusely pointed and wider than long (Fig. 2e). flank. Cnemaspis heteropholis: heterogeneous dorsal Ventrals, 94; scales around vent and base of tail scalation. Cnemaspis jerdonii: spine-like tubercles smooth, imbricate. No preanal pores; 5 femoral present on flank. Cnemaspis nairi: no femoral pores. pores (Fig. 2f). Cnemaspis otai and C. sisparensis: paired postmentals Forelimbs relatively short, slender (LAL 12.2% in contact. Cnemaspis wynadensis: subcaudals in SVL, UAL 9.8% SVL); hind limbs moderately long a regular enlarged (widened) series. Cnemaspis (TBL 18.4% SVL, FEL 18.7% SVL); tibia short. Dorsal yercaudensis: 2 preanal pores. Cnemaspis boiei: scales on both fore and hind limbs with 1–3 keel- no preanal or femoral pores. Cnemaspis gracilis: like ridges; ventral scales on upper and lower arm postmentals in contact. Cnemaspis ornata: dorsal smooth, a few of them carinate. Scales on anterior tubercles arranged in 15 longitudinal rows, dark- surface of thigh carinate, those on posterior surface brown collar on nape, no femoral pores, preanal smooth (Fig. 2f). Scales on dorsal and ventral surface pores 6–8. Cnemaspis australis: subcaudals pointed, of shank mostly smooth, a few carinate. Dorsal strongly keeled. Cnemaspis nilagirica: dorsal scales scales on manus carinate, ventrals smooth. Digits isolated, scattered; subcaudals on median row elongate, slender, all bearing slightly recurved claws; enlarged, subequal. subdigital lamellae unnotched, entire, except for 2 Cnemaspis tropidogaster: gular, pectoral and or 3 fragmented scales at base; subdigital lamellae abdominal scales keeled, spine-like tubercles on finger I, 8, finger II, 11, finger III, 11, finger IV, 11, present on flank. Cnemaspis kandiana, C. podihuna, finger V, 9; toe I, 9, toe II, 12, toe III, 11, toe IV, 14, C. scalpensis, C. gemunu and C. phillipsi: spine-like toe V, 9; interdigital webbing absent; relative length tubercles present on flank. Cnemaspis alwisi: 9 of digits (fingers): III (2.9 mm) > IV (2.5 mm) > II femoral pores, spine-like tubercles present on flank. (2.4 mm) > V (2.1 mm) > I (1.8 mm); (toes) > III (3.8 Cnemaspis molligodai: 5 preanal and 8 or 9 femoral mm) > IV (3.7 mm) > II (3.3 mm) > V (3.0 mm) > I pores, spine-like tubercles present on flank. Cnemaspis (2.1 mm). samanalensis: gular, pectoral and abdominal scales Tail tapering (damaged); tail base distinctly tricarinate, subcaudals keeled, spine-like tubercles swollen. Enlarged tubercles absent on base of tail. present on flank. Cnemaspis retigalensis: dorsal Dorsal scales on mid-tail imbricate, carinate, directed scales heterogeneous, gulars carinate, pectoral and backwards (Fig. 2g); a conical post-cloacal spur abdominal scales smooth, a single preanal pore and present on each side; subcaudals smooth, those on 3 or 4 femoral pores, spine-like tubercles present median row slightly enlarged (Fig. 2h). on flank. Cnemaspis punctata: 5–7 femoral pores, spine-like tubercles present on flank, distinctive Variation. Five infralabials and 2 internasals in spotted dorsal Coloration. Cn emaspis kumarasinghei, paralectotype BMNH 46.11.22.22a; 3–5 femoral pores; C. menikay and C. latha: presence of preanal pores, pores absent in females; female paralectotype BMNH tubercles present. Cnemaspis silvula, C. pulchra, C. pava 46.11.22.22c with enlarged endolymphatic sacs, and C. clivicola: pectoral and abdominal scales keeled, suggestive of the specimen having been collected spine-like tubercles present on flank. Cnemaspis during the breeding season; SVL range 29.3–30.4 upendrai: some gular scales tricarinate, pectoral and mm (3 ex.). abdominal scales keeled, spine-like tubercles present

18 Zeylanica Re v i s i o n o f Cn e m a s p i s

on flank. Cnemaspis kallima: gular scales smooth, ventral surface of neck smooth, imbricate. Dorsal spine-like tubercles present on flank; subcaudals surfaces of both fore and hind limbs covered by on distal half of tail carinate. Cnemaspis amith: gular imbricate scales. scales carinate; pectoral and abdominal scales This is consistent with the original description of smooth; dorsal granules heterogeneous; subcaudals C. jerdonii (Theobald, 1868), which was as follows: on median row enlarged, irregular. “Male. Back uniformly granular. Tail with a single row of large subcaudal plates or scuta, with a few Remarks large scales along their edges. A strong double fold Cnemaspis indica is distinguished from all its on the throat. Scales of the belly small, rhomboid. peninsular Indian and Sri Lankan congeners by Along the sides two rows of distant erect spines, possessing imbricate dorsal scales. become obsolete on the tail. Sides keeled. Pubic region covered with same scales as the belly. No large Cnemaspis jerdonii (Theobald, 1868) præanals. Femoral pores eight on each thigh; large, Gymnodactylus jerdonii Theobald, 1868 cup-shaped, placed along the hind edge of limb, bordered in font by rather largish scales, and behind Syntypes. ZSI 6179 (adult), 9.6 mm HL; ZSI 6180 by the granular scales of the back. Length, head (juvenile), 3.4 mm HL, no specific locality; both and body 1.50, tail 1.50==3.00. Colour (in spirits) specimens too badly damaged and fragmented to above, greenish grey mottled with brown, beneath, facilitate measurements. greenish white. Nine upper and seven lower labials. Nostrils close behind the rostral. Lower rostral large, Note. The syntypes of this species are in too poor just separates a pair of small triangular chin plates. a state to facilitate detailed description, for which Allied to G. Mysoriensis by its spines, but not to be reason we also refrain from selecting a lectotype. identified with any species described by Jerdon” Fortunately, Theobald’s original description is (Theobald, 1868). remarkably detailed for its time: the following diagnosis and description, based in part on the Comparisons. Cnemaspis jerdonii differs from each syntypes, is supplemented by data provided in the of its peninsular Indian and Sri Lankan congeners, original description. respectively, by the following opposing suites of characters: Diagnosis Cnemaspis beddomei: spine-like tubercles absent Cnemaspis jerdonii differs from its peninsular Indian on flank, preanal pores 6–9, femoral pores absent. and Sri Lankan congeners by a combination of Cnemaspis goaensis: postmentals widely separated by the following characters. Maximum SVL 38.1 mm; a medial scale. Cnemaspis littoralis: a black marking paired postmentals separated by a medial scale; on nape. Cnemaspis mysoriensis: femoral pores 2 each postmental bounded by 2 scales including or 3. Cnemaspis indraneildasii: paired postmentals medial scale; dorsal scales homogeneous; spine-like separated by three small scales. Cnemaspis indica, tubercles present on flank; ventrals rhomboid; no C. nairi, C. otai, C. sisparensis and C. yercaudensis: erect spines laterally on tail; tail laterally keeled; spine-like tubercles absent on flank. Cnemaspis boiei: scales on dorsal and ventral surface of neck imbricate, no preanal or femoral pores. Cnemaspis gracilis: smooth; dorsal scales on both fore and hind limbs postmentals in contact. Cnemaspis ornata: dorsal imbricate. Preanal pores absent; 8 femoral pores on tubercles arranged in 15 longitudinal rows, dark- each side; median series of subcaudals enlarged; brown collar on nape, femoral pores absent, preanal supralabials and infralabials to angle of jaws, 9 and pores 6–8. Cnemaspis wicksii, C. monticola and C. 7 respectively. andersonii: gular scales carinate. Cnemaspis australis: subcaudals pointed, strongly keeled, in a regular Redescription of syntype ZSI 6179 enlarged median series. Cnemaspis nilagirica: gular Pupil rounded. Granules on snout smooth; scales scales carinate; enlarged caudal tubercles present. on interorbital, supercilium and ventral surface of Cnemaspis tropidogaster: gular, pectoral and head, smooth. Mental subpentagonal, wider than abdominal scales keeled. Cnemaspis kandiana and C. long, posteriolaterally in contact with two enlarged menikay: gular scales carinate. Cnemaspis podihuna: postmentals, medially with a single hexagonal preanal pores present. Cnemaspis scalpensis, C. postmental scale; postmentals bordered posteriorly gemunu and C. phillipsi: 12–16 femoral pores. by 5 smooth scales including medial scale. Ear Cnemaspis alwisi: caudal tubercles present. Cnemaspis opening elongate, oblique. Scales on dorsal and molligodai: 5 preanal pores. Cnemaspis samanalensis:

Vol. 7, No. 1 19 Ma n a m e n d r a -Ar a c h c h i e t a l .

gular, pectoral and abdominal scales tricarinate, divided by a medial groove, in contact with first subcaudals keeled. Cnemaspis retigalensis: dorsal supralabial (Fig. 3a). Nares separated by two slightly scales heterogeneous, gulars carinate. Cnemaspis enlarged supranasals and a single internasal scale; punctata: 5–7 femoral pores, distinctive spotted dorsal internasal extends towards snout tip (Fig. 3a). Nares coloration. Cnemaspis kumarasinghei and C. latha: rounded, dorsally orientated, not in contact with subcaudal scales in an enlarged irregular median first supralabial. Two postnasals, the lower one in series. Cnemaspis silvula, C. pulchra, C. pava and broad contact with first supralabial (Fig. 3a). Mental C. clivicola: pectoral and abdominal scales keeled. subtriangular, wider than long, posteriolaterally in Cnemaspis upendrai: some gular scales tricarinate; contact with two enlarged postmentals; postmentals pectoral and abdominal scales keeled. Cnemaspis broadly in contact with each other, bordered kallima: subcaudals on distal half of tail carinate. posteriorly by 5 smooth scales (Fig. 3b). Supralabials Cnemaspis amith: gular scales carinate; dorsal 7, decreasing in size towards gape; infralabials 6, granules heterogeneous. decreasing in size towards gape (Fig. 3c). Ear opening oval, higher than wide, oblique (TW 66.6% of TH). Scales on ventral surface of neck smooth. Seventeen Cnemaspis gracilis (Beddome, 1870) scales between anterior margin of ear opening and Gymnodactylus gracilis Beddome, 1870 posterior margin of eye. Conical granules laterally on (Figures 3a–g; Table 2; Plate 1e, f) neck, larger than those of interorbital region. Body slender, short. Mid-dorsal granules Lectotype (here designated). BMNH 74.4.29.393 heterogeneous, keeled or smooth, as large as scales (male), 29.7 mm SVL, “Palghat Hills” (Kerala State, of occipital region; large, keeled tubercles (Fig. 3d) in India), coll. Colonel R. H. Beddome. 11 or 12 rows on dorsum; spine-like tubercles present on flank; ventrals, 111, imbricate, smooth (Fig. 3e); Paralectotypes. BMNH 74.4.29.394 (female), 32.9 2 preanal pores separated by 2 unpored scales; 4 mm SVL; BMNH 74.4.29.395 (female), 27.9 mm SVL; femoral pores on each side (Fig. 3f). BMNH 74.4.29.396 (female), 25.4 mm SVL; BMNH Forelimbs moderately long, slender (LAL 14.4% 74.4.29.397 (male), 30.6 mm SVL; BMNH 74.4.29.398 SVL, UAL 14.8% SVL); hind limbs long (TBL 18.1% (male), 27.6 mm SVL, “Palghat hills” (Kerala State, SVL, FEL 21.2% SVL); tibia short. Dorsal scales on India), coll. Colonel R. H. Beddome. both fore and hind limbs smooth, a few of them carinate; ventral scales on upper and lower arm Diagnosis smooth. Dorsal scales on thigh carinate, a few of Cnemaspis gracilis differs from its peninsular Indian them smooth. Scales on dorsal and ventral surface of and Sri Lankan congeners by a combination of the thigh (Fig. 3f), shank, manus and pes, smooth. Digits following characters. Maximum SVL 32.9 mm; paired elongate, slender, all bearing slightly recurved claws; postmentals broadly in contact, each postmental subdigital lamellae entire, unnotched; subdigital bounded by 2 scales; ventrals, 111; dorsal scales lamellae on finger I, 10, finger II, 14, finger III, 16, heterogeneous with large, keeled tubercles in 11 or finger IV, 17, finger V, 14; toe I, 10, toe II, 16, toe III, 12 rows on dorsum; spine-like tubercles present on 20, toe IV, 22, toe V, 17; interdigital webbing absent; flanks; ventrals smooth, imbricate; 2 preanal pores, relative length of digits (fingers): IV (2.9 mm) > III separated by 2 unpored scales; 3–5 femoral pores (2.7 mm) > V (2.6 mm) > II (2.2 mm) > I (2.0 mm); on each side; subcaudals smooth, imbricate, not (toes) IV (3.9 mm) > III (3.5 mm) ~ V (3.5 mm) > II enlarged; supralabials to angle of jaws, 7; subdigital (3.1 mm) > I (2.1 mm). lamellae on digit IV of pes, 22. Tail tapering, with 6 large, conical tubercles at its base, about as large as tubercles on mid-dorsum. Description of lectotype Dorsal scales on mid-tail similar to scales on mid- An adult male, 29.7 mm SVL (for measurements, see body, directed backwards (Fig. 3g). Tail with about Table 2). Head long (HL 27.9% of SVL), broad (HW 6 oval, keeled tubercles separated by 2–6 scales; 20.8% of SVL, HW 74.6% of HL), distinct from neck. a conical post-cloacal spur present on each side; Snout length somewhat greater than eye diameter subcaudals smooth, not enlarged, imbricate (Fig. (ED 64.1% of ES); interorbital broad (IO 12.0% of HL); 3f). pupil rounded. Granules on snout smooth, larger than those of occipital region; scales on interorbital Coloration. In preservative: (see Plate 1e, f) dorsally and supercilium conical, granular; scales on ventral pinkish brown; dark spots along mid-dorsum, from surface of head smooth. Rostral scale partially nape to sacrum; two black streaks behind eye; mid-

20 Zeylanica Re v i s i o n o f Cn e m a s p i s

a b

c d

f

e g

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 region of mid-body, showing enlarged tubercles; e, preanal and hind-limb regions, showing arrangement of femoral and preanal 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.

Vol. 7, No. 1 21 Ma n a m e n d r a -Ar a c h c h i e t a l .

dorsal tubercles pale; limb with pale tubercles; digits and C. latha: postmentals separated medially by a banded; venter whitish, head with dark streaks. small scale. Cnemaspis silvula, C. pulchra, C. pava and C. clivicola: pectoral and abdominal scales keeled. Variation. Supralabials 6 or 7; infralabials 6 or 7; 3–5 Cnemaspis upendrai: some gular scales tricarinate, femoral pores; pores absent in females. SVL range pectoral and abdominal scales keeled. Cnemaspis 25.4–32.9 mm (7 ex.). kallima: subcaudals on distal half of tail carinate. Cnemaspis amith: gular scales carinate; subcaudals Comparisons on median row enlarged, irregular. Cnemaspis gracilis differs from each of its peninsular Indian and Sri Lankan congeners, respectively, by the Remarks following opposing suites of characters: Kluge (2001) listed Cnemaspis gracilis (Beddome, 1870) Cnemaspis beddomei: spine-like tubercles absent (= Gymnodactylus gracilis) in synonymy of C. kandiana on flank, no interanasals, subcaudals on median row (Kelaart, 1852). Examination of the respective types enlarged, in a series of alternating wider and narrower show that they are clearly two distinct taxa (see scales, preanal pores 6–9, no femoral pores. Cnemaspis respective diagnoses and comparisons). goaensis: postmentals widely separate by a medial scale. Cnemaspis littoralis: dorsal scales homogeneous, Cnemaspis ornata (Beddome, 1870) median series of subcaudals enlarged, hexagonal. Gymnodactylus ornatus Beddome, 1870 Cnemaspis mysoriensis: femoral pores 2 or 3. Cnemaspis (Figures 4a–j; Table 2; Plate 2c–f) indraneildasii: paired postmentals separated by small scale. Cnemaspis heteropholis: paired postmentals Lectotype (here designated). BMNH 74.4.29.400 separated by three small scales, no post-cloacal spurs. (male), 42.0 mm SVL, “South Tinnevelly Hills” Cnemaspis jerdonii: no preanal pores. Cnemaspis nairi: (Tirunelveli, Tamil Nadu State, India). no femoral pores. Cnemaspis otai: 4 preanal pores. Cnemaspis sisparensis: 7 or 8 femoral pores. Cnemaspis Paralectotypes. BMNH 74.4.29.401 (male), 41.7 wynadensis: subcaudals enlarged, subequal; no preanal mm SVL; BMNH 74.4.29.402 (male), 37.6 mm pores, 5 femoral pores, postmentals widely separated. SVL; BMNH 74.4.29.403 (juvenile), 25.6 mm SVL; Cnemaspis yercaudensis: postmentals separated by a BMNH 74.4.29.404 (juvenile), 26.6 mm SVL; BMNH single scale. Cnemaspis boiei: no preanal or femoral 74.4.29.405 (female), 37.2 mm SVL; BMNH 74.4.29.406 pores, postmentals widely separated by 2 medial (juvenile), 25.6 mm SVL; BMNH 74.4.29.407 (juvenile), scales. Cnemaspis indica: no preanal pores. Cnemaspis 26.7 mm SVL; BMNH 74.4.29.408 (juvenile), 27.4 mm ornata: dorsal tubercles arranged in 15 longitudinal SVL; BMNH 74.4.29.409 (juvenile), 22.6 mm SVL, rows, a dark-brown collar on nape, no femoral pores, “South Tinnevelly Hills” (Tirunelveli, Tamil Nadu preanal pores 6–8. Cnemaspis wicksii, C. andersonii State, south-western India). and C. monticola: gulars carinate. Cnemaspis australis: subcaudals pointed, strongly keeled, those on median Diagnosis row of slightly enlarged. Cnemaspis nilagirica: dorsal Cnemaspis ornata differs from its peninsular Indian scales homogeneous; pectoral and abdominal scales and Sri Lankan congeners by a combination of the weakly keeled; subcaudals on median row enlarged, following characters. Maximum SVL 42.0 mm; paired subequal. postmentals widely separated by a medial scale; each Cnemaspis tropidogaster: gular, pectoral and postmental bounded by 3 or 4 scales including medial abdominal scales keeled. Cnemaspis kandiana and scale; ventrals, 161; dorsal scales heterogeneous with C. menikay: gulars carinate. Cnemaspis podihuna, C. enlarged tubercles; spine-like tubercles absent on scalpensis, C. gemunu and C. phillipsi, homogeneous flank; ventrals smooth, subimbricate; 6–8 preanal dorsal scalation. Cnemaspis alwisi, 9 femoral pores. pores; no femoral pores; subcaudals smooth, Cnemaspis molligodai: 5 preanal and 8 or 9 femoral median series with individual large, divided scales pores, dorsal scales homogeneous; subcaudals alternating with pairs of smaller undivided ones; on median row enlarged, subequal. Cnemaspis supralabials to angle of jaws, 7–9; subdigital lamellae samanalensis: gular, pectoral and abdominal scales on digit IV of pes, 30. tricarinate, subcaudals keeled. Cnemaspis retigalensis: gulars carinate, a single preanal pore and 3 or 4 femoral Description of lectotype pores; subcaudals on median row enlarged, irregular. An adult male, 42.0 mm SVL (for measurements, see Cnemaspis punctata: 5–7 femoral pores; distinctive Table 2). Head long (HL 29.0% of SVL), broad (HW spotted dorsal coloration. Cnemaspis kumarasinghei 20.0% of SVL, HW 68.8% of HL), distinct from neck.

22 Zeylanica Re v i s i o n o f Cn e m a s p i s

Snout length greater than eye diameter (ED 59.0% of on each side (Fig. 4f). Subcaudals on median row ES); interorbital broad (IO 11.4% of HL); pupil rounded. enlarged, smooth, with individual large, divided Granules on snout smooth, larger than those of occipital scales alternating with pairs of smaller undivided region; scales on interorbital and supercilium granular; ones (Fig. 4h). scales on ventral surface of head smooth. Rostral scale partially divided by a medial groove, in contact Variation. Supralabials 7–9; infralabials 7 or 8; with first supralabial (Fig. 4a). Nares separated by postmentals bordered posteriorly by 5 smooth scales two enlarged scales, in contact with supranasals; no in BMNH 74.4.29.401 and in BMNH 74.4.29.402; 6–8 internasals (Fig. 4a). Nares rounded, dorsally orientated, preanal pores; pores absent in females. SVL range not in contact with first supralabial. Two post-nasals, 26.6–42.0 mm (9 ex.). the upper one enlarged, the lower one in contact with first supralabial (Fig. 4a). Mental subpentagonal, wider Coloration. In preservative: colour variable (see Plate than long, truncate posteriorly; mental posteriolaterally 2c–f), dorsally pinkish brown, with a pair of dark in contact with two enlarged postmentals, medially spots along mid-dorsal line; several pale spots form with a single hexagonal postmental scale; postmentals an uninterrupted mid-dorsal line behind and between widely separate, bordered posteriorly by 6 smooth dark spots in juveniles. A pale chevron on head, with scales including a medial scale (Fig. 4b). Supralabials a dark spot at its angle; another pale semicircular 8, infralabials 8, decreasing in size towards gape (Fig. marking originates behind each eye and extends 4c). Ear opening elliptical, much higher than wide, to angle of chevron. Neck with a pale band (which oblique (TW 12.5% of TH). Dorsal and lateral scales precedes a darker band in some adult specimens). on neck granular, intermixed with enlarged tubercles. Venter dusky white with dark markings. Twenty-three granules between anterior margin of ear opening and posterior margin of eye. A large conical Comparisons. Cnemaspis ornata differs from each scale laterally on mid-neck. of its peninsular Indian and Sri Lankan congeners, Body robust, short. Mid-dorsal granules isolated, respectively, by the following opposing suites of scattered, not keeled, intermixed with large tubercles characters: (Fig. 4d); spine-like tubercles on flank; ventrals, 161, Cnemaspis beddomei: subdigital lamellae on subimbricate, longer than wide, smooth (Fig. 4e); 6 proximal series entire. Cnemaspis goaensis: presence preanal pores; no femoral pores (Fig. 4f). of femoral pores. Cnemaspis littoralis: body slender, Forelimbs long, slender (LAL 16.9% SVL, UAL dorsal scales homogeneous, subcaudals in median 13.0% SVL); hind limbs moderately long (TBL 18.3% series enlarged, hexagonal. Cnemaspis mysoriensis: SVL, FEL 18.3% SVL); tibia short. Dorsal scales on femoral pores 2 or 3. Cnemaspis indraneildasii: irregular both fore and hind limbs smooth; ventral scales on rows of dorsal tubercles. Cnemaspis heteropholis: upper arm and lower arm smooth. Scales on thigh enlarged conical tubercles absent on neck, no post- dorsally and ventrally smooth (Fig. 4f). Scales on cloacal spurs. Cnemaspis jerdonii: no preanal pores. dorsal and ventral surface of shank smooth. Scales Cnemaspis nairi: subdigital lamellae on proximal series on manus and pes smooth, rounded. Digits elongate, entire. Cnemaspis otai: paired postmentals in broad slender, all bearing slightly recurved claws (Fig. 4j); contact, 4 preanal and 3 femoral pores. Cnemaspis subdigital lamellae fragmented on proximal portion; sisparensis: postmentals in contact, no preanal pores, 7 an enlarged scansor near middle of digits, more than or 8 femoral pores. Cnemaspis wynadensis: subcaudals twice width of other lamellae; 2 or 3 fragmented enlarged, subequal; no preanal pores, 5 femoral scales behind each enlarged scansor; distal lamellae pores. Cnemaspis yercaudensis: 2 preanal and 3 femoral entire (Fig. 4i); subdigital lamellae on finger I, 21, pores. Cnemaspis boiei: no preanal or femoral pores. finger II, 21, finger III, 28, finger IV, 29, finger V, 22; Cnemaspis gracilis: postmentals in contact. Cnemaspis toe I, 14, toe II, 23, toe III, 28, toe IV, 30, toe V, 27; indica: postmentals in contact. Cnemaspis wicksii, C. interdigital webbing absent; relative length of digits andersonii and C. monticola: gulars carinate, spine- (fingers): IV (5.0 mm) III (4.6 mm) > V (4.2 mm) ~ II like tubercles present on flank. Cnemaspis australis: (4.2 mm) > I (2.7 mm); (toes) IV (6.0 mm) > III (5.1 subcaudals pointed, strongly keeled scales. Cnemaspis mm) ~ V (5.1 mm) > II (4.5 mm) > I (2.8 mm). nilagirica: dorsal scales homogeneous; pectoral and Tail (damaged), tapering; tail base distinctly abdominal scales weakly keeled; subcaudals on swollen. Enlarged tubercles absent at base of tail. median row enlarged, subequal. Dorsal scales on mid-tail imbricate, smooth, directed Cnemaspis tropidogaster: gular, pectoral and backwards, with 2–6 rows of enlarged tubercles abdominal scales keeled. Cnemaspis kandiana, C. (Fig. 4g); one conical post-cloacal spur present podihuna, C. scalpensis, C. gemunu, C. phillipsi, C.

Vol. 7, No. 1 23 Ma n a m e n d r a -Ar a c h c h i e t a l .

a b

c d

e f g

h i j

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 and ventral 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 left pes; j, lateral aspect of claw of digit IV of left pes. Scale bars: 1 mm.

24 Zeylanica Re v i s i o n o f Cn e m a s p i s

meni­kay, C. alwisi and C. molligodai: presence of fe­ than eye diameter (ED 72.2% of ES); interorbital moral pores. Cnemaspis samanalensis: gular, pectoral narrow (IO 13.1% of HL); pupil rounded. Granules and abdominal scales tricarinate, subcaudals keeled. on snout smooth; scales on ventral surface of head Cnemaspis retigalensis: gulars carinate, a single smooth. Rostral scale partially divided by a medial preanal pore and 3 or 4 femoral pores. Cnemaspis groove, in contact with first supralabial (Fig. 5a). punc­tata: 5–7 femoral pores, distinctive spotted dorsal Nares separated by two enlarged supranasals; small coloration. Cnemaspis upendrai, C. kumarasinghei, C. internasal scales separate supranasals posteriorly latha, C. pulchra, C. pava, C. kallima, C. silvula and C. (Fig. 5a). Nares rounded, dorsally orientated, not clivicola: presence of femoral pores. Cnemaspis amith: in contact with first supralabial (Fig. 5a). About gular scales carinate. four postnasals, the uppermost one enlarged (Fig. 5a). Mental subpentagonal, wider than long, Cnemaspis wynadensis (Beddome, 1870) posteriolaterally in contact with two widely Gymnodactylus wynadensis Beddome, 1870 separated, enlarged postmentals and medially with (Figures 5a–g; Table 2) three small postmental scales; postmentals bordered posteriorly by 9 smooth (2 enlarged lateral and 7 small Lectotype (here designated). BMNH 74.4.29.355 medial) scales (Fig. 5b). Supralabials 5, infralabials 6, (male), 33.4 mm SVL, “Wynaad” (Kerala, India), coll. decreasing in size towards gape (Fig. 5c). Ear opening Colonel Beddome. oval, higher than wide, oblique (TW 40.0% of TH). Scales on ventral surface of neck smooth. Paralectotypes. BMNH 74.4.29.356 (male), 36.6 mm Body robust. Mid-dorsal granules homogeneous, SVL; BMNH 74.4.29.357 (male), 34.2 mm SVL; BMNH isolated and scattered, smooth (Fig. 5d); spine-like 74.4.29.358 (male), 39.4 mm SVL; BMNH 74.4.29.359 tubercles absent on flank; abdominal scales smooth, (male), 40.3 mm SVL; BMNH 74.4.29.360 (male), 30.7 subimbricate, longer than wide, obtuse (Fig. 5e). mm SVL; BMNH 74.4.29.361 (male), 31.3 mm SVL; Scales around vent and base of tail smooth, imbricate BMNH 74.4.29.362 (male), 40.6 mm SVL; BMNH (Fig. 5f). No preanal pores, 5 femoral pores (Fig. 74.4.29.363 (male), 38.9 mm SVL; BMNH 74.4.29.364 5f). (male), 34.3 mm SVL; BMNH 74.4.29.365 (male), Forelimbs relatively short, slender (LAL 14.0% 33.7 mm SVL; BMNH 74.4.29.366 (female), 42.1 mm SVL, UAL 10.1% SVL); hind limbs long (TBL 18.2% SVL; BMNH 74.4.29.367 (female), 41.2 mm SVL; SVL, FEL 20.6% SVL); tibia short. Dorsal scales on BMNH 74.4.29.368 (female), 40.8 mm SVL; BMNH both fore and hind limbs smooth; ventral scales on 74.4.29.369 (subadult male), 25.3 mm SVL; BMNH upper arm and lower arm smooth. Scales on anterior 74.4.29.370 (subadult female), 25.5 mm SVL; BMNH surface of thigh and ventral side smooth (Fig. 5f); 74.4.29.371 (juvenile), 18.4 mm SVL, Wynaad, coll. those on ventral surface of shank, and on manus Colonel Beddome. and pes, smooth. Digits elongate, slender, all bearing slightly recurved claws; subdigital lamellae entire, Diagnosis unnotched; subdigital lamellae on finger I, 9, finger Cnemaspis wynadensis differs from its peninsular II, 12, finger III, 16, finger IV, 15, finger V, 13; toe I, 10, Indian and Sri Lankan congeners by a combination toe II, 13, toe III, 18, toe IV, 18, toe V, 17; interdigital of the following characters. Maximum SVL 40.6 mm; webbing absent; relative length of digits (fingers): III paired postmentals separated by 3 medial scales; each (3.0 mm) > IV (2.8 mm) > II (2.7 mm) > V (2.6 mm) > postmental bounded by 4 scales including medial I (1.8 mm); (toes) IV (3.5 mm) > III (3.4 mm) ~ V (3.4 scale; dorsal scales homogeneous, smooth, isolated mm) > II (2.8 mm) > I (2.1 mm). and scattered; spine-like tubercles absent on flank; Tail tapering, its length greater than SVL (TAL abdominal scales smooth, subimbricate; no preanal 112.8% of SVL); tail base slightly swollen, lacking pores; 4–6 femoral pores on each side; subcaudals enlarged tubercles. Dorsal scales on tail smooth, im­ on median series enlarged, subequal; supralabials bricate, directed backwards (Fig. 5g); no post-cloacal to angle of jaws, 5; subdigital lamellae on digit IV spurs; subcaudals smooth, median series regular, of pes, 18. enlarged, widened, circular or hexagonal (Fig. 5f).

Description of lectotype Variation. Femoral pores, 4–6; pores absent in An adult male, 33.4 mm SVL (for measurements, females; SVL range from 18.4 to 42.1 mm (16 ex.). see Table 2). Head moderately long (HL 27.2% of SVL), broad (HW 19.1% of SVL, HW 70.3% of HL), Comparisons distinct from neck. Snout length a little greater Cnemaspis wynadensis differs from each of its

Vol. 7, No. 1 25 Ma n a m e n d r a -Ar a c h c h i e t a l .

a b

c d

e f

g

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 and ventral regions, respectively, of mid-body; f, preanal, hind limb and tail regions, showing arrangement of femoral pores and arrangement of subcaudal scales; g, arrangement of scales on dorsal aspect of tail. Scale bars: 1 mm.

26 Zeylanica Re v i s i o n o f Cn e m a s p i s

peninsular Indian and Sri Lankan congeners, Paralectotypes. ZSI 5866 (male), 27.9 mm SVL; ZSI respectively, by the following opposing suites of 5867 (female), 26.6 mm SVL, from ‘Preparis Island’, characters: in the Andaman Archipelago, Bay of Bengal. Cnemaspis beddomei: preanal pores 6–9, no femoral pores. Cnemaspis goaensis: presence of preanal pores. Diagnosis Cnemaspis littoralis: body slender, dorsal scales Cnemaspis wicksii differs from its peninsular Indian homogeneous. Cnemaspis mysoriensis: spine-like and insular congeners by a combination of the tubercles present on flank, femoral pores 2 or 3. following characters. Maximum SVL 28.6 mm; Cnemaspis indraneildasii: spine-like tubercles present paired postmentals separated by a medial scale, on flank. Cnemaspis heteropholis and C. sisparensis: each bounded by 3 scales including medial scale; subcaudals on median series divided. Cnemaspis dorsal scales heterogeneous; gulars weakly carinate; jerdonii: spine-like tubercles present on flank. other ventrals smooth, imbricate; dorsal scales on Cnemaspis nairi: no femoral pores. Cnemaspis otai: both fore and hind limbs smooth; ventral scales on paired postmentals in broad contact, 4 preanal and arm smooth; scales on anterior and ventral surfaces 3 femoral pores. Cnemaspis yercaudensis: 2 preanal of thigh smooth; supralabials to angle of jaws, 5; 4 and 3 femoral pores. Cnemaspis boiei: no preanal or preanal pores; 4 or 5 femoral pores on each side; femoral pores. Cnemaspis gracilis: postmentals in subcaudals smooth, 2 smaller obtuse scales between contact, subcaudals in median series not enlarged. single larger rounded ones in median series. Cnemaspis indica: postmentals in contact. Cnemaspis ornata: dorsal tubercles arranged in 15 longitudinal Description of lectotype rows, dark-brown collar on nape, no femoral pores, An adult female, 28.6 mm SVL (for measurements, preanal pores 6–8. Cnemaspis wicksii, C. andersonii see Table 2). Head moderately long (HL 26.9% of and C. monticola: gulars carinate. Cnemaspis australis: SVL), moderately broad (HW 15.3% of SVL, HW subcaudals pointed, strongly keeled. Cnemaspis 57.1% of HL), distinct from neck. Pupil rounded. nilagirica: pectoral and abdominal scales weakly Granules on snout smooth; scales on ventral keeled. surface of head feebly keeled. Mental subtriangular, Cnemaspis tropidogaster: gular, pectoral and wider than long, posteriolaterally in contact with abdominal scales keeled, spine-like tubercles two enlarged postmentals, medially with a single present on flank. Cnemaspis kandiana, C. podihuna, hexagonal postmental scale; postmentals bordered C. scalpensis, C. gemunu and C. phillipsi: spine- posteriorly by 5 smooth scales including medial scale like tubercles present on flank. Cnemaspis alwisi: (Fig. 6a). Supralabials 5, infralabials 7, decreasing in 9 femoral pores, spine-like tubercles present on size towards gape. Scales on ventral surface of neck flank. Cnemaspis molligodai: 5 preanal and 8 or feebly carinate. 9 femoral pores, spine-like tubercles present on Body slender. Mid-dorsal scales granular; flank. Cnemaspis samanalensis: gular, pectoral and abdominal scales smooth, those around vent and abdominal scales tricarinate, subcaudals keeled, base of tail smooth, imbricate; 4 preanal and 4 (left spine-like tubercles present on flank. Cnemaspis side) and 5 (right side) femoral pores (Fig. 6b). retigalensis: a single preanal pore and 3 or 4 femoral Dorsal scales on both fore and hind limbs smooth; pores; spine-like tubercles present on flank; ventral scales on upper arm and lower arm smooth. subcaudals of median row enlarged, irregular. Scales on both anterior and ventral surfaces of thigh Cnemaspis punctata: spine-like tubercles present smooth (Fig. 6b). Scales on ventral surface of shank on flank. Cnemaspis kumarasinghei, C. menikay, C. feebly carinate, unnotched; subdigital lamellae on pulchra, C. pava, C. upendrai, C. latha, C. kallima, C. finger I, 10, finger II, 12, finger III, 16, finger IV, 14, silvula and C. clivicola: presence of preanal pores. finger V, 11. Cnemaspis amith: gular scales carinate; subcaudals Tail broken (a broken tail remains in the specimen on median row enlarged, irregular. jar). Enlarged tubercles absent on base of tail. Dorsal scales on tail oval, directed backwards; subcaudals Cnemaspis wicksii (Stoliczka, 1873) smooth, 2 smaller obtuse scales between single larger Gymnodactylus wicksii Stoliczka, 1873 rounded ones in median series (Fig. 6c). (Figures 6a–c) Measurements of lectotype (those of paralectotypes Lectotype (here designated). ZSI 5868 (male), ZSI 5866 and ZSI 5867 in parentheses; measurements 28.6 mm SVL, from “Preparis Island” (Andamans in mm): HL, 7.7 (7.6, 6.9); HW, 4.4 (4.4, 4.1); SVL, 28.6 Archipelago, Bay of Bengal). (27.9, 26.6).

Vol. 7, No. 1 27 Ma n a m e n d r a -Ar a c h c h i e t a l . a

b

c

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

Coloration. In preservative: general body colour tubercles absent on flank, gulars smooth. Cnemaspis greyish-dark brown. No prominent markings on boiei: no preanal or femoral pores, spine-like tubercles body. absent on flank. Cnemaspis gracilis: postmentals in contact, subcaudals in median series not enlarged. Variation. Preanal and femoral pores absent in Cnemaspis indica: postmentals in contact, spine-like females. SVL ranges from 26.6 to 28.6 mm (3 ex.). tubercles absent on flank. Cnemaspis ornata: dorsal tubercles arranged in 15 longitudinal rows, dark- Comparisons brown collar on nape, no femoral pores, preanal Cnemaspis wicksii differs from each of its peninsular pores 6–8. Cnemaspis andersonii: 7 supralabials, Indian and Sri Lankan congeners, respectively, by the scales on ventro-lateral region carinate, subcaudals following opposing suites of characters: in median series not enlarged. Cnemaspis monticola: Cnemaspis beddomei: spine-like tubercles absent on subcaudals carinate. Cnemaspis australis: subcaudals flank, preanal pores 6–9, no femoral pores. Cnemaspis pointed, strongly keeled. Cnemaspis nilagirica: dorsal goaensis: subcaudals in median series not enlarged. scales homogeneous; pectoral and abdominal scales Cnemaspis littoralis: dorsal scales homogeneous, weakly keeled; subcaudals on median row enlarged, median series of subcaudals enlarged, hexagonal. subequal. Cnemaspis mysoriensis: femoral pores 2 or 3. Cnemaspis Cnemaspis tropidogaster: gular, pectoral and indraneildasii and C. heteropholis: gulars smooth. abdominal scales keeled. Cnemaspis kandiana: Cnemaspis jerdonii: no preanal pores. Cnemaspis nairi: except in the median series, subcaudals carinate. spine-like tubercles absent on flank, no femoral pores. Cnemaspis podihuna, C. scalpensis, C. gemunu and C. Cnemaspis otai: paired postmentals in broad contact. phillipsi: 12–16 femoral pores. Cnemaspis alwisi: no Cnemaspis sisparensis: postmentals in contact, spine- preanal pores. Cnemaspis molligodai: 5 preanal and like tubercles absent on flank, no preanal pores, 7 or 8 or 9 femoral pores, gular, pectoral and abdominal 8 femoral pores. Cnemaspis wynadensis: subcaudals scales smooth; subcaudals on median row enlarged, enlarged, subequal; no preanal pores, 5 femoral subequal. Cnemaspis samanalensis: gular, pectoral and pores, spine-like tubercles absent on flank. Cnemaspis abdominal scales tricarinate, subcaudals keeled. yercaudensis: 2 preanal and 3 femoral pores, spine-like Cnemaspis retigalensis: a single preanal pore and 3 or

28 Zeylanica Re v i s i o n o f Cn e m a s p i s

4 femoral pores. Cnemaspis punctata: no preanal pores, isolated and scattered; 7 preanal pores; no femoral distinctive spotted dorsal coloration. Cnemaspis pores; subcaudals on median row widened, irregular, kumarasinghei and C. latha: gular scales smooth. some scales divided; supralabials to angle of jaws, 6; Cnemaspis silvula, C. pulchra, C. pava, C. clivicola and subdigital lamellae on digit IV of pes, 16. C. upendrai: pectoral and abdominal scales keeled. Cnemaspis kallima: gular scales smooth, subcaudals on Description of lectotype distal half of tail carinate. Cnemaspis menikay: dorsal Adult male, 46.6 mm SVL (for measurements, see granules conical, 1 or 2 preanal pores. Cnemaspis Table 2). Head long (HL 28.1% of SVL), broad (HW amith: dorsal granules unkeeled. 20.3% of SVL, HW 72.5% of HL), distinct from neck. Snout length greater than eye diameter (ED 57.3% Remarks of ES); interorbital broad (IO 16.7% of HL); pupil We have included a description of Cnemaspis wicksii rounded. Granules on snout smooth, larger than here, even though its range is outside our region of those of occipital region; scales of occipital region interest (peninsular India and Sri Lanka), because it granular, those on interorbital and supercilium was previously listed in the synonymy of C. kandaiana conical, and those on ventral surface of head by Smith (1935), where it has hitherto remained. granular, smaller than those on abdominal. Rostral Based on an examination of their name-bearing scale partially divided by a medial groove, in contact types, however, Cnemaspis wicksii is immediately with first supralabial (Fig. 7a). Nares separated by distinguishable from C. kandaiana by having the two enlarged supranasals; no internasals (Fig. 7a). scales of the dorsal surface of both limbs smooth Nares rounded, dorsally orientated, not in contact (vs carinate in C. kandaiana); all subcaudals smooth with first supralabial. Two enlarged postnasals (vs subcaudals carinate except for those on median present, the lower one in broad contact with first row, which are smooth); 5 supralabial scales (vs 8 or supralabial (Fig. 7a). Mental subpentagonal, truncate 9); and no tubercles on tail base (vs about 6 enlarged posteriorly, as wide as long, posteriolaterally tubercles on tail base). We consider Cnemaspis wicksii in contact with two widely separated quadrate to be a valid species, clearly distinguishable from all postmentals and medially in contact with a single its Sri Lankan and peninsular Indian congeners. elongate postmental scale; postmentals bordered posteriorly by 8 smooth scales including medial scale Cnemaspis beddomei (Theobald, 1876) (Fig. 7b). Supralabials 6, infralabials 7, decreasing Gymnodactylus beddomei Theobald, 1876 in size towards gape (Fig. 7c). Ear opening oval, Gymnodactylus marmoratus Beddome, 1870 higher than wide, oblique. Scales on ventral surface (Figures 7a–g; Table 2; Plate 1g, h) of neck granular. Granules between anterior margin of ear opening and posterior margin of eye 29. Lectotype (here designated). BMNH 1946.9.4.83 Conical granules on sides of neck larger than those (male), 46.6 mm SVL, “South Tinnevelly (Tirunelveli, of occipital region. southern Tamil Nadu State, India) and Travankor Body robust, short. Mid-dorsal scales granular, Hills 3000–5000 feet” (Travancore, Kerala State, isolated and scattered, conical, intermixed with India), coll. Colonel R. H. Beddome. large scales (Fig. 7d); spine-like tubercles absent on flank; abdominal scales smooth (Fig. 7e). Ventrals Paralectotypes. BMNH 1946.9.4.82 (female), 48.2 mm 154, isolated and scattered (Fig. 7e); scales around SVL; BMNH 1946.9.4.84 (male), 42.2 mm SVL; BMNH preanal area enlarged; 7 preanal pores, no femoral 1946.9.4.85 (male), 43.9 mm SVL; “South Tinnevelly pores (Fig. 7f). [Tirunelveli, southern Tamil Nadu State, India] and Forelimbs moderately long, slender (LAL 13.5% Travankor Hills 3000–5000 feet” [Travancore, Kerala SVL, UAL 13.9% SVL); hind limbs long (TBL 17.3% State, India], coll. R. H. Colonel Beddome. SVL, FEL 24.4% SVL); tibia short. Dorsal scales on both fore and hind limbs granular, conical; ventral Diagnosis scales on upper arm and lower arm granular and Cnemaspis beddomei differs from its peninsular Indian smaller than dorsals. Dorsal scales of thigh and and Sri Lankan congeners by a combination of the shank granular (Fig. 7f); ventral scales on shank following characters. Maximum SVL 50.6 mm; imbricate, smooth, those on thigh enlarged, conical. paired postmentals separated by a medial scale; each Dorsal sales on manus and pes granular, ventrals postmental bounded by 4 or 5 scales including medial imbricate, smooth. Digits elongate, slender, all scale; ventrals, 154; dorsal scales heterogeneous; bearing slightly recurved claws; subdigital lamellae spine-like tubercles absent on flank; ventrals smooth, entire, unnotched; subdigital lamellae on finger I,

Vol. 7, No. 1 29 Ma n a m e n d r a -Ar a c h c h i e t a l . a b

c d

f e

g

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 and ventral 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.

30 Zeylanica Re v i s i o n o f Cn e m a s p i s

11, finger II, 14, finger III, 18, finger IV, 19, finger V, presence of femoral pores. Cnemaspis monticola: spine- 15; toe I, 17, toe II, 19, toe III, 18, toe IV, 16, toe V, 10; like tubercles present on flank. Cnemaspis australis: interdigital webbing absent; relative length of digits subcaudals pointed, strongly keeled. Cnemaspis (fingers): V (5.6 mm) > IV (5.2 mm) > III (5.0 mm) > nilagirica: dorsal scales homogeneous; pectoral and II (4.7 mm) > I (3.6 mm); (toes) IV (6.7 mm) > III (6.1 abdominal scales weakly keeled; subcaudals on mm) > V (5.8 mm) > II (5.6 mm) > I (3.4 mm). median row enlarged, subequal. Regenerated tail tapering, its length less than Cnemaspis tropidogaster: gular, pectoral and SVL (TAL 74.6% of SVL); tail base distinctly abdominal scales keeled, spine-like tubercles swollen. Scales on tail base conical to imbricate; present on flank. Cnemaspis kandiana, C. podihuna, enlarged tubercles absent on tail. Dorsal scales on C. scalpensis, C. menikay, C. gemunu and C. phillipsi: tail imbricate; a single enlarged, oval-shaped, post- spine-like tubercles present on flank. Cnemaspis cloacal spur present on each side; subcaudal scales alwisi: no preanal pores, spine-like tubercles present on tail base imbricate to subimbricate, increasing in on flank. Cnemaspis molligodai: 8 femoral pores, spine- size towards tail tip (Fig. 7g). like tubercles present on flank; subcaudals in median row enlarged, subequal. Cnemaspis samanalensis: Variation. Supralabials, 6 or 7; infralabials, 6 or 7; gular, pectoral and abdominal scales tricarinate, preanal pores 6–8; femoral pores absent; pores absent subcaudals keeled, spine-like tubercles present on in females; SVL range 42.2–46.6 mm (4 ex.). flank. Cnemaspis retigalensis: gulars carinate, a single Unregenerated tail of paralectotype BMNH preanal pore and 3 or 4 femoral pores, spine-like 1946.9.4.86 with a median series of enlarged, divided tubercles present on flank. Cnemaspis punctata: no subcaudals, with individual large scales between preanal pores, spine-like tubercles present on flank, pairs of smaller ones (see Fig. 4h of C. ornata), most distinctive spotted dorsal coloration. Cnemaspis subcaudal scales divided on distal portion of tail. kumarasinghei: spine-like tubercles present on flank. Cnemaspis latha: ventrals 111; gular scales smooth, Coloration. In preservative, dorsally light brown abdominal scales and subcaudal scales smooth, with dark mottling in region of head, pale mottling spine-like tubercles present on flank. Cnemaspis on body; tail with pale bands; venter brownish white; silvula, C. pulchra and C. pava: pectoral and abdominal two dark, V-shaped streaks on chin and gular region scales keeled, spine-like tubercles present on flank. (see Plate 1h). Cnemaspis clivicola pectoral and abdominal scales keeled, spine-like tubercles present on flank, Comparisons. Cnemaspis beddomei differs from each subcaudals in median series not enlarged. Cnemaspis of its peninsular Indian and Sri Lankan congeners, upendrai: some gular scales tricarinate, pectoral and respectively, by the following opposing suites of abdominal scales keeled, spine-like tubercles present characters: on flank. Cnemaspis kallima: gular scales smooth, Cnemaspis goaensis: presence of femoral pores. spine-like tubercles present on flank, subcaudals Cnemaspis littoralis: body slender, subcaudals of on distal half of tail carinate. Cnemaspis amith: gular median series enlarged, hexagonal. Cnemaspis scales carinate; pectoral and abdominal scales mysoriensis: spine-like tubercles present on flank, smooth; dorsal granules heterogeneous; subcaudals femoral pores 2 or 3. Cnemaspis indraneildasii on median row enlarged, irregular. and C. heteropholis: dorsal scales heterogeneous. Cnemaspis jerdonii: no preanal pores. Cnemaspis Cnemaspis sisparensis (Theobald, 1876) nairi: a dark gular band present. Cnemaspis otai: Gymnodactylus sisparensis Theobald, 1876 paired postmentals in broad contact, 4 preanal and Gymnodactylus maculatus Beddome, 1870 3 femoral pores. Cnemaspis sisparensis: postmentals Gonatodes bireticulatus Annandale, 1915 in contact, no preanal pores, 7 or 8 femoral pores. Cnemaspis anaikattiensis Mukherjee, Bhupathy & Cnemaspis wynadensis: subcaudals enlarged, subequal; Nixon, 2005 no preanal pores, 5 femoral pores. Cnemaspis (Figures 8a–g, 9 a–f; Table 2; Plate 2a, b) yercaudensis: 2 preanal and 3 femoral pores. Cnemaspis boiei: no preanal or femoral pores. Cnemaspis gracilis: Holotype. BMNH 74.4.29.383 (male) (fragmented), postmentals in contact, subcaudals in median series “Sholakal, the foot of Sispara Ghat”, coll. Colonel not enlarged. Cnemaspis indica: postmentals in contact, Beddome. no preanal pores. Cnemaspis ornata: dorsal tubercles arranged in 15 longitudinal rows, a dark-brown Other material. ZSI 17970 (male), 42.7 mm SVL, collar on nape. Cnemaspis wicksii and C. andersonii: holotype of Gonatodes bireticulatus Annandale,

Vol. 7, No. 1 31 Ma n a m e n d r a -Ar a c h c h i e t a l .

1915, “Karalai 1300–3000 ft, Cochin State”, coll. F. Tail tapering, enlarged tubercles absent at its base. H. Gravely (see Mukherjee et. al., 2005); ZSI 25601, Dorsal scales on tail imbricate, directed backwards (male), 61.0 mm SVL, ZSI 25602, (female), 58.0 mm (Fig. 8f). No post-cloacal spurs. Subcaudals smooth; (respectively, holotype and paratype of Cnemaspis scales on median series enlarged with single large anaikattiensis Mukherjee, Bhupathy & Nixon, 2005), scales alternating between pairs of smaller ones Anaikatti Hills, Western Ghats, Coimbatore district, (Fig. 8h). Tamil Nadu, India, 11º05’30.9’’N, 76º47’36.2’’E, coll. Measurements of holotype (in mm): HL, 13.1; A. M. A. Nixon & D. Mukherjee, 17 September, 2003 HW, 8.6; TAL, 55.2 (other measurements could not (not examined). be taken because the specimen is fragmented: see Plate 2e, f). Diagnosis Cnemaspis sisparensis differs from its peninsular Variation (based on ZSI 17970): an adult male, 42.7 Indian and Sri Lankan congeners by a combination mm SVL. Head long (HL 28.1% of SVL), moderately of the following characters. Maximum SVL 42.7 broad (HW 20.3% of SVL, HW 72.5% of HL), distinct mm; paired postmentals widely separated by two from neck. Snout length greater than eye diameter medial scales; each postmental bounded by 4 scales (ED 69.5% of ES). No internasals; lower postnasals in including medial scale; dorsal scales homogeneous, broad contact with first supralabial (Fig. 9a). Mental isolated and scattered; spine-like tubercles absent subtriangular, as wide as long, posteriolaterally on flank; ventrals smooth, imbricate; supralabials to in contact with two widely separated, enlarged angle of jaws 7 or 8; no preanal pores, 7 or 8 femoral postmentals and medially with 2 small medial scales; pores; subcaudals smooth; those on median series postmentals bordered posteriorly by 8 smooth scales enlarged, with individual large scales alternating (2 enlarged lateral scales and 6 small medial scales) between pairs of smaller ones. (Fig. 9b). Supralabials 8, infralabials, 10, decreasing in size towards gape (Fig. 9c). Ear opening oval, higher Description of holotype than wide, oblique (TW 70.0% of TH) (Fig. 9c). Scales An adult male. Head broad (HW 65.6% of HL), on ventral surface of neck smooth, tuberculate. Body pupil rounded. Granules on snout smooth, larger robust. Femoral pores on each side, 7 (Fig. 9d). Scales than those of occipital region; scales on ventral on manus and pes smooth, rounded. Tail tapering, surface of head with smooth tubercles. Rostral scale its length greater than SVL (TAL 129.7% of SVL); partially divided by a medial groove, in contact enlarged tubercles present on base of tail. Dorsal with first supralabial (Fig. 8a). Nares separated by scales on tail imbricate, directed backwards (Fig. 9e). two enlarged supranasals and three internasals, Subcaudals smooth; scales of median series enlarged, the median one of which extends towards snout tip with individual large scales alternating between pairs (Fig. 8a). Nares rounded, dorsally orientated, not of smaller ones (Fig. 9f). in contact or in contact with first supralabial (Fig. Measurements of ZSI 17970 (in mm): ED, 3.2; ES 8a). Three postnasals, the upper one enlarged, the 4.6; HL, 12.0; HW, 8.7; IN, 2.5; SVL, 42.7; TAL, 55.4; lower one in contact with first supralabial (Fig. 8a). TYD, 3.6; TYE, 0.7×1.0. Mental subtriangular, truncate posteriorly, as wide as long, posteriolaterally in contact with two widely Coloration. In preservative: dorsally light to dark separate, enlarged postmentals (damaged) and brown with dark-brown mottling on dorsum (see medially with a small, hexagonal postmental scale; Plate 2a, b); nape with a dark-brown marking; tail postmentals bordered posteriorly by several small with pale bands; digits banded; venter brownish scales (damaged) (Fig. 8b). Supralabials 7, infralabials white with indistinct light-brown patches. 8, decreasing in size towards gape (Fig. 8c). Mid-dorsal granules homogeneous, isolated Comparisons. Cnemaspis sisparensis differs from each and scattered; spine-like tubercles absent on flank; of its peninsular Indian and Sri Lankan congeners, abdominal scales smooth, imbricate; scales around respectively, by the following opposing suites of vent and base of tail smooth, imbricate (Fig. 8d). No characters: preanal pores; 8 femoral pores (right side); left side Cnemaspis beddomei: preanal pores 6–9, no femoral damaged (Fig. 8e). pores. Cnemaspis goaensis: presence of preanal pores. Dorsal scales on both fore and hind limbs smooth; Cnemaspis littoralis: subcaudals of median series ventral scales on upper arm and lower arm smooth. enlarged, hexagonal. Cnemaspis mysoriensis: spine- Scales on anterior surface of thigh (Fig. 8e) and like tubercles present on flank, femoral pores 2 or 3. ventral surface of shank smooth. Cnemaspis indraneildasii: spine-like tubercles present

32 Zeylanica Re v i s i o n o f Cn e m a s p i s a b

c

d

f

e g

Figure 8. Cnemaspis sisparensis: BMNH 74.4.29.383, holotype, male, a, dorsal aspect of snout; b, ventral aspect of chin (dotted line indicates damaged area); c, lateral aspect of gape, showing arrangement of supralabials and infralabials; d, arrangement of 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.

Vol. 7, No. 1 33 Ma n a m e n d r a -Ar a c h c h i e t a l . a b

c

d

e f

Figure 9. Cnemaspis sisparensis: holotype of Gonatodes bireticulatus Annandale, 1915: ZSI 17970, male, a, dorsal aspect of snout; b, ventral aspect of chin; c, lateral aspect of head, showing arrangement of supralabials and infralabials; d, preanal and right hind-limb regions, showing arrangement of femoral pores; e, arrangement of scales on dorsal aspect of tail; f, arrangement of subcaudal scales. Scale bars: 1 mm. on flank. Cnemaspis heteropholis: heterogeneous dorsal subcaudals pointed, strongly keeled. Cnemaspis scalation. Cnemaspis jerdonii: spine-like tubercles nilagirica: dorsal scales homogeneous; pectoral and present­ on flank. Cnemaspis nairi: no femoral abdominal scales weakly keeled; subcaudals on pores. Cnemaspis otai: paired postmentals in broad median row enlarged, subequal. contact, 4 preanal and 3 femoral pores, ventrals Cnemaspis tropidogaster: gular, pectoral and smooth. Cnemaspis wynadensis: subcaudals enlarged, abdominal scales keeled, spine-like tubercles subequal. Cnemaspis yercaudensis: 2 preanal and 3 present on flank. Cnemaspis kandiana, C. podihuna, femoral pores. Cnemaspis boiei: no preanal or femoral C. scalpensis, C. allauwa, C. gemunu and C. phillipsi: pores. Cnemaspis gracilis: postmentals in contact. spine-like tubercles present on flank. Cnemaspis Cnemaspis indica: postmentals in contact. Cnemaspis alwisi: 9 femoral pores, spine-like tubercles present ornata: dorsal tubercles arranged in 15 longitudinal on flank. Cnemaspis molligodai: 5 preanal and 8 or 9 rows, a dark-brown collar on nape, no femoral pores, femoral pores, spine-like tubercles present on flank. preanal pores 6–8. Cnemaspis wicksii, C. andersonii Cnemaspis samanalensis: gular, pectoral and abdominal and C. monticola: gulars carinate. Cnemaspis australis: scales tricarinate; subcaudals keeled; spine-like tub­

34 Zeylanica Re v i s i o n o f Cn e m a s p i s

ercles present on flank. Cnemaspis retigalensis: single Diagnosis preanal pore and 3 or 4 femoral pores, spine-like Cnemaspis andersonii differs from its peninsular tubercles present on flank. Cnemaspis punctata: 5–7 Indian and Sri Lankan congeners by a combination femoral pores, spine-like tubercles present on flank, of the following characters. Maximum SVL 31.5 mm; distinctive spotted dorsal coloration. Cnemaspis paired postmentals separated by a medial scale; each upendrai, C. kumarasinghei, C. pulchra, C. pava, C. postmental bounded by 3 scales including medial latha, C.kallima, C. silvula and C. clivicola: presence of scale; dorsal scales heterogeneous; gulars carinate; preanal pores. Cnemaspis amith: gular scales carinate; other ventrals smooth, imbricate; supralabials to dorsal granules unkeeled. angle of jaws, 7; preanal pores 3, femoral pores 4 on each side; no median series of enlarged subcaudals. Remarks Mukherjee et al. (2005) described Cnemaspis anai­ Redescription of holotype kattiensis­ from the Anaikatti Hills (11º05’30.9”N An adult male, 31.5 mm SVL (for measurements, see and 76º47’36.2”E), Western Ghats, a part of the Table 2). Head relatively short (HL 25.3% of SVL), Nil­giri Biosphere Reserve in Tamil Nadu. While moderately broad (HW 14.5% of SVL, HW 57.5% of acknowledging that of its Western-Ghats congeners HL), distinct from neck. Snout length greater than the new species most closely resembled C. sisparensis, eye diameter (ED 63.8% of ES), pupil rounded. Scales Mukherjee et al. (2005) compared it with what they on ventral surface of head and neck weakly carinate. considered to be “The holotype of C. sisparensis Rostral scale partially divided by a medial groove, (Theobald) [= Gonatodes bireticulatus] Annd. collected in contact with first supralabial (Fig. 10a). Nares by F. H. Gravely from ‘Kavalai, 1300–3000 ft, Co­ separated by two enlarged supranasals (damaged on chin State’ ZSI 17970, Zoological Survey of India, left side) and a single internasal, as large as supranasal Kolkata”. The holotype of C. sisparensis is in fact (Fig. 10a). Nares rounded, dorsally orientated, in BMNH 74.4.29.383, a male specimen collected from contact with first supralabial (Fig. 10a). Mental “Sholakal, the foot of Sispara Ghat”. subpentagonal, wider than long, posteriolaterally in We examined the types of C. bireticulatus at ZSI contact with two enlarged postmentals and medially and C. sisparensis at the BMNH. There is no doubt that with a single small hexagonal postmental scale; these two species are conspecific, the distinguishing postmentals bordered posteriorly by 5 smooth scales characters used by Mukherjee et al. (2005)—”no including medial scale (Fig. 10b). Supralabials 7, internasal scale in C. sisparensis (vs presence in infralabials 7, both series decreasing in size towards the new species), nostril not in contact with first gape; ear opening oval, higher than wide, oblique supralabial (vs in contact in C. anaikattiensis)—being (TW 60.0% of TH) (Fig. 10c). extremely variable in Cnemaspis. The internasal scale Body slender. Mid-dorsal granules heterogeneous; is often present or absent in different individuals pectoral and abdominal scales smooth; scales on within a single population, and the nostril in contact ventro-lateral region weakly carinate; 3 preanal and or not in contact on either side of the same individual 4 femoral pores (Fig. 10d). Tail tapering; a single (e.g., see Remarks under C. podihuna, below). The enlarged, conical, post-cloacal spur present on each type localities of C. anaikattiensis (Anaikatti Hills near side; subcaudals in median series not enlarged. Coimbatore) and C. sisparensis (Sholakal, at the foot of Sispara Ghat, 11º12’N, 76º28’E, at the northeast end Measurements of holotype (in mm): ED, 2.3; ES, of Silent Valley National Park), are also relatively 3.6; HL, 8.0; HW, 4.6; SVL, 31.5; TAL, 40.2; TYD, 2.4; close to each other. In any event, the description TYE, 0.5×0.3. and diagnosis of C. anaikattiensis provided by Mukherjee et al. (2005) exactly match the holotype Coloration. In preservative: overall body colour of C. sisparensis, and we therefore here consider the dark-greyish brown. former to be a junior synonym of the latter. Comparisons. Cnemaspis andersonii differs from each Cnemaspis andersonii (Annandale, 1904) of its peninsular Indian and Sri Lankan congeners, Gonatodes andersonii Annandale, 1904 respectively, by the following opposing suites of (Figures 10a–d) characters: Cnemaspis beddomei: preanal pores 6–9, no Holotype. ZSI 15012 (male), 31.5 mm SVL; “Nar­ femoral pores. Cnemaspis goaensis: subcaudals on condum”, Andaman Archipelago (India), Bay of median row enlarged. Cnemaspis littoralis: dorsal Bengal. scales homogeneous, median series of subcaudals

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d c

Figure 10. Cnemaspis andersonii: ZSI 15012, holotype, male, a, dorsal aspect of snout (left side damaged); b, ventral aspect of chin; c, lateral aspect of head, showing arrangement of supralabials and infralabials; d, preanal and right hind-limb regions, showing arrangement of femoral and preanal pores. Scale bars: 1 mm. enlarged, hexagonal. Cnemaspis mysoriensis: femoral and C. phillipsi: 12–16 femoral pores. Cnemaspis pores 2 or 3. Cnemaspis indraneildasii: dorsal scales of alwisi: 7–9 femoral pores. Cnemaspis molligodai: 5 both limbs smooth; ventral, gular and caudal scales preanal and 8 or 9 femoral pores, dorsal scales smooth. Cnemaspis heteropholis: gulars smooth, no homogeneous; subcaudals on median row enlarged, post-cloacal spurs. Cnemaspis jerdonii and C. nairi: no subequal. Cnemaspis samanalensis: gular, pectoral and femoral pores. Cnemaspis indica, C. otai, C. sisparensis, abdominal scales tricarinate, subcaudals keeled. C. wynadensis and C. yercaudensis: spine-like tubercles Cnemaspis retigalensis: single preanal pore and 3 or 4 absent on flank. Cnemaspis boiei: no preanal or femoral femoral pores; subcaudals on median row enlarged, pores. Cnemaspis gracilis: 2 preanal pores separated irregular. Cnemaspis punctata: 5–7 femoral pores, by 2 unpored scales, postmentals broadly in contact. distinctive spotted dorsal coloration. Cnemaspis Cnemaspis ornata: no femoral pores, preanal pores kumarasinghei: gular scales smooth. Cnemaspis latha: 6–8, dark brown collar on nape. Cnemaspis wicksii: 5 gular scales smooth. Cnemaspis silvula, C. pulchra supralabials, scales on ventro-lateral region smooth, and C. pava: pectoral and abdominal scales keeled. subcaudals in median series enlarged. Cnemaspis Cnemaspis clivicola: pectoral and abdominal scales monticola: no large, conical tubercles at base of keeled. Cnemaspis upendrai: some gular scales tail, spine-like tubercles present on flank, pectoral tricarinate, pectoral and abdominal scales keeled. and abdominal scales smooth. Cnemaspis australis: Cnemaspis kallima: gular scales smooth. Cnemaspis subcaudals pointed, strongly keeled. Cnemaspis menikay: dorsal granules conical, 1 or 2 preanal nilagirica: dorsal scales homogeneous; subcaudals pores. Cnemaspis amith: dorsal granules unkeeled; on median row enlarged, subequal. subcaudals on median row enlarged, irregular. Cnemaspis tropidogaster: gular, pectoral and abdominal­ scales keeled. Cnemaspis kandiana: Remarks sub­caudals on median row enlarged, irregular. Cnemaspis andersonii was listed in the synonymy Cnemaspis­ podihuna: subcaudals on median row of C. kandiana by Smith (1935), where it has since enlarged, subequal. Cnemaspis scalpensis, C. gemunu remained. Examination of its holotype (the only

36 Zeylanica Re v i s i o n o f Cn e m a s p i s

known specimen representing this taxon) shows, 8 each, decreasing in size towards gape (Fig. 11c). however, that it is distinguished from C. kandiana Ear opening oval, higher than wide, oblique (TW by having the subcaudals of the median row not 66.6% of TH) (Fig. 11c). Scales on ventral surface of enlarged (vs enlarged in C. kandiana); and scales of neck feebly carinate. the ventro-lateral region of body carinate (vs smooth). Body robust. Mid-dorsal granules heterogeneous, We therefore consider Cnemaspis andersonii to be a mixed with large keeled tubercles from head to valid species. sacrum; ventral abdominal scales smooth; scales around vent and base of tail smooth, imbricate; 3 Cnemaspis goaensis Sharma, 1976 preanal and 4 femoral pores (Fig. 11d). Cnemaspis goaensis Sharma, 1976 Forelimbs moderately long, slender (LAL 13.4% (Figures 11a–e; Table 2) SVL, UAL 12.3% SVL); hind limbs long (TBL 18.1% SVL, FEL 29.0% SVL); tibia short. Dorsal scales on Holotype. ZSI 22110 (male), 27.5 mm SVL; “ca. 3 km both fore and hind limbs weakly carinate; ventral S. of Forest Rest House, Canacona (Poinguinim), scales on upper arm and lower arm smooth. Scales on Goa” (south-western India). thigh and shank smooth. Digits elongate, slender, all bearing slightly recurved claws; subdigital lamellae Paratypes. ZSI 22215 (male), 28.3 mm SVL; same entire, unnotched; subdigital lamellae on finger I, locality data as holotype. 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; Diagnosis interdigital webbing absent; relative length of digits Cnemaspis goaensis differs from its peninsular Indian (fingers): IV (2.5 mm) ~ II (2.5 mm) > III (2.3 mm) > V and Sri Lankan congeners by a combination of the (2.0 mm) > I (1.4 mm); (toes) II (3.5 mm) > I (2.8 mm) following characters. Maximum SVL 28.3 mm; > III (2.7 mm) > IV (2.5 mm) > V (1.8 mm). paired postmentals separated by a medial scale; Tail tapering (its length less than SVL in paratype, each postmental bounded by 3 scales including ZSI 22215: TAL 85.0% of SVL); tail base distinctly medial scale; dorsal scales heterogeneous mixed with swollen; 4–6 enlarged, conical, keeled tubercles on large keeled tubercles; gulars weakly carinate; other base of tail, about as large as granules on mid-body. ventral scales smooth, imbricate; 3 preanal pores and Dorsal scales on mid-tail imbricate, carinate, directed 2–4 femoral pores on each side; subcaudals slightly backwards; a conical post-cloacal spur present enlarged; supralabials to angle of jaws, 8; subdigital on each side; median row of subcaudals slightly lamellae on digit IV of pes, 16. enlarged on proximal half of tail, subquadrangular, smooth (Fig. 11e); scales on distal half carinate. Redescription of holotype An adult male, 27.5 mm SVL (for measurements, Coloration. In preservative: in general body colour see Table 2). Head long (HL 30.1% of SVL), broad greyish dark brown. (HW 19.2% of SVL, HW 63.8% of HL), distinct from neck. Snout length about equal to eye diameter (ED Variation. Seven supralabials and 7 infralabials in 96.5% of ES); interorbital narrow (IO 6.0% of HL); paratype ZSI 22215. Femoral pores, 2–4 on each side pupil rounded. Granules on snout keeled; scales (2 ex.); pores absent in females. on interorbital and supercilium granular; scales on ventral surface of head weakly carinate. Rostral scale Comparisons. Cnemaspis goaensis differs from each partially divided by a medial groove, in contact with of its peninsular Indian and Sri Lankan congeners, first supralabial (Fig. 11a). Nares separated by two respectively, by the following opposing suites of enlarged supranasals, pentagonal internasal and characters: three small scales; internasal extends towards snout Cnemaspis beddomei: no interanasals, subcaudals in tip (Fig. 11a). Nares rounded, dorsally orientated, a series of alternating wide and narrow enlarged scales, not in contact or in contact with first supralabial. no femoral pores. Cnemaspis littoralis: body slender, Two postnasals, the lower one in broad, in contact dorsal scales homogeneous, subcaudals on median with first supralabial (Fig. 11a). Mental subtriangular, series enlarged, hexagonal. Cnemaspis mysoriensis: wider than long, posteriolaterally in contact with femoral pores 2 or 3. Cnemaspis indraneildasii and C. two enlarged postmentals and medially with a heteropholis: gulars smooth. Cnemaspis jerdonii: no single hexagonal postmental scale; postmentals preanal pores. Cnemaspis nairi: no femoral pores. bordered posteriorly by 5 smooth scales including Cnemaspis otai: paired postmentals in broad contact, medial scale (Fig. 11b). Supralabials and infralabials gulars smooth. Cnemaspis sisparensis: postmentals

Vol. 7, No. 1 37 Ma n a m e n d r a -Ar a c h c h i e t a l .

a b

c d

e

Figure 11. Cnemaspis goaensis: ZSI 22110, holotype, male, a, dorsal aspect of snout; b, ventral aspect of chin; c, lateral aspect of head, showing arrangement of supralabials and infralabials (mouth distorted); d, preanal region right hind-limb regions, showing arrangement of femoral and preanal pores; e, arrangement of subcaudal scales. Scale bars: 1 mm. in contact, no preanal pores, 7 or 8 femoral pores. Cnemaspis monticola: Dorsal scales on mid-tail keeled, Cnemaspis wynadensis: subcaudals enlarged, subequal; isolated and granular. Cnemaspis australis: subcaudals no preanal pores, 5 temporal pores, postmentals pointed, strongly keeled, those on; median row widely separated. Cnemaspis yercaudensis: gulars slightly enlarged. Cnemaspis nilagirica: dorsal scales smooth. Cnemaspis boiei: no preanal or femoral pores, homogeneous; subcaudals on median row enlarged, postmentals widely separated by 2 medial scales. subequal. Cnemaspis gracilis: postmentals in contact, subcaudals Cnemaspis tropidogaster: gular, pectoral and abdominal in median series not enlarged. Cnemaspis indica: scales keeled. Cnemaspis kandiana: scales on thigh and postmentals in contact, no preanal pores. Cnemaspis shank carinate. Cnemaspis podihuna, C. kumarasinghei ornata: dorsal tubercles arranged in 15 longitudinal and C. latha: gulars smooth. Cnemaspis scalpensis, rows, dark-brown collar on nape, no femoral pores, C. gemunu and C. phillipsi: 12–16 femoral pores. preanal pores 6–8. Cnemaspis wicksii: 5 supralabials, Cnemaspis alwisi: 9 femoral pores. Cnemaspis subcaudals enlarged. Cnemaspis andersonii: ventro- molligodai: 5 preanal and 8 or 9 femoral pores, gular, lateral region carinate, subcaudals in median series pectoral and abdominal scales smooth, dorsal scales not enlarged, internasal as large as supranasal. homogeneous; subcaudals on median row enlarged,

38 Zeylanica Re v i s i o n o f Cn e m a s p i s

subequal. Cnemaspis samanalensis: gular, pectoral and in contact with first supralabial (Fig. 12a). Nares abdominal scales tricarinate, subcaudals keeled. separated by two enlarged supranasals and a single Cnemaspis retigalensis: single preanal pore and 3 or 4 internasal scale, which is extended towards tip of femoral pores. Cnemaspis punctata: 5–7 femoral pores, snout (Fig. 12a). Nares rounded, dorsally orientated, distinctive spotted dorsal coloration. Cnemaspis not in contact with first supralabial. Two postnasals, silvula, C. pulchra, C. pava and C. clivicola: pectoral the lower one in broad contact with first supralabial and abdominal scales keeled. Cnemaspis upendrai: (Fig. 12a). Mental subpentagonal, wider than long, some gular scales tricarinate, pectoral and abdominal posteriolaterally in contact with two enlarged scales keeled. Cnemaspis kallima: gular scales smooth. postmentals and medially with a single hexagonal Cnemaspis menikay: dorsal granules conical, 1 or postmental scale; postmentals bordered posteriorly 2 preanal pores. Cnemaspis amith: dorsal granules by 5 smooth scales including medial scale (Fig. 12b). unkeeled; subcaudals on median row enlarged, Supralabials 8 and infralabials 7, decreasing in size irregular. towards gape (Fig. 12c). Ear opening oval, higher than wide, oblique. Scales on ventral surface of neck carinate. Twenty-three granular scales between Cnemaspis monticola new species anterior margin of ear opening and posterior margin (Figures 12a–h; Table 2) of eye. Two large, conical tubercles laterally on neck on each side. Holotype. BMNH 74.4.29.372 (female), 26.0 mm Body elongate. Mid-dorsal granules with both SVL, “Wynaad” (Kerala State, south-western India), smooth and keeled scales, intermixed with large, purchased from Colonel Beddome (formerly a conical, keeled tubercles (Fig. 12d); 5 spine-like syntype of C. tropidogaster). tubercles on flank; ventral pectoral and abdominal scales smooth, imbricate, circular (Fig. 12e). Ventrals, Paratypes. BMNH 74.4.29.373 (female), 33.1 mm SVL; 129; scales around vent and base of tail smooth, BMNH 74.4.29.374 (female), 18.3 mm SVL; “Wynaad” imbricate; no preanal and femoral pores. (Kerala State, south-western India), purchased Forelimbs moderately long, slender (LAL 14.2% from Colonel Beddome (formerly syntypes of C. SVL, UAL 11.9% SVL); hind limbs moderately tropidogaster). long (TBL 16.9% SVL, FEL 19.2% SVL); tibia short. Dorsal scales on both fore and hind limbs carinate; Diagnosis ventral scales on upper arm and lower arm smooth. Cnemaspis monticola is distinguished from its pen­ Scales on anterior surface of thigh carinate, those insular Indian and Sri Lankan congeners by a on ventral surface smooth. Granules on posterior combination of the following characters: maximum surface of thigh smooth. Scales on ventral surface SVL 33.1 mm; paired postmentals separated by a of shank carinate. Scales on manus and pes carinate, small scale; each postmental bounded by 3 scales rounded. Digits elongate, slender, all bearing including medial scale; dorsal scales heterogeneous; 5 slightly recurved claws; subdigital lamellae entire, spine-like tubercles present on flank; gulars carinate, unnotched; subdigital lamellae on finger I, 8, finger other ventrals smooth, rounded, ventral surface of II, 10, finger III, 15, finger IV, 14, finger V, 13; toe I, 9, thigh smooth; preanal and femoral pores absent toe II, 14, toe III, 16, toe IV, 18, toe V, 16; interdigital in females; subcaudals enlarged, intermixed with webbing absent; relative length of digits (fingers): IV smooth, carinate scales; supralabials to angle of jaws, (2.5 mm) > III (2.4 mm) > V (2.2 mm) > II (2.1 mm) > 8; subdigital lamellae on digit IV of pes, 18. I (1.4 mm); (toes) IV (3.5 mm) > III (3.0 mm) > V (2.9 mm), II and I (broken). Description of holotype Regenerated tail tapering, its length less than An adult female, 26.0 mm SVL (for measurements, see SVL (TAL 67.3% of SVL); tail base distinctly swollen. Table 2). Head long (HL 29.6% of SVL), moderately Enlarged tubercles present on base of tail (Fig. 12f). broad (HW 17.6% of SVL, HW 59.7% of HL), distinct Dorsal scales on tail keeled, directed backwards (Fig. from neck. Snout length greater than eye diameter 12g); one conical post-cloacal spur present on each (ED 58.3% of ES); interorbital broad (IO 10.3% of HL); side; subcaudals enlarged, intermixed with smooth pupil rounded. Granules on snout keeled, larger than and carinate scales (Fig. 12h). those of occipital region; scales on interorbital and supercilium intermixed with carinate and smooth Coloration. In preservative: general body colour light scales; scales on dorsal surface of neck smooth. brown; head with irregular dark brown blotches. Rostral scale partially divided by a medial groove, Colour in life not known.

Vol. 7, No. 1 39 Ma n a m e n d r a -Ar a c h c h i e t a l .

a b

c d

e f

g h

Figure 12. Cnemaspis monticola: BMNH 74.4.29.372, holotype, female, 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 and ventral regions, respectively, of mid-body; f, arrangement of dorsal scalation of scarum region including base of tail; g, h, arrangement of scales on dorsal and ventral regions, respectively, of mid-tail. Scale bars: 1 mm.

40 Zeylanica Re v i s i o n o f Cn e m a s p i s

Variation. Ventral pectoral and abdominal scales C. menikay and C. phillipsi: presence of smooth carinate, scales on preanal region smooth; endo­ subcaudals. Cnemaspis alwisi: 9 femoral pores. lymph­atic sacs present in paralectotype, BMNH Cnemaspis molligodai: 5 preanal and 8 or 9 femoral 74.4.29.374. pores, gular, pectoral and abdominal scales smooth, dorsal scales homogeneous; subcaudals on median Natural history. Not known. row enlarged, subequal. Cnemaspis samanalensis: gular, pectoral and abdominal scales tricarinate. Etymology. The species name, ‘monticola’, is Latin Cnemaspis retigalensis: pectoral and abdominal scales for ‘inhabitant of the mountains’, an allusion to smooth, a single preanal pore and 3 or 4 femoral the type locality, the Wynaad Hills of Kerala State, pores. Cnemaspis kumarasinghei: dorsal scales on India. body and subcaudals smooth. Cnemaspis silvula, C. pulchra and C. pava: pectoral and abdominal scales Comparisons. Cnemaspis monticola differs from each keeled. Cnemaspis clivicola: median subcaudal scale- of its peninsular Indian and Sri Lankan congeners, row not enlarged. Cnemaspis upendrai: pectoral and respectively, by the following opposing suites of abdominal scales keeled. Cnemaspis latha: ventrals characters: 111, gular scales smooth. Cnemaspis kallima: gular Cnemaspis beddomei: spine-like tubercles absent scales smooth. Cnemaspis amith: dorsal granules on flank, 2 large supranasals, no interanasals, unkeeled. subcaudals on median series comprise alternating wide and narrow enlarged scales. Cnemaspis Remarks goaensis: postmentals widely separate by a medial See Remarks under account of C. tropidogaster. scale; subcaudals enlarged, subequal; subdigital lamellae on 4th toe, 16. Cnemaspis littoralis: dorsal Cnemaspis australis new species scales homogeneous, subcaudals on median series (Figures 13a–g; Table 2) enlarged, hexagonal. Cnemaspis mysoriensis, dorsal scales homogeneous. Cnemaspis indraneildasii: dorsal Holotype. BMNH 82.5.22.67 (male), 30.4 mm SVL; scales of both limbs smooth. Cnemaspis heteropholis: “Tinnevelly” (= Tirunelveli District, Tamil Nadu State, paired postmentals separated by three small scales; south-western India), purchased from Col. Beddome enlarged tubercles absent on neck; nostril not in (formerly a syntype of Cnemaspis tropidogaster). contact with first supralabial; no post-cloacal spurs. Cnemaspis jerdonii, C. nairi, C. otai, C. sisparensis, C. Diagnosis wynadensis, and C. yercaudensis: spine-like tubercles Cnemaspis australis differs from its peninsular Indian absent on flank. Cnemaspis boiei: postmentals widely and Sri Lankan congeners by a combination of the separated by 2 medial scales; spine-like tubercles following characters. Maximum SVL 30.4 mm; paired absent on flank. Cnemaspis gracilis: postmentals in postmentals separated by 1 or 2 small scales; each contact; subcaudals enlarged, smooth. Cnemaspis postmental bounded by 3 scales including medial indica: postmentals in contact, spine-like tubercles scale; dorsal scales heterogeneous with smooth absent on flank. Cnemaspis ornata: dorsal tubercles and carinate scales; spine-like tubercles absent on arranged in 15 longitudinal rows, dark-brown flank; ventrals weakly keeled; 3 preanal and 4 or 5 collar on nape. Cnemaspis wicksii: 5 supralabials, 14 femoral pores on each side; tail with enlarged conical subdigital lamellae on 4th toe, upper arm and lower scales forming whorls; subcaudals keeled, 2 smaller arm, thigh and tibia dorsally covered by smooth scales between single larger ones in median series; scales. Cnemaspis andersonii: subcaudals in median supralabials to angle of jaws, 8; subdigital lamellae series not enlarged; heterogeneous dorsal scales; on digit IV of pes, 15. scales on ventro-lateral region carinate; internasal as large as supranasal; Cnemaspis australis: no flank Description of holotype. tubercles; subcaudals pointed, strongly keeled; An adult male, 30.4 mm SVL (for measurements, see median row of subcaudals slightly enlarged. Table 2). Head long (HL 28.2% of SVL), moderately Cnemaspis nilagirica: dorsal scales homogeneous; broad (HW 17.4% of SVL, HW 61.6% of HL), distinct pectoral and abdominal scales weakly keeled; from neck. Snout length greater than eye diameter subcaudals on median row enlarged, subequal. (ED 57.5% of ES); interorbital broad (IO 11.6% of Cnemaspis tropidogaster: gular, pectoral and HL); pupil rounded. Granules on snout intermixed abdominal scales keeled. Cnemaspis kandiana, C. with carinate and smooth scales and few enlarged podihuna, C. scalpensis, C. gemunu, C. punctata, tubercles, larger than those of occipital region;

Vol. 7, No. 1 41 Ma n a m e n d r a -Ar a c h c h i e t a l .

scales on interorbital and supercilium smooth; scales Coloration. In preservative: general body colour light on ventral surface of head smooth, few carinate. to dark brown. Colour in life not known. Rostral scale partially divided by a medial groove, in contact with first supralabial (Fig. 13a). Nares Natural history. Not known. separated by two enlarged supranasals and a single internasal scale (Fig. 13a). Nares rounded, dorsally Etymology. The species name ‘australis’ is Latin orientated, not in contact with first supralabial. Two for ‘southern’; an allusion to the type locality being postnasals, the lower one in broad contact with very nearly at the southernmost tip of the Indian first supralabial (Fig. 13a). Mental subtriangular, as peninsula. wide as long, posteriolaterally in contact with two enlarged postmentals and medially with 2 small Comparisons. Cnemaspis australis differs from each postmental scales; postmentals bordered posteriorly of its peninsular Indian and Sri Lankan congeners, by 6 smooth scales, including 2 medial scales (Fig. respectively, by the following opposing suites of 13b). Supralabials 8, infralabials 7, decreasing in size characters: towards gape. Ear opening oval, higher than wide, Cnemaspis beddomei: ventrals smooth, preanal oblique (TW 50.0% of TH). Scales on ventral surface pores 6–9, no femoral pores. Cnemaspis goaensis: of neck carinate, laterally smooth, intermixed with ventrals smooth. Cnemaspis littoralis: dorsal scales large, conical granules. 20 scales between anterior homogeneous, subcaudals on median series margin of ear opening and posterior margin of eye. enlarged, hexagonal. Cnemaspis mysoriensis: dorsal Five conical granules larger than those of occipital scales homogeneous. Cnemaspis indraneildasii: region laterally behind ear. dorsal scales of both limbs smooth; ventral, gular Body slender, elongate. Mid-dorsal granules and caudal scales smooth. Cnemaspis heteropholis: smooth and carinate, intermixed with enlarged paired postmentals separated by three small scales, conical keeled tubercles (Fig. 13c); spine-like enlarged tubercles absent on neck, nostril not in tubercles absent on flank; abdominal scales weakly contact with first supralabial and no post-cloacal keeled (Fig. 13d). Ventrals, 121, bluntly pointed (Fig. spurs. Cnemaspis jerdonii: no preanal pores. Cnemaspis 13d); scales around vent and base of tail smooth, nairi: no femoral pores. Cnemaspis otai: ventrals imbricate; 3 preanal and 5 (left side) and 4 (right side) smooth. Cnemaspis sisparensis: ventrals smooth, femoral pores (Fig. 13e). preanal pores absent, femoral pores 7 or 8. Cnemaspis Forelimbs relatively short, slender (LAL 13.8% wynadensis: subcaudals enlarged, subequal; no SVL, UAL 10.8% SVL); hind limbs long (TBL 17.7% preanal pores, 5 femoral pores, postmentals widely SVL, FEL 20.3% SVL); tibia short. Scales on anterior separated, ventrals smooth. Cnemaspis yercaudensis: surface of thigh carinate, those on ventral surface ventrals smooth. Cnemaspis boiei: no preanal or smooth. Scales on dorsal surface of manus and pes femoral pores. Cnemaspis gracilis, C. wicksii and C. keeled; on ventral surface smooth, granular. Digits indica: ventrals smooth. Cnemaspis ornata: no femoral elongate, slender, all bearing slightly recurved claws; pores, preanal pores 6–8, dark brown collar on nape. subdigital lamellae entire, unnotched (Fig. 13g); Cnemaspis andersonii: gulars carinate; other ventral subdigital lamellae on finger I, 11, finger II, 12, finger scales smooth. Cnemaspis monticola: no large, conical III, 13, finger IV, 13, finger V, 12; toe I, 10, toe II, 12, tubercles on base of tail, spine-like tubercles present toe III, 12, toe IV, 15, toe V, 15; interdigital webbing on flank; pectoral and abdominal scales smooth. absent; relative length of digits (fingers): IV (2.7 mm) Cnemaspis nilagirica: dorsal scales homogeneous; > V (2.4 mm) > III (2.5 mm) > II (2.5 mm) > I > (2.0 subcaudals on median row enlarged, subequal. mm); (toes) IV (3.5 mm) > III (3.1 mm) > II (3.0 mm) Cnemaspis tropidogaster: spine-like tubercles > I (1.9 mm) > V (3.2 mm). present on flank. Cnemaspis kandiana, C. podihuna, Tail tapering, its length greater than SVL (TAL C. scalpensis, C. gemunu and C. phillipsi: presence of 1.15% of SVL); flattened; tail base distinctly swollen. smooth subcaudals. Cnemaspis alwisi: 9 femoral pores. Six large, conical tubercles on base of tail, about as Cnemaspis molligodai: dorsal scales homogeneous; large as granules on mid-body. Dorsal scales on tail subcaudals on median row enlarged, subequal. pointed, keeled, directed backwards. Tail with about Cnemaspis samanalensis: gular, pectoral and abdominal 6–8 broad, pointed, keeled tubercles, separated by scales tricarinate. Cnemaspis retigalensis: pectoral and about 2 or 3 scales; one conical post-cloacal spur abdominal scales smooth. Cnemaspis punctata: 5–7 present on each side; subcaudals pointed, keeled, femoral pores; distinctive spotted dorsal coloration. median row with pairs of smaller scales between Cnemaspis kumarasinghei, C. kallima and C. latha: individual larger ones (Fig. 13f). abdominal scales and subcaudal scales smooth.

42 Zeylanica Re v i s i o n o f Cn e m a s p i s

a b

c d

e f

g

Figure 13. Cnemaspis australis: BMNH 82.5.22.67, holotype, male, a, dorsal aspect of snout; b, ventral aspect of chin; c, d, arrangement of scales on dorsal and ventral regions, respectively, of mid-body; e, preanal and hind-limb regions showing arrangement of femoral and preanal pores; f, arrangement of subcaudal scales; g, ventral aspect of digit I of left manus. Scale bars: 1 mm.

Vol. 7, No. 1 43 Ma n a m e n d r a -Ar a c h c h i e t a l .

Cnemaspis silvula, C. clivicola and C. upendrai: spine- on right side); postmentals bordered posteriorly like tubercles present on flank. Cnemaspis amith: by 5 smooth scales including medial scales (Fig. pectoral and abdominal scales smooth; subcaudals 14b). Supralabials, 8, infralabials, 6, decreasing in on median row enlarged, irregular. size towards gape (Fig. 14c). Ear opening elliptical, higher than wide, oblique (TW 25.0% of TH). Scales Remarks on ventral side of neck carinate. Granules between See Remarks under account of C. tropidogaster. anterior surface of ear opening and posterior margin of eye, 22. Body elongate. Mid-dorsal granules homogeneous, Cnemaspis nilagirica isolated and scattered, with a few large, carinate (Figures 14a–h; Table 2) scales (Fig. 14d); 7 spine-like tubercles on flank; abdominal scales imbricate, rounded, feebly carinate Material examined. BMNH 74.4.29.729 (female), 42.3 (Fig. 14e). Ventrals, 137, scales around vent and at mm SVL; “Nilgiries” (Nilgiri District, Tamil Nadu base of tail weakly carinate, imbricate; no preanal State, south-western India), purchased from Colonel or femoral pores. Beddome (formerly a syntype of C. tropidogaster). Forelimbs moderately long, slender (LAL 13.2% SVL, UAL 13.7% SVL); hind limbs relatively long Diagnosis (TBL 16.5% SVL, FEL 18.9% SVL); tibia short. Cnemaspis nilagirica differs from its peninsular Dorsal scales on both fore and hind limbs carinate Indian and Sri Lankan congeners by a combination or smooth; ventral scales on upper and lower arms of the following characters. Maximum SVL at least smooth. Scales on anterior surface of thigh carinate, 42.3 mm; paired postmentals separated by a small those on ventral surface smooth. Granules on scale; each postmental bounded by 2 scales; dorsal posterior surface of thigh smooth. Scales on dorsal scales homogeneous; spine-like tubercles present and ventral surface of shank smooth. Scales on on flanks; ventrals smooth; no preanal and femoral dorsal side of manus and pes carinate, on ventral pores in female holotype; tail with enlarged pointed side smooth. Digits elongate, slender, all bearing scales forming whorls; subcaudals enlarged, smooth; claws that are slightly recurved; subdigital lamellae supralabials to angle of jaw, 8; subdigital lamellae entire, unnotched (Fig. 14h); subdigital lamellae on on digit IV, 16. finger I, 11, finger II, 12, finger III, 15, finger IV, 15, finger V, 14; toe I, 11, toe II, 14, toe III, 15, toe IV, Description of typical example 16, toe V, 15; interdigital webbing absent; relative An adult female, 42.3 mm SVL, BMNH 74.4.29.729 length of digits (fingers): IV (4.2 mm) ~ III (4.2 mm) (for measurements, see Table 2). Head relatively > V (4.0 mm) > II (3.7 mm) > I (3.0 mm); (toes) III short (HL 25.0% of SVL), broad (HW 17.0% of (5.5 mm) ~ IV (5.5 mm) > V (4.7 mm) > II (4.5 mm) SVL, HW 67.9% of HL), distinct from neck. Snout > I (2.7 mm). length greater than eye diameter (ED 59.6% of ES); Regenerated tail tapering, its length less than interorbital wide (IO 11.3% of HL); pupil rounded. SVL (TAL 77.3% of SVL). Dorsal scales on mid-tail Granules on snout larger than those in occipital oval, keeled, directed backwards. About 6 enlarged, region, carinate, intermixed with few smooth scales; pointed tubercles on dorsum, separated from one scales immediately above supralabials smooth; another by 6 scales (Fig. 14f); subcaudals smooth, scales on interorbital and supercilium smooth; scales with an enlarged median series of pentagonal scales on ventral side of head smooth, a few carinate. (Fig. 14g). Rostral scale partially divided by a medial groove, in contact with first supralabial (Fig. 14a). Nares Coloration. In preservative: overall body colour separated by two enlarged supranasals and a single brown; head with irregular yellow markings; an elongated internasal scale (Fig. 14a). Two enlarged uninterrupted yellow vertebral line present; limbs scales behind internasal-supranasal scale row. Nares brown; throat dark brown, venter yellow. In life: rounded, dorsally orientated, not in contact with not known. first supralabial (Fig. 14a). Two postnasals, the lower one in broad contact with first supralabial. Mental Etymology. The species epithet ‘nilagirica’ is subtriangular, as wide as long, posteriolaterally in a Latinized compound noun derived from the contact with two enlarged postmentals and medially Sanskrit roots for the type locality, the Nilgiris [Blue with a single small pentagonal postmental scale Mountains] of southern India (nil = blue, giri = (mental in narrow contact with second medial scale mountains), applied here as a noun in apposition.

44 Zeylanica Re v i s i o n o f Cn e m a s p i s

a b

c d

e f

g h

Figure 14. Cnemaspis nilagirica: BMNH 74.4.29.729, holotype, female, 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 and ventral regions, respectively, of mid-body; f, g, arrangement of scales on dorsal and ventral regions, respectively, of mid-tail; h, ventral aspect of digit V of right pes. Scale bars: 1 mm.

Vol. 7, No. 1 45 Ma n a m e n d r a -Ar a c h c h i e t a l .

Comparisons. Cnemaspis nilagirica differs from each along the tail; chin and throat covered with of its peninsular Indian and Sri Lankan congeners, small hexagonal scales, gradually changing respectively, by the following opposing suites of to rounded imbricated scales which cover the characters: abdomen; scales of tail imbricate, with 3 rows Cnemaspis beddomei, C. nairi, C. otai, C. sisparensis, of larger nail-shaped scutæ beneath; scales C. wynadensis and C. yercaudensis: spine-like tub­ of the basal joints of the fingers and toes, ercles absent on flank. Cnemaspis goaensis: post­ enlarged, nail-shaped. Body above greyish- mentals widely separate by medial scale. Cnemaspis brown, with a light stripe down the centre of littoralis: subcaudals on median series enlarged, the back, and a series of dark brown marks hexagonal. Cnemaspis mysoriensis: three rows of on the head, back and sides; legs and feet enlarged subcaudals. Cnemaspis indraneildasii and banded, beneath bluish-white; chin, throat and C. heteropholis: dorsal scalation heterogeneous. anterior portion of palpabræ bright yellow. Cnemaspis jerdonii: gular and throat scales smooth. Length of one 2½ inches, of which the tail 1 Cnemaspis boiei: postmentals (widely) separated by 2 is 1 /4. I have quite recenty found this small medial scales, spine-like tubercles absent. Cnemaspis and very distinctly characterized species of gracilis and C. indica: postmentals in contact. Gecko at Bangalore, frequenting rocks and Cnemaspis ornata: dorsal tubercles arranged in 15 also entering out-houses. The young has the longitudinal rows. Cnemaspis wicksii, C. andersonii tail flesh-coloured: 2 femoral pores on one side and C. monticola: gular scales carinate. Cnemaspis and 3 on the other. australis: subcaudals pointed, strongly keeled. Cnemaspis tropidogaster, C. kandiana, C. samanalensis, Dispite references to the species in the non-taxon­ C. silvula, C. upendrai, C. pava, and C. clivicola: omic literature (e.g., Biswas, 2005), no specimens dorsal scales heterogeneous. Cnemaspis podihuna, attributable to C. mysoriensis have, to our knowledge, C. scalpensis, C. gemunu, C. alwisi, C. molligodai, C. been recorded since the original description. The punctata, and C. phillipsi: caudal scales smooth. holotype of C. nilagirica shows a few characters Cnemaspis kumarasinghei, C. retigalensis and C. latha: not mentioned by Jerdon, which are potentially of subcaudals on median row enlarged, irregular. generic value, e.g., keeled ventral scales. Given that Cnemaspis kallima: subcaudals keeled. Cnemaspis C. nilagirica is known only from a single female, the menikay: dorsal scales heterogeneous, conical. presence and number of preanal and femoral pores Cnemaspis amith: pectoral and abdominal scales in this population, too, is unknown. The locality from smooth; dorsal granules heterogeneous; subcaudals which the specimen of C. nilagirica was collected, the on median row enlarged, irregular. Nilgiri Hills, is ~ 250 km distant from Bangalore. What is more, Bangalore is located at an altitude ~ 900 Remarks m, whereas the Nilgiri Hills rise to 2,637 m at Doda Cnemaspis nilagirica, as shown above, is clearly Betta. Given this distance and difference in elevation, distinct from all peninsular Indian and Sri Lankan we consider it likely that C. nilagirica will prove to species of Cnemaspis known from recent populations. be distinct from C. mysoriensis, and hope that future We are unable to eliminate, however, the possibility exploration by herpetologists in India will serve to that it is identical with C. mysoriensis (Jerdon, 1853), clarify the identity of these taxa. the types of which are evidently lost: there has been See also Remarks under account of C. tro­pido­ no record of them in the century-and-a-half since gaster. the original description. Although no material of the species has survived, Jerdon provided what was for his time a relatively detailed description, Key to the peninsular Indian species of Cnemaspis together with a precise type locality, Bangalore, which is today a sprawling city of some six million 1a. Spine-like tubercles on flanks...... 2 inhabitants: 1b. Spine-like tubercles absent on flanks...... 3

Scales of back uniform, granulose, with 2 2a. Dorsal scales on mid-body homogeneous (Fig. 1d)...... 4 or 3 rows of spines on each side, and on the 2b. Dorsal scales on mid-body tail; tail beneath with a triple series of large heterogeneous (Fig. 3d)...... 5 uniform scutæ. Head and back covered with small granulose scales, and two or three distant 3a. Preanl pores absent (Fig. 5f)...... 6 rows of spines on each side and extending 3b. Preanal pores present (Fig. 4f)...... 7

46 Zeylanica Re v i s i o n o f Cn e m a s p i s

4a. Subcaudals on median row SRI LANKAN SPECIES enlarged (Fig. 5f)...... 8 4b. Subcaudals on median row Cnemaspis kandiana (Kelaart, 1852) not enlarged...... C. mysoriensis / C. nilagirica Gymnodactylus kandianus Kelaart, 1852 Gymnodactylus humei Theobald, 1876 5a. Subcaudals on median row not enlarged (Fig. 3g)...... C. gracilis (Figures 15a–g, 16; Tables 3, 6; Plate 3a–d) 5b. Subcaudals on median row enlarged, irregular (Fig. 8g)...... 9 Lectotype (here designated). BMNH 53.4.1.1 (female), 33.2 mm SVL, “Ceylon” (Sri Lanka) purchased from 6a. Femoral pores present (Fig. 5f)...... 10 Dr. Kelaart. 6b. Femoral pores absent (Fig. 1f)...... C. boiei Paralectotypes. BMNH 80.2.2.119 (male), 29.6 mm 7a. Femoral pores present...... 11 SVL; BMNH 80.2.2.119A (male), 31.6 mm SVL; BMNH 7b. Femoral pores absent...... 12 80.2.2.119B (female), 32.8 mm SVL; BMNH 80.2.2.119 8a. Femoral pores, 8...... C. jerdonii (female), 31.4 mm SVL; BMNH 80.2.2.119 (female), 8b. Femoral pores, 14–18...... C. littoralis 34.6 mm SVL, “Ceylon” (Sri Lanka) purchased from Dr. Kelaart.. 9a. Gular scales smooth...... 13 9b. Gular scales carinate...... 14 Other material. ZSI 5971 (male), 36.0 mm SVL, holotype of Gymnodactylus humei Theobald, 1876; 10a. Dorsal scales on mid-body WHT 7305, 7307, 7308, 7310, Gannoruwa Forest granular (Fig. 5d)...... 15 Reserve, near Peradeniya, Kandy District, 07º17’10” 10b. Dorsal scales on mid-body N, 80º35’30” E, 700 m, coll. S. Batuwita, 22 November, imbricate (Fig. 2d)...... C. indica 1997; WHT 7267, 7313, 7319, 7322, same location data 11a. Pectoral and abdominal as above, coll. S. Batuwita, A. Silva & K. Maduwage, scales smooth (Fig. 2e)...... 16 02 November, 2002; WHT 7212, 7213, Hantana 11b. Pectoral and abdominal Mountain near Peradeniya, 07º14’30” N, 80º37’E, scales keeled (Fig. 13d)...... C. australis n. sp. 762 m, coll. S. Batuwita & D. A. Karunaratne, 15 November, 2005. 12a. Subdigital lamellae on proximal portion entire (Fig. 13g)...... 17 Diagnosis 12b. Subdigital lamellae on proximal Cnemaspis kandiana differs from its peninsular Indian portion fragmented (Fig. 4i)...... C. ornata and Sri Lankan congeners by a combination of the 13a. Mid-dorsal enlarged following characters. Maximum SVL 36.0 mm; tubercles numerous...... C. heteropholis paired postmentals separated by a small scale; each 13b. Mid-dorsal enlarged postmental bounded by 3 scales including medial tubercles moderate...... C. indraneildasii scale; ventrals, 121–138; ventral scales across mid- body, 19; dorsal scales heterogeneous, isolated and 14a. Subcaudals smooth (Fig. 11e)...... C. goaensis scattered; 6 spine-like tubercles present on flank; 14b. Subcaudals keeled (Fig. 12h)...... C. monticola n. sp. ventrals smooth, scales on ventral surface of thigh carinate, smooth; 2–4 preanal pores; 3 or 4 femoral 15a. Femoral pores, 7 or 8; median series of subcaudals in an pores on each side; tail with enlarged conical scales enlarged irregular series...... C. sisparensis forming whorls; subcaudals of median series 15b. Femoral pores, 4–6; median enlarged, irregular; supralabials to angle of jaws, 8 series of subcaudals in an or 9; subdigital lamellae on digit IV of pes, 18–20; enlarged regular series...... C. wynadensis (in life) dorsum dark brown, with about seven black V-shaped markings, a light brown blotch behind 16a. Preanal pores, 2; postmentals each marking. not in contact...... C. yercaudaensis 16b. Preanal pores, 4; postmentals Description of lectotype in contact...... C.otai An adult female, 33.2 mm SVL (for measurements, 17a. Tail subcylindrical, elongate...... C. nairi see Table 3). Head relatively short (HL 25.6% of SVL), 17b. Tail dorso-ventrally flattened moderately broad (HW 16.9% of SVL, HW 65.9% of and lanceolate (Plate 1g, h)...... C. beddomei HL), distinct from neck. Snout length greater than

Vol. 7, No. 1 47 Ma n a m e n d r a -Ar a c h c h i e t a l .

eye diameter (ED 59.1% of ES); interorbital moderate Dorsal scales on tail oval, directed backwards. Tail (IO 9.4% of HL); pupil rounded. Granules on snout with about 6 broad, keeled conical tubercles which keeled; scales immediately above supralabials are separated from one another by about 6 scales; a smooth, larger than those of occipital region; scales single conical post-cloacal spur present on each side; on interorbital and supercilium conical; scales subcaudals on median row enlarged, smooth, with on ventral surface of head smooth. Rostral scale pairs of smaller scales between individual larger partially divided by a medial groove, in contact with ones; other scales carinate (Fig. 15f). first supralabial (Fig. 15a). Nares separated by two enlarged supranasals and a single internasal scale Coloration. In preservative: dorsally light brown (Fig. 15a). Nares rounded, dorsally orientated, not with indistinct irregular pale markings on dorsum. in contact with first supralabial. Two postnasals, Limbs with pale spots; endolymphatic sacs brownish the lower one in broad contact with first supralabial yellow; venter whitish (see Plate 3a, b). (Fig. 15a). Mental subtriangular, as wide as long, In life: dorsum dark-brown, with about seven posteriolaterally in contact with two elongate black V-shaped markings; a light-brown blotch postmentals and medially with a single small behind each of these (see Plate 3c). pentagonal postmental scale; postmentals bordered posteriorly by 5 smooth scales including medial scale Variation. Paralectotype BMNH 80.2.2.119 followed (Fig. 15b). Supralabials 9, infralabials 8, decreasing in by paralectotypes BMNH 80.2.2.119A, BMNH size towards gape (Fig. 15c). Ear opening oval, higher 80.2.2.119B, BMNH 80.2.2.119, BMNH 80.2.2.119 in than wide, oblique (TW 50.0% of TH). Scales on parentheses: supralabials 8 (8, 8, 8, 8), infralabials ventral surface of neck feebly carinate. Twenty-three 7 (8, 7, 7, 8). Preanal and femoral pores in males: granules between anterior margin of ear opening and preanal pores, 2 or 4; femoral pores, 3 or 4; pores posterior margin of eye. Conical granules on side of absent in females; ventrals, 121–138; SVL ranges neck larger than those of occipital region. A large 29.6–36.0 mm. conical scale at back of ear. Body slender, short. Mid-dorsal granules isolated Distribution and natural history. The species is and scattered, intermixed with large conical tubercles endemic to the moist forests (rainfall > 2,500 mm (Fig. 15d); 6 spine-like tubercles on flank; abdominal yr-1) of the Central Province of Sri Lanka, where it scales wider than long, circular or bluntly pointed, appears restricted to altitudes in the range 400–700 smooth (Fig. 15e). Scales around vent and base of m. Cnemaspis kandiana is a diurnal-crepuscular gecko, tail smooth, imbricate; no preanal and femoral pores most often observed on rock faces and occasionally (Fig. 15f). on tree trunks in shaded forest habitats. On Hantana Forelimbs moderately long, slender (LAL 13.3% mountain, some individuals were observed under SVL, UAL 11.5% SVL); hind limbs long (TBL 18.7% decaying logs. These geckos were also seen in SVL, FEL 19.6% SVL); tibia short. Dorsal scales on edificarian habitats including abandoned houses both fore and hind limbs carinate; ventral scales on close to the forest, two or more individuals often upper arm narrow, pointed, smooth; ventral scales occurring together. It is evidently a communal- on lower arm smooth. Scales on anterior surface nesting species (see Plate 3d). We recorded C. kandiana of thigh carinate , those on ventral surface smooth from following localities: Gannoruwa, Hantana (Fig. 15f). Granules on posterior surface of thigh Mountain, Kandy, Pahala-Kadugannawa, Polgolla conical. Scales on ventral surface of shank carinate, (Fig. 16). The following lizards were found to occur in those ventrally on manus and pes smooth, rounded. sympatry with C. kandiana: C. scalpensis, Hemidactylus Digits elongate, slender, all bearing slightly recurved depressus, Cyrtodactylus fraenatus, Geckollea triedra, claws; subdigital lamellae entire, unnotched (Fig. Lyriocephalus scutatus, Otocryptis wiegmanni, Calotes 15g); subdigital lamellae on finger I, 11, finger II, liolepis, Lankascincus taylori and L. gansi. 14, finger III, 15, finger IV, 9, finger V, 14; toe I, 11, toe II, 14, toe III, 15, toe IV, 18, toe V, 17; interdigital Comparisons. Cnemaspis kandiana differs from each webbing absent; relative length of digits (fingers): IV of its Sri Lankan and peninsular Indian congeners, (2.8 mm) > V (2.6 mm) ~ III (2.6 mm) > II (2.3 mm) respectively, by the following opposing suites of > I (1.8 mm); (toes) III (3.8 mm) > II (3.6 mm) > I (3.3 characters: mm) > IV (3.0 mm) > V (1.9 mm). Cnemaspis tropidogaster: keeled ventrals and Tail tapering, its length a little greater than SVL heterogeneous dorsal scales. Cnemaspis podihuna: (TAL 102.4% of SVL). Six large, conical tubercles on a median row of regular, enlarged, hexagonal base of tail, about as large as granules on mid-body. subcaudals. Cnemaspis scalpensis, C. gemunu and C.

48 Zeylanica Re v i s i o n o f Cn e m a s p i s 4.4 1.3 – 5.1 3.3 1.6 2.7 3.4 3.9 3.0 2.5 3.8 2.3 2.5 2.9 3.7 4.9 1.5 2.3 2.7 3.0 2.0 6.3 8.1 5.3 0.8 0.6 3.9 2.9 4.6 7.2 0.8 4.0 5×3 7236 13.2 29.2 WHT 4.3 1.0 – 5.5 3.6 1.7 2.8 3.4 3.8 2.9 2.2 3.9 2.6 2.4 3.3 4.7 5.5 1.6 2.6 3.0 3.3 2.5 6.4 8.9 5.8 1.0 0.8 4.4 3.0 5.3 8.3 1.0 4.2 8×4 14.5 33.3 7238 WHT 6.3 1.3 5.7 4.2 1.8 2.7 3.3 3.9 3.1 3.0 4.5 2.5 2.8 3.3 4.4 5.9 1.8 2.6 3.0 3.0 2.3 6.5 9.3 6.0 1.0 1.1 4.2 3.5 5.5 8.4 1.0 4.3 5×3 16.7 33.9 36.6 C. phillipsi 7237 WHT 5.5 1.5 5.9 3.6 1.9 2.7 3.2 3.6 2.9 2.2 4.3 2.3 2.0 2.8 4.3 6.5 1.8 2.6 3.0 3.2 2.3 5.9 8.9 5.7 1.0 0.7 4.6 3.2 3.7 8.0 0.9 4.4 5×3 7248 14.9 33.9 33.5 WHT WHT 7248 ) ( WHT holotype and podihuna ; C. 2.8 1.1 8.9 – 3.4 2.5 2.0 2.7 2.2 1.9 1.2 2.4 2.9 1.7 2.0 2.0 2.8 4.8 1.0 1.6 1.9 2.2 1.7 3.9 6.6 4.2 1.8 0.3 3.0 2.1 3.8 5.3 0.7 1.9 4×2 23.3 7334 WHT

R 2.8 0.5 2.3 4.1 2.1 1.2 2.4 2.6 2.6 2.3 1.8 1.8 2.2 1.9 2.5 3.8 1.3 2.0 1.9 2.0 1.7 4.3 5.8 3.7 0.6 0.8 3.2 2.4 3.7 5.3 0.7 3.0 C. podihuna 2×1 11.4 22.8 23.6 NMSL (uncat.)

8×5 3.7 1.3 – 4.7 6.8 1.9 3.3 3.5 4.3 3.6 3.7 2.9 3.6 2.6 3.3 4.1 6.4 2.2 2.7 2.9 2.9 2.6 6.0 8.6 5.5 1.1 0.8 4.4 3.7 5.8 8.0 0.7 3.7 31.4 BMNH C. tropidogaster 71.12.14.49 holotype (NMSL, uncat.) and 1 example (WHT 7334) of 7334) (WHT example 1 and uncat.) (NMSL, holotype 4.7 1.1 4.2 5.9 3.8 1.7 2.8 3.4 4.0 3.2 3.0 2.7 2.7 3.3 4.3 5.4 1.7 2.5 2.8 3.1 2.4 6.4 9.1 6.1 1.0 0.6 4.4 3.2 5.3 8.3 1.0 4.0 6×4 15.0 7265 35.6 34.4 WHT Cnemaspis kandiana ; neotype (NMSL 2004.1.1) and three examples ( WHT 7268, WHT 7277, WHT

R 6×4 3.8 1.2 3.7 5.6 3.5 1.6 2.7 3.1 3.9 3.0 2.5 2.3 2.7 2.8 4.0 5.8 1.6 2.3 2.4 2.7 2.7 6.0 8.5 5.5 0.9 0.5 4.2 3.0 4.9 7.5 0.9 3.8 24.0 12.7 31.0 7277 B WHT C. tropidogaster; tropidogaster; C. 4.5 1.1 3.4 5.2 3.6 1.6 2.7 3.4 4.1 3.2 2.5 2.5 2.8 2.9 3.9 5.7 1.9 2.6 3.0 3.0 2.6 6.7 8.4 5.5 1.0 0.5 4.5 3.2 5.0 7.7 0.9 4.2 3.5,36.0 7×3 C. scalpensis 13.3 7268 31.3 WHT

4.7 1.0 2.9 5.3 3.3 1.7 2.7 3.1 3.9 3.2 2.8 2.3 2.6 2.8 3.5 4.4 1.6 2.3 2.4 2.5 2.0 5.8 7.7 5.0 1.0 0.6 4.3 2.7 4.7 7.0 0.8 3.4 5×4 13.0 29.7 37.0 NMSL 2004.1.1

1.2 3.8 3.9 6.2 – 3.3 3.6 3.8 3.0 1.9 2.9 3.6 2.6 3.4 4.3 6.5 1.8 2.3 2.6 2.8 2.6 5.7 8.5 5.6 1.1 0.8 4.4 3.6 5.3 8.0 0.8 3.8 6×3 33.2 34.0 BMNH 53.4.1.1 C. kandiana holotype (BMNH 71.12.14.49) of 71.12.14.49) (BMNH holotype C. scalpensis; C.

UAL UEW Table 3. Measurements (in Table mm) of lectotype (BMNH 53.4.1.1) of of 7265 ) AG B indicates tail broken; R indicates tail regenerated; – indicates tail lost . R indicates tail regenerated; and 3 paratypes ( WHT 7237, 7238, 7236 ) of C. phillipsi. B indicates tail broken;

DBE TBL TBW TL I TL TL II TL TL III TL TL IV TL TL V TL TYD TYE (×10) DFE ED EN ES FEL FL I FL FL II FL FL III FL FL IV FL FL V FL FOL HL HW IN IO LAL MBE MFE MN NS PAL SVL TAL

Vol. 7, No. 1 49 Ma n a m e n d r a -Ar a c h c h i e t a l .

a b

c d

e f

g

Figure 15. Cnemaspis kandiana: BMNH 53.4.1.1, lectotype, female, 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 and ventral regions, respectively, of mid-body; f, preanal, right hind limb and tail regions, showing absence of pores and arrangement of subcauidal scales; g, ventral aspect of right manus. Scale bars: 1 mm.

50 Zeylanica Re v i s i o n o f Cn e m a s p i s

phillipsii: 12–16 femoral pores. Cnemaspis alwisi: 9 survived (Pethiya­­goda & Mana­­­mendra-Arachchi, femoral pores. Cnemaspis molligodai: 5 preanal and 1997). Eight of the syntypes of his Gymnodactylus 8 or 9 femoral pores, gular, pectoral and abdominal kandianus, however, did survive­ in the BMNH col­ scales smooth, dorsal scales homogeneous; median lection, but these had evidently not until now been row of subcaudals enlarged, subequal. Cnemaspis reported on, helping to verify the suspicion expressed samanalensis: gular, pectoral and abdominal scales by Das & Leong (2004) and Das (2005) that C. kandiana tricarinate, subcaudals keeled. Cnemaspis retigalensis: is in fact a restricted-range Sri Lankan highland single preanal pore and 3 femoral pores. Cnemaspis endemic. punctata: 5–7 femoral pores and a distinct spotted Kluge (2001) listed Cnemaspis malabarica (Jerdon, dorsal colour pattern. Cnemaspis kumarasinghei 1853) in the synonymy of C. kandiana. According to and C. latha: gular scales and subcaudals smooth. Jerdon (1853), however, C. malabarica (type locality Cnemaspis silvula, C. pulchra, C. pava, C. clivicola and Kerala State, southern India) has homogeneous C. upendrai: pectoral and abdominal scales keeled. dorsal scalation: “Scales of the back uniform, small Cnemaspis kallima: subcaudals on distal half of tail granulose…” (vs dorsal scalation heterogeneous in C. carinate. Cnemaspis menikay: dorsal granules conical, kandiana). For this reason and for the reason that C. subcaudals smooth. Cnemaspis amith: dorsal granules kandiana shows a very restricted range in Sri Lanka, heterogeneous, unkeeled. we consider these species distinct. No type material Cnemaspis beddomei: preanal pores 6–9, no femoral of Cnemaspis malabarica is known to survive and it is pores. Cnemaspis goaensis: subcaudals on median row important that the identity of this species be stabilized slightly enlarged. Cnemaspis littoralis: body slender, through the designation and description of a neotype dorsal scales homogeneous; subcaudals in median from an extant population in the type locality. series enlarged, hexagonal. Cnemaspis mysoriensis: Smith (1935) also placed Cnemaspis wicksii dorsal scales homogeneous, three rows of enlarged (Stoliczka, 1873) and Cnemaspis andersonii (Annandale, subcaudal scales. Cnemaspis indraneildasii: dorsal 1904) in the synonymy of C. kandiana. Based on an scales of both limbs smooth, gular scales smooth. examination of their types, however, we recognize Cnemaspis heteropholis: gular scales smooth. Cnemaspis these species as distinct and valid (see Remarks sisparensis and C. wynadensis: spine-like tubercles under the respective species accounts). absent on flank. Cnemaspis boiei: no preanal or femoral pores. Cnemaspis gracilis: 2 preanal pores separated by 2 scales. Cnemaspis indica: spine-like tubercles absent on flank. Cnemaspis ornata: no femoral pores, dorsal tubercles arranged in 15 longitudinal rows, dark brown collar on nape. Cnemaspis wicksii: thigh and tibia dorsally covered in smooth scales, 4 preanal pores. Cnemaspis andersonii: scales along ventro-lateral region carinate. Cnemaspis jerdonii: no preanal pores. Cnemaspis nairi, C. otai and C. yercaudensis: spine-like tubercles absent on flank. Cnemaspis monticola: scales on ventral surface of thigh smooth. Cnemaspis australis: subcaudals on median row pointed, strongly keeled. Cnemaspis nilagirica: dorsal scales homogeneous; pectoral and abdominal scales weakly keeled; subcaudals on median row enlarged, subequal.

Remarks Cnemaspis kandiana has served as a catchall species for a number of almost certainly distinct geckos from across South Asia (Boulenger, 1890; De Rooij, 1915; Brongersma, 1934; Smith, 1935; Annandale, 1909; Abdulali, 1955; Taylor, 1963; Dring et al., 1990; Thakur, 1998; Thomas & Easa, 1997; Rösler, 1981; Cox et al., 1998). It had long been supposed that few, if any, of Figure 16. Distribution of Cnemaspis kandiana in Sri Lanka. the types of species described by E. F. Kelaart had The type locality is indicated by a solid symbol.

Vol. 7, No. 1 51 Ma n a m e n d r a -Ar a c h c h i e t a l .

The eight syntypes of C. kandiana belong to two no preanal pores; 13–15 femoral pores on either distinct species, one of which (C. amith) we here side; subcaudals smooth, scales on median row describe as new. The only other species of Cnemaspis enlarged, subequal; supralabials to angle of jaws, we have found to be sympatric with C. kandiana is C. 7–9; subdigital lamellae on digit IV of pes, 19–21; scalpensis. Cnemaspis kandiana differs from C. amith, (in life) dorsum light-brown, with about six light- however, by the presence of keeled dorsal granules brown confluent blotches; nape with a distinct short (absent in C. amith, whose dorsal granules are black marking. unkeeled). It is also distinguished from C. scalpensis by the presence of preanal pores and lacking a row Description of neotype of subequal median subcaudals. We suspect therefore An adult male, 29.7 mm SVL (for measurements, see that while the syntypes of C. kandiana sensu stricto Table 3). Head moderately long (HL 25.9% of SVL) came from the environs of Kandy (“Kandyan Hills”, and broad (HW 16.8% of SVL, HW 64.9% of HL), to which the species is even now restricted), they dis­tinct from neck. Snout length greater than eye included also examples from elsewhere in Sri Lanka dia­meter (ED 74.2% of ES); interorbital moderate (i.e., C. amith) that Kelaart (1852) mistakenly assigned (IO 7.7% of HL); pupil rounded. Granules on snout to C. kandiana. smooth, larger than those of occipital region; scales Wick­rama­singhe & Munindra­dasa (2007) listed on interorbital and supercilium granular; enlarged four ‘syntypes’ of Cnemaspis kandiana from NMSL; conical tubercles present on top and sides of head Pethiyagoda (2007), however, has shown that these and on sides of neck; scales on ventral surface are not in fact syntypes of this species. of head and neck smooth. Rostral scale partially divided by a medial groove, in contact with first Cnemaspis scalpensis (Ferguson, 1877) supralabial (Fig. 17a). Nares separated by two Gymnodactylus scalpensis Ferguson, 1877 enlarged supranasals and a single internasal scale; Cnemaspis ranwellai Wickrama­ ­singhe, 2006 an enlarged scale behind each supranasal (Fig. (Figures 17a–i, 18; Tables 3, 6; Plate 4a–c) 17a). Nares rounded, dorsally orientated, not in contact with first supralabial. Two postnasals, the Neotype (here designated), NMSL 2004.1.1 (male), lower one in broad contact with first supralabial 29.7 mm SVL (holotype of Cnemaspis ranwellai Wick­ (Fig. 17a). Mental subpentagonal, constricted by rama­singhe, 2006), Gannoruwa Forest Reserve, near first infralabials, wider than long, posteriolaterally Peradeniya, Kandy District, 07º16’56.7”N, 80º35’ in contact with two quadrate postmentals and 54.0”E, 576 m, coll. L. J. Mendis Wick­rama­singhe, medially with two postmental scales; postmentals 27 December 2003. bordered posteriorly by 8 smooth scales including medial scale (Fig. 17b). Supralabials 9, infralabials Other material. NMSL 2004.2.1, NMSL 2004.3.1, 7, decreasing in size towards gape (Fig. 17c). Ear NMSL 2004.4.1 (three paratypes of Cnemaspis opening oval, higher than wide, oblique (TW 80.0% ranwellai Wickrama­ singhe,­ 2006), Gannoruwa, of TH). Kandy, Sri Lanka, same collection data as neotype; Body slender, relatively long (AG 43.7% SVL). WHT 7265, 7274, 7275, 7276, 7320, Gannoruwa Mid-dorsal granules homogeneous isolated and Forest Reserve, near Peradeniya, Kandy District, scattered (Fig. 17d), intermixed laterally with large 07º17’10” N, 80º35’30” E, 700 m, coll. S. Batuwita, conical tubercles; 10 bluntly pointed tubercles on 05 March 1998; WHT 7268, WHT 7269, same locality flank; abdominal scales smooth (Fig. 17e). Ventrals, data as above, coll. S. Batuwita, 13 August 2003. 130; ventral scales across midbody, 17; scales around vent and base of tail smooth, imbricate; no preanal Diagnosis pores; 15 (left side) and 14 (right side) femoral pores Cnemaspis scalpensis differs from its peninsular (Fig. 17f). Indian and Sri Lankan congeners by a combination Forelimbs moderately long, slender (LAL 14.4% of the following characters. Maximum SVL 36.9 SVL, UAL 15.8% SVL); hind limbs long (TBL 14.8% mm; paired postmentals in contact or separated SVL, FEL 17.8% SVL); tibia short. Dorsal scales on by a medial scale; each postmental bounded both fore and hind limbs smooth; ventral scales by 4 scales including medial scale/s; ventrals, on upper arm narrow, pointed, unkeeled; those on 122–130; ventral scales across midbody, 17–19; lower arm smooth. Scales on anterior surface and dorsal scales homogeneous; dorsal tubercles ventral surface of thigh smooth. Scales on ventral isolated, scattered; spine-like or bluntly pointed surface of shank, manus and pes smooth. Digits tubercles present on flanks; ventrals smooth; elongate, slender, all bearing slightly recurved

52 Zeylanica Re v i s i o n o f Cn e m a s p i s

claws; subdigital lamellae entire, unnotched; Lyrocephalus scutatus, Otocryptis wiegmanni and subdigital lamellae on finger I, 12, finger II, 15, Calotes liolepis. finger III, 18, finger IV, 18, finger V, 15; toe I, 11, toe II, 15, toe III, 19, toe IV, 20, toe V, 18; interdigital Comparisons. Cnemaspis scalpensis differs from each webbing absent; relative length of digits (fingers): of its Sri Lankan and peninsular Indian congeners, IV (2.5 mm) ~ III (2.4 mm) > V (2.0 mm) ~ II (2.3 respectively, by the following opposing suites of mm) > I (1.6 mm); (toes) IV (3.9 mm) > III (3.1 mm) characters: > V (3.2 mm) > II (2.7 mm) > I (1.7 mm). Cnemaspis tropidogaster: keeled ventrals and Tail tapering, its length greater than SVL (TAL heterogeneous dorsal scales. Cnemaspis kandiana, C. 124.5% of SVL); tail base distinctly swollen. Base menikay and C. podihuna: presence of preanal pores. of tail with 6–8 rows of large tubercles, decreasing Cnemaspis gemunu: each postmental bounded by to 2 rows towards tail tip. Dorsal scales on tail three scales, nostril in contact with first supralabial, smooth, oval, directed backwards; a small, conical, basal-most subdigital lamellae of distal series post-cloacal spur present on each side; subcaudals sometimes fragmented. Cnemaspis alwisi: 9 femoral smooth, with a median regular series of enlarged pores. Cnemaspis molligodai: presence of preanal scales (Fig. 17g). pores. Cnemaspis samanalensis: gular, pectoral and abdominal scales tricarinate, subcaudals keeled. Coloration. In preservative: dorsally olive with six Cnemaspis retigalensis: dorsal scales heterogeneous indistinct light-brown patches. Nape with a dark gulars carinate, a single preanal pore and 3 femoral making. Limbs and tail with dark bands; venter pores; median row of subcaudals enlarged, irregular. pale olive (see Plate 4a). Cnemaspis phillipsi: ventrals, 129–141; enlarged In life: dorsum light-brown, with about six tubercles absent on dorsal surface of tail. Cnemaspis confluent light-brown blotches; nape with a distinct punctata: 5–7 femoral pores; distinctive spotted black marking (see Plate 4b, c). dorsal coloration. Cnemaspis kumarasinghei, C. kallima and C. latha: presence of preanal pores. Cnemaspis Variation. WHT 7268, WHT 7277 and WHT 7265 in silvula, C. pulchra, C. pava, C. clivicola and C. upendrai: parentheses. Supralabials (9, 8, 7); infralabials (8, 7, subcaudals, and pectoral and abdominal scales 8); subdigital lamellae on finger I (13, 11, 11), finger keeled. Cnemaspis amith: gular scales carinate; dorsal II (13, 14, 13), finger III (16, 15, 15), finger IV (17, 17, granules heterogeneous; subcaudals on median row 16), finger V (16, 13, 13); toe I (12, 11, 12), toe II (15, enlarged, irregular. 15, 16), toe III (18, 18, 19), toe IV (21, 19, 19), toe V Cnemaspis beddomei: no femoral pores. Cnemaspis (17, 16, 17). Ventrals, 122–130; ventral scales across goaensis: preanal pores 2 or 3, gulars keeled. Cnemaspis mid-body, 17–19; no preanal pores, femoral pores, littoralis: a black spot on the nape. Cnemaspis 13–15; pores absent in females. SVL ranges from 25.2 mysoriensis: three rows of enlarged subcaudals, 2 or 3 to 36.6 mm (13 ex.). Supranasals in contact anteriorly, femoral pores. Cnemaspis indraneildasii: dorsal scales posteriorly separated by internsal in WHT 7268. heterogeneous. Cnemaspis heteropholis: dorsal scales Postmentals separated by a single medial scale in heterogeneous, paired postmentals separated by WHT 7268 (Fig. 17h) and broadly in contact in WHT three small scales, no post-cloacal spurs. Cnemaspis 7265 (Fig. 17i). jerdonii: lacks caudal tubercles, 8 femoral pores. Cnemaspis indica, C. nairi, C. otai, C. sisparensis, C. Distribution and natural history. The species wynadensis, and C. yercaudensis: spine-like tubercles endemic to Sri Lanka, evidently restricted to the absent on flank. Cnemaspis boiei: lacks preanal and Kandy valley, especially Gannoruwa Forest Reserve femoral pores. Cnemaspis gracilis: 2 preanal pores (altitiude ~500 m; rainfall > 2,500 mm yr-1) (Fig. separated by 2 scales, 4 femoral pores. Cnemaspis 18). A fast-moving gecko, usually seen in pairs but ornata: lacks femoral pores, dorsal tubercles arranged sometimes in groups of four or five individuals. It in 15 longitudinal rows. Cnemaspis wicksii and C. was observed mostly on the trunks of large trees andersonii: gulars carinate. Cnemaspis monticola: (> 1 m gbh), usually 2–3 m above ground level in gulars and caudals carinate. Cnemaspis australis: well-shaded areas within the forest. The geckos were subcaudals pointed, strongly keeled. Cnemaspis seen near branch-forks of large trees and close to tree nilagirica: pectoral and abdominal scales weakly holes, in which the geckos took refuge when alarmed. keeled. The following lizards were observed in sympatry with Cnemaspis scalpensis: C. kandiana, Hemidactylus Remarks. In describing C. ranwellai, Wickrama­ ­ depressus, Cyrtodactylus fraenatus, Geckollea triedra, singhe (2006: 26–28) admitted that “the new species

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c d

e f

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Figure 17. Cnemaspis scalpensis: NMSL 2004.01.1, neotype, male (holotype of Cnemaspis ranwellai Wickramasinghe, 2006), 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 and ventral regions, respectively, of mid-body; f, preanal and right hind-limb regions, showing arrangement of femoral pores; g,arrangement of subcaudal scales; h, i, ventral aspects of chin respectively of, WHT 7268 and WHT 7265. Scale bars: 1 mm.

54 Zeylanica Re v i s i o n o f Cn e m a s p i s

is similar to” C. scalpensis (which, however, he spelt stabilize its taxonomic status, and so that its type “scalapensis”). He identified C. scalpensis (the types locality could be defined. We therefore designate of which are lost: see below) not on the basis of NMSL 2004.1.1 (male), 29.7 mm SVL, the holotype topotypes, but from a population of Cnemaspis from of Cnemaspis ranwellai Wickrama­ ­singhe, 2006, as the Gammaduwa (35 km NNE of Kandy), well outside neotype of Gymnodactylus scalpensis Ferguson, 1877. the type locality defined in the original description Further, we fix the type locality of C. scalpensis as the of C. scalpensis (see discussion in Pethiyagoda, 2007). Gannoruwa Forest Reserve, near Peradeniya, Kandy Then, with no explanation for their change of heart, District, which lies within and is consistent with Wick­rama­singhe & Munindra­dasa (2007) selected the type locality defined by Ferguson (1877). The a neotype and three new ‘syntypes’ (sic) from neotype is entirely consistent with what is known Agarapatana, 50 km south (n.b.) of Kandy. These of the former name-bearing type. neotype and ‘new syntype’ designations have been Cnemaspis ranwellai Wick­rama­singhe, 2006 is shown to be invalid by Pethiyagoda (2007). now a junior objective synonym of Gymnodactylus The type locality of C. ranwellai is Gannoruwa, scalpensis Ferguson, 1877. within the suburbs of Kandy and therefore within the type locality of C. scalpensis. Further, C. ranwellai Cnemaspis tropidogaster (Boulenger, 1885) is entirely consistent with the original description of Gonatodes kandianus var. tropidogaster Boulenger, 1885 C. scalpensis, and is the only species of Cnemaspis in (Figures 19a–f; Tables 3, 6) the vicinity of Kandy to have 12–16 femoral pores, which character was listed for C. scalpensis in its Lectotype. BMNH 71.12.14.49 (male), 31.4 mm SVL, original description. “Ceylon” (= Sri Lanka), Presented by W. Thwaites. In view of the type specimens of C. scalpensis never having been traced or referred to in the 130 years that Diagnosis have elapsed since its description, we are certain that Cnemaspis tropidogaster differs from its peninsular these are lost. We searched for these specimens in Indian and Sri Lankan congeners by a combination collections that might contain material collected by of the following characters. Maximum SVL 31.4 Ferguson—NMSL, BMNH, ZSI, BNHS—but without mm; paired postmentals separated medially by a success. It is necessary and desirable that a neotype small scale; each postmental bounded by 3 scales be designated for C. scalpensis so as to clarify and including medial scale; ventrals, 133–144; dorsal scales heterogeneous; spine-like tubercles present on flanks; ventrals including pectoral, abdominal, precloacal and tail base area, carinate; 3 preanal and 4 or 5 femoral pores on each side; no enlarged tubercles on tail base; supralabials to angle of jaws, 7; subdigital lamellae on digit IV of pes, 18.

Description of lectotype An adult male, 31.4 mm SVL (for measurements, see Table 3). Head moderately long (HL 27.3% of SVL), moderately broad (HW 17.5% of SVL, HW 63.9% of HL), distinct from neck. Snout length moderately greater than eye diameter (ED 63.4% of ES); interorbital moderate (IO 9.0% of HL); pupil rounded. Granules on snout keeled, larger than those of occipital region. Rostral scale partially divided by a medial groove, in contact with first supralabial (Fig. 19a). Nares separated by two enlarged supranasals and a single internasal scale that extends towards snout tip (Fig. 19a). Nares rounded, dorsally orientated, not in contact or contact with first supralabial. Postnasals asymmetrical, 3 (left side) and 2 (right side), the lower one in broad contact Figure 18. Distribution of Cnemaspis scalpensis in south- with first supralabial (Fig. 19a). Mental subtriangular, western Sri Lanka. as wide as long, posteriolaterally in contact with

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two postmentals and medially with a single small molligodai: 5 preanal and 8 or 9 femoral pores, gular, hexagonal postmental scale; postmentals bordered pectoral and abdominal scales smooth, dorsal scales posteriorly by 5 smooth scales including medial scale homogeneous; median row of subcaudals enlarged, (Fig. 19b). Supralabials 7, infralabials 7, decreasing subequal. Cnemaspis samanalensis: gular, pectoral and in size towards gape. Ear opening oval, higher than abdominal scales tricarinate, subcaudals keeled. wide (TW 62.5% of TH). Scales on ventral surface Cnemaspis retigalensis: pectoral and abdominal scales of neck carinate. Twenty scales between anterior smooth, a single preanal pore, 3 or 4 femoral pores. margin of ear opening and posterior margin of eye. Cnemaspis punctata: 5–7 femoral pores; distinctive No conical enlarged granules laterally on neck. spotted dorsal coloration. Cnemaspis kumarasinghei: Body slender, elongate. Mid-dorsal granules pectoral and abdominal scales smooth. Cnemaspis mixed with large keeled tubercles (Fig. 19c); 6 spine- silvula, C. pulchra and C. pava: conical enlarged like tubercles on flank; abdominal scales imbricate, tubercles laterally on neck from ear to forelimb pointed, strongly carinate (Fig. 19d). Ventrals, 133, insertion. Cnemaspis clivicola: median row of scales around vent imbricate and carinate (Fig. 19e) subcaudals in median series not enlarged. Cnemaspis and base of tail imbricate and carinate; 3 preanal upendrai: some gulars tricarinate, precloacal and and 5 (left side) (Fig. 19e) and 4 (right side) femoral ventral scales on tail base region smooth, 15 spine- pores. like tubercles on flank, enlarged conical tubercles Forelimbs moderately long, slender (LAL 14.0% present on base of tail. Cnemaspis kallima and C. latha: SVL, UAL 11.7% SVL); hind limbs long (TBL 21.6% pectoral and abdominal scales smooth. Cnemaspis SVL, FEL 20.3% SVL); tibia long. Dorsal scales on amith: pectoral and abdominal scales smooth. both fore and hind limbs carinate; ventral scales on Cnemaspis beddomei and C. goaensis: ventrals upper and lower arm carinate. Scales on anterior smooth. Cnemaspis littoralis: dorsal scales homo­ surface and ventral surface of thigh, especially geneous. Cnemaspis mysoriensis: dorsal scales around knees, carinate. Granules on posterior surface homogeneous, 2 or 3 femoral pores. Cnemaspis of thigh conical. Scales on dorsal and ventral surfaces indraneildasii: dorsal scales of both limbs, and of shank carinate. Scales on manus and pes carinate. ventral, gular and caudal scales smooth. Cnemaspis Digits elongate, slender, all bearing slightly recurved heteropholis: paired postmentals separated by claws; subdigital lamellae entire, unnotched (Fig. three small scales, nostril not in contact with first 19f); subdigital lamellae on finger I, 11, finger II, supralabial, no post-cloacal spurs, ventral, gular 13, finger III, 14, finger IV, 16, finger V, 14; toe I, 9, and caudal scales smooth. Cnemaspis sisparensis toe II, 13, toe III, 16, toe IV, 18, toe V, 17; interdigital and C. wynadensis: spine-like tubercles absent on webbing absent; relative length of digits (fingers): IV flank. Cnemaspis boiei: ventrals smooth, no preanal (2.9 mm) ~ III (2.9 mm) > II (2.7 mm) > V (2.6 mm) > or femoral pores. Cnemaspis gracilis: ventrals smooth, I (2.2 mm); (toes) IV (4.3 mm) > V (3.6 mm) > III (3.5 2 preanal pores separated by 2 scales. Cnemaspis mm) > II (3.3 mm) > I (1.9 mm). indica: spine-like tubercles absent on flank. Cnemaspis Tail broken; tail base distinctly swollen. Enlarged ornata: no femoral pores, dorsal tubercles arranged tubercles absent on base of tail; a single post-cloacal in 15 longitudinal rows, ventrals smooth. Cnemaspis spur present on each side (Fig. 19e). wicksii: ventrals smooth. Cnemaspis andersonii: pectoral and abdominal scales smooth. Cnemaspis Coloration. In preservative: body dorsally dark jerdonii: no preanal pores. Cnemaspis nairi: spine- brown with unequal light-brown patches; venter like tubercles absent on flank, no femoral pores. brown; neck and flank laterally with light-brown Cnemaspis otai: spine-like tubercles absent on flank. markings. In life: Not known. Cnemaspis yercaudensis: spine-like tubercles absent on flank, ventrals smooth. Cnemaspis monticola: pectoral Comparisons. Cnemaspis tropidogaster differs and abdominal scales smooth. Cnemaspis australis: from each of its Sri Lankan and peninsular Indian scales of preanal area smooth; enlarged tubercles congeners, respectively, by the following opposing present on base of tail. Cnemaspis nilagirica: dorsal suites of characters: scales homogeneous. Cnemaspis kandiana and C. menikay: pectoral, abdominal and preanal area smooth. Cnemaspis Remarks podihuna: all ventral scales smooth; median row of In describing Gonatodes kandianus var. tropidogaster, subcaudals enlarged, hexagonal, subequal. Cnemaspis Boulenger (1885) had before him seven specimens scalpensis, C. gemunu and C. phillipsi: 12–16 femoral drawn from a vast area: Sri Lanka, Tinnevelly (Tamil pores. Cnemaspis alwisi: 9 femoral pores. Cnemaspis Nadu State, India), the Nilgiri Hills (Tamil Nadu

56 Zeylanica Re v i s i o n o f Cn e m a s p i s a b

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Figure 19. Cnemaspis tropidogaster: BMNH 71.12.14.49, holotype, 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 and ventral regions, respectively, of mid-body; f, preanal and hind-limb regions, showing arrangement of femoral and preanal pores; i, ventral aspect of digit III of right pes. Scale bars: 1 mm.

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State, India) and Wynaad (Kerala State, India). Cnemaspis podihuna Deraniyagala, 1944 Having compared these with C. kandiana, Boulenger Cnemaspis podihuna Deraniyagala, 1944 remarked, “The following specimens agree so well in (Figures 20a–h, 21a–h, 22; Tables 3, 6; Plate 5a, b) structure and their different modes of coloration with G. kandianus that I cannot consider them specifically Holotype. NMSL uncatalogued (in poor condition) distinct from the latter. However, as their ventral (male), 23.6 mm SVL, collected 8 km from sea at scales are keeled, they should be kept distinct from Lahugala in the Eastern Province, 06º50’N, 81º41’E, the typical form. They may be called the tropidogaster 7 April, 1944. form.” Boulenger (1885) overlooked, however, several Recent material. WHT 7334 (male), 23.3 mm SVL, key characters that serve to distinguish the species Maligawila near Okkampitiya, Moneragala District, he included in his series of syntypes, e.g., the Uva Province, 06º43’N, 81º17’E, 130 m, coll. S. homogeity/heterogeneity of dorsal scutellation; Batuwita, 30 April, 2006; NMSL 20061002, (male), 24.1 relative sizes of subcaudal scales and their carination; mm SVL; 20061003, (male), 24.7 mm SVL; 20061004, presence/absence of conical tubercles on side of neck (female), 21.8 mm SVL; Lahugala; NMSL RG 21, 22.8 and enlarged tubercles on tail base; and presence/ mm SVL, Kalkanna, Koslanda. absence of spine-like tubercles on flank. Based on these characters, we distinguish four Diagnosis species within his syntype series, three of which (C. Cnemaspis podihuna differs from its peninsular Indian australis, C. monticola, C. nilagirica) we here describe and Sri Lankan congeners by a combination of the as new. Cnemaspis tropidogaster differs from C. aus­ following characters. Maximum SVL 24.7 mm; tralis by lacking conical tubercles on side of neck (vs paired postmentals separated by a medial scale; each present, in C. australis); having smooth (vs keeled) postmental bounded by 3 or 4 scales including medial dorsal granules; having keeled (vs smooth) scales on scale; ventrals, 113–117; ventral scales across mid- preanal area; and lacking (vs possessing) enlarged body, 13–20; dorsal scales heterogeneous; 5 spine-like tubercles on base of tail. It differs from C. monticola tubercles present on flank; ventrals smooth; 3 or 4 by the absence of conical tubercles on side of neck preanal pores and 3–6 femoral pores on each side; (vs present in C. monticola); having smooth dorsal diastema scales, 5 or 7; subcaudals smooth, median granules (vs keeled); having keeled pectoral and row enlarged, subequal; supralabials to angle of abdominal scales (vs smooth); having the scales of jaws, 8; subdigital lamellae on digit IV of pes, 18 or the preanal area keeled (vs smooth); and enlarged 19; (in life) dorsum light-brown, with about six light- tubercles absent on base of tail (vs present). brown confluent blotches; nape with a distinct black Cnemaspis tropidogaster sensu stricto is known marking; venter light yellow. only from its lectotype (Boulenger, 1885, noted the presence of a female specimen from Sri Lanka in his Description of holotype series of syntypes, but this has subsequently been (Data for WHT 7334 within square brackets.) Adult lost or misplaced): we have been unable to locate male, 23.6 [23.3] mm SVL (for measurements, see a wild population of this species. Our survey of Sri Table 3). Head long (HL 24.5 [28.3]% of SVL), broad Lankan Cnemaspis is far from complete, however, and (HW 15.6 [18.0]% of SVL, HW 63.7 [63.6]% of HL), more species almost certainly await discovery. We distinct from neck. Snout length a little greater than are hopeful that a living population of this species eye diameter (ED 88.0 [71.4]% of ES; [interorbital will be discovered in the future. narrow (IO 4.5% of HL)]; pupil rounded. Granules We attempted to ascertain the identity of the on snout smooth; scales on occipital, interobital, geckos assigned to C. topidogaster by previous supercilium, region of ear, and dorsum and sides of authors, e.g., Deraniya­ ­gala (1953), Wickrama­ ­singhe neck granular, not keeled; those on chin, gular and (2006) and Wick­rama­singhe & Munindra­dasa (2007). throat granular, not keeled. Rostral scale partially Although these specimens are registered at NMSL divided by a medial groove, in contact with first (RG 08 and RG 09), L. Kariyawasam, the collection supralabial (Fig. 20a, 21a). Nares separated by two manager, was unable to locate them for us despite enlarged supranasals [a single internasal scale (Fig. an extensive search. 21a)]. Nares rounded, dorsally orientated; nostril not The ‘syntypes’ assigned to C. tropidogaster in contact on right side and in contact on left side with by Wick­rama­singhe & Munindra­dasa (2007) are first supralabial (Fig. 21c). [nostril not in contact on erroneous and invalid (see Pethiyagoda, 2007). left side (Fig. 21c)]. Two postnasals, the lower one in

58 Zeylanica Re v i s i o n o f Cn e m a s p i s

broad contact with first supralabial (Fig. 21c). Mental In life: dorsum light-brown, with about six subpentagonal, truncate posteriorly, wider than long, confluent light-brown blotches; nape with a distinct posteriolaterally in contact with two slightly enlarged black marking; venter light yellow (see Plate 5b). postmentals and medially with a single hexagonal postmental scale; postmentals bordered posteriorly Variation. Postmentals separated medially by a by 6 [5] smooth scales including medial scale (Fig. somewhat larger scale in NMSL 20061001. Ventrals, 20b, 21b). Supralabials 8, decreasing in size towards 113–117; 3 or 4 preanal pores and 3–6 femoral pores gape; 7 [6] infralabials, decreasing in size towards on each side; scales on diastema, 5 or 7; pores absent gape (Fig. 20c, 21c). [Ear opening oval, higher than in females. SVL range from 21.8 to 24.7 mm (6 ex.). wide, oblique (TW 50.0 [50.0%] of TH).] Body slender, relatively short (AG 48.3 [38.1]% Distribution and natural history. The species is SVL). Mid-dorsal granules homogeneous, isolated endemic to Sri Lanka, apparently being restricted and scattered (Fig. 20d, 21d); 5 [5] spine-like tubercles to the eastern coastal floodplain. According to on flank; abdominal scales smooth (Fig. 20e, 21e). Deraniyagala (1944), C. podihuna is essentially Ventrals, 118 [115]; scales across midbody, 20 [13]; arboreal and generally frequents the boles of trees scales around vent and base of tail smooth, imbricate; with buttress roots. The specimen collected from 4 preanal, 4 [6] (left side) (Fig. 20f, 21f) and 4 [5] Maligawila (WHT 7334, in the Eastern Province, (right side) femoral pores; diastema with enlarged altitude ~130 m, rainfall < 2,000 mm yr-1, in a 7 [5] smooth scales, separates preano-femoral row well-shaded monsoon-forest habitat) was found (Fig. 20f, 21f). on a large tree trunk densely covered by roots of Forelimbs moderately short, slender (LAL 13.5 a species of Ficus. When alarmed, the geckos took [12.8]% SVL, UAL 11.8 [12.0]% SVL); hind limbs refuge among these roots (their coloration offers relatively long (TBL 16.1 [14.5]% SVL, FEL 17.3 excellent camouflage against the tree bark). We found [20.6]% SVL); tibia short. Dorsal, lateral and ventral C. podihuna to be abundant wherever it occurred. scales of upper arm smooth, imbricate; lower arm Populations of this gecko are known from Lahugala dorsally with unkeeled granules, ventrally with National Park (the type locality), Maligawila near smooth imbricate scales. Thigh dorsally covered by Okkampitiya (Fig. 22), Mihintale, Kuruwekotha, smooth scales and unkeeled granules; rear edge of Kukulagoda and Kalakanna near Koslanda. thigh with unkeeled granules; front edge ventrally with smooth, imbricate scales. Scales on ventral Comparisons. Cnemaspis podihuna differs from each surface of shank, palm and sole, smooth. Digits of its Sri Lankan and peninsular Indian congeners, elongate, slender, all bearing slightly recurved respectively, by the following opposing suites of claws; subdigital lamellae entire, unnotched (Fig. characters: 21g); subdigital lamellae on finger I, 9[10, finger II, Cnemaspis tropidogaster: keeled ventrals and het­ 11[13], finger III, 13[14], finger IV, 11[15], finger V, ero­geneous dorsal scales. Cnemaspis kandiana and C. 12[14]; toe I, 10[11], toe II, 12[14], toe III, 13[16], toe menikay: gular scales keeled. Cnemaspis scalpensis, C. IV, 15[19], toe V, 13[17]; interdigital webbing absent; gemunu and C. phillipsii: 12–16 femoral pores. Cnemas­ relative length of digits (fingers): IV (2.0 mm) ~ II pis alwisi: 7–9 femoral pores. Cnemaspis molligodai: 5 (2.0 mm) > III (1.9 mm) > V (1.7 mm) > I (1.3 mm); preanal and 8 or 9 femoral pores. Cnemaspis samanal­ (toes) IV (2.6 mm) ~ III (2.6 mm) > II (2.4 mm) > V ensis: gular, pectoral and abdominal scales tricarinate, (2.3 mm) > I (1.2 mm). sub­caudals keeled. Cnemaspis retigalensis: dorsal Partially regenerated tail tapering, its length a scales heterogeneous gulars carinate, pectoral and ab­ little less than SVL (TAL 96.6% of SVL); tail base dom­inal scales smooth, a single preanal pore and 3 or distinctly swollen; scales on tail base granular. 4 femoral pores; median row of subcaudals enlarged, Dorsal scales on tail granular or imbricate, directed irregular. Cnemaspis punctata: 5–7 femoral pores; backwards. Tail with about 4–6 enlarged conical or distinctive (spotted) dorsal coloration. Cnemaspis oval granules forming whorls (Fig. 20g); a single kumarasinghei and C. latha: subcaudals on median row blunt post-cloacal spur present on each side (Fig. enlarged, irregular. Cnemaspis silvula, C. upendrai, C. 21h); subcaudals smooth, with a regular median pulchra, C. pava and C. clivicola: gular, subcaudal and series of, enlarged widened scales (Fig. 20h, 21h). ventral scales, and those on pectoral and abdominal, carinate. Cnemaspis kallima: subcaudals on distal Coloration. In preservative: dorsum olive, with dis­ half of tail carinate. Cnemaspis amith: gular scales tinct dorsal bands interrupted medially by mid-dors­ carinate; dorsal granules heterogeneous; subcaudals al line; tail banded; venter whitish (see Plate 5a). on median row enlarged, irregular.

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Figure 20. Cnemaspis podihuna: NMSL uncatalouged holotype, male, a, dorsal aspect of snout; b, ventral aspect of chin; c, lateral aspect of head showing supralabials and infralabials; d, e, dorsal and ventral scalation, respectively, of mid-body; f, preanal and left hind limb regions showing arrangement of femoral and preanal pores; g, lateral aspect of tail, showing whorls of conical enlarged tubercles (arrowheads indicate conical tubercles); h, arrangement of subcaudal scales. Scale bars: 1 mm. 60 Zeylanica Re v i s i o n o f Cn e m a s p i s a b

d c

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Figure 21. Cnemaspis podihuna: WHT 7334, male, a, dorsal aspect of snout; b, ventral aspect of chin; c, lateral aspect of head showing supralabials and infralabials; d, e, dorsal and ventral aspects, respectively, of mid-body; f, preanal and left hind limb regions showing arrangement of femoral and preanal pores; g, ventral aspect of digit II of right manus; h, arrangement of subcaudal scales. Scale bars: 1 mm.

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Cnemaspis beddomei: no femoral pores. Cnemaspis Remarks goaensis: gulars keeled. Cnemaspis littoralis: body Pethiyagoda (2007) showed that the designation of slender, median series of subcaudals enlarged, NMSL 20061001 as the neotype of Cnemaspis podihuna hexagonal, a black spot on the nape. Cnemaspis by Wick­rama­singhe & Munindra­dasa (2007) is mysoriensis: three rows of enlarged subcaudals, 2 erroneous and invalid. The latter authors rejected or 3 femoral pores. Cnemaspis indraneildasii and C. the NMSL holotype of C. podihuna on the grounds heteropholis: dorsal scales heterogeneous. Cnemaspis that it “differs in morphological detail to [the] type sisparensis and C. wynadensis: spine-like tubercles description”. In support of this statement, they stated absent on flank. Cnemaspis boiei: no preanal or that the putative holotype of C. podihuna possessed femoral pores. Cnemaspis gracilis: 2 preanal pores the following characters: separated by 2 scales, 4 femoral pores. Cnemaspis indica: spine-like tubercles absent on flank. Cnemaspis large snout to eye distance; postmentals separated ornata: no femoral pores, dorsal tubercles arranged by a small scale; nostrils are in contact with first in 15 longitudinal rows, a dark brown collar on supralabial; throat scales smooth; dorsal tubercles nape. Cnemaspis wicksii: subcaudals in median series granulated; absence of groups of carinated large scales in dorsal body; spine-like tubercles absent enlarged, irregular. Cnemaspis andersonii: gulars on flanks; ventral smooth and subimbricate; carinate, subcaudals in median series not enlarged; subcaudals extremely large; slightly large, absence dorsals heterogeneous. Cnemaspis jerdonii: no preanal of conical tubercles in ventrolateral margin of the pores. Cnemaspis nairi, C. otai and C. yercaudensis: tail; smooth and segmented scales intermixed with spine-like tubercles absent on flank. Cnemaspis subconicals in dorsal tail. monticola: subcaudals keeled. Cnemaspis australis: subcaudals pointed, strongly keeled. Cnemaspis They then pointed out that the the above nilagirica: pectoral and abdominal scales weakly characters were at variance with the illustration of keeled. C. podihuna provided in the original description by Deraniyagala (1944) in which “the nostrils are not in contact with the first supralabial and the ventrolateral margin of the tail contains conical tubercles.” Contact between nostril and first supralabial is extremely variable in Cnemaspis and not a reliable diagnostic character. Indeed, in the holotype itself, there is contact on the left side of the specimen while there is no contact on the right side (in keeping with convention Deraniyagala described bilateral characters only from the left side, which is also the side he delineated) (see Fig. 20c). This is therefore not a useful character in diagnosing the species. As for the ventrolateral margin of the tail, it does contain conical tubercles although with age these have become soft and less obvious (see Fig. 20g), apparently for which reason they were overlooked by Wick­rama­singhe & Munindra­dasa (2007). These authors also incorrectly noted the absence of spine- like tubercles on the flanks: spine-like tubercles are indeed present on the flanks though again, owing to the age and state of preservation of the specimen, these are not as obvious as in a fresh example. Wick­rama­singhe & Munindra­dasa (2007) went on to compare and contrast C. tropidogaster with C. podihuna and C. molligodai, on which discussion we do not comment because they had not examined the lectotype of C. tropidogaster and, having erroneously and unnecessarily designated four ‘syntypes’ Figure 22. Distribution of Cnemaspis podihuna in Sri Lanka; for it (Pethiya­goda, 2007), had evidently entirely the type locality is indicated by a solid symbol. misconceived the identity also of this species.

62 Zeylanica Re v i s i o n o f Cn e m a s p i s

Cnemaspis gemunu Bauer, de Silva, Comparisons. Cnemaspis gemunu differs from each Greenbaum & Jackman, 2007 of its Sri Lankan and peninsular Indian congeners, (Plate 5c, Table 6) respectively, by the following opposing suites of characters: Holotype. AMB 7495, (male), 34.0 mm SVL; Sri Cnemaspis tropidogaster: C. kandiana, C. menikay Lanka, Central Province, Nuwara Eliya District, and C. podihuna: presence of preanal pores. Cnemaspis Hakgala, Hakgala Botanical Gardens (06o55’30’’N, 80 salpensis: ventrals, 122–130. Cnemaspis alwisi: 9 o49’15’’E, 1660 m, coll. A. M. Bauer, A. de Silva & C. A. femoral pores. Cnemaspis molligodai: presence of Austin, 15 November, 2002 (not examined). preanal pores. Cnemaspis samanalensis: gular, pectoral and abdominal scales tricarinate, subcaudals keeled. Paratype. AMB 7507, (female), 2nd milepost from Cnemaspis retigalensis: dorsal scales heterogeneous, Borangamuwa, 25 km N Balangoda, Province of gulars carinate, a single preanal pore and 3 femoral Sabaragamuwa, Ratnapura District, Balangoda Rd., pores. Cnemaspis phillipsi: no enlarged tubercles Sri Lanka, 06o44’39’’ N, 80o42’28’’E, 1250 m, coll. A. M. on dorsal surface of tail. Cnemaspis punctata: 5–7 Bauer, A. de Silva & C. A. Austin, 17 November, 2002 femoral pores, distinctive spotted dorsal coloration. (not examined). Cnemaspis kumarasinghei, C. kallima and C. latha: presence of preanal pores. Cnemaspis silvula, C. Other material. WHT 7221, WHT 7347, WHT 7348, clivicola, C. pulchra, C. pava and C. upendrai: pectoral, Hakgala, near Hakgala Botanical Garden, Nuwara abdominal and subcaudal scales scales keeled. Eliya District, Central Province, Sri Lanka, 06o55’N, Cnemaspis amith: gular scales carinate; dorsal 80o49’E, 1830 m, coll. S. Batuwita, 02 October, 2006; granules heterogeneous; subcaudals on median row NMSL 2006.11.01, NMSL 20061102, NMSL 20061103, enlarged, irregular. NMSL 20061104, Agarapatana, Hatton, Sri Lanka, Cnemaspis beddomei: preanal pores 6–9, no 06º50’58.1’’N, 80º40’35.0’’E, 1524m, coll. L. J. Mendis femoral pores. Cnemaspis goaensis: preanal pores Wick­rama­singhe & D. A. I. Munindradasa, 17 2 or 3, gulars keeled. Cnemaspis littoralis: body September, 2006. slender. Cnemaspis mysoriensis: 2 or 3 femoral pores. Cnemaspis indraneildasii and C. heteropholis: dorsal Diagnosis scales heterogeneous. Cnemaspis jerdonii: 8 femoral Cnemaspis gemunu differs from its peninsular Indian pores. Cnemaspis indica, C. nairi, C. otai, C. sisparensis, and Sri Lankan congeners by a combination of the C. wynadensis and C. yercaudensis: spine-like tubercles following characters. Maximum SVL 34.0 mm; paired absent on flank. Cnemaspis boiei: lacks preanal and postmentals in contact or separated by a medial scale; femoral pores. Cnemaspis gracilis: 2 preanal pores each postmental bounded by 2 or 3 scales; ventrals, separated by 2 scales, 4 femoral pores. Cnemaspis 113–115; ventral scales across mid-body, 13–16; ornata: lacks femoral pores, dorsal tubercles arranged dorsal scales homogeneous; irregular row of spine- in 15 longitudinal rows. Cnemaspis wicksii and C. like tubercles present on flank; ventrals smooth; andersonii: gulars carinate. Cnemaspis monticola: no preanal pores, 11–14 femoral pores per side; gulars and caudals carinate. Cnemaspis australis: subcaudals smooth, median row enlarged, subequal; subcaudals pointed, strongly keeled. Cnemaspis supralabials to angle of jaws, 8 or 10; subdigital nilagirica: pectoral and abdominal scales weakly lamellae on digit IV of pes, 19; (in life) dorsal pattern keeled. of dark chevrons, with a pale, broad, discontinuous vertebral stripe from nape on to tail; venter of male Remarks yellow in life, with a white throat (see Plate 5c). Wick­rama­singhe & Munindra­dasa (2007) designated a neotype and three new ‘syntypes’ [sic] for C. Distribution and natural history notes. Endemic scalpensis, implicitly fixing the type locality of this to the central highlands of Sri Lanka, C. gemunu species as Agarapatana. While this designation and is sympatric with following lizards: Ceratophora type-locality are erroneous and invalid in terms of the stoddartii, Cophotis ceylanica, Lankascincus taprobanensis. Insternational Code of Zoological Nomenclature (see This gecko was often noticed on moss-covered tree also Remarks under C. scalpensis, above, and reasons trunks and in crevices in the barks of large trees given in Pethiyagoda, 2007), we note in passing that in montane rainforests. Populations are known at the ‘neotype’ and ‘syntypes’ incorrectly designated present from Hakgala, Balangoda and Agarapatana by Wick­rama­singhe & Munindradasa­ (2007) for C. near Hatton in the Nuwara Eliya District. scalpensis are in fact examples of C. gemunu.

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Cnemaspis alwisi Wickramasinghe patric lizards included Cnemaspis retigalensis­ , Calo­ & Munindradasa, 2007 dactylodes illingworthorum, Geckollea triedra, G. yakhuna, (Figure 23; Plate 6a, b; Table 6) Hemidactylus depressus, H. maculatus hunae, Lanka­ scincus taylori, L. fallax, Eutropis macularius macularius, Holotype. NMSL 2004.9.1, (male), 39.92 mm SVL, Calotes liolepis and Otocryptis nigristigma. Dolukanda, Kurunegala, Sri Lanka, (07º37’07.8’’N, 80º24’50.3’’E, 152m), coll. L. J. Mendis Wickrama­ ­ Comparisons. Cnemaspis alwisi differs from each singhe & D. A. I. Munindra­dasa, 23 December, of its Sri Lankan and peninsular Indian congeners, 2003. respectively, by the following opposing suites of characters: Paratypes. NMSL 2004.9.2, (female), 37.84 mm SVL; Cnemaspis tropidogaster: keeled ventrals and NMSL 2004.9.3, (male), 32.99 mm, same collection heterogeneous dorsal scales. Cnemaspis kandiana, data as holotype. C. menikay, C. podihuna and C. molligodai: presence of preanal pores. Cnemaspis gemunu, C. scalpensis Other material. WHT 7336, 7337, 7338, Warakawehera, and C. phillipsii: 12–16 femoral pores. Cnemaspis near Ridigama, Kurunegala District, Northwestern samanalensis: gular, pectoral and abdominal scales Province of Sri Lanka, 07o30’N, 80o29’E, 100 m, coll. S. tricarinate, subcaudals keeled. Cnemaspis retigalensis: Batuwita & S. Ranasinghe, 27 February, 2006; WHT dorsal scales heterogeneous, gulars carinate, a single 7343, WHT 7345, WHT 7346, WHT 7344, Waduwawa, preanal pore and 3 femoral pores; median row of near Allauwa, Kegalle District, Sabaragamuwa subcaudals enlarged, irregular. Cnemaspis punctata: Province of Sri Lanka, 07o15’N, 80o16’E, 300m, 5–7 femoral pores, distinct spotted dorsal coloration. coll. S. Batuwita, 19 December, 2006; WHT 5918 Cnemaspis clivicola, C. pulchra, C. pava, C. silvula and C. Kumaradola Estate, Moneragala District, Uva upendrai: gulars, pectoral, abdominal and subcaudal Province of Sri Lanka, 06o53’N, 81o22’E, 305 m, coll. M. scales keeled. Cnemaspis kumarasinghei, C. kallima and Meegaskumbura & M. M. Bahir, 04 December, 2003; C. latha: presence of preanal pores. Cnemaspis amith: WHT 6518, 6519, same location data as above, coll. M. M. Bahir & S.V.Nanayakkara, 30 April, 1998.

Diagnosis Cnemaspis alwisi differs from its peninsular Indian and Sri Lankan congeners by a combination of the following characters. Maximum SVL 39.9 mm; paired postmentals separated by a medial scale; each postmental bounded by 2 scales; ventrals, 135–152; ventral scales across mid-body, 27–31; dorsal scales homogeneous; spine-like tubercles present on flank; ventrals smooth; no preanal pores, 7–9 femoral pores per side; subcaudals smooth, median row enlarged, subequal; supralabials to angle of jaws, 8–10; subdigital lamellae on digit IV of pes, 17–21; (in life) dorsum light-brown, with about six pale confluent blotches; nape with a distinct black marking.

Distribution and natural history notes. Cnemaspis alwisi was observed mostly on rock faces within caves and overhangs. It is a fast-moving species, tak­ing cover in narrow crevices when alarmed. It was observed to be a communal egg layer (Plate 6b). Cnemaspis alwisi was recorded from forests in the intermediate climatic zone of Sri Lanka, at Waraka­ wehera and Dolukanda (Kurunegala District), Al­ lauwa (Kegalle District), Kumaradola rubber estate and Maragalawatta (Moneragala District), and Figure 23. Distribution of Cnemaspis alwisi in Sri Lanka; the Ritigala (Anuradhapura District) (Fig. 23). Sym­ type locality is indicated by a solid symbol.

64 Zeylanica Re v i s i o n o f Cn e m a s p i s

gular scales carinate; subcaudals on median row been unable to deduce what this statistic means; did enlarged, irregular. the authors actually examine more than 90% of the Cnemaspis beddomei: ventrals keeled, preanal pores living individuals of C. ranwellai in the type locality? 6–9, no femoral pores. Cnemaspis goaensis: preanal How exhaustively could they have sampled, to have pores 2 or 3, gulars keeled. Cnemaspis littoralis: arrived at this satistic? In any event, there is no body slender, a black spot on the nape. Cnemaspis mention in the Methods declared in either Wick­rama­ mysoriensis: three rows of enlarged subcaudals, singhe (2006) or Wick­rama­singhe & Munindra­dasa 2 or 3 femoral pores. Cnemaspis indraneildasii (2007) to support this claim. and C. heteropholis: heterogeneous dorsal scales. Cnemaspis boiei: lacks preanal or femoral pores. Cnemaspis kumarasinghei Wickramasinghe Cnemaspis gracilis: 2 preanal pores separated by 2 & Munindradasa, 2007 scales, 4 femoral pores. Cnemaspis indica: ventrals, (Figure 24; Plate 6c; Table 6) 95 and postmentals in contact. Cnemaspis ornata: lacks femoral pores, dorsal tubercles arranged Holotype. NMSL 20061301, (male), 31.61 mm SVL, in 15 longitudinal rows. Cnemaspis jerdonii: lacks Maragala, Monaragala, Sri Lanka, 06º52’55.8’’N, caudal tubercles. Cnemaspis nairi, C. otai and C. 81º23’09.7’’E, 910 m, coll. L. J. Mendis Wickrama­ ­ yercaudensis: scales on the dorsal surface of limbs singhe & R. H. S. S. Fernando, 27 April, 2004. keeled. Cnemaspis sisparensis: heterogeneous dorsal scalation. Cnemaspis wynadensis: subcaudals Paratypes. NMSL 20061302, (male), 28.96 mm SVL, enlarged, subequal; no preanal pores, 5 femoral same collection data as holotype. pores, postmentals widely separated, 4–6 femoral pores and no cloacal spurs. Cnemaspis wicksii and Other material. WHT 7330, 7331, 7332, Sandagala C. andersonii: presence of preanal pores. Cnemaspis near Tissamaharama, Hambantota District, Southern monticola: subcaudals keeled, spine-like tubercles Sri Lanka, 06o20’30’’N, 80o16’ E, 100 m, coll. S. present on flank. Cnemaspis australis: subcaudals Batuwita & J. R. De Lile, 29 July, 2005; WHT 7211, pointed, strongly keeled scales. Cnemaspis nilagirica: Kariwilakotuwa, Buttala, Moneragala District, pectoral and abdominal scales weakly keeled. Uva Province of Sri Lanka, 06o40’N, 81o15’E, 160 m, coll. S. Batuwita, 05 November, 2005; WHT 7335, Remarks Kariwilakotuwa, Buttala, Moneragala District, Uva In their Remarks on C. alwisi, Wick­rama­singhe & Province of Sri Lanka, 06o40’N, 81o15’E, 160 m, coll. Munindra­dasa (2007) stated as follows: “C. alwisi S. Batuwita, 05 November, 2005; WHT 7024, 7025, sp. nov. is congener [sic] with C. ranwellai and C. 7026, Mihintale, Anuradhapura District, 08o21’N, scalpensis from morphological characters. However, 80o30’E, 150 m, coll. M. M. Bahir & D. Gabadage, 10 C. alwisi can easily be distinguished from both by October, 1995. having two internarsals [sic] and low count of lamella in forth [sic] finger and toe, and femoral pores, and Diagnosis from C. ranwellai by the separated postmental [sic] Cnemaspis kumarasinghei differs from its peninsular (more than 90% of C. ranwellai population in the type Indian and Sri Lankan congeners by a combination locality shows contacted postmental [sic])...”. of the following characters. Maximum SVL 31.61 mm; Even noting that the species they refer to as C. paired postmentals separated by a medial scale; each ranwellai is C. scalpensis, and the one they call C. postmental bounded by 4 scales; ventrals, 121–133; scalpensis is in fact C. gemunu, C. alwisi is not usefully ventral scales across mid-body, 17–21; dorsal scales diagnosed from C. scalpensis and C. gemunu by these homogeneous; 8 spine-like or blunt tubercles on characters. The presence of internasal scale/s is flanks; ventrals smooth; 2 preanal pores, 3–5 femoral variable within these species, for which subdigital- pores per side; subcaudals smooth, the median row lamellae meristics too, overlap (see descriptions, enlarged, irregular; supralabials to angle of jaws, 7 above). Interestingly, Wick­rama­singhe & Munindra­ or 8; subdigital lamellae on digit IV of pes,16–18; (in dasa (2007) overlooked the presence of spine-like life) dorsum olive green, with about five black bands; tubercles on flank of C. alwisi, a useful character in nape with a black spot, occipital region with two dark the identification of this species. markings; venter pinkish white. We have grappled with the statement by Wick­ rama­singhe & Munindra­dasa (2007) that “more Distribution and natural history notes. We observed than 90% of C. ranwellai population in the type C. kumarasinghei to be most active around 1600–1800 locality shows contacted postmental”, but have h, with specimens being sighted both in crevices

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and on rock faces. The species inhabits dense forest subequal. Cnemaspis samanalensis: gular, pectoral in Sri Lanka’s dry zone (rainfall < 2,000 mm yr-1), and abdominal scales tricarinate, subcaudals keeled. at altitudes ranging from 100–900 m (Fig. 24). A Cnemaspis retigalensis: gulars carinate, a single female at Sandagala measuring 30.0 mm SVL laid preanal pore, 3 femoral pores. Cnemaspis punctata: 5–7 two hard-shelled eggs 6.0×5.0 mm, which hatched femoral pores; distinctive spotted dorsal coloration. after 53 days. The hatchlings were 11.9 and 12.2 mm Cnemaspis silvula, C. pulchra, C. pava, C. upendrai and SVL. Cnemaspis kumarasinghei was observed to occur C. clivicola: gular, pectoral and abdominal scales in sympatry with the following lizards: Calodactylodes keeled. Cnemaspis kallima: subcaudals on distal half illingworthorum, Hemidactylus depressus, H. frenatus, H. of tail carinate. Cnemaspis latha: ventrals 111; lacks leschenaultii, H. maculatus hunae and H. triedrus lankae, distinct cross-bars on dorsum. Cnemaspis amith: gular Lankascincus fallax, Sitana ponticeriana, Otocryptis scales carinate. nigristigma. Cnemaspis beddomei: preanal pores 6–9, no femoral pores. Cnemaspis goaensis: gulars keeled. Cnemaspis Comparisons. Cnemaspis kumarasinghei differs littoralis: median series of subcaudals enlarged, from each of its Sri Lankan and peninsular Indian subcaudals hexagonal. Cnemaspis mysoriensis: 3 congeners, respectively, by the following opposing rows of enlarged subcaudals, 2 or 3 femoral pores. suites of characters: Cnemaspis indraneildasii: dorsal scales heterogeneous. Cnemaspis tropidogaster: dorsal scales hetero­ Cnemaspis heteropholis: dorsal scales heterogeneous. geneous, with pointed, keeled tubercles; ventrals Cnemaspis sisparensis and C. wynadensis: spine-like keeled. Cnemaspis kandiana and C. menikay: gular tubercles absent on flank. Cnemaspis boiei: no preanal scales carinate. Cnemaspis podihuna: a median or femoral pores. Cnemaspis gracilis: 2 preanal pores series of enlarged subcaudals and 6 femoral pores. separated from each other by 2 scales, 4 femoral Cnemaspis scalpensis, C. gemunu and C. phillipsii: pores. Cnemaspis indica: spine-like tubercles absent 12–16 femoral pores. Cnemaspis alwisi: 9 femoral on flank. Cnemaspis ornata: no femoral pores, dorsal pores. Cnemaspis molligodai: 5 preanal and 8 or 9 tubercles arranged in 15 longitudinal rows. Cnemaspis femoral pores; median row of subcaudals enlarged, wicksii: scales on dorsal surface of thigh carinate, 4 preanal pores. Cnemaspis andersonii: gulars carinate. Cnemaspis jerdonii: no preanal pores. Cnemaspis nairi, C. otai and C. yercaudensis: spine-like tubercles absent on flank. Cnemaspis monticola: subcaudals keeled. Cnemaspis australis: subcaudals keeled. Cnemaspis nilagirica: pectoral and abdominal scales weakly keeled; subcaudals on median row enlarged, subequal.

Remarks In their Remarks under C. kumarasinghei, Wickrama­ singhe & Munindra­dasa (2007) distinguished this species “from C. kandiana by [its] having smooth gula [sic] scales and, from C. retigalensis by having preanal scales, and also from morphometric analysis.” We have pored over this perplexing statement with bewilderment and can only conclude that by “by having preanal scales” these authors intended to convey that its preanal scales are pored (stated as 2–3 pored scales in their Diagnosis). However, they overlooked the fact that the holotype and three paratypes of C. retigalensis each have a single preanal pore (see Fig. 25a): the presence/absence of preanal pores therefore does not serve to separate C. kumarasinghei from C. retigalensis. Additionally, they also overlooked the presence of spine-like Figure 24. Distribution of Cnemaspis kumarasinghei in Sri tubercles on flank in both Cnemaspis retigalensis and Lanka; the type locality is indicated by a solid symbol. C. kumarasinghei.

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Cnemaspis molligodai Wickramasinghe granules heterogeneous; subcaudals on median row & Munindradasa, 2007 enlarged, irregular. (Plate 7a; Table 6) Cnemaspis beddomei: femoral pores absent. Cnemaspis goaensis: gulars keeled. Cnemaspis littoralis: Holotype. NMSL 20061401, (male), 26.62 mm SVL, a black spot on nape. Cnemaspis mysoriensis: three Waratalgoda, Rathnapura, Sri Lanka, (06º32’19.4’’N, rows of enlarged subcaudals, 2 or 3 femoral pores. 80º18’27.4’’E, 387 m), coll. L. J. Mendis Wickrama­ ­ Cnemaspis indraneildasii, C. heteropholis: dorsal scales singhe, 10 April, 2000. heterogeneous. Cnemaspis sisparensis, C. nairi, C. otai, C. yercaudensis, C. indica and C. wynadensis: spine-like Paratypes. NMSL 20061402, (male), 27.77 mm SVL; tubercles absent on flank. Cnemaspis boiei: no preanal NMSL 20061403, (male), 25.82 mm SVL; NMSL or femoral pores. Cnemaspis gracilis: 2 preanal pores 20061404, (male), 27.21 mm SVL; NMSL 20061405, separated by 2 scales, 4 femoral pores. Cnemaspis (female), 27.11 mm SVL, same collection data as ornata: no femoral pores, dorsal tubercles arranged holotype. in 15 longitudinal rows, a dark brown collar on nape. Cnemaspis wicksii and C. andersonii: gulars carinate, Diagnosis dorsals heterogeneous. Cnemaspis jerdonii: preanal Cnemaspis molligodai differs from its peninsular pores absent. Cnemaspis monticola: subcaudals keeled. Indian and Sri Lankan congeners by a combination Cnemaspis australis: subcaudals pointed, strongly of the following characters. Maximum SVL 29.0 keeled. Cnemaspis nilagirica: pectoral and abdominal mm; paired postmentals in contact or separated scales weakly keeled. by a medial scale/s; each postmental bounded by 2 scales; ventrals, 129–134; ventral scales across Remarks midbody, 15–19; dorsal scales homogeneous; spine- In their Remarks section under C. molligodai, Wick­ like tubercles present on flank; gular, pectoral and rama­singhe & Munindra­dasa (2007) state that this abdominal scales smooth; 5 preanal pores, 8 or 9 species “is congener [sic] with C. podihuna from femoral pores per side; subcaudals smooth, those morphological characters. However, C. molligodai on median row enlarged, subequal; supralabials to can easily be distinguished from C. podihuna by angle of jaws, 8–10; subdigital lamellae on digit IV having narsal [sic] connected to [sic] first supralabial, of pes, 19–23; caudal tubercles present. intraorbital [sic] count, preannal [sic] pores count, dorsal tubercle count and snout to eye distance, Comparisons. Cnemaspis molligodai differs from each and also from morphometric analysis…”. It is not of its Sri Lankan and peninsular Indian congeners, clear what these authors intended by “narsal”, even respectively, by the following opposing suites of though we accept this to mean “nasal” (i.e., in the characters: context of a substantive use of the adjective ‘nasal’ Cnemaspis tropidogaster: ventrals keeled, dorsal adopted by by convention in reptile taxonomy). We scales heterogeneous. Cnemaspis kandiana and C. therefore conclude that what Wick­rama­singhe & menikay: gular scales keeled. Cnemaspis podihuna: Munindradasa (2007) mean by “narsal” is the naris 3 or 4 preanal pores, 3–6 femoral pores, 113–117 or nostril opening, which by convention is referred ventrals. Cnemaspis scalpensis, C. gemunu and C. to as ‘nostril’ in gecko taxonomy: e.g., Bauer et al. phillipsii: 12–16 femoral pores, no preanal pores. (2007). Contact between the nostril and the first Cnemaspis alwisi: absence of preanal pores. Cnemaspis supralabial, however, is extremely variable within samanalensis: gular, pectoral and abdominal scales species of Cnemaspis: indeed, while we found this tricarinate, subcaudals keeled. Cnemaspis retigalensis: to be variable within all the species we examined, dorsal scales heterogeneous, gulars carinate, a single we nevertheless included this character-state in preanal pore, 3 femoral pores, subcaudals on median our descriptions—but not our diagnoses—only row enlarged, irregular. Cnemaspis punctata: 5–7 for the sake of completeness. In this respect, the femoral pores; absence of preanal pores; distinctive other characters listed by Wick­rama­singhe & (spotted) dorsal coloration. Cnemaspis kumarasinghei Munindradasa (2007) for distinguishing C. molligodai and C. latha: subcaudals on median row enlarged, from C. podihuna too, are variable and not useful in irregular. Cnemaspis silvula, C. pulchra, C. pava, C. facilitating consistent differential diagnosis. upendrai and C. clivicola: gular, subcaudal, ventral, In our opinion, Cnemaspis podihuna can most pectoral and abdominal scales carinate. Cnemaspis reliably be separated from C. molligodai by the kallima: subcaudals on distal half of tail carinate. following characters: 3 or 4 preanal pores (vs 5 in Cnemaspis amith: gular scales carinate; dorsal C. molligodai), 3–6 femoral pores (vs 8 or 9 in C.

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molligodai) and 113–117 ventral scales (vs 129–134 Cnemaspis kumarasinghei, C. kallima and C. latha: in C. molligodai). Wick­rama­singhe & Munindradasa gular scales smooth. Cnemaspis silvula, C. pulchra, (2007) also overlooked the presence of spine-like C. pava, C. clivicola and C. upendrai: pectoral and tubercles on flank—a useful diagnostic character—in abdominal scales keeled. Cnemaspis menikay: dorsal Cnemaspis molligodai. granules conical. Cnemaspis amith: dorsal granules heterogeneous and unkeeled. Cnemaspis beddomei: preanal pores 6–9, femoral Cnemaspis retigalensis Wickramasinghe pores absent. Cnemaspis goaensis: 3, preanal pores. & Munindradasa, 2007 Cnemaspis littoralis: subcaudals on median row (Figure 25a; Plate 7b; Table 6) enlarged, hexagonal. Cnemaspis mysoriensis: dorsal scales homogeneous, three rows of enlarged Holotype. NMSL 20061201, (male), 28.76 mm SVL, subcaudals. Cnemaspis indraneildasii and C. heteropholis: Weweltenna, Ritigala, Sri Lanka, 08º06’40.3’’N, gular scales smooth. Cnemaspis sisparensis, C. indica, 80º39’31.4’’E, 710m, coll. L. J. Mendis Wick­rama­singhe C. nairi, C. otai, C. yercaudensis and C. wynadensis: & D. A. I. Munindra­dasa, 30 October, 2005. spine-like tubercles absent on flank. Cnemaspis boiei: no preanal or femoral pores. Cnemaspis gracilis: Paratypes. NMSL 20061202, (female), 30.87 mm 2 preanal pores separated by 2 scales. Cnemaspis SVL; NMSL 20061203, (female), 26.56 mm; NMSL ornata: no femoral pores, dorsal tubercles arranged 20061204, (male), 27.67 mm SVL; same locality data as holotype, coll. L. J. Mendis Wickramasinghe & Roshan Rodrigo, 27 August, 2006. a Diagnosis Cnemaspis retigalensis differs from its peninsular Indian and Sri Lankan congeners by a combination of the following characters. Maximum SVL 30.8 mm; paired postmentals in contact or separated by a medial scale; each postmental bounded by 2 or 3 scales; ventrals, 121–128; ventral scales across midbody, 16–20; dorsal scales heterogeneous; spine- like tubercles present on flank; gular scales carinate, other ventrals smooth; a single preanal pore, 3 or 4 femoral pores (Fig. 25a); subcaudals smooth, median row enlarged, irregular; supralabials to angle of jaws, 7 or 8; subdigital lamellae on digit IV of pes, 16–20; caudal tubercles present. b Comparisons. Cnemaspis retigalensis differs from each of its Sri Lankan and peninsular Indian congeners, respectively, by the following opposing suites of characters: Cnemaspis kandiana: 2–4 preanal pores. Cnemaspis tropidogaster: gular, pectoral and abdominal scales keeled. Cnemaspis podihuna: subcaudals on median row enlarged, subequal. Cnemaspis scalpensis, C. gemunu and C. phillipsii: 12–16 femoral pores. Cnemaspis alwisi: 9 femoral pores. Cnemaspis molligodai: 5 preanal and 8 or 9 femoral pores, gular, pectoral and abdominal scales smooth, Figure 25. a, Cnemaspis retigalensis Wickramasinghe and dorsal scales homogeneous; subcaudals on median Munindradasa, 2007: NMSL 2006.12.01, holotype, male, row enlarged, subequal. Cnemaspis samanalensis: preanal region and right hind limb regions, showing gular, pectoral and abdominal scales tricarinate, arrangement of femoral and preanal pores; b, Cnemaspis samanalensis Wickramasinghe and Munindradasa, 2007: subcaudals keeled. Cnemaspis punctata: 5–7 femoral NMSL 2006.15.01, holotype, male, arrangement of tricarinate pores and a distinct spotted dorsal colour pattern. mid-ventral scales. Scale bars: 1 mm.

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in 15 longitudinal rows, a dark-brown collar on Cnemaspis tropidogaster, C. silvula, C. pulchra, nape. Cnemaspis wicksii: 4 preanal pores. Cnemaspis C. pava, C. upendrai and C. clivicola, pectoral and andersonii: 3 preanal pores. Cnemaspis jerdonii: no abdominal scales unicarinate. Cnemaspis kandiana preanal pores. Cnemaspis monticola: subcaudals and C. menikay, pectoral and abdominal scales carinate. Cnemaspis australis: subcaudals pointed, smooth. Cnemaspis podihuna, gular, pectoral and strongly keeled. Cnemaspis nilagirica: dorsal scales abdominal scales smooth; subcaudals on median row homogeneous; pectoral and abdominal scales weakly enlarged, subequal. Cnemaspis scalpensis, C. gemunu keeled. and C. phillipsii, 12–16 femoral pores. Cnemaspis alwisi, 9 femoral pores. Cnemaspis molligodai, gular, Remarks pectoral and abdominal scales smooth, dorsal scales (See Remarks under account of Cnemaspis kumara­ homogeneous; subcaudals on median row enlarged, singhei.) irregular. Cnemaspis retigalensis, pectoral and abdominal scales smooth; a single preanal pore; 3 or 4 femoral pores. Cnemaspis punctata, 5–7 femoral pores; Cnemaspis samanalensis Wickramasinghe distinctive spotted dorsal coloration. Cnemaspis & Munindradasa, 2007 kumarasinghei, C. kallima and C. latha, pectoral and (Figure 25b; Plate 7c; Table 6) abdominal scales smooth. Cnemaspis amith: pectoral and abdominal scales smooth. Holotype. NMSL 20061501, (male), 36.91 mm SVL, Cnemaspis beddomei, femoral pores absent. Samanala upper region (Adam’s Peak), Ratnapura, Cnemaspis goaensis, ventrals smooth. Cnemaspis Sri Lanka, (06º45’47.1’’N, 80º29’30.1’’E, 1430 m, coll. L. littoralis, body slender, dorsal scales homogeneous, J. Mendis Wickramasinghe, Mahesh Chathuranga, D. subcaudals on median row enlarged, hexagonal. A. I. Munindradasa, 11 February, 2005. Cnemaspis mysoriensis, dorsal scales homogeneous, three rows of enlarged subcaudals. Cnemaspis Paratypes. NMSL 20061502, (male), 32.48 mm SVL; indraneildasii and C. heteropholis, gular, ventral, and NMSL 20061503, (male), 34.37 mm; NMSL 20061504, caudal scales smooth. Cnemaspis indica, C. nairi, C. (female), 36.53 mm SVL; NMSL 20061505, (female), otai, C. yercaudensis, C. sisparensis and C. wynadensis, 36.18 mm SVL; same collection data as holotype. spine-like tubercles absent on flank. Cnemaspis boiei, ventrals smooth, no preanal or femoral pores. Other material. WHT 0691, Eratne, Adam’s Peak, Cnemaspis gracilis, ventrals smooth. Cnemaspis Ratnapura, Sabaragamuwa Province of Sri Lanka, ornata, no femoral pores, dorsal tubercles arranged 06º47’N, 80º23’E, 460m, coll. D. Gabadage, 29 January, in 15 longitudinal rows. Cnemaspis wicksii and C. 1994. andersonii, pectoral and abdominal scales smooth. Cnemaspis jerdonii, no preanal pores. Cnemaspis Diagnosis monticola, pectoral and abdominal scales smooth. Cnemaspis samanalensis differs from its peninsular Cnemaspis australis, scales on preanal region smooth. Indian and Sri Lankan congeners by a combination Cnemaspis nilagirica: subcaudals on median row of the following characters. Maximum SVL 37.5 enlarged, subequal. mm; paired postmentals in contact or separated by a medial scale; each postmental bounded by 3 scales; Remarks ventrals, 128–144; ventral scales across midbody, Wickramasinghe & Munindradasa (2007) had 22–26; dorsal scales heterogeneous, keeled; spine- misidentified­ C. tropidogaster (the type material like tubercles present on flank; gular, pectoral and of which they did not examine and which they abdominal scales tricarinate (Fig. 25b); 3 preanal evidently imagined did not exist), for which species pores, 3 or 5 femoral pores per side; subcaudals they erroneously designated four new ‘syntypes’ keeled, median row enlarged, irregular; supralabials from south-western Sri Lanka. These, however, to angle of jaws, 8 or 10; subdigital lamellae on digit are specimens of C. silvula, one of the species we IV of pes,19; caudal tubercles present. describe here as new. Cnemaspis tropidogaster sensu stricto is distinguished from C. silvula by the absence Comparisons. Cnemaspis samanalensis differs of enlarged tubercles on lateral surface of neck and from each of its Sri Lankan and peninsular Indian base of tail (present in C. silvula); and the presence congeners, respectively, by the following opposing of smooth mid-dorsal granules (vs keeled granules suites of characters: in C. silvula). Wickramasinghe & Munindradasa

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(2007) also failed to observe that C. samanalensis enlarged postmentals and medially with a single is distinguished from both C. tropidogaster and C. quadrate postmental scale; postmentals bordered silvula by having the gular, chest and ventral scales posteriorly by 5 smooth scales including medial scale tricarinate (see Fig. 25a) (vs unicarinate in C. tropido­ (Fig. 26b). Supralabials 9, infralabials 8, decreasing gaster and C. silvula). in size towards gape (Fig. 26c). Ear opening oval, higher than wide, oblique (TW 60.0% of TH). Scales Cnemaspis phillipsi new species on ventral surface of neck smooth; 2 or 3 enlarged (Figures 26a–h, 27; Tables 3, 6; Plate 8a–c) tubercles laterally on neck. Body robust, relatively long (AG 44.4% SVL). Holotype. WHT 7248 (male), 33.5 mm SVL, Mid-dorsal granules homogeneous, isolated and Gammaduwa Estate, Gammaduwa near Rattota, scattered (Fig. 26d); 5 spine-like tubercles on flank; Matale District, 07º34’N, 80º42’ E, 760 m, coll. S. abdominal scales smooth (Fig. 26e). Ventrals, 133; Batuwita & P. Wickramathilaka, 23 March 2007. ventral scales across midbody, 25; scales around vent and base of tail smooth, imbricate. Preanal pores Paratypes. WHT 7236 (male), 29.2 mm SVL, WHT absent; 16 femoral pores on each side (Fig. 26f). 7237, 7238 (2 females), 36.6 mm and 33.3 SVL, same Forelimbs moderately long, slender (LAL 13.7% collection data as holotype. SVL, UAL 16.4% SVL); hind limbs moderately long (TBL 17.6% SVL, FEL 19.4% SVL); tibia short. Dorsal Diagnosis scales on both fore and hind limbs carinate; ventral Cnemaspis phillipsi differs from its peninsular Indian scales on upper arm narrow, pointed, smooth; on and Sri Lankan congeners by a combination of the lower arm smooth. Scales on anterior and ventral sur­ following characters. Maximum SVL 36.6 mm; paired faces of thigh and shank smooth; those on palm and postmentals in contact or separated by a medial sole smooth, rounded. Digits elongate, slender, all scale; each postmental bounded by 2 scales including bearing slightly recurved claws; subdigital lamellae medial scale; ventrals, 129–141; ventral scales across entire, unnotched; subdigital lamellae on finger I, midbody, 18–25; dorsal scales homogeneous; spine- 11, finger II, 12, finger III, 15, finger IV, 17, finger V, like tubercles present on flank; ventrals smooth; no 13; toe I, 9, toe II, 15, toe III, 18, toe IV, 17, toe V, 18; preanal pores; 15 or 16 femoral pores on each side; interdigital webbing absent; relative length of digits subcaudals smooth, median row enlarged, subequal; (fingers): IV (3.2 mm) > III (3.0 mm) > V (2.3 mm) > supralabials to angle of jaws, 8 or 9; subdigital II (2.6 mm) > I (1.8 mm); (toes) IV (3.6 mm) > III (3.2 lamellae on digit IV of pes 17–19; no caudal tubercles; mm) > V (2.9 mm) > II (2.7 mm) > I (1.9 mm). (in life) dorsum dusky brown, with about eight Tail tapering, its length a little greater than SVL narrowly separated rusty-brown blotches; nape with (TAL 101.1% of SVL); tail base distinctly swollen. a distinctive broad, long, black marking. Enlarged, conical tubercles absent on base of tail. Dorsal scales on tail unkeeled, granular, directed Description of holotype backwards; a small conical post-cloacal spur present An adult male, 33.5 mm SVL (for measurements, see on each side (Fig. 26f); subcaudals on median row Table 3). Head relatively long (HL 26.5% of SVL), smooth, enlarged, subequal (Fig. 26g ). moderately broad (HW 17.0% of SVL, HW 64.0% of HL), distinct from neck. Snout length more than Coloration. In preservative: dorsum olive brown; twice eye diameter (ED 46.5% of ES); interorbital nape with a black mark; venter dusky white (see Plate moderate (IO 7.8% of HL); pupil rounded. Granules 8a). In life: dorsum dusky brown, with about eight on snout smooth, larger than those of occipital rusty-brown narrowly separated blotches; nape with region; scales on interorbital and supercilium oval; a distinct broad and long black marking; ventrally scales on ventral surface of head smooth. Rostral bright yellow (see Plate 8b). scale partially divided by a medial groove, in contact with first supralabial (Fig. 26a). Nares separated Variation. In the three paratypes (WHT 7237, WHT by two enlarged supranasals and a single enlarged 7238 and WHT 7236): supralabials (8, 8, 9); infra­labials internasal scale that extends towards snout tip (Fig. (9, 8, 9); subdigital lamellae on finger I (10, 10, 11), 26a). Nares rounded, dorsally orientated, in contact finger II (14, 13, 15), finger III (16, 15, 17), fing­er IV (17, with first supralabial. Two postnasals, the lower 18, 17), finger V (14, 14, 15); toe I (12, 11, 12), toe II (15, one in broad contact with first supralabial (Fig. 26a). 16, 15), toe III (17, 18, 17), toe IV (19, 18, 19), toe V (15, Mental subpentagonal, truncate posteriorly, wider 16, 15). Ventrals, 129–141; ventral scales across mid- than long, posteriolaterally in contact with two body, 18–25. Femoral pores, 15 or 16; pores absent

70 Zeylanica Re v i s i o n o f Cn e m a s p i s a b

c d

e f

g h

Figure 26. Cnemaspis phillipsi: WHT 7248, holotype, 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 and ventral regions, respectively, of mid-body; f, preanal region and right hind limb regions, showing arrangement of femoral pores; g, arrangement of subcaudal scales; h, ventral aspects of chin of paratype, WHT 7237. Scale bars: 1 mm.

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in females. SVL range 29.2–36.6 mm. Postmentals respectively, by the following opposing suites of broadly in contact in WHT 7237 (Fig. 26h) characters: Cnemaspis tropidogaster: ventrals keeled, dorsal Distribution and natural history notes. Cnemaspis scales heterogeneous. Cnemaspis kandiana: gular scales phillipsi is endemic to Sri Lanka and evidently keeled. Cnemaspis podihuna and C. molligodai: preanal restricted to vicinity of Gammaduwa (see Fig. 27) pores present. Cnemaspis gemunu: each postmental in the northern Knuckles Range (altitiude ~760 m; bounded by three scales, nostril in contact with rainfall ~ 2,500 mm yr-1). It is a fast-moving gecko, first supralabial, basal-most lamella of distal series usually seen in pairs. It was observed mostly on the sometimes fragmented. Cnemaspis alwisi: 9 femoral trunks of large trees (>1 m gbh), usually 3–5 m above pores. Cnemaspis scalpensis: ventrals, 122–127; conical ground level, in well-shaded areas within secondary tubercles along dorsolateral region of body and forest (see Plate 8c). Unlike many arboreal species of on tail. Cnemaspis samanalensis: gular, pectoral and Sri Lankan Cnemaspis, it appears to prefer smooth- abdominal scales tricarinate, subcaudals keeled. barked trees. The following lizards were observed Cnemaspis retigalensis: dorsal scales heterogeneous, in sympatry with Cnemaspis phillipsi: C. punctata, C. gulars carinate, a single preanal pore, 3 or 4 femoral kallima, Cyrtodactylus soba, Geckollea triedra, Otocryptis pores; subcaudals on median row enlarged, irregular. wiegmanni, and Calotes liocephalus. Cnemaspis punctata: 5–7 femoral pores, distinctive spotted dorsal coloration. Cnemaspis clivicola, C. Etymology. The species name is a patronym, Latinized silvula, C. pulchra, C. pava and C. upendrai: pectoral, in the possessive singular case, honouring W. W. A. abdominal and subcaudal scales keeled. Cnemaspis Phillips (1892–1981), who apparently collected the kallima, C. kumarasinghei, C. menikay and C. latha: first specimens of this species from his tea plantation preanal pores present. Cnemaspis amith: gular scales at Gammaduwa, subsequently identified by P. E. P. carinate; dorsal granules heterogeneous; subcaudals Deraniyagala (1953) as C. jerdonii scalpensis. on median row enlarged, irregular. Cnemaspis beddomei: ventrals keeled, preanal Comparisons. Cnemaspis phillipsi differs from each pores 6–9, no femoral pores. Cnemaspis goaensis: of its Sri Lankan and peninsular Indian congeners, preanal pores 2–3, gulars keeled. Cnemaspis littoralis: body slender, a black spot on nape. Cnemaspis mysoriensis: three rows of enlarged subcaudals, 2 or 3 femoral pores. Cnemaspis indraneildasii: dorsal scales heterogeneous. Cnemaspis jerdonii: 8 femoral pores. Cnemaspis heteropholis: dorsal scales heterogeneous. Cnemaspis indica, C. nairi, C. otai, C. sisparensis, C. wynadensis and C. yercaudensis: spine-like tubercles absent on flank. Cnemaspis boiei: preanal and femoral pores absent. Cnemaspis gracilis: 2 preanal pores separated by 2 scales, 4 femoral pores. Cnemaspis ornata: femoral pores absent, dorsal tubercles arranged in 15 longitudinal rows. Cnemaspis wicksii: subcaudals on median series irregular. Cnemaspis andersonii: gulars carinate, subcaudals in median series not enlarged, dorsals heterogeneous. Cnemaspis monticola: gulars and subcaudals carinate. Cnemaspis australis: subcaudals pointed, strongly keeled; scales on median row slightly enlarged. Cnemaspis nilagirica: pectoral and abdominal scales weakly keeled.

Remarks Cnemaspis phillipsi is the species previously identified from Gammaduwa as C. jerdonii scalpensis by Figure 27. Distribution of Cnemaspis phillipsi: the Knuckles Deraniyagala (1932, 1953) and C. scalpensis by Hills, Sri Lanka. Wickramasinghe (2006).

72 Zeylanica Re v i s i o n o f Cn e m a s p i s

Cnemaspis punctata new species elongate postmentals and medially with a single (Figures 28a–g, 29; Tables 4, 6; Plate 8 c, 9a, b) hexagonal postmental scale; postmentals bordered posteriorly by 5 smooth scales including medial scale Holotype. WHT 7256 (male), 37.1 mm SVL, (Fig. 28b). Supralabials 10, infralabials 7, decreasing Gammaduwa Estate, Gammaduwa near Rattota, in size towards gape (Fig. 28c). Ear opening oval, Matale District, 07º34’N, 80º42’ E, 760 m, coll. S. higher than wide, oblique (TW 37.5% of TH). Scales Batuwita & P. Wickramathilaka, 23 March 2007. on ventral surface of neck smooth. Conical granules laterally on neck as large as those of occipital region Paratypes. WHT 7223 (male), 35.2 mm SVL, WHT and body. 7226 (male), 36.0 mm SVL, WHT 7243 (male), 34.0 Body robust, relatively long (AG 43.9% SVL). mm SVL, NMSL (formerly WHT 7244) (male), 33.8 Dors­al granules homogeneous (Fig. 28d); a row of mm SVL, same collection data as holotype. 12 irregular spine-like tubercles on flank; ventrals, 135; pectoral and abdominal scales smooth (Fig. 28e). Other materials. WHT 7224, 7225, 7247, 7250 (4 Ventral scales across midbody, 23; scales around males) WHT 7240, 7241 (2 females) 7231, 7232, 7234, vent smooth, imbricate (Fig. 28f); those on base of 7235, 7242, 7246 (6 subadult females), same collection tail smooth, imbricate. No preanal pores; 5 (left side) data as holotype. and 6 (right side) (Fig. 28f) femoral pores. Forelimbs moderately long, robust (LAL 12.6% Diagnosis SVL, UAL 15.6% SVL); hind limbs moderately Cnemaspis punctata differs from its peninsular Indian long (TBL 18.0% SVL, FEL 18.3% SVL); tibia short. and Sri Lankan congeners by a combination of the Dorsal surface of both fore and hind limbs covered fol­low­ing characters. Maximum SVL 36.0 mm; by granular scales; scales on ventral surface of both paired postmentals separated by a medial scale; upper and lower arm smooth. Scales on anterior each postmental bounded by 4 scales including surface of thigh smooth, those on posterior surface medial scale; ventrals, 131–135; ventral scales across granular. Scales on ventral surface of shank smooth; midbody,­ 20–29; dorsal scales homogeneous; dorsal those on manus and pes smooth, rounded. Digits tubercles isolated, scattered; spine-like tubercles elongate, slender, all bearing slightly recurved claws; present on flanks; ventrals smooth; preanal pores subdigital lamellae entire, unnotched; subdigital ab­sent; 5–7 femoral pores on each side; subcaudals lamellae on finger I, 12, finger II, 16, finger III, 17, smooth, scales on median row enlarged, subequal; finger IV, 18, finger V, 17; toe I, 13, toe II, 16, toe III, supra­labials to angle of jaws, 7–10; subdigital lam­ 20, toe IV, 20, toe V, 16; interdigital webbing absent; ellae on digit IV of pes, 17–23; (in life) dorsum dark, relative length of digits (fingers): IV (3.5 mm) > III with minute olive-coloured spots forming an ir­ (3.2 mm) > II (2.9 mm) > V (2.6 mm) > I (1.9 mm); regular pattern; two longitudinal series of distinct (toes) IV (4.1 mm) > III (3.8 mm) > V (3.5 mm) > II dusky-white spots on dorsum; tail with about nine (3.0 mm) > I (1.7 mm). dusky-white blotches. Tail tapering, its length greater than SVL (TAL 111% of SVL); tail base distinctly swollen; five rows Description of holotype. An adult male, 37.1 mm SVL of enlarged, rounded tubercles on base of tail. Dorsal (for measurements, see Table 4). Head relatively short scales on tail homogeneous, directed backwards; (HL 25.3% of SVL), broad (HW 18.3% of SVL, HW about 5 broad, rounded tubercles on tail, forming 72.3% of HL), distinct from neck. Snout length more whorls; 2 conical post-cloacal spurs present on each than twice eye diameter (ED 42.8% of ES); interorbital side; subcaudals smooth, those on median series moderate (IO 9.0% of HL); pupil rounded. Granules regular, enlarged, quadrate to hexagonal (Fig. 28g). on snout smooth, larger than those of occipital region; scales on interorbital and supercilium rounded, those Coloration. In preservative: dorsally greyish, with on ventral surface of head smooth. Rostral scale dis­tinct, scattered, dark and pale spots from nape to partially divided by a medial groove, in contact with base of tail; venter dusky white; tail with pale blotch­ first supralabial (Fig. 28a). Nares rounded, dorsally es. In life: dorsum dark, with minute olive spots orientated, separated by two enlarged supranasals forming an irregular pattern; two longitudinal series and a single subequal internasal scale (Fig. 28a). of distinct, dusky-white spots on dorsum; tail with Nostril not in contact with first supralabial. Two about nine dusky-white blotches (see Plate 9a, b). postnasals, the lower one enlarged, in broad contact with first supralabial (Fig. 28a). Mental subtriangular, Variation. In the four paratypes (WHT 7244, 7223, as wide as long, posteriolaterally in contact with two 7226 and 7243): supralabials (9, 9, 9, 9); infralabials

Vol. 7, No. 1 73 Ma n a m e n d r a -Ar a c h c h i e t a l .

a b

c d

e f

g

Figure 28. Cnemaspis punctata: WHT 7256, holotype, 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 and ventral regions, respectively, of mid-body; f, preanal region and right hind limb regions, showing arrangement of femoral pores; g, arrangement of subcaudal scales. Scale bars: 1 mm.

74 Zeylanica Re v i s i o n o f Cn e m a s p i s

(7, 7, 7, 9); ventrals (133, 132, 131, 133); ventral scales respectively, by the following opposing suites of across mid-body (29, 21, 20, 26); subdigital lamellae characters: on finger I, 12 (12, 11, 12, 11), finger II (15, 14, 14, 13), Cnemaspis tropidogaster: ventrals keeled, dorsal finger III (18, 17, 17, 16), finger IV (19, 18, 20, 17), scales heterogeneous. Cnemaspis kandiana, C. podihuna finger V (17, 17, 17, 16); toe I (12, 12, 12, 11), toe II and C. molligodai: preanal pores present. Cnemaspis (16, 15, 16, 14), toe III (21, 18, 19, 17), toe IV (23, 17, gemunu: 12 femoral pores. Cnemaspis alwisi: ventrals, 21, 18), toe V (19, 11, 16, 15); SVL range from 22.1 to 146–152, femoral pores 9. Cnemaspis samanalensis: 39.9 mm (17 ex.). Femoral pores, 5–7 in males; pores gular, pectoral and abdominal scales tricarinate, absent in females. subcaudals keeled. Cnemaspis retigalensis: gulars carinate, a single preanal pore, 3 or 4 femoral pores; Distribution and natural history. This fast-moving subcaudals on median row enlarged, irregular. gecko appears to be restricted to the vicinity of Cnemaspis scalpensis and C. phillipsii: 13–16 femoral Gammaduwa, at the northern extremity of the pores. Cnemaspis kumarasinghei: presence of preanal Knuckles Range (see Fig. 29, Plate 8c). It is locally pores. Cnemaspis clivicola, C. pulchra, C. pava, C. silvula abundant in edificarian habitats (e.g., an abandoned and C. upendrai: subcaudals, pectoral and abdominal cardamom-processing warehouse). In nearby second­ scales keeled. Cnemaspis kallima, C. menikay and C. ary forest, these geckos were observed most commonly latha: preanal pores present. Cnemaspis amith: gular on rock surfaces and crevices. The following lizards scales carinate; subcaudals on median row enlarged, were observed in sympatry: Cnemaspis phillipsii, C. irregular. kallima, Cyrtodactylus soba, Geckollea triedra, Otocryptis Cnemaspis beddomei: ventrals keeled, preanal wiegmanni, Calotes liocephalus. pores 6–9, femoral pores absent. Cnemaspis goaensis: preanal pores 2–3, gulars keeled. Cnemaspis littoralis: Etymology. The species name is a Latin adjective body slender. Cnemaspis mysoriensis: three rows of (gender feminine) meaning ‘dotted’, a reference to enlarged subcaudals, 2 or 3 femoral pores. Cnem­ the distinctive dorsal colour pattern of this gecko. aspis indraneildasii: dorsal scales heterogeneous. Cnemaspis jerdonii: caudal tubercles absent. Cnemaspis Comparisons. Cnemaspis punctata differs from each heteropholis: dorsal scales heterogeneous. Cnemaspis of its Sri Lankan and peninsular Indian congeners, indica, C. nairi, C. otai, C. sisparensis, C. wynadensis and C. yercaudensis: spine-like tubercles absent on flank. Cnemaspis boiei: preanal and femoral pores absent. Cnemaspis gracilis: 2 preanal pores separated by 2 scales, 4 femoral pores. Cnemaspis ornata: femoral pores absent, dorsal tubercles arranged in 15 longitudinal rows. Cnemaspis wicksii: subcaudals on median row enlarged, irregular. Cnemaspis andersonii: gulars carinate, subcaudals in median series not enlarged, dorsal scales heterogeneous. Cnemaspis monticola: gulars carinate. Cnemaspis australis: subcaudals pointed, strongly keeled. Cnemaspis nilagirica: pectoral and abdominal scales weakly keeled.

Cnemaspis upendrai new species (Figures 30a–g, 31; Tables 4, 6; Plate 11c, d)

Holotype. WHT 7189 (male), 31.5 mm SVL; Pussellawa, Kandy District, Central Province of Sri Lanka, 07º05’N, 80º30’E, 900 m, coll. S. Batuwita & K. Wewelwala, 27 November, 2002.

Paratypes. WHT 7184, 31.0 mm SVL, 7186, 29.2 mm SVL, 7187, 31.2 mm SVL, 7188, 32.7 mm SVL, NMSL Figure 29. Distribution of Cnemaspis punctata: the Knuckles (formerly WHT 7181), 26.0 mm SVL (5 males), NMSL hills, Sri Lanka. (formerly WHT 7182), 32.5 mm SVL, WHT 7183, 33.5

Vol. 7, No. 1 75 Ma n a m e n d r a -Ar a c h c h i e t a l .

7×4 4.2 0.7 8.1 5.5 3.6 4.0 1.1 1.0 5.5 8.6 5.4 2.4 2.8 2.8 1.3 2.2 3.9 1.8 6.2 4.2 3.5 2.7 2.5 2.7 3.4 3.5 4.1 3.1 2.9 1.8 4.1 5.2 – 32.0 BMNH C. amith 63.3.19.1066A holotype (WHT holotype pulchra; C. 5×2 3.6 1.0 7.1 4.8 2.9 4.1 0.7 1.0 5.1 8.0 5.8 2.7 2.7 2.5 1.4 2.2 3.9 1.7 6.1 3.8 2.5 2.8 2.5 3.3 2.5 3.6 3.8 3.2 2.6 1.7 3.3 5.2 – 30.4 14.2 7214 WHT C. latha Cnemaspis punctata; holotype (WHT 5×5 4.9 1.1 8.1 5.8 3.5 4.9 0.5 1.1 5.8 9.0 7.3 1.8 2.8 3.2 1.5 3.4 4.5 2.9 6.3 4.2 3.7 3.1 3.4 3.7 2.6 4.6 4.2 3.9 3.2 1.8 3.6 6.5 33.6 13.5 36.3 WHT 1573a 3×1 4.6 1.0 8.5 5.7 3.1 4.8 0.5 1.1 5.7 9.5 7.6 1.7 2.7 3.3 1.4 3.5 3.9 3.0 6.8 3.9 3.0 3.2 2.9 3.9 2.6 4.8 3.9 4.0 3.5 2.1 3.8 6.5 C. pulchra 7023 34.2 14.9 41.6 WHT 4×3 4.4 1.0 7.2 4.5 3.0 4.2 0.5 1.1 5.4 7.9 6.4 2.4 3.4 2.9 1.5 2.4 4.8 1.8 5.9 3.8 2.3 2.7 2.2 4.2 2.2 4.2 3.8 3.5 2.7 1.7 3.3 5.6 7184 31.0 14.3 33.1 WHT 5×3 4.9 0.9 7.4 5.0 2.7 4.3 0.5 1.1 5.5 8.3 7.0 2.6 3.1 3.0 1.5 2.6 5.0 2.1 5.7 3.8 2.3 2.7 2.7 3.7 2.3 4.3 3.5 3.9 3.4 1.9 3.5 5.8 31.2 14.3 34.8 7187 WHT 5×4 4.7 1.0 8.0 5.3 3.3 4.8 0.5 1.1 5.7 8.9 7.3 2.8 3.4 3.4 1.5 2.8 4.6 2.0 6.1 4.1 2.5 3.1 2.8 3.4 2.6 4.2 4.0 3.5 3.0 1.9 3.7 6.2 32.7 14.0 7188 30.8 WHT C. upendrai ; holotype (WHT 7023) and paratype (WHT 1573a) of 1573a) (WHT paratype and 7023) (WHT holotype upendrai ; C. 6×4 4.1 1.0 7.1 4.8 2.5 4.0 0.4 1.1 5.1 7.7 6.5 2.5 3.0 2.8 1.5 2.4 3.7 1.7 5.1 3.8 2.5 2.6 2.6 3.4 2.2 3.9 3.4 3.3 2.9 1.5 3.1 5.4 – 29.2 14.7 7186 WHT 6×2 4.5 0.9 7.5 5.0 2.7 4.3 0.5 1.0 5.6 8.6 6.8 2.7 3.4 3.1 1.4 2.5 4.1 1.8 6.4 3.8 2.8 2.9 2.8 3.6 2.4 4.1 3.7 3.5 2.9 1.7 3.5 6.0 7189 31.5 13.8 40.7 WHT 5×6 4.6 1.0 8.3 5.5 3.5 4.3 0.6 1.0 6.4 9.1 7.2 3.0 3.6 3.1 1.5 2.5 5.5 1.8 6.6 4.0 3.1 3.4 1.8 3.6 2.5 4.2 3.9 4.0 3.2 1.8 4.4 6.4 – 34.0 14.9 7243 WHT

R 2×6 4.7 1.0 8.4 5.5 3.4 5.0 0.6 1.0 6.2 8.9 7.3 2.8 3.3 3.3 1.8 2.8 4.9 2.0 6.3 4.2 3.1 3.5 1.7 3.8 2.3 4.5 4.1 3.9 3.2 1.8 4.4 6.5 36.0 17.0 27.4 7226 WHT

R 5×6 4.6 0.9 8.1 5.5 3.4 4.0 0.7 1.0 6.0 8.9 7.3 2.9 3.6 3.2 1.5 2.7 4.9 1.9 6.4 4.0 2.8 3.2 1.7 4.0 2.3 4.4 4.2 3.6 3.1 1.8 4.3 6.5 22.4 35.2 14.8 7223 WHT C. punctata 4×6 – 4.6 1.0 8.0 5.5 3.5 4.0 0.8 1.0 6.0 8.9 7.1 2.9 3.4 3.1 1.6 2.4 5.1 1.9 6.3 4.0 2.8 3.3 1.8 3.6 2.3 4.4 4.3 3.9 3.1 1.9 4.5 6.1 33.8 15.0 7244 WHT 3×8 4.4 1.0 8.4 5.9 3.7 4.7 0.9 1.1 6.8 9.4 7.0 2.6 3.5 3.2 1.3 2.9 5.8 1.9 6.8 4.2 2.8 3.7 1.8 3.5 2.8 4.1 4.5 3.8 3.0 1.7 4.8 6.7 41.2 37.1 16.3 7256 WHT

TAL SVL PAL NS MN MFE MBE LAL IO IN HW HL FOL FL V FL FL IV FL Table 4. Table Measurements (in mm) of holotype (WHT 7256) and 4 paratypes (WHT7244, WHT 7223, WHT 7226, WHT of 7184) 7243) WHT 7187, of WHT 7188, WHT 7186, (WHT paratypes 4 and 7189) – indicates tail lost . R indicates tail regenerated; 7214) of C. scitula; and holotype amith. B indicates tail broken; FL III FL UEW FL II FL UAL FL I FL FEL ES TYE (×10) EN TYD ED TL V TL DFE TL IV TL DBE TL III TL AG TL II TL TL I TL

TBW TBL

76 Zeylanica Re v i s i o n o f Cn e m a s p i s a b

c d

e

f

g

Figure 30. Cnemaspis upendrai: WHT 7189, holotype, 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 and ventral regions, respectively, of mid-body; f, preanal and right hind-limb regions, showing arrangement of femoral and preanal pores; g, arrangement of subcaudal scales. Scale bars: 1 mm.

Vol. 7, No. 1 77 Ma n a m e n d r a -Ar a c h c h i e t a l .

mm SVL, 7185, 33.9 mm SVL, 7190, 34.2 mm SVL, tail rusty brown; venter white. Some individuals 7191, 35.2 mm SVL, 7192, 33.9 mm SVL, 7193, 33.6 uni­form yellowish brown, without prominent dorsal mm SVL, 7194, 33.9 mm SVL, 7195, 33.5 mm SVL (9 markings. females), WHT 7196, 16.4 mm SVL, 7197, 17.9 mm SVL (2 juveniles), same collection data as holotype; Description of holotype WHT 7260 (male), 28.3 mm SVL, same location data An adult male, 31.5 mm SVL (for measurements, see as holotype, coll. S. Batuwita & P. Wickramathilaka, Table 4). Head moderately long (HL 27.3% of SVL), 26 March, 2007. moderately broad (HW 17.7% of SVL, HW 65.1% of HL), distinct from neck. Snout length a little greater Diagnosis than eye diameter (ED 73.6% of ES); interorbital Cnemaspis upendrai differs from its peninsular Indian narrow (IO 5.8% of HL); pupil rounded. Granules and Sri Lankan congeners by a combination of the on snout keeled, larger than those of occipital region; following characters. Maximum SVL 35.2 mm; scales of occipital region, interobital, supercilium paired postmentals separated by a medial scale; each and ventral surface of head keeled, granular. Rostral postmental bounded by 4 scales including medial scale partially divided by a medial groove, in contact scale; ventrals, 112–128; ventral scales across mid- with first supralabial (Fig. 30a). Nares separated by body, 16–25; dorsal scales heterogeneous; a row of two enlarged supranasals, upper postnasal (left side) 15 irregular, spine-like tubercles present on flank; and a single oval internasal scale (Fig. 30a). Nares ventrals smooth; 2 or 3 preanal pores, 4 or 5 femoral rounded, dorsally orientated, not in contact with pores on each side; subcaudals keeled, median row first supralabial. Two postnasals, the lower one in enlarged, irregular; supralabials to angle of jaws, 7 broad contact with first supralabial (Fig. 30a). Mental or 8; subdigital lamellae on digit IV of pes, 17–21; (in sub­pentagonal, wider than long, posteriolaterally in life) dorsum light-brown, with about eight median con­tact with two enlarged quadrate postmentals and black markings extending from neck and on to tail; medi­ally with a single small pentagonal postmental much larger, irregular, dark-brown blotches on sides; scale; postmentals bordered posteriorly by 5 smooth occipital region with two black chevrons, a light scales including medial scale (Fig. 30b). Supralabials yellow marking between them, followed by a black 7, infralabials 7, decreasing in size towards gape (Fig. spot on neck; a narrow interorbital band present; 30c). Ear opening oval, higher than wide, oblique (TW 33.3% of TH). Scales on ventral surface of neck uni­carinate, bicarinate or tricarinate. Large, conical, keeled granules behind ear and laterally on neck, the latter intermixed with larger conical granules. Body slender, relatively long (AG 43.8% SVL). Mid-dorsal granules keeled, heterogeneous, intermixed with enlarged, conical, keeled tubercles (Fig. 30d); 15 spine-like tubercles on flank; abdominal scales keeled. Ventrals, 128; imbricate, pointed (Fig. 30e); ventral scales across midbody, 25, wider than long; scales around preanal region and base of tail smooth, imbricate; 3 preanal pores, 4 femoral pores on each side (Fig. 30f). Forelimbs moderately long, slender (LAL 13.6% SVL, UAL 13.0% SVL); hind limbs long (TBL 19.0% SVL, FEL 20.3% SVL); tibia short. Dorsal scales on fore limbs and thigh carinate; ventral scales on upper arm smooth, lower arm with both smooth and weakly keeled scales. Scales on anterior surface of thigh carinate, on ventral surface both smooth and keeled; those on posterior surface of thigh granular. Dorsal surface of shank with keeled scales, smooth scales and granular scales; scales on ventral surface keeled. Dorsal scales on manus and pes keeled, ventrally granular, not keeled. Digits elongate, slender, all Figure 31. Distribution of Cnemaspis upendrai in Sri Lanka’s central highlands. bearing slightly recurved claws; subdigital lamellae

78 Zeylanica Re v i s i o n o f Cn e m a s p i s

entire, unnotched; subdigital lamellae on finger I, respectively, by the following opposing suites of 12, finger II, 14, finger III, 17, finger IV, 18, finger V, characters: 14; toe I, 13, toe II, 16, toe III, 19, toe IV, 21, toe V, 13; Cnemaspis tropidogaster: nostril in contact with interdigital webbing absent; relative length of digits first supralabial, ventral scales on precloacal and tail (fingers): IV (3.4 mm) > III (3.1 mm) > V (2.7 mm) > base area carinate, 6 spine-like tubercles on flank, II (2.5 mm) > I (1.8 mm); (toes) IV (4.1 mm) > V (3.6 enlarged tubercles absent on base of tail. Cnemaspis mm) > III (3.5 mm) > II (2.9 mm) > I (1.7 mm). kandiana and C. menikay: pectoral and abdominal Tail tapering, its length greater than SVL (TAL scales smooth. Cnemaspis podihuna: gular, pectoral and 129.2% of SVL); tail base distinctly swollen. Enlarged, abdominal scales smooth; subcaudals on median row conical keeled tubercles present on base of tail, about enlarged, subequal. Cnemaspis scalpensis, C. gemunu as large as tubercles on mid-dorsum. Dorsal scales on and C. phillipsii: 12–16 femoral pores. Cnemaspis tail keeled, directed backwards. Six rows of pointed alwisi: 9 femoral pores. Cnemaspis molligodai: gular, tubercles forming whorls on tail; a single conical pectoral and abdominal scales smooth, dorsal scales post-cloacal spur present on each side; subcaudals homogeneous; subcaudals on median row enlarged, pointed, keeled, median row with pairs of smaller subequal. Cnemaspis samanalensis: gular, pectoral and scales between individual larger ones (Fig. 30g). abdominal scales tricarinate. Cnemaspis retigalensis: pectoral and abdominal scales smooth, a single Coloration. In preservative: dorsally dark brown preanal pore and 3 or 4 femoral pores. Cnemaspis with indistinct darker bands interrupted medially; punctata: 5–7 femoral pores and distinctive spotted nape with a dark marking; venter dusky white with dorsal coloration. Cnemaspis kumarasinghei: pectoral darker markings. and abdominal scales smooth. Cnemaspis silvula In life: dorsum light brown, with about six and C. pava: ventral scales on tail base carinate. confluent light-brown blotches; nape with a distinct Cnemaspis clivicola: subcaudals in median series not black marking (see Plate 11c). enlarged. Cnemaspis kallima and C. latha: pectoral and abdominal scales smooth. Cnemaspis pulchra: gular Variation. Paratypes (WHT 7188, WHT 7187, 7184 scales unicarinate. Cnemaspis amith: pectoral and and WHT 7186). Supralabials (8, 7, 7, 7); infralabials abdominal scales smooth. (7, 7, 7, 8); subdigital lamellae on finger I (11, 10, 10, Cnemaspis beddomei: femoral pores absent. Cnemas­ 9), finger II (13, 12, 12, 12), finger III (17, 15, 14, 16), pis goaensis: ventrals smooth. Cnemaspis littoralis­ : finger IV (17, 15, 15, 14), finger V (15, 13, 13, 13); toe dorsal scales homogeneous, subcaudals on median I (10, 10, 10, 9), toe II (14, 13, 13, 13), toe III (18, 15, row enlarged, hexagonal. Cnemaspis mysor­iensis: 15, 17), toe IV (19, 18, 18, 17), toe V (17, 15, 16, 16). dorsal scales homogeneous, three rows of enlarged Ventrals, 112–128; ventral scales across mid-body, subcaudals. Cnemaspis indraneildasii and C. hetero­ 16–25; SVL ranges from 16.4 to 35.2 mm (17 ex.). pholis: ventral, gular and caudal scales smooth. Preanal pores 2 or 3, femoral pores, 4 or 5 in males; Cnemas­pis indica, C. nairi, C. otai, C. yercaudensis, C. sis­ pores absent in females. parensis and C. wynadensis: spine-like tubercles absent on flank. Cnemaspis boiei: ventrals smooth, no pre­anal Distribution and natural history. Cnemaspis upendrai or femoral pores. Cnemaspis gracilis: ventrals smooth. appears to be restricted to the vicinity of Pussellawa Cnemaspis ornata: no femoral pores, dorsal tub­ercles and Ramboda in the up-country wet zone (altitiude arranged in 15 longitudinal rows. Cnemaspis wicksii ~900 m; rainfall > 2,500 mm yr-1) (Fig. 31). The geckos and C. andersonii: pectoral and abdominal scales were observed on moist rock faces and in crevices, smooth. Cnemaspis jerdonii: no preanal pores. Cnemas­ as well as in edificarian habitats. Communal egg pis monticola: pectoral and abdominal scales smooth. laying too, was observed (Plate 11d). At Pussellawa, Cnemaspis australis: scales on preanal region smooth. C. upendrai occurs in sympatry with the lizards Hemi­ Cnemaspis nilagirica: dorsal scales homogeneous; phyllodactylus typus, Cyrtodactylus ramboda, Lanka­ subcaudals on median row enlarged, subequal. scincus fallax and L. taylori. Cnemaspis clivicola new species Etymology. The species name is a patronym Latinized (Figures 32a–g, 33; Tables 5, 6; Plate 11a, b) in the genitive singular honouring the Sri Lankan prehistorian Dr Siran Upendra Deraniyagala. Holotype. WHT 7204 (male), 36.6 mm SVL, Nanu Oya, Nuwara Eliya District, Central Province of Comparisons. Cnemaspis upendrai differs from each Sri Lanka, 06º56’30”N, 80º44’30”E, 1623 m, coll. S. of its Sri Lankan and peninsular Indian congeners, Batuwita, 11 August, 2005.

Vol. 7, No. 1 79 Ma n a m e n d r a -Ar a c h c h i e t a l . a b

c d

e

f

g

Figure 32. Cnemaspis clivicola: WHT 7204, holotype, 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 and ventral regions, respectively, of mid-body; f, preanal and right hind-limb regions, showing arrangement of femoral and preanal pores; g, arrangement of subcaudal scales. Scale bars: 1 mm.

80 Zeylanica Re v i s i o n o f Cn e m a s p i s

Paratypes. NMSL (formerly WHT 7178) (male), 34.2 distinct from neck. Snout length moderately greater mm SVL, 7180 (male), 33.9 mm SVL, same location than eye diameter (ED 68.1% of ES); interorbital and collection data as holotype, 31 May, 2003; WHT moderate (IO 7.5% of HL); pupil rounded. Granules 7179 (male), 34.6 mm SVL, 7205 (male), 44.3 mm on snout and interorbital keeled; occipital region, SVL, 7215 (female), 40.9 mm SVL, same collection area around ear, and top and sides of neck covered in data as holotype. granules similar in size to those on snout, intermixed with enlarged conical tubercles; granules around Diagnosis gape, scales bordering infralabials, and postmentals, Cnemaspis clivicola differs from its peninsular Indian smooth; chin and throat with keeled granules. Rostral and Sri Lankan congeners by a combination of scale slightly divided by a medial groove, in contact the following characters. Maximum SVL 40.9 mm; with first supralabial (Fig. 32a). Nares separated paired postmentals separated by a medial scale; by two enlarged supranasals and two pentagonal each postmental bounded by 4 scales including internasal scales, the anterior one extending medial scale; ventrals, 128–144; ventral scales across towards snout (Fig. 32a). Nares rounded, dorsally mid-body, 20–26; dorsal scales heterogeneous; a row orientated; nostril with rather narrow contact with of 10–15 irregular spine-like tubercles present on first supralabial (Fig. 32a). Two postnasals, the flank; ventrals keeled; 3 or 4 preanal pores, 4 femoral lower one in broad contact with first supralabial pores on each side; subcaudals in median series not (Fig. 32a). Mental subpentagonal, wider than long, enlarged, keeled; supralabials to angle of jaws 7 or posteriolaterally in contact with two enlarged 8; subdigital lamellae on digit IV of pes 17–19; (in postmentals and medially with a single small life) dorsum grey, with a series of seven W-shaped pentagonal postmental scale; postmentals bordered markings; a narrow interorbital band. posteriorly by 6 (3 on left side, 2 on right side and 1 medial) smooth scales (Fig. 32b). Supralabials 9, Description of holotype infralabials 7, decreasing in size towards gape (Fig. An adult male, 36.6 mm SVL (for measurements, see 32c). Ear opening oval, higher than wide, oblique Table 5). Head relatively short (HL 25.4% of SVL), (TW 60.0% of TH); 5 conical tubercles laterally be­ flattened (HW 15.5% of SVL, HW 61.2% of HL), tween ear and insertion of forelimb. Body robust, relatively long (AG 43.9% SVL). Mid-dorsal granules heterogeneous, keeled, intermixed with large conical keeled tubercles (Fig. 32d); a row of 12–15 irregular spine-like tubercles on flank; abdominal scales strongly keeled (Fig. 32e). Ventrals, 144, bluntly pointed, longer than wide; scales across midbody, 26; scales around vent and base of tail smooth, imbricate; 4 preanal and 4 femoral pores on each side (Fig. 32f). Forelimbs moderately long, slender (LAL 13.6% SVL, UAL 13.6% SVL); hind limbs long (TBL 18.0% SVL, FEL 20.2% SVL); tibia short. Dorsal scales on both fore and hind limbs carinate; ventral scales on upper arm unkeeled, granular, those on lower arm weakly keeled. Scales on anterior surface of thigh keeled; dorsal and posterior surface of thigh with conical granules; scales on ventral surface carinate (Fig. 32f). Dorsal and posterior surfaces of shank with conical granules, those on ventral surface carinate. Scales dorsally on manus and pes keeled, on ventral surface smooth, imbricate and granular. Digits elongate, slender, all bearing slightly recurved claws; subdigital lamellae entire, unnotched; subdigital lamellae on finger I, 10, finger II, 13, finger III, 14, finger IV, 15, finger V, 12; toe I, 10, toe II, 13, toe III, Figure 33. Distribution of Cnemaspis clivicola in the central 16, toe IV, 18, toe V, 15; interdigital webbing absent; highlands of Sri Lanka. relative length of digits (fingers): IV (3.8 mm) > III

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(3.5 mm) > V (3.0 mm) > II (2.8 mm) > I (1.9 mm); tricarinate. Cnemaspis retigalensis: pectoral and ab­ (toes) IV (5.2 mm) > III (4.2 mm) > V (4.1 mm) > II dom­inal scales smooth, a single preanal pore and 3 or (3.4 mm) > I (2.0 mm). 4 femoral pores; subcaudals on median row enlarged, Tail tapering, its length greater than SVL (TAL subequal. Cnemaspis punctata: 5–7 femoral pores, 123.3% of SVL) in paratype WHT 7205 (tail of distinctive spotted dorsal coloration. Cnemaspis holotype partially regenerated); tail base distinctly kumarasinghei: pectoral and abdominal scales smooth. swollen, with enlarged, keeled, conical tubercles. Cnemaspis silvula, C. pulchra, C. pava and C. upendrai: Dorsal scales on tail keeled, directed backwards. Tail subcaudals on median row enlarged. Cnemaspis with 6–8 whorls of pointed keeled scales; a single kallima: abdominal scales smooth. Cnemaspis latha: conical post-cloacal spur on each side; subcaudals pectoral and abdominal scales smooth. Cnemaspis strongly keeled, pointed, not enlarged (Fig. 32g). amith: pectoral and abdominal scales smooth; subcaudals on median row enlarged, irregular. Coloration. In preservative: dorsally dusky brown; Cnemaspis beddomei: femoral pores absent. nape with a dark marking; venter dusky white. In Cnemaspis goaensis: ventrals smooth. Cnemaspis life: dorsum grey, with a series of seven W-shaped littoralis: dorsal scales homogeneous, subcaudals markings; a narrow black interorbital band (see on median row enlarged, hexagonal. Cnemaspis, Plate 11a). mysoriensis: dorsal scales homogeneous, three rows of enlarged subcaudals, 2 or 3 femoral pores. Cnemaspis Variation. Paratypes WHT 7178, WHT 7215, indraneildasii: dorsal scales of both limbs smooth; WHT 7179 and WHT 7180: supralabials (8, 8, 7, ventral, gular and caudal scales smooth. Cnemaspis 7); infralabials (7, 8, 7, 8); ventrals (128, 131, 143, heteropholis: dorsal scales heterogeneous, ventral and 141); ventral scales across mid-body (20, 21, 21, 22); caudal scales smooth. Cnemaspis indica, C. sisparensis subdigital lamellae on finger I (10, 10, 11, 11), finger and C. wynadensis: spine-like tubercles absent on II (12, 13, 13, 13), finger III (14, 16, 16, 15), finger IV flank. Cnemaspis boiei: ventrals smooth, no preanal (15, 16, 17, 16), finger V (13, 13, 13, 13); toe I (9, 10, or femoral pores. Cnemaspis gracilis: ventrals smooth. 10, 9), toe II (13, 13, 14, 13), toe III (15, 17, 17, 16), toe Cnemaspis ornata: no femoral pores, dorsal tubercles IV (17, 18, 19, 17), toe V (15, 15, 16, 14); SVL range arranged in 15 longitudinal rows. Cnemaspis wicksii: 33.9–40.9 mm (6 ex.). Preanal pores 3 or 4; 4 femoral upper arm and lower arm, thigh and tibia covered pores in males; pores absent in females. dorsally by smooth scales. Cnemaspis andersonii: pectoral and abdominal scales smooth, scales of Distribution and natural history. The type series ventro-lateral region carinate. Cnemaspis jerdonii: was collected from an edificarian habitat at Nanu no preanal pores. Cnemaspis nairi, C. otai and C. Oya (altitiude ~1600 m; rainfall > 2,500 mm yr-1) (Fig. yercaudensis: spine-like tubercles absent on flank; 33). Communal egg laying site observed at the type ventrals smooth. Cnemaspis monticola: pectoral locality (Plate 11b). The only other lizard observed and abdominal scales smooth. Cnemaspis australis: in the vicinity was Lankascincus deignani. subcaudals on median row enlarged. Cnemaspis nilagirica: dorsal scales homogeneous; subcaudals Etymology. The species name is Latin for ‘inhabitant on median row enlarged, subequal. of the hills’, an allusion to the montane type locality of this species. Remarks The known range of C. clivicola lies wholly within that Comparisons. Cnemaspis clivicola differs from each of C. gemunu. The two species, however, are easily of its Sri Lankan and peninsular Indian congeners, distinguished: C. clivicola has 3 or 4 preanal pores respectively, by the following opposing suites of (vs preanal pores absent in C. gemunu); pectoral and characters: abdominal scales keeled (vs smooth); subcaudals on Cnemaspis tropidogaster: no enlarged tubercles on median row not enlarged (vs enlarged); and dorsal tail base. Cnemaspis kandiana, C. menikay, C. podihuna, granules keeled (vs smooth). We have, however, been C. scalpensis, C. gemunu and C. phillipsi: presence of unable to locate a population of C. clivicola except for smooth ventrals and subcaudals. Cnemaspis alwisi: 9 the one around Nanu Oya. femoral pores. Cnemaspis molligodai: 5 preanal and 8 or Cnemaspis clivicola and C. gemunu are the only 9 femoral pores, gular, pectoral and abdominal scales species in this genus known from altitudes above smooth, dorsal scales homogeneous; subcaudals 1,600 m in Sri Lanka. The only other gecko presently on median row enlarged, subequal. Cnemaspis known from similar elevation in Sri Lanka is Geckoella samanalensis: gular, pectoral and abdominal scales triedra.

82 Zeylanica Re v i s i o n o f Cn e m a s p i s 3.5 0.8 6.5 4.5 2.6 3.6 0.5 1.0 4.9 7.2 5.3 2.0 2.3 1.4 2.4 4.1 2.1 1.5 5.5 2.5 3.5 2.8 2.3 3.1 2.6 2.6 1.5 2.3 3.5 1.4 5.0 3.5 – 3×1 7349 11.4 26.9 WHT C. menikay 4×1 3.6 0.9 6.7 4.4 2.8 4.1 0.5 0.9 4.6 7.1 6.0 2.0 2.5 1.4 2.4 4.0 2.3 1.7 4.8 2.6 3.4 3.0 2.3 3.5 2.4 2.8 1.4 2.5 3.3 1.5 6.8 3.6 – 26.8 12.6 7219 WHT ; holotype clivicola ; C. 4×1 4.3 1.1 7.7 5.1 3.1 4.6 0.5 1.1 5.5 8.5 6.5 3.1 3.5 1.5 3.3 4.0 2.9 1.9 6.7 3.0 4.2 4.0 3.0 4.7 2.6 4.0 2.6 3.2 3.9 2.1 5.9 3.7 32.4 14.1 7297 39.4 WHT

B C. pava 5×1 4.5 0.9 7.4 4.9 2.7 4.5 0.5 1.0 5.5 8.3 6.5 2.7 3.4 1.7 3.2 4.2 2.7 2.1 6.9 3.1 4.0 4.0 2.6 4.7 2.7 4.1 2.2 3.5 3.9 4.0 2.2 6.4 31.8 28.8 15.4 7286 WHT 4×4 4.1 1.0 7.8 5.2 3.2 4.5 0.6 1.1 5.5 8.6 6.5 2.5 2.9 1.4 2.8 4.1 2.3 1.3 6.3 3.0 3.9 3.5 2.8 3.9 2.9 3.5 2.4 3.0 3.8 3.7 1.9 – 5.5 31.9 14.9 7228 WHT C. kallima; holotype (WHT 7286) and paratype (WHT 7297) C. kallima 5×3 4.0 0.9 7.0 4.7 2.6 4.1 0.5 1.1 5.1 7.8 6.2 2.4 2.9 1.1 2.8 3.8 2.4 1.5 5.8 2.7 3.5 3.1 2.6 3.7 2.6 3.2 2.4 2.6 3.2 3.4 1.5 – 5.4 28.8 12.9 7245 WHT 3×1 3.9 1.0 7.2 5.1 3.0 3.8 0.4 0.9 4.9 8.2 5.7 2.3 2.9 1.5 2.8 3.7 2.4 1.6 6.4 2.9 3.7 3.5 2.6 3.9 3.0 3.1 2.0 2.5 3.9 3.2 1.7 5.5 7209 WHT 30.3 33.6 12.6 C. silvula 3.7 0.9 7.0 4.7 2.5 4.2 0.4 1.0 4.8 7.7 5.8 2.1 2.5 1.5 2.3 3.5 2.1 1.4 6.2 2.8 3.5 2.8 2.7 3.4 2.7 3.0 2.0 2.5 3.7 – 1.5 5.4 5×2 7217 28.6 12.2 35.1 WHT

R 7×4 5.2 1.1 8.4 5.5 3.6 5.2 0.7 1.2 6.0 9.4 8.0 3.2 3.9 1.6 3.7 4.8 3.2 2.2 8.1 2.8 4.9 4.3 3.1 5.1 2.8 4.2 3.0 3.4 4.5 4.4 2.0 6.9 40.9 18.2 32.5 7215 WHT 7180 4.9 1.0 7.5 5.1 2.9 4.8 0.5 1.1 5.6 8.5 7.2 3.1 3.6 1.4 3.3 4.7 2.9 2.1 6.5 2.5 4.0 4.3 2.9 5.2 2.4 4.2 2.4 3.2 3.6 3.9 1.9 – 6.1 6×3 WHT 33.9 18.2 C. silvula; holotype (WHT 7245) and paratype (WHT 7228) of 4.4 1.0 7.7 5.1 3.0 4.5 0.6 1.0 5.2 8.3 7.0 2.8 3.6 1.3 3.3 4.4 2.7 2.1 5.8 2.8 4.1 4.2 3.0 4.7 2.5 4.0 2.3 3.0 3.9 3.5 1.9 5.9 6×5 34.2 17.5 39.2 7178 WHT C. clivicola

B 4.7 1.0 7.9 5.2 2.9 4.8 0.5 1.2 5.8 8.7 6.9 3.0 3.7 1.1 3.4 4.5 2.7 2.0 5.8 2.9 4.3 4.1 3.1 4.8 2.6 3.8 2.3 3.3 3.8 3.7 1.7 6.0 7×3 7179 WHT 34.6 16.5 39.2 R 4.9 1.1 8.5 5.7 3.2 5.0 0.7 1.2 5.7 9.3 7.9 3.0 3.8 1.5 3.5 5.0 2.8 1.9 7.4 3.3 4.4 4.1 3.0 5.2 3.0 4.2 2.8 3.4 4.2 4.3 2.0 6.6 5×3 34.2 36.6 16.1 7204 WHT

TAL SVL PAL NS MN MFE MBE LAL IO IN HW HL FOL FL V FL Measurements (in mm) of holotype (WHT 7204) and 4 paratypes (WHT 7179, WHT 7178 (now NMSL uncat.), WHT 7180, WHT 7215) of 7215) WHT 7180, WHT uncat.), NMSL (now 7178 WHT 7179, (WHT paratypes 4 and 7204) (WHT holotype of mm) (in Measurements 5. Table (WHT 7217) and paratype (WHT 7209) of – indicates tail lost . R indicates tail regenerated; B indicates tail broken; of C. pava ; and holotype (WHT 7219) paratype 7349) menikay. FL IV FL UEW FL III FL UAL FL II FL FL I FL TYE (×10) FEL TYD ES TL V TL EN TL IV TL ED TL III TL DFE TL II TL DBE TBW TL I TL AG

TBL

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Cnemaspis silvula new species see Table 5). Head relatively short (HL 26.9% of (Figures 34a–g, 35; Tables 5, 6; Plate 10d) SVL), flattened (HW 16.7% of SVL, HW 62.3% of HL), distinct from neck. Snout length a little greater Holotype. WHT 7208 (male), 28.6 mm SVL, Kombala- than eye diameter (ED 77.1% of ES); interorbital Kottawa Forest Reserve, Kottawa, Galle District, narrow (IO 5.1% of HL); pupil rounded. Granules Southern Province of Sri Lanka, 06º06’N, 80º20’E, 60 on snout keeled, larger than those of interorbital m, coll. S. Batuwita, 24 October, 2005. region; conical tubercles on occipital. Rostral scale slightly divided by a medial groove, in contact with Paratypes. WHT 7206, (female), 31.4 mm SVL, WHT first supralabial (Fig. 34a). Nares separated by two 7210 (female), 31.9 mm SVL, NMSL (formerly WHT enlarged supranasals and three internasal scales, 7207), (male), 26.0 mm SVL, WHT 7217, (male), the anterior one enlarged, extending towards snout 30.9 mm SVL, WHT 7209, (male), 30.3 mm SVL, (Fig. 34a). Nares rounded, dorsally orientated, not WHT 7216 (male), 19.0 mm SVL, same location and in contact with first supralabial. Two postnasals, collection data as holotype, 10 October, 2004. the lower one in broad contact with first supralabial on left side, a single scale separates first supralabial Other material. WHT 7220 (female), Kanneliya and lower post-nasal on right side (Fig. 34a). Mental Forest Reserve, Galle District, Southern Province, subpentagonal, wider than long, truncate posteriorly; 06º15’N, 80º20’E, 150 m, coll. S. Batuwita, 24 July, posteriolaterally in contact with two enlarged 2005; WHT 7333, Nawinna, Galle District, Southern postmentals and medially with a single hexagonal Province, 06º04’N, 80º12’E, 5 m, coll. S. Batuwita, 10 postmental scale; postmentals bordered posteriorly February, 2001; WHT 7018, Rumassala, near Galle, by 7 smooth scales including medial scale (Fig. 34b). Galle District, Southern Province, 06º01’N, 80º14’E, 5 Supralabials 7, infralabials 8, decreasing in size m, coll. M. M. Bahir, 04 November, 1995; WHT 7202, towards gape (Fig. 34c). Ear opening oval, higher 7203, Pituwala near Elpitiya, Beraliya Forest Reserve, than wide, oblique (TW 40.0% of TH). Scales on top Galle District, Southern Province, 06º15’N, 80º12’E, and sides of neck, area around ear, chin, and throat, 80 m, coll. S. Batuwita & M. Meegaskumbura, 04 except for mental and postmentals, granular, weakly June, 2005; WHT 7354, Pitadeniya, Sinharaja World carinate; 5 conical tubercles laterally between ear and Heritage Site, 06º22’30”N, 80º28’E, 300 m, coll. S. insertion of forelimb. Batuwita, 04 October, 2001; WHT 7027, Koskulana, Body slender, relatively long (AG 42.6% SVL). Ratnapura District, Sabaragamuwa Province, Mid-dorsal granules heterogeneous, unkeeled, few 06º25’N, 80º 27’E, 460 m, coll. D. Gabadage, 09 April, scales keeled mixed with slightly enlarged conical 1994; NMSL RG 17, (17 ex., 12 males and 5 females), tubercles (Fig. 34d); a row of 10–15 spine-like Hunuwela, no collection data; NMSL 011, Atweltota, irregular tubercles on flank; pectoral and abdominal coll. Mendis Wickramasinghe. scales strongly keeled, wider than long, pointed (Fig. 34e). Ventrals, 138; scales across midbody, 20; scales Diagnosis around vent (Fig. 34f) and on base of tail keeled, Cnemaspis silvula differs from its peninsular Indian imbricate; 3 preanal and 4 femoral pores on each and Sri Lankan congeners by a combination of side (Fig. 34f). the following characters. Maximum SVL 28.6 mm; Forelimbs moderately long, slender (LAL 14.6% paired postmentals separated by a medial scale; each SVL, UAL 12.2% SVL); hind limbs long (TBL 18.8% postmental bounded by 4 scales including medial SVL, FEL 21.6% SVL); tibia short. Scales on dorsal scale; ventrals, 132–138; ventral scales across mid- and anterior surfaces of upper arm keeled, those body, 20; dorsal scales heterogeneous; a row of 10–15 on posterior surface with both smooth and carinate irregular spinous tubercles on flank; ventrals, 138, scales; unkeeled granules on ventral surface. Scales keeled; 3 preanal pores, 4 femoral pores on each side; on anterior and ventral surfaces of thigh carinate subcaudals keeled, those on median row enlarged, (Fig. 34f); unkeeled granules on posterior surface. irregular; supralabials to angle of jaws, 7; subdigital Scales on dorsal surface of shank mostly conical, lamellae on digit IV of pes, 18; (in life) dorsum dark- some scales keeled; those on posterior and ventral brown to light-brown, with about seven indistinct, surfaces carinate. Scales on dorsal surface of lighter, V-shaped markings; occipital region with manus and pes keeled, on ventral surface smooth, lighter spots. sometimes granular. Digits elongate, slender, all bearing slightly recurved claws; subdigital lamellae Description of holotype entire, except for 2 or 3 fragmented scales at base, An adult male, 28.6 mm SVL (for measurements, unnotched; subdigital lamellae on finger I, 10, finger

84 Zeylanica Re v i s i o n o f Cn e m a s p i s a b

c d

e f

g

Figure 34. Cnemaspis silvula: WHT 7208, holotype, 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 and ventral regions, respectively, of mid-body; f, preanal and right hind-limb regions, showing arrangement of femoral and preanal pores; g,arrangement of subcaudal scales. Scale bars: 1 mm.

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II, 12, finger III, 15, finger IV, 15, finger V, 14; toe I, 10, Variation. In paratype, WHT 7209: supralabials, 8; toe II, 15, toe III, 16, toe IV, 18, toe V, 17; interdigital infralabials, 7; subdigital lamellae on finger I, 11, webbing absent; relative length of digits (fingers): IV finger II, 13, finger III, 15, finger IV, 16, finger V, 14; (2.5 mm) > III (2.3 mm) > V (2.1 mm) ~ II (2.1 mm) > toe I, 11, toe II, 14, toe III, 16, toe IV, 18, toe V, 15; 2 I (1.4 mm); (toes) IV (3.4 mm) > III (3.0 mm) > V (2.8 preanal pores and 5 femoral pores in each side; pores mm) > II (2.5 mm) > I (1.5 mm). absent in females. Ventrals, 132–138 (5 ex.); SVL range Tail tapering, its length greater than SVL (TAL from 19.0 to 31.9 mm. NMSL .011, with 4 preanal 122.7% of SVL); tail base distinctly swollen, dorsally pores separated medially by an unpored scale. and laterally covered with smooth, unequal, weakly carinate granules; large, conical, keeled tubercles Distribution and natural history. Cnemaspis silvula present on base of tail. Dorsal scales on tail keeled, is distributed throughout Sri Lanka’s lowland ‘wet imbricate, directed backwards. Tail with 6–8 conical zone’ (rainfall > 2,000 mm yr-1) from Kalutara to the enlarged tubercles; about 13 whorls of enlarged Galle and Matara Districts. Specimens have been tubercles on unregenerated tail; a single conical recorded from Kottawa, Kaneliya, Beraliya and post-cloacal spur present on each side. Subcaudal Sinharaja Foreset Reserves (including Koskulana and scales strongly keeled, pointed, those on median Pitadeniya) (Fig. 35). Cnemaspis silvula appears to be row enlarged, with pairs of smaller scales between most active around 1000 h, specimens being seen individual larger ones (Fig. 34g). on tree trunks, especially the bark of Dipterocarpus hispidus, and on moist rock faces. A female measuring Coloration. In preservative: dorsally dusky brown 35.0 mm SVL in Kottawa Forest Reserve laid two with distinct dark markings on paravertebral region hard-shelled eggs, each of diameter 5.0 mm. One of and on to tail; light-brown patches on limbs; venter these hatched after 56 days. The following lizards dusky brown. were observed in sympatry: Hemidactylus frenatus, H. In life: dorsum dark brown to light brown, with about depressus, Calotes liolepis, Otocryptis wiegmanni, Lyrio­ seven indistinct, V-shaped markings; occipital region cephalus scutatus, Lankascincus fallax, L. gansi, Eutropis with a light-yellow spot between two chevrons; a carinata lankae, E. macularius macularius. black spot on neck; venter white (see Plate 10d). Etymology. The species name is the Latin diminutive of ‘forest’, a reference to the habitats in which the species occurs, applied here as a noun in ap­ position.

Comparisons. Cnemaspis silvula differs from each of its Sri Lankan and peninsular Indian congeners, respectively, by the following opposing suites of characters: Cnemaspis tropidogaster: dorsal scales heterogeneous with pointed, keeled tubercles. Cnemaspis kandiana and C. menikay: pectoral and abdominal scales smooth. Cnemaspis podihuna: subcaudals on median row enlarged, subequal; 6 femoral pores. Cnemaspis scalpensis, C. gemunu and C. phillipsii: 12–16 femoral pores. Cnemaspis alwisi: 9 femoral pores. Cnemaspis molligodai: 5 preanal and 8 or 9 femoral pores, gular, pectoral and abdominal scales smooth, dorsal scales homogeneous; subcaudals on median row enlarged, subequal. Cnemaspis samanalensis: gular, pectoral and abdominal scales tricarinate. Cnemaspis retigalensis: pectoral and abdominal scales smooth, a single preanal pore and 3 or 4 femoral pores. Cnemaspis punctata: 5–7 femoral pores; distinctive, spotted dorsal coloration. Cnemaspis kumarasinghei, Figure 35. Distribution of Cnemaspis silvula in south-western pectoral and abdominal scales smooth. Cnemaspis Sri Lanka; the type locality is indicated by a solid symbol. clivicola: subcaudals in median series not enlarged.

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Cnemaspis upendrai and C. pulchra: ventral scales on Description of holotype tail base smooth. Cnemaspis kallima: abdominal scales An adult male, 30.4 mm SVL (for measurements, see smooth. Cnemaspis latha: pectoral and abdominal Table 4). Head relatively short (HL 26.3% of SVL), scales smooth. Cnemaspis pava: dorsal enlarged flattened (HW 16.7% of SVL, HW 63.7% of HL), tubercles subtrihedral. Cnemaspis amith: pectoral and distinct from neck. Snout length moderately greater abdominal scales smooth. than eye diameter (ED 65.7% of ES); interorbital Cnemaspis beddomei: femoral pores absent. moderate (IO 8.7% of HL); pupil rounded. Granules Cnemaspis goaensis: ventral scales smooth. Cnemaspis on snout keeled, larger than those on interorbital littoralis: dorsal scales homogeneous, subcaudals region; scales on interorbital area keeled; occipital, on median row enlarged, hexagonal. Cnemaspis top and sides of neck, and area around ear covered my­soriensis: dorsal scales homogeneous, three rows by conical tubercles; some scales on ventro-lateral of enlarged subcaudals. Cnemaspis indraneildasii: region keeled; scales on ventral surface of head ventral, gular and caudal scales smooth. Cnemaspis granular, unkeeled. Rostral scale partially divided by heteropholis: dorsal scales heterogeneous, ventral, a medial groove, in contact with first supralabial (Fig. gular and caudal scales smooth. Cnemaspis indica, 36a). Nares separated by two enlarged supranasals C. nairi, C. otai, C. yercaudensis, C. sisparensis and and two internasal scales, the anterior one enlarged C. wy­nadensis: spine-like tubercles absent on flank. and extending towards snout tip (Fig. 36a). Nares Cnemaspis boiei: ventrals smooth, no preanal or rounded, dorsally orientated, not in contact with femoral pores. Cnemaspis gracilis: ventrals smooth. first supralabial (Fig. 36a). Two postnasals, the lower Cnemaspis ornata: no femoral pores, dorsal tubercles one in broad contact with first supralabial (Fig. 36a). arranged in 15 longitudinal rows. Cnemaspis wicksii: Mental subtriangular, wider than long, truncate upper arm and lower arm, thigh and tibia dorsally posteriorly, posteriolaterally in contact with two covered by smooth scales. Cnemaspis andersonii: enlarged postmentals and medially with a single pectoral and abdominal scales smooth. Cnemaspis small hexagonal postmental scale; postmentals jerdonii: no preanal pores. Cnemaspis monticola: bordered posteriorly by 5 smooth scales including pec­toral and abdominal scales smooth. Cnemaspis medial scale (Fig. 36b). Supralabials 7, infralabials australis: scales on preanal region smooth. Cnemaspis 7, decreasing in size towards gape (Fig. 36c). Ear nilagirica: dorsal scales homogeneous; subcaudals on opening oval, higher than wide, oblique (TW 40.0% median row enlarged, subequal. of TH). Scales on ventral surface of neck smooth, imbricate; 3 conical tubercles laterally between ear and forelimb insertion region. Cnemaspis latha new species Body slender, long (AG 46.7% SVL). Mid-dorsal (Figures 36a–g, 37; Tables 4, 6; Plate 9c) granules homogeneous, isolated and scattered, very few carinate granules and enlarged tubercles Holotype. WHT 7214 (male), 30.4 mm SVL; (Fig. 36d); 6 spine-like tubercles on flank; pectoral Bandarawela, Badulla District, Central Province of and abdominal scales smooth (Fig. 36e). Ventrals, Sri Lanka, 06º50’N, 80º59’E, 1220 m, coll. S. Batuwita, 111, longer than wide, rounded or bluntly pointed 01 May, 2006. (Fig. 36e); ventral scales across midbody, 13; scales around vent smooth (Fig. 36f), those on base of tail Diagnosis smooth, imbricate; 2 preanal and 4 femoral pores on Cnemaspis latha differs from its peninsular Indian each side (Fig. 36f). and Sri Lankan congeners by a combination of Forelimbs moderately long, slender (LAL 13.4% the following characters. Maximum SVL 30.4 mm; SVL, UAL 12.8% SVL); hind limbs long (TBL 17.1% paired postmentals separated by a medial scale; each SVL, FEL 20.0% SVL); tibia short. Scales on dorsal postmental bounded by 3 scales including medial and anterior surfaces of upper arm keeled; rear scale; ventrals, 111; ventral scales across mid-body, surface of upper arm with both smooth and carinate 13; dorsal scales homogeneous; 6 spine-like tubercles scales, ventral surface with unkeeled granules. Scales present on flank; ventrals smooth; 2 preanal pores, on dorsal surface of thigh smooth, weakly carinate; 4 femoral pores on each side; subcaudals smooth, those on anterior surface carinate; posterior and median row enlarged, irregular; supralabials to ventral surfaces with unkeeled granules. Scales on angle of jaws, 7; subdigital lamellae on digit IV of dorsal surface of shank granular; those on anterior pes, 17; (in life) dorsum dusky-brown, with about and posterior surfaces weakly keeled, smooth, five irregular bands with a middle paler spot; nape granular. Dorsal scales on manus and pes weakly with a dark spot. keeled; ventral scales granular, imbricate. Digits

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Figure 36. Cnemaspis latha: WHT 7214, holotype, 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 and ventral regions, respectively, of mid-body; f, preanal and right hind-limb regions, showing arrangement of femoral and preanal pores; g, arrangement of subcaudal scales. Scale bars: 1 mm.

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elongate, slender, all bearing slightly recurved claws; Distribution and natural history. Cnemaspis latha was subdigital lamellae entire, unnotched; subdigital observed in edificarian habitats at the type locality lamellae on finger I, 9, finger II, 11, finger III, 14, (altitiude ~1200 m; rainfall > 2,500 mm yr-1), although finger IV, 15, finger V, 13; toe I, 9, toe II, 12, toe III, we suspect it occurs also in the forested areas around 15, toe IV, 17, toe V, 13; interdigital webbing absent; Bandarawela (Fig. 37). relative length of digits (fingers): IV (2.7 mm) ~ V (2.7 mm) > III (2.5 mm) > II (2.2 mm) > I (1.7 mm); Etymology. The species name is a feminine noun (toes) IV (3.6 mm) > V (3.3 mm) > III (3.2 mm) > II transliterated from the Sinhala, meaning pretty, (2.6 mm) > I (1.7 mm). elegant, beautiful: often used also as a proper noun, Tail tapering, its length greater than SVL; tail base it is here applied as a substantive in apposition. distinctly swollen; dorsal and lateral scales granular, unkeeled. Enlarged, conical, keeled tubercles present Comparisons. Cnemaspis latha differs from each of on base of tail, about as large as granules on mid-body. its Sri Lankan and peninsular Indian congeners, Dorsal scales on tail heterogeneous, weakly keeled, respectively, by the following opposing suites of directed backwards. Tail with about 6–8 enlarged, characters: pointed tubercles forming whorls; a single conical Cnemaspis tropidogaster: dorsal scales hetero­ post-cloacal spur present on each side; subcaudals geneous with pointed, keeled tubercles; ventrals smooth, median row with pairs of smaller obtuse keeled. Cnemaspis kandiana and C. menikay: gular scales between single larger subpentagonal or obtuse scales carinate. Cnemaspis podihuna: subcaudals on ones (Fig. 36g). median row enlarged, hexagonal; 6 femoral pores. Cnemaspis scalpensis, C. gemunu and C. phillipsii: Coloration. In preservative: dorsally light brown 12–16 femoral pores. Cnemaspis alwisi: 9 femoral with dark, irregular blotches; area around nape area pores. Cnemaspis molligodai: 5 preanal and 8 or 9 with dark markings (a dark spot on nape itself); gular femoral pores; subcaudals on median row enlarged, region dark; venter dusky white. In life: dorsum subequal. Cnemaspis samanalensis: gular, pectoral and dusky-brown, with about five irregular bands and a abdominal scales tricarinate, subcaudals keeled. paler spot near mid-dorsum (see Plate 9c). Cnemaspis retigalensis: gulars carinate, a single preanal pore and 3 or 4 femoral pores. Cnemaspis punctata: 5–7 femoral pores; distinctive spotted dorsal coloration. Cnemaspis kumarasinghei: ventrals, 121–133; 5 distinct cross-bars on dorsum. Cnemaspis silvula, C. pulchra, C. pava, C. clivicola and C. upendrai: pectoral and abdominal scales keeled. Cnemaspis kallima: subcaudals on distal half of tail carinate. Cnemaspis amith: gular scales carinate. Cnemaspis beddomei: preanal pores 6–9, no femoral­ pores. Cnemaspis goaensis: gulars keeled. Cnemaspis­ littoralis: subcaudals on median row en­larged, hexagonal. Cnemaspis mysoriensis: three rows of enlarged subcaudals, 2 or 3 femoral pores. Cnemas­pis indraneildasii: dorsal scales of both limbs smooth; ventral, gular and caudal scales smooth; 20 mid-ventral scales. Cnemaspis heteropholis: dorsal scales heterogeneous. Cnemaspis indica, C. nairi, C. otai, C. yercaudensis, C. sisparensis and C. wynadensis: spine-like tubercles absent on flank. Cnemaspis boiei: no preanal or femoral pores. Cnemaspis gracilis: 2 preanal­ pores separated by 2 scales, 4 femoral pores. Cnemas­pis ornata: no femoral pores, dorsal tubercles ar­ranged in 15 longitudinal rows. Cnemaspis wicksii: dorsal scales of upper arm and lower arm smooth. Cnemaspis andersonii: gulars carinate. Cnemaspis Figure 37. Distribution of Cnemaspis latha in Sri Lanka. jerdonii: no preanal pores. Cnemaspis monticola: gulars carinate. Cnemaspis australis: subcaudals pointed,

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strongly keeled. Cnemaspis nilagirica: dorsal scales un­ life) dorsum dark to light brown, with about three keeled; pectoral and abdominal scales weakly keeled; quadrate dusky-yellow markings; tail with several subcaudals on median row enlarged, subequal. pale bands; venter dusky white or yellow.

Remarks Description of holotype Although we have described it on the basis of the An adult male, 28.8 mm SVL (for measurements, see holotype alone, we are confident that C. latha is a Table 5). Head relatively short (HL 27.0% of SVL), valid species given that it is clearly diagnosable flattened (HW 17.7% of SVL, HW 65.3% of HL), from all other Sri Lankan Cnemaspis. Of its insular distinct from neck. Snout length a little greater than congeners, the ones it resembles most closely are C. eye diameter (ED 74.2% of ES); interorbital narrow kumarasinghei and C. kallima. While the type localities (IO 6.4% of HL); pupil rounded. Granules on snout of C. latha and C. kumarasinghei are separated by keeled, larger than those on interorbital region; scales ~80 km, the two species can also be distinguished of interorbital region keeled; granules intermixed by the former having 111 ventrals (vs 121–133 in C. with enlarged conical tubercles on occipital, top kumarasinghei); and lacking cross-bars (vs 5 distinct and sides of neck, and around ear. Chin and throat cross-bars present on dorsum). Cnemaspis latha is with unkeeled granules; scales of gular area weakly distinguished from C. kallima by the absence of keeled keeled, imbricate. Rostral scale partially divided by a subcaudals on median row of the distal half of the tail medial groove, in contact with first supralabial (Fig. (vs keeled subcaudals present in C. kallima); and by 38a). Nares separated by two enlarged supranasals having smooth gular scales (vs keeled gular scales); and a single internasal scale; internasal extends its type locality, Bandarawela, is also ~115 km distant towards snout; 4 slightly enlarged scales behind from that of C. kallima (Rattota). supranasal-internasal row (Fig. 38a). Nares rounded, dorsally orientated; naris not in contact with first Cnemaspis kallima new species supralabial­ (Fig. 38a). Two postnasals, the lower (Figures 38a–h, 39; Tables 5, 6; Plate 10a–c) one in broad contact with first supralabial (Fig. 38a). Mental subpentagonal, truncate posteriorly; wider Holotype. WHT 7245 (male), 28.8 mm SVL; than long, posteriolaterally in contact with two Gammaduwa Estate, Gammaduwa near Rattota, enlarged postmentals and medially with a single Matale District, 07º34’N, 80º42’ E, 760 m, coll. S. small hexagonal postmental scale; postmentals Batuwita & P. Wickramathilaka, 23 March 2007. bordered posteriorly by 7 smooth scales including medial scale (Fig. 38b). Supralabials 7, infralabials, Paratypes. WHT 7239, (female), 29.8 mm SVL, 7255, 7, decreasing in size towards gape (Fig. 38c). Ear (female), 27.6 mm SVL, WHT 7222, (female), 33.5 mm opening oval, higher than wide, oblique (TW 60.0% SVL, 7253, (female), 32.0 mm SVL, 7227, (female), 23.2 of TH). Scales on ventral surface of neck feebly mm SVL, 7254, (male), 24.2 mm SVL, 7252, (female), carinate; 6 conical tubercles laterally, between ear 28.7 mm SVL, 7230, (female), 34.2 mm SVL, 7229, and insertion of forelimb. (female), 35.1 mm SVL, 7249, (male), 25.2 mm SVL, Body slender, relatively long (AG 44.7% SVL). 7251, (male), 34.4 mm SVL, 7228, (male), 31.9 mm Mid-dorsal granules heterogeneous, isolated and SVL, same collection data as holotype. scattered, intermixed with large, conical, weakly- keeled or unkeeled tubercles (Fig. 38d); 15 irregular Diagnosis rows of spine-like tubercles on flank; ventro-lateral Cnemaspis kallima differs from its peninsular Indian scales enlarged, flattened, smooth; pectoral and and Sri Lankan congeners by a combination of abdominal scales smooth (Fig. 38e). Ventrals, 133, the following characters. Maximum SVL 35.1 mm; circular, wider than long; scales across midbody, paired postmentals separated by a medial scale; each 21; scales around vent and base of tail smooth, postmental bounded by 4 scales including medial imbricate; 3 preanal and 4 femoral pores on each scale; ventrals, 133–135; ventral scales across mid- side (Fig. 38f). body, 21; dorsal scales heterogeneous; an irregular Forelimbs moderately long, slender (LAL 14.2% row of 15 spine-like tubercles present on flank; SVL, UAL 13.1% SVL); hind limbs long (TBL 18.7% ventrals smooth; 3 preanal pores, 4 femoral pores SVL, FEL 20.1% SVL); tibia short. Dorsal surface on each side; anterior half of subcaudals smooth, of upper arm with both keeled and smooth scales; posterior half carinate, median row enlarged, posterior surface with mostly smooth scales and irregular; supralabials to angle of jaws, 7 or 8; few carinate ones; ventral scales unkeeled, granular. subdigital lamellae on digit IV of pes, 18 or 20; (in Scales on dorsal and anterior surfaces of lower arm

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Figure 38. Cnemaspis kallima: WHT 7245, holotype, 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 and ventral regions, respectively, of mid-body; f, preanal and left hind limb regions, showing arrangement of femoral and preanal pores; g, h, arrangement of smooth and keeled subcaudal scales respectively of, proximal and distal portions of tail. Scale bars: 1 mm.

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keeled; those on ventral surface unkeeled, granular. Variation. In paratype, WHT 7228: supralabials, 8; Scales on dorsal surface of thigh smooth, a (very) few infralabials, 8; subdigital lamellae on finger I, 10, weakly carinate; those on anterior surface imbricate, finger II, 12, finger III, 15, finger IV, 15, finger V, 14; carinate; those on posterior surface unkeeled, toe I, 10, toe II, 13, toe III, 17, toe IV, 18, toe V, 16; 3 granular; scales on ventral surface of thigh smooth, preanal and 4 femoral pores in each side. SVL range imbricate. Dorsal surface of tibia with unkeeled 23.2–35.1 mm (14 ex); pores absent in females. granules; scales on anterior, posterior and ventral Tail length greater than SVL in paratype WHT surfaces imbricate, carinate. Dorsal scales on manus 7254; tail base distinctly swollen, dorsally with and pes keeled, those on ventral surface imbricate, enlarged, elevated or conical scales, laterally with smooth, granular. Dorsal surface of digits with unkeeled granules. Dorsal scales on tail similar to weakly-keeled scales. Digits elongate, slender, all those on tail base, directed backwards; tail with bearing slightly recurved claws. Subdigital lamellae about 6–8 enlarged conical, pointed scales forming entire, unnotched; subdigital lamellae on finger I, 13 whorls in paratype WHT 7254; a single conical 11, finger II, 13, finger III, 16, finger IV, 17, finger V, post-cloacal spur on each side (Fig. 38g); subcaudals 15; toe I, 11, toe II, 14, toe III, 16, toe IV, 20, toe V, 18; on anterior half of tail smooth, those on posterior half interdigital webbing absent; relative length of digits carinate, with pairs of smaller scales between single (fingers): IV (2.9 mm) > V (2.4 mm) > III (2.8 mm) > larger obtuse ones in median series (Fig. 38h). II (2.4 mm) > I (1.5 mm); (toes) IV (3.7 mm) > III (3.2 mm) > V (3.1 mm) > II (2.6 mm) > I (1.5 mm). Distribution and natural history. Cnemaspis kallima Tail broken (see Variation for description of appears to be a ground-dwelling gecko. Individuals tail). were observed mostly under moist rocks, never on trees or on rock faces at Gammaduwa Estate in the Coloration. In preservative: dorsally olive brown vicinity of Knuckles Range (Fig. 39, Plate 10b). This with indistinct dark markings prominent on nape gecko exhibits communal egg laying (Plate 10c). (some individuals with a pale paravertebral stripe); The following lizards were observed in sympatry: gular area dark, venter dusky white. In life: dorsum Cyrtodactylus soba, Geckollea triedra, Cnemaspis punc­ light brown, with about six confluent light-brown tata, C. phillipsi, L. taylori, Otocryptis wiegmanni, and blotches; nape with a distinct black marking (Plate Calotes liocephalus. 10a). Etymology. The species name is derived from the Greek noun καλλοσ (kallos, = beauty), applied here as a feminine diminutive noun in apposition.

Comparisons. Cnemaspis kallima differs from each of its Sri Lankan and peninsular Indian congeners, respectively, by the following opposing suites of characters: Cnemaspis tropidogaster: pectoral and abdominal scales keeled. Cnemaspis kandiana: gular scales carinate. Cnemaspis podihuna: subcaudals on median series enlarged, hexagonal. Cnemaspis scalpensis, C. gemunu and C. phillipsii: 12–16 femoral pores. Cnemaspis alwisi: 9 femoral pores. Cnemaspis molligodai: 5 preanal and 8 or 9 femoral pores, gular, pectoral and abdominal scales smooth, dorsal scales homogeneous; subcaudals on median row enlarged, subequal. Cnemaspis samanalensis: gular, pectoral and abdominal scales tricarinate. Cnemaspis retigalensis: gulars carinate, a single preanal pore and 3 or 4 femoral pores. Cnemaspis punctata: 5–7 femoral pores; distinctive spotted dorsal coloration. Cnemaspis kumarasinghei­ and C. latha: subcaudals smooth. Figure 39. Distribution of Cnemaspis kallima: the Knuckles Cnemaspis silvula, C. clivicola, C. pulchra, C. pava and Hills, Sri Lanka. C. upendrai: pectoral and abdominal scales keeled.

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Cnemaspis menikay: subcaudals smooth. Cnemaspis subcaudals in median series not enlarged, keeled; amith: gular scales carinate. supralabials to angle of jaws, 7 or 8; subdigital Cnemaspis beddomei: preanal pores 6–9, no femoral lamellae on digit IV of pes, 17–20. pores. Cnemaspis goaensis: gulars keeled. Cnemaspis littoralis: body slender, dorsal scales homogeneous; Description of holotype subcaudals on median row enlarged, hexagonal. An adult male, 34.2 mm SVL (for measurements, Cnemaspis mysoriensis: dorsal scales homogeneous, see Table 4). Head long (HL 27.7% of SVL), broad three rows of enlarged subcaudals. Cnemaspis indra­ (HW 16.6% of SVL, HW 60.0% of HL), distinct neildasii: dorsal scales heterogeneous. Cnemaspis from neck. Snout length a little greater than eye heteropholis: no post-cloacal spurs, dorsal scales diameter (ED 74.3% of ES); interorbital narrow (IO heterogeneous.­ Cnemaspis indica, C. nairi, C. otai, 5.2% of HL); pupil rounded. Granules on snout C. yercaudensis, C. sisparensis and C. wynadensis: oval, keeled; scales immediately above supralabials spine-like tubercles absent on flank. Cnemaspis boiei: carinate, larger than those of occipital region; scales no preanal or femoral pores. Cnemaspis gracilis: on interobital, supercilium and ventral side of head 2 preanal pores separated by 2 scales, 4 femoral carinate, granular. Rostral scale partially divided by pores. Cnemaspis indica: spine-like tubercles absent a medial groove, in contact with first supralabial (Fig. on flank. Cnemaspis ornata: no femoral pores, 40a). Nares separated by two enlarged supranasals dorsal tubercles arranged in 15 longitudinal rows. (Fig. 40a). Nares rounded, dorso-laterally orientated, Cnemaspis wicksii: subcaudals smooth. Cnemaspis not in contact with first supralabials (Fig. 40a). Two andersonii: gulars carinate. Cnemaspis jerdonii: no postnasals, the lower one in broad contact with first preanal pores. Cnemaspis monticola: gulars carinate. supralabial (Fig. 40a). Mental subpentagonal, wider Cnemaspis australis: no spine-like tubercles on flanks; than long, truncate posteriorly, posteriolaterally subcaudals strongly keeled. Cnemaspis nilagirica: in contact with two elongate postmentals and dorsal scales homogeneous; pectoral and abdominal medially with a single pentagonal postmental scale; scales weakly keeled; subcaudals on median row postmentals bordered posteriorly by 5 smooth scales enlarged, subequal. including medial scale (Fig. 40b). Supralabials 8, infralabials 7, decreasing in size towards gape (Fig. Cnemaspis pulchra, new species 40c). Ear opening oval, higher than wide, oblique (Figures 40a–g, 41; Tables 4, 6; Plate 12a) (TW 33.3% of TH). Scales on ventral side of chin, throat and neck carinate. Obtuse tubercles (a few of Holotype. WHT 7023, (male), 34.2 mm SVL, them keeled) on side of neck, between ear and eye, Rakwana, Ratnapura District, 06o30’N, 80o34’E, 458m, 6; between ear and forelimb insertion, 11. M. M. Bahir & D. Gabadage, 06 March, 1996. Body slender, elongate (AG 43.5% SVL). Mid- dorsal granules conical, weakly keeled, mixed with Paratypes. WHT 1573a, (male), 33.6 mm SVL; WHT large conical keeled tubercles (Fig. 40d); 6 spine- 7021, (male), 32.1 mm SVL, 7022, (male), 29.0 mm like tricarinate tubercles on flank; pectoral scales SVL, same location data as holotype, M. M. Bahir & strongly keeled; abdominal scales weakly keeled D. Gabadage, 14 March, 1996; WHT 7011, (male), 32.1 (Fig. 40e). Ventrals, 133; scales across midbody, 24; mm SVL, Morningside Forest Reserve, adjacent to scales around vent weakly keeled; those on base of east of the Sinharaja World Heritage Site, Ratnapura tail smooth, imbricate. Four preanal pores separated District, 06o24’N, 80o38’E, 1060 m, coll. D. Gabadage medially by a single unpored scale; 4 femoral pores & Dharmasiri, 21 February, 1996. on each side (Fig. 40f). Forelimbs moderately long, slender (LAL 14.0% Diagnosis SVL, UAL 11.4% SVL); hind limbs relatively long Cnemaspis pulchra differs from its peninsular Indian (TBL 19.0% SVL, FEL 19.8% SVL); tibia long. Dorsal and Sri Lankan congeners by a combination of scales on both fore and hind limbs carinate; ventral the following characters. Maximum SVL 34.2 mm; scales on upper arm granular, not keeled (a few scales paired postmentals separated by a medial scale; carinate); insertion of upper arm with tricarinate each postmental bounded by 2 scales; ventrals, scales; lower arm ventrally with keeled, imbricate 121–133; ventral scales across mid-body, 24–27; dorsal scales. Scales on anterior surface of thigh imbricate, scales heterogeneous, isolated, scattered; 6 spine- unicarinate or tricarinate; those on ventral surface like tubercles on flanks; pectoral and abdominal unicarinate (Fig. 40f). Scales on posterior surface of scales smooth; ventral scales of tail base smooth; 3 thigh conical, unkeeled, granular. Scales on ventral or 4 preanal pores, 4–6 femoral pores on each side; surface of shank keeled, imbricate; those on dorsal

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Figure 40. Cnemaspis pulchra: WHT 7023, holotype, 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 and ventral regions, respectively, of mid-body; f, preanal and left hind limb regions, showing arrangement of femoral and preanal pores; g, arrangement of subcaudal scales. Scale bars: 1 mm.

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surface of manus and pes smooth, on ventral surface Variation. In paratype, WHT 1573a: supralabials, 9; smooth, imbricate, unkeeled, granular. Digits infralabials, 8; ventrals, 27; ventral scales across mid- elongate, slender, all bearing slightly recurved claws; body, 121; 3 preanal pores; 3 femoral pores on each subdigital lamellae entire, unnotched; subdigital side; subdigital lamellae on finger I, 11, finger II, 14, lamellae on finger I, 12, finger II, 13, finger III, 15, finger III, 16, finger IV, 16, finger V, 15, and toe I, 11, finger IV, 16, finger V, 14; toe I, 11, toe II, 14, toe III, toe II, 16, toe III, 19, toe IV, 20, toe V, 19. SVL range 16, toe IV, 17, toe V, 17; interdigital webbing absent; 29.0–34.2 mm (6 ex.); 3–5 and 4–6 femoral pores on relative length of digits (fingers): II (3.5 mm) > III each side. (3.3 mm) > I (3.0 mm) > IV (2.7 mm) > V (1.7 mm); (toes) IV (4.8 mm) > III (4.0 mm) > V (3.9 mm) > II Distribution and natural history notes. Cnemaspis (3.5 mm) > I (2.1 mm). pulchra was observed on tree trunks and rock faces at Tail tapering, its length greater than SVL (TAL Rakwana and Morningside Forest Reserve (altitiude 121.6% of SVL); tail base distinctly swollen. Eight ~1,000 m) on the eastern side of the Sinharaja World enlarged, conical, keeled tubercles on base of Heritage Site (Fig. 41). Exploration of this low- tail, about as large as granules on mid-dorsum. altitude cloud forest has brought to light a large Caudal scales at mid-tail keeled, imbricate, directed number of new vertebrate species, including the backwards. Tail with about 6 enlarged, keeled, gecko Cyrtodactylus subsolanus and the agamid lizard pointed tubercles separated from one another by Calotes desilvai, which together with C. liolepis, occur 2–5 scales. A single conical post-cloacal spur present in sympatry with C. pulchra. on each side. Subcaudals strongly keeled, those on median row slightly enlarged, with pairs of smaller Etymology. The species name is a Latin (feminine) scales between individual larger ones (Fig. 40g). adjective meaning beautiful, pretty, elegant. Coloration. In preservative: dorsum dusky brown with four W-shaped markings; similar marking Comparisons. Cnemaspis pulchra differs from each behind tail base; tail banded; nape with a distinct of its Sri Lankan and peninsular Indian congeners, spot; venter dusky white (see Plate 12a). respectively, by the following opposing suites of In life: Not known. characters: Cnemaspis tropidogaster: ventral scales on pre­ cloacal and tail base area carinate, no enlarged conic­al tubercles laterally on neck and dorsally on tail base. Cnemaspis kandiana, C. podihuna, C. scalpensis, C. retigalensis, C. gemunu, C. punctata, C. phillipsii, C. molligodai, C. kumarasinghei, C. kallima C. menikay and C. latha: pectoral and abdominal scales smooth. Cnemaspis samanalensis: gular, pectoral and abdominal scales tricarinate. Cnemaspis clivicola: subcaudals in median series not enlarged. Cnemaspis pava and C. silvula: ventral scales on precloacal and tail-base area carinate. Cnemaspis upendrai: some gul­ars tricarinate. Cnemaspis amith: pectoral and abdominal scales smooth. Cnemaspis beddomei: femoral pores absent. Cnemaspis goaensis: ventrals smooth. Cnemaspis littoralis: dorsal scales homogeneous, subcaudals on median series enlarged, hexagonal. Cnemaspis mysoriensis: dorsal scales homogeneous, three rows of enlarged subcaudals. Cnemaspis indraneildasii and C. heteropholis: ventral, gular and caudal scales smooth. Cnemaspis indica, C. nairi, C. otai, C. yercaudensis, C. sisparensis and C. wynadensis: spine-like tubercles absent on flank. Cnemaspis boiei: ventrals smooth, preanal and femoral pores absent. Cnemaspis gracilis: Figure 41. Distribution in Sri Lanka of Cnemaspis pulchra; ventrals smooth. Cnemaspis ornata: femoral pores the type locality is indicated by a solid symbol. absent, dorsal tubercles arranged in 15 longitudinal

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rows. Cnemaspis wicksii and C. andersonii: pectoral two enlarged supranasals and a single internasal and abdominal scales smooth. Cnemaspis jerdonii: scale (Fig. 42a). Nares rounded, dorsally orientated, preanal pores absent. Cnemaspis monticola: pectoral not in contact with first supralabial (Fig. 42a). Two and abdominal scales smooth. Cnemaspis australis: postnasals, the lower one in narrow contact with scales on preanal region smooth. Cnemaspis nilagirica: first supralabial (Fig. 42a). Mental subpentagonal, dorsal scales homogeneous; subcaudals on median longer than wide, posteriolaterally in contact with row enlarged, subequal. two elongate postmentals and medially with a single small postmental scale; postmentals bordered Cnemaspis menikay, new species posteriorly by 7 smooth scales (Fig. 42b). Supralabials (Figures 42a–h, 43; Tables 5, 6; Plate 12b) 7, infralabials 8, decreasing in size towards gape (Fig. 42c). Ear opening elliptical, higher than wide, oblique Holotype. WHT 7219, (male), 26.8 mm SVL, Ihala- (TW 25.0% of TH). Scales on ventral side of neck Kalugala, Allauwa, Kegalle District, Sabaragamuwa feebly carinate. Twelve obtuse enlarged tubercles Province of Sri Lanka, 07o15’N, 80o16’E, 300m, coll. S. between anterior margin of ear opening and posterior Batuwita, 19 April, 2006 margin of eye; 15 laterally on neck, larger than those of occipital region. Paratypes. WHT 7349, (male), 26.9 mm SVL, same Body slender, elongate (AG 47.0% SVL). Mid- location data as holotype, coll. A. S. N. Batuwita, 05 dorsal granules conical, mostly unkeeled (a few November, 2005; WHT 7218, (male), 28.0 mm SVL, keeled granules present), larger than those of same data as holotype. occipital region, mixed with conical enlarged, keeled tubercles (Fig. 42d); enlarged tubercles prominent on Diagnosis dorso-lateral region of body; 25 spine-like tubercles Cnemaspis menikay differs from its peninsular Indian on ventro-lateral region of body; pectoral and and Sri Lankan congeners by a combination of the abdominal scales as wide as long, imbricate, smooth following characters. Maximum SVL 28.0 mm; (Fig. 42e). Ventrals, 129; scales across midbody, 26; paired postmentals separated by a medial scale; scales around vent and at ventral tail-base smooth, each postmental bounded by 3 scales; ventrals, imbricate. A single preanal pore and 4 femoral pores 126–138; ventral scales across mid-body, 20–26; present (Fig. 42f). dorsal scales heterogeneous; 15 spine-like tubercles Forelimbs moderately long, slender (LAL 15.2% on flanks; ventrals smooth; 1 or 2 preanal pores, 3 SVL, UAL 14.9% SVL); hind limbs long (TBL 25.3% or 4 femoral pores per side; subcaudals in median SVL, FEL 17.9% SVL); tibia long. Dorsal scales on series not enlarged, keeled; supralabials to angle of both fore and hind limbs carinate; ventral scales on jaws, 7 or 8; subdigital lamellae on digit IV of pes, upper arm and lower arm smooth. Scales on anterior 15; (in life) dorsum dark-brown, with yellowish spots surface of thigh carinate, those on ventral surface laterally; uninterrupted dusky yellow mid-dorsal smooth (Fig. 42f); scales on posterior surface of thigh line; occipital region with three yellow spots. granular. Scales on ventral surface of shank carinate, granular; those on posterior surface keeled; ventral Description of holotype surface of shank with both smooth and keeled scales. An adult male, 26.8 mm SVL (for measurements, Dorsal scales on manus and pes carinate, ventral see Table 5). Head relatively long (HL 26.4% of ones granular, unkeeled. Digits elongate, slender, all SVL), broad (HW 17.1% of SVL, HW 64.7% of HL), bearing slightly recurved claws; subdigital lamellae distinct from neck. Snout length a little greater than entire, unnotched; subdigital lamellae on finger I, eye diameter (ED 70.5% of ES); interorbital moderate 9, finger II, 11, finger III, 13, finger IV, 14, finger V, (IO 7.0% of HL); pupil rounded. Granules on snout 12; toe I, 9, toe II, 12, toe III, 15, toe IV, 15, toe V, 14; keeled; scales immediately above supralabials interdigital webbing absent; relative length of digits smooth, larger than those of occipital region; scales (fingers): IV (2.5 mm) > III (2.4 mm) > V (2.0 mm) > on interorbital and supercilium keeled, smaller II (2.3 mm) > I (1.7 mm); (toes) IV (3.5 mm) > V (3.0 than those on snout; occipital region and area mm) > III (2.8 mm) > II (2.5 mm) > I (1.5 mm). around tympanum with conical granules, mixed Tail detached, tapering; regenerated tail in with enlarged conical tubercles; scales on ventral paratype WHT 7218 subequal to SVL (TAL 100.0% surface of chin unkeeled, granular; throat with of SVL); tail base distinctly swollen. Eight conical, conical granules, a few of them keeled. Rostral scale keeled, enlarged tubercles on base of tail, about partially divided by a medial groove, in contact as large as those on mid-dorsum. Caudal scales with first supralabial (Fig. 42a). Nares separated by at mid-tail conical, keeled, directed backwards; a

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Figure 42.Cnemaspis menikay: WHT 7219, holotype, 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 and ventral regions, respectively, of mid-body; f, preanal and left hind limb regions, showing arrangement of femoral and preanal pores; g, arrangement of scales on dorsal and ventral regions, respectively, of mid-tail; h, arrangement of subcaudal scales; h, dorsal aspect of snout of paratype, WHT 7349. Scale bars: 1 mm.

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single conical post-cloacal spur present on each side; The species was also observed in edificarian habitats subcaudals smooth, those on median row enlarged, and in home gardens in the vicinity of the type with pairs of smaller scales between individual larger locality (Fig. 43). It occurs in sympatry with the ones (Fig. 42g). following lizards: Hemidactylus depressus, Lanka­ scincus fallax. Coloration. In preservative: dorsum dusky brown with distinct scattered pale spots; venter dusky Comparisons. Cnemaspis menikay differs from each white. In life: dorsum dark-brown, with yellowish of its Sri Lankan and peninsular Indian congeners, spots laterally; an uninterrupted dusky yellow mid- respectively, by the following opposing suites of dorsal stripe; occipital region with three yellow spots characters: (see Plate 12b). Cnemaspis kandiana: subcaudals keeled except for those on median row. Cnemaspis tropidogaster: Variation. In paratypes WHT 7349 and WHT 7218, enlarged tubercles at tail base, abdominal scales respectively, supralabials (8, 9); infralabials (7, 7). keeled. Cnemaspis podihuna: subcaudals on median Subdigital lamellae on finger I, (13, 9,), finger II, (14, row enlarged, subequal. Cnemaspis scalpensis, C. 11), finger III, (14, 12), finger IV, (12, 13,), finger V, (11, gemunu and C. phillipsii: 12–16 femoral pores. 13); toe I, (9, 9), toe II, (12, 12), toe III, (14, 14), toe IV, Cnemaspis alwisi: 9 femoral pores. Cnemaspis (15, 15), toe V, (16, 15); SVL range from 26.8 to 28.0 molligodai: 5 preanal and 8 or 9 femoral pores; gular, mm (3 ex.). Preanal pores, 1 or 2; femoral pores, 3 pectoral and abdominal scales smooth, dorsal scales or 4. In paratype WHT 7349, an enlarged internasal homogeneous; subcaudals on median row enlarged, scale present (Fig. 42h); granules on chin and throat, subequal. Cnemaspis samanalensis: gular, pectoral and ventro-lateral region and a few subcaudals keeled. abdominal scales tricarinate, subcaudals keeled. Cnemaspis retigalensis: pectoral and abdominal Etymology. The species name is a transliteration scales smooth. Cnemaspis punctata: 5–7 femoral of the Sinhala word for jewel or gem, an object of pores; a distinct spotted dorsal colour pattern. great value, a term of endearment by which women Cnemaspis kumarasinghei and C. latha: gular scales and are sometimes addressed; applied here as a noun in subcaudals smooth. Cnemaspis silvula, C. pulchra, C. apposition. pava, C. clivicola and C. upendrai: abdominal scales keeled. Cnemaspis kallima: subcaudals on posterior Natural history notes. Cnemaspis menikay was ob­ half of tail carinate. Cnemaspis amith: dorsal granules served mostly on tree trunks in secondary forest close heterogeneous and unkeeled. to rubber plantations in the Sabaragamuwa Province. Cnemaspis beddomei: preanal pores 6–9; femoral pores absent. Cnemaspis goaensis: pectoral and abdominal scales smooth. Cnemaspis littoralis: dorsal scales homogeneous, subcaudals on median series enlarged, hexagonal. Cnemaspis mysoriensis: dorsal scales homogeneous, three rows of enlarged subcaudals. Cnemaspis indraneildasii and C. heteropholis: gular scales smooth. Cnemaspis indica, C. sisparensis, C. nairi, C. otai, C. yercaudensis and C. wynadensis: spine-like tubercles absent on flank. Cnemaspis boiei: preanal and femoral pores absent. Cnemaspis gracilis: 2 preanal pores separated by a pair of scales. Cnemaspis ornata: femoral pores absent; a dark brown collar present on nape. Cnemaspis wicksii: thigh and tibia covered dorsally with smooth scales; 4 preanal pores. Cnemaspis andersonii: abdominal scales smooth, subcaudals in median series not enlarged. Cnemaspis jerdonii: preanal pores absent. Cnemaspis monticola: scales on ventral surface of thigh smooth. Cnemaspis australis: subcaudals strongly keeled, pointed. Cnemaspis nilagirica: dorsal scales Figure 43. Distribution of Cnemaspis menikay in south- homogeneous; subcaudals on median row enlarged, western Sri Lanka. subequal.

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Remarks Description of holotype Cnemaspis menikay is the only Sri Lankan member of An adult male, 31.8 mm SVL (for measurements, this genus to have the dosal scutellation comprising see Table 5). Head relatively long (HL 26.1% of entirely of spine-like granules (Fig. 42d). SVL), broad (HW 17.2% of SVL, HW 66.2% of HL), distinct from neck. Snout length moderately greater than eye diameter (ED 67.5% of ES); interorbital Cnemaspis pava new species narrow (IO 4.8% of HL); pupil rounded. Granules (Figures 44a–g, 45; Tables 5, 6; Plate 12c) on snout keeled, larger than those of occipital region. Interorbital, occipital region and neck with conical Holotype. WHT 7286, (male), 31.8 mm SVL, granules. Five enlarged conical tubercles extend from Nawalapitiya, Kandy District, Central Province of behind tynpanum to forelimb insertion. Granules Sri Lanka, 07o03’N, 80o28’E, 600 m, coll. S. Batuwita, on neck intermixed with enlarged conical tubercles. 12 August, 2005. Rostral scale partially divided by a medial groove, in contact with first supralabial (Fig. 44a). Nares Paratypes. WHT 7297, (male), 32.4 mm SVL, WHT separated by two enlarged supranasals and a single 7282, (female), 34.0 mm SVL, WHT 7285, (female), internasal scale (Fig. 44a). Nares rounded, dorsally 33.0 mm SVL, WHT 7290, (male), 32.0 mm SVL, orientated, not in contact with first supralabial. Two WHT 7299, (male), 32.4 mm SVL, same collection postnasals, the lower one in broad contact with first data as holotype; WHT 7289, (male), 32.4 mm SVL, supralabial (Fig. 44a). WHT 7291, (male), 30.6 mm SVL, WHT 7292, (male), Mental subpentagonal, wider than long, truncate 29.1 mm SVL, WHT 7293, (female), 33.1 mm SVL, posteriorly, posteriolaterally in contact with two WHT 7296, (male), 33.9 mm SVL, WHT 7301, (male), postmentals and medially with a single hexagonal 33.0 mm SVL, WHT 7302, (female), 32.9 mm SVL, postmental scale; postmentals bordered posteriorly same location data as holotype, coll. S. Batuwita, 19 by 7 smooth scales including medial scale (Fig. January, 1997. WHT 7281, (male), 31.0 mm SVL, WHT 44b). Three rows of granules around mental on 7283, (female), 30.5 mm SVL, WHT 7288, (male), 32.2 chin. Supralabials 8, infralabials, 7, decreasing in mm SVL, WHT 7294, (male), 32.8 mm SVL, WHT size towards gape (Fig. 44c). Ear opening elliptical, 7295, (male), 30.6 mm SVL, WHT 7298, (female), higher than wide (TW 20.0% of TH). Scales on chin 32.2 mm SVL, WHT 7300, (male), 31.5 mm SVL, granular, keeled; throat with weakly-keeled conical same location data as holotype, coll. S. Batuwita, 22 granules; scales on ventral surface of neck carinate, November, 2002. imbricate. Body slender, long (AG 48.4% of SVL). Mid-dorsal Other materials. NMSL 2004.11.04, NMSL 2004.11.05; granules keeled, mixed with larger keeled tubercles Kitulgala, coll. Mendis Wickramasinghe, 11 May, (Fig. 44d); 11 irregular row of spine-like tubercles 2003. on flank; pectoral and abdominal scales imbricate, pointed, strongly carinate (Fig. 44e). Ventrals, 141; Diagnosis scales aross mid-body, 22; scales around vent and Cnemaspis pava differs from its peninsular Indian base of tail keeled, imbricate; 5 preanal pores; 6 (left and Sri Lankan congeners by a combination of the side) and 5 (right side) femoral pores (Fig. 44f). following characters. Maximum SVL 32.4 mm; paired Forelimbs moderately long, slender (LAL 14.1% postmentals separated medially by a small scale; SVL, UAL 13.2% SVL); hind limbs long (TBL 20.1% each postmental bounded by 3 scales; dorsal scales SVL, FEL 21.6% SVL); tibia long. Dorsal scales on heterogeneous; spine-like tubercles present on flank; both fore and hind limbs carinate; ventral scales on ventrals, 141–143; ventral scales across mid-body, upper arm smooth; lower arm with both smooth 22–25; ventrals (including pectoral, abdominal, and carinate scales. Scales on anterior and ventral precloacal and scales on tail-base) carinate; 2 or 4 surfaces of thigh carinate (Fig. 44f); those on posterior preanal pores and 5 femoral pores on each side; tail surface granular, unkeeled. Scales on anterior and base without enlarged tubercles; supralabials to angle dorsal surfaces of shank weakly carinate; ventral of jaws, 7 or 8; subdigital lamellae on digit IV of pes, surface of shank with large, keeled scales. Scales 18 or 19; (in life) dorsum yellowish brown, with about on manus and pes carinate on both dorsal and six V-shaped markings; a distinct W-shaped mark on ventral sides. Digits elongate, slender, all bearing occipital region and an uninterrupted mid-dorsal slightly recurved claws; subdigital lamellae entire, line present. unnotched; subdigital lamellae on finger I, 10, finger

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Figure 44. Cnemaspis pava: WHT 7286, holotype, 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 and ventral regions, respectively, of mid-body; f, preanal and right hind-limb regions, showing arrangement of femoral and preanal pores; g, arrangement of subcaudal scales. Scale bars: 1 mm.

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II, 13, finger III, 15, finger IV, 16, finger V, 14; toe I, 11, Distribution and natural history notes. Although toe II, 14, toe III, 17, toe IV, 18, toe V, 16; interdigital we found Cnemaspis pava in crevices on moist rocks webbing absent; relative length of digits (fingers): IV at the type locality, we observed this gecko to be (3.4 mm) ~ III (3.2 mm) > II (2.7 mm) ~ V (2.7 mm) > more abundant in edificarin habitats. Its range I (2.1 mm); (toes) IV (4.7 mm) > III (4.1 mm) > V (4.0 appears restricted to the environs of Nawalapitiya mm) > II (3.5 mm) > I (2.2 mm). and Gampola (Fig. 45). Tail tapering (tip detached), its length a little less than SVL; unregenerated tail in paratype, WHT Etymology. The species name, pava, is a Latin 7297 longer than SVL (TAL 121.6% of SVL); tail adjective meaning ‘little’, an allusion to the diminutive base distinctly swollen. Fourteen rows of enlarged habit of this gecko. conical tubercles on tail base. Dorsal scales on tail keeled, granular, directed backwards; 4–6 enlarged Comparisons. Cnemaspis pava differs from each of conical tubercles forming whorls on tail; a single its Sri Lankan and peninsular Indian congeners, post-cloacal spur present on each side. Subcaudals respectively, by the following opposing suites of keeled, 2 smaller scales between individual larger characters: ones in median series (Fig. 44g) Cnemaspis tropidogaster: dorsal scales smooth, no enlarged tubercles on base of tail. Cnemaspis Coloration. In preservative: dorsum dusky brown kandiana: pectoral, abdominal and preanal scales with a cluster of pale spots either side of paravertebral smooth. Cnemaspis podihuna: all ventral scales smooth. line; venter dusky white. In life: dorsum yellowish Cnemaspis scalpensis, C. gemunu and C. phillipsi: 12–16 brown, with about six V-shaped markings; a distinct femoral pores. Cnemaspis alwisi: 7–9 femoral pores. W-shaped mark on occipital region; an uninterrupted Cnemaspis molligodai: 5 preanal and 8 or 9 femoral mid-dorsal stripe present (see Plate 12c). pores; gular, pectoral and abdominal scales smooth; dorsal scales homogeneous; subcaudals on median Variation. In paratype, WHT 7297: supralabials, 7; row enlarged, subequal. Cnemaspis samanalensis: infralabials, 7; subdigital lamellae on finger I, 11, gular, pectoral and abdominal scales tricarinate, finger II, 15, finger III, 17, finger IV, 16, finger V, 14; toe subcaudals keeled. Cnemaspis retigalensis: pectoral and I, 11, toe II, 15, toe III, 19, toe IV, 19, toe V, 16; 2 preanal abdominal scales smooth; a single preanal pore and pores. SVL range from 29.1 to 34.0 mm (20 ex.). 3 or 4 femoral pores present. Cnemaspis punctata: 5–7 femoral pores; a distinctive spotted dorsal coloration. Cnemaspis kumarasinghei: pectoral and abdominal scales smooth. Cnemaspis clivicola: subcaudals on median row not enlarged. Cnemaspis kallima and C. latha: pectoral and abdominal scales smooth. Cnemaspis silvula: ventrals, 138. Cnemaspis upendrai and C. pulchra: ventrals, 112–133; ventral scales on tail base smooth; some gular scales tricarinate. Cnemaspis amith: pectoral and abdominal scales smooth; dorsal granules heterogeneous and unkeeled. Cnemaspis beddomei and C. goaensis: pectoral and abdominal scales smooth. Cnemaspis littoralis: dorsal scales homogeneous, subcaudals on median series enlarged, hexagonal. Cnemaspis mysoriensis: dorsal scales homogeneous, three rows of enlarged subcaudals. Cnemaspis indraneildasii: gular and caudal scales smooth. Cnemaspis heteropholis: no post-cloacal spurs; gular, ventral, and caudal scales smooth. Cnemaspis indica, C. nairi, C. otai, C. yercaudensis, C. sisparensis and C. wynadensis: spine-like tubercles absent on flank. Cnemaspis boiei: preanal and femoral pores absent. Cnemaspis gracilis: ventrals smooth, 2 preanal pores separated by 2 scales. Cnemaspis Figure 45. Distribution of Cnemaspis pava in central Sri ornata: no femoral pores; dorsal tubercles arranged Lanka. in 15 longitudinal rows; ventrals smooth. Cnemaspis

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wicksii, C. andersonii and C. monticola: pectoral and length somewhat greater than eye diameter (ED abdominal scales smooth. Cnemaspis jerdonii: preanal 59.5% of ES); interorbital distance broad (IO 12.7% pores absent. Cnemaspis australis: scales of preanal of HL); pupil rounded. Granules on snout keeled, region smooth. Cnemaspis nilagirica: dorsal scales larger than those of occipital region; scales on inter- homogeneous; subcaudals on median row enlarged, orbital, supercillium and ventral surface of head subequal. granular. Rostral scale partially divided by a medial groove, in contact with first supralabial (Fig. 46a). Remarks Nares separated by two enlarged supranasals and Of its Sri Lankan congeners, Cnemaspis pava most a single internasal scale. Nares rounded, dorsally closely­ resembles C. silvula, fromt which it is orientated, in contact with first supralabial. Two distinguished, however, by having 141–143 ventrals postnasals, the lower one in broad contact with (vs 132–138 ventrals in C. silvula); mid-dorsal first supralabial. Mental subtriangular, truncate tubercles subtriehedral, neumerous (vs mid-dorsal posteriorly, wider than long, posteriolaterally in tubercles rounded, scattered). The ranges of the contact with two enlarged postmentals and medially two species are also widely separated, lying ~70 km with a single postmental scale; postmentals bordered apart (C. pava is known from the mid-hills around posteriorly by 5 smooth scales including medial scale. Nawalapitiya and Gampola, whereas the nearest Supralabials 11, infralabials, 9, decreasing in size location in the range of C. silvula, which is known towards gape (Fig. 46a). Ear opening oval, higher from the south-western wet-zone lowlands, is than wide, oblique (TW 57.1% of TH). Scales on Koskulana in the Sinharaja WHS). ventral surface of neck carinate. Enlarged, bluntly- pointed tubercles on side of neck, of similar size to those on mid-dorsum. Cnemaspis amith new species Body robust, elongate. Mid-dorsal granules (Figures 46a–d; Tables 5, 6; Plate 2g, h) homogeneous, smooth, intermixed with smooth enlarged tubercles (Fig. 46b); 5 spine-like tubercles Holotype. BMNH 63.3.19.1066A (female), 32.0 mm on flank. Ventrals, 131, smooth, imbricate, bluntly SVL, ‘purchased from Dr. Kelaart, Ceylon’ (formerly a pointed or rounded (Fig. 46c); scales around vent and syntype of Gymnodactulus kandianus Kelaart, 1852). on base of tail smooth, imbricate (Fig. 46d). Preanal and femoral pores absent. Paratypes. BMNH 63.3.19.1066B (male), 33.0 mm Forelimbs moderately short, slender (LAL SVL; BMNH 63.3.19.1066C (female), 32.4 mm SVL, 12.5% SVL, UAL 12.1% SVL); hind limbs relatively ‘purchased from Dr. Kelaart, Ceylon’ (formerly long (TBL 16.2% SVL, FEL 19.3% SVL); tibia short. syntypes of Gymnodactulus kandianus Kelaart, Dorsal scales on both fore and hind limbs carinate; 1852). ventral scales on upper arm and lower arm smooth, imbricate. Scales on dorsal surface of thigh and shank Diagnosis carinate, those on ventral surface smooth (Fig. 46d). Cnemaspis amith differs from its peninsular Indian Scales on manus and pes smooth, rounded. Digits and Sri Lankan congeners by a combination of elongate, slender, all bearing slightly recurved claws; the following characters. Maximum SVL 33.0 mm; subdigital lamellae entire, unnotched; subdigital paired postmentals separated by a medial scale; each lamellae on finger I, (broken), finger II, 15, finger III, postmental bounded by 2 scales; ventrals, 123–131; 17, finger IV, 17, finger V, 14; toe I, 8, toe II, 17, toe III, dorsal scales homogeneous, granular unkeeled, with 17 (broken), toe IV, 9 (broken), toe V, 19; interdigital enlarged tubercles; 5 spine-like tubercles on flanks; webbing absent; relative length of digits (fingers): IV gular scales carinate, pectoral and abdominal scales (2.8 mm) ~ III (2.8 mm) > V (2.4 mm) > II (2.2 mm) > smooth; 3 preanal pores and 3 femoral pores on each I (1.8 mm); (toes) IV (3.5 mm) > III (3.1 mm) > II (2.9 side; subcaudals with a median series of smooth, mm) > V (2.7 mm) > I (1.8 mm). enlarged scales; supralabials to angle of jaws, 8; Tail broken; tail base swollen. Enlarged tubercles lamellae under digit IV of pes, 18 or 19. absent on base of tail. Dorsal scales on tail keeled, directed backwards. Tail with about 8 pointed, Description of holotype keeled, enlarged tubercles forming whorls; a single An adult female, 32.0 mm SVL (for other conical post-cloacal spur present on each side; Measurements, see Table 4). Head moderately long subcaudals smooth, scales of median series enlarged, (HL 26.8% of SVL), moderately broad (HW 17.1% of with pairs of obtuse scales between individual larger SVL, HW 63.9% of HL), distinct from neck. Snout rounded ones.

102 Zeylanica Re v i s i o n o f Cn e m a s p i s a b

c d

Figure 46. Cnemaspis amith: BMNH 63.3.19.1066A, lectotype, female, a, lateral aspect of gape, showing arrangement of supralabials and infralabials; b, c, arrangement of dorsal and ventral regions, respectively, of mid-body; d, preanal and right hind-limb regions, showing arrangement of femoral and preanal pores of BMNH 63.3.19.1066B, paralectotype, male. Scale bars: 1 mm.

Coloration. In preservative: dorsally olive with a respectively, by the following opposing suites of series of lighter confluent blotches on dorsum; nape characters: with two dark chevrons, a pale spot between them Cnemaspis kandiana and C. retigalensis: dorsal and a dark spot posterior to them. Venter dusky granules keeled. Cnemaspis tropidogaster: pectoral white with prominent dark markings on gular region, and abdominal scales keeled. Cnemaspis podihuna throat and ventro-lateral regions (see Plate 2g, h). and C. molligodai: median row of regular series Colour in life not known. of enlarged hexagonal subcaudals. Cnemaspis scalpensis, C. gemunu and C. phillipsi: 12–16 femoral Variation. Variation in paralectotypes (BMNH pores. Cnemaspis alwisi: 9 femoral pores Cnemaspis 63.3.19.1066B and BMNH 63.3.19.1066 C) indicated punctata: 5–7 femoral pores and distinct spotted in parentheses: supralabials (8, 7); infralabials (7, 7); dorsal colouration. C. kumarasighei, and C. latha: ventrals (123, 131); SVL (32.0, 33.0 mm). Three preanal gular and subcaudals smooth. Cnemaspis silvula, C. and femoral pores in male paralectotype BMNH clivicola, C. samanalensis, C. pulchra, C. pava and C. 63.3.19.1066B (Fig. 46d). upendrai: chest, abdomen scales keeled. Cnemaspis kallima: subcaudals keeled. Cnemaspis menikay: dorsal Etymology. The species-name is a patronym applied granules conical in shape. as a noun in apposition, honouring the late Dr Amith Cnemaspis beddomei: preanal pores 6–9, no femoral Munindradasa, co-author of Wickramasinghe & pores. Cnemaspis goaensis: subcaudals enlarged Munindradasa (2007). median series. Cnemaspis littoralis: median series of subcaudals enlarged and hexagonal. Cnemaspis Comparisons. Cnemaspis amith differs from each mysoriensis: subcaudal in a triple series enlarged of its Sri Lankan and peninsular Indian congeners, scales. Cnemaspis indraneildasii: dorsal scales of both

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limbs smooth, gular scales smooth. Cnemaspis hetero­ 4a. Preanal pores absent (Fig. 28f)...... 7 pholis: gular scales smooth. Cnemaspis sisparensis 4b. Preanal pores present (Fig. 20f)...... 8 and C. wynadensis: no spine-like tubercles on flanks. Cnemaspis boiei: no preanal or femoral pores. 5a. Scales on gular or ventro-lateral region carinate...... 9 Cnemaspis gracilis: 2 preanal pores separated by 2 5b. Scales on gular and scales. Cnemaspis indica: no spine-like tubercles on ventro-lateral region smooth...... 10 flanks. Cnemaspis ornata: no femoral pores, dorsal tubercles arranged in 15 longitudinal rows, dark 6a. Subcaudals on median row enlarged...... 11 brown collar on nape. Cnemaspis wicksii: thigh and 6b. Subcaudals on median row tibia dorsally covered by smooth scales. Cnemaspis not enlarged (Fig. 32g)...... C. clivicola andersonii: scales along ventro-lateral region carinate. Cnemaspis jerdonii: no preanal pores. Cnemaspis nairi, 7a. Femoral pores, 5–9...... 12 C. otai and C. yercaudensis: no spine-like tubercles 7b. Femoral pores, 12–16...... 13 on flanks. Cnemaspis monticola and C. australis: 8a. Preanal pores, 3 or 4; femoral dorsal granules keeled. Cnemaspis nilagirica: dorsal pores, 3–6; ventrals, 113–117...... C. podihuna scales homogeneous; pectoral and abdominal scales 8b. Preanal pores, 5; femoral weakly keeled; subcaudals on median row enlarged, pores, 8 or 9; ventrals, 129–134...... C. molligodai subequal. 9a. Gular scales carinate; subcaudals on Remarks entire median row smooth (Fig. 36g)...... 14 Cnemaspis amith is known only from its holotype 9b. Gular scales smooth; subcaudals on and two paratypes, which were formerly part of the distal half of median row carinate (Fig. 38h)...... C. kallima syntype series of C. kandiana. A living population of this species has not yet been found. The collector, E. 10a. Ventrals, 111; ventral scales across mid- F. Kelaart, explored widely in Sri Lanka, at various body, 13; dorsum lacking black bands...... C. latha times being stationed at Colombo, Trincomalee and 10b. Ventrals, 121–133; ventral scales across Nuwara Eliya. It is possible that he did not collect mid-body, 17–21; dorsum with 5 these three specimens together with C. kandiana, but black bands...... C. kumarasinghei found them elsewhere and (mis)-identified them as belonging to that species. We hope that future 11a. Pectoral and abdominal scales exploration will serve to uncover a living population unicarinate (Fig. 19d)...... 15 11b. Pectoral and abdominal scales of C. amith in Sri Lanka. tricarinate (Fig. 25b)...... C. samanalensis Cnemaspis amith differs from C. kandiana in several respects: having the mid-dorsal granules smooth (vs 12a. Femoral pores, 5–7; spotted dorsal carinate in C. kandiana); subcaudals smooth (vs keeled coloration...... C. punctata except for those in the median row). 12b. Femoral pores, 7–9; white blotches along mid-dorsal line...... C. alwisi

Key to the species of Cnemaspis in Sri Lanka 13a. Ventrals, 122–141; basal-most lamellae of distal series entire...... 16 1a. Abdominal scales smooth (Fig. 17e); 13b. Ventrals, 113–115; basal-most lamellae dorsal scales on body homogeneous of distal series fragmented...... C. gemunu (Fig. 20d) or heterogeneous (Fig. 19c)...... 2 1b. Abdominal scales keeled (Fig. 19d); 14a. Dorsal granules obtuse; enlarged dorsal scales on body heterogeneous...... 3 tubercles prominent on whole body including dorsum...... 17 2a. Subcaudals on median row subequal 14b. Dorsal granules spinous; enlarged (Fig. 21h); dorsal scales on body homogeneous...... 4 tubercles prominent only on 2b. Subcaudals on median row irregular dorso-lateral surface of body...... C. menikay (Fig. 34g); dorsal scales on body heterogeneous...... 5 15a. Gular scales unicarinate; 3a. Tail base with enlarged, ventrals, 138–141...... 18 tubercles (Fig. 12f)...... 6 15b. Some gular scales tricarinate; 3b. Tail base without ventrals, 112–128...... C. upendrai enlarged tubercles...... C. tropidogaster

104 Zeylanica Re v i s i o n o f Cn e m a s p i s

16a. Conical tubercles present on dorso- homogeneous dorsal scalation; carination of ventral lateral region of body and on tail; scales; presence/absence of caudal tubercles; ventrals, 122–130...... C. scalpensis presence/absence of spine-like tubercles on flank; 16b. Enlarged tubercles absent on nature of subcaudal scalation. Even when large dorso-lateral region of body and on tail; series of specimens have been involved, we have ventrals, 129–141...... C. phillipsi found these character states to be consistent within 17a. Dorsal granules keeled (Fig. 15d)...... 19 individual species. 17b. Dorsal granules smooth (Fig. 46b)...... C. amith New species in this group are also not unexpected from Sri Lanka. For example, the limited phylogeny 18a. Ventral scales on base of tail carinate...... 20 presented by Bauer et al. (2007) showed that the taxa 18b. Ventral scales on base of tail smooth...... C. pulchra they identified as C. kandiana and C. tropidogaster on morphological grounds in fact comprised 19a. A single preanal pore (Fig. 25a)...... C. retigalensis clusters of paraphyletic species. These authors 19b. Preanal pores 2–4...... C. kandiana also demonstrated that the (insular) Sri Lankan 20a. Mid-dorsal tubercles numerous; Cnemaspis are not monophyletic; an elucidation of ventrals, 141–143...... C. pava their relationships, however, will have to await the 20b. Mid-dorsal tubercles moderate; availability of material from India. In any event, ventrals, 132–138...... C. silvula Fig. 7 of Bauer et al. (2007) shows that some species identified on the basis of external morphology alone (e.g., ‘C. tropidogaster’ sensu Deraniyagala, 1955), Discussion in fact comprise distinct—not necessarily closely- related—lineages whose superficial similarity is The new species described here are based mostly evidently the result of convergence. Indeed, it on a collection of ca 250 specimens gathered in the would appear that some of these have already course of a survey of saurians commissioned by been named as new species both herein (e.g., C. the Department of Wildlife Conservation in 1998. clivicola from Kondagala (cf. C. tropidogaster)) and Specimens were collected from representative by Wickramasinghe & Munindradasa, 2007 (e.g., C. habitats in some 50 localities. kumarasinghei from Mihintale (cf. C. kandiana) and The most recent previous reviews of the C. alwisi from Ritigala (cf. C. scalpensis)). Though Sri Lankan geckos were by Taylor (1953) and limited, the phylogeny of Bauer et al. (2007) also Deraniyagala (1955), since which there had been suggests that many ‘cryptic’ species await discovery no attempt to evaluate the Sri Lanks’s gekkonid among the Sri Lankan Cnemaspis—a prediction that lizard diversity until the description of several new is at least partially fulfilled by the 11 new species species in the last few years (Batuwita & Bahir, described herein. 2005; Wickramasinghe, 2006; Bauer et. al., 2007; Two nominal species, Gymnodactylus malabaricus Wickramasinghe & Munindradasa, 2007). The Jerdon, 1853 (type locality ‘Forests of Malabar’) and island’s gecko fauna now stands at 42 valid species, Gymnodactylus littoralis Jerdon, 1853 (type locality including the 11 new species described herein. ‘Sea Coast in Malabar’) are not represented either With 21 species, Cnemaspis is now by far the most by types or extant material, and their identity is speciose gekkonid genus in Sri Lanka, followed somewhat in doubt. In his original description, by Hemidactylus and Cyrtodactylus, each of which Jerdon described C. malabarica as having: “Scales of contains seven and six species, respectively (Kluge, the back uniform, small granulose, a few at the root 2001; Batuwita & Bahir, 2005). of the tail between the two hind legs smaller, those in The description of a large number of new species the tail above larger, imbricated; a few small spines at from a single locality (in this case, Sri Lanka) the root of the tail: dark brown above, marbled with inevitably raises the question, “Are the authors black spots and a white spot on the nape. Length of 4 simply splitting a small number of old species into one 2 /10 inches [61 mm], of which the tail is about a large number of new ones?” The character-states half.” Subsequent to being placed in the synonymy we have used here to separate species are ones that of C. kandiana by Smith (1935), C. malabarica has recent authors on this group of geckos (e.g., Bauer, been overlooked by authors of this genus; it appears 2002; Bauer et al., 2007; Das, 2005; Das & Bauer, 2000; nevertheless to be a valid species (see Remarks Das & Sengupta, 2000; Das & Grismer, 2003; Das & under account of C. kandiana). Jerdon also provided Leong, 2004) have considered reliable, e.g.: presence/ useful data on habitat and distinctive coloration of C. absence of preanal and femoral pores; heterogenous/ malabarica that should facilitate the recollection of this

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species from the forests of Kerala: “it frequents large fore remains to be re-defined on the basis of fresh trees and rocks. It is generally to be found on those collections from the coastal districts of southern large trees, whose base is protected by buttresses, in Kerala. the inequalities of which it finds a secure retreat, and We note in passing that the description provided occassinally it hides itself under a cleft in the bark. for C. nairi by Inger et al. (1984) is entirely consistent The dark markings on its back are edged lighter, with C. beddomei (type locality south Tirunelveli, and it is of a bluish white colour beneath. It is of 08º40’N, 77º40’E, alt. 50 m, ~ 90 km from the coast, a more slender habit than the proceeding species” Tamil Nadu: see redescription of C. beddomei, above, (Jerdon, 1853). and Fig. 7b, 7f, Plate 1g, 1h, especially the dark gular Cnemaspis littoralis, too, lacks type material, band). Given the difference in altitude between Pon but was considered valid by Smith (1935) because Mudi and Tirunelveli, however, further examination Jerdon’s original description included useful of material is necessary before a determination diagnostic characters: “[A] very slender form, scales could be made as to whether these populations are of back small, equal; those on the muzzle larger; conspecific. beneath the tail a series of large six-sided scales. Boulenger (1890) also synonymized Gymnodactylus Colour pale brown, with a series of paler marks along planipes Beddome, 1870 (type locality “Nellicootah, the back and tail, a black spot on the nape, the chin below the Nilgiris (on the western side)” [which 3 and throat pale yellow. Length 2 /10 inches [58.5 mm], we presume to be Nellakota, 11º30’N, 76º25’E, alt. 2 th of which tail 1 /10 … pale brown, with a series of 1,500 m, ~ 80 km from the west coast, Kerala, rather paler marks along the back and tail, a black spot on than Nilakkottai, 10º10’N, 77º50’E, alt. 600 m, ~ 200 the nape, the chin and throat pale yellow.” He added km from the west coast, Tamil Nadu State] with C. that the type was collected in an edificarian habitat: littoralis. Given that Boulenger’s specimens of C. “I procured a single specimen of this well marked littoralis came from Nilambur and Nellakota, and species of Gecko in a ware-house on the Sea Coast not from the western littoral of Malabar (Kerala), in Malabar, and have not yet been fortunate enough we consider it unlikely that C. planipes is in fact a to find another” (Jerdon, 1853). synonym of C. littoralis. In his original description, Boulenger (1890) added to the definition of C. Beddome (1870) listed several important characters littoralis on the basis of specimens he received from for C. planipes, including: body and tail uniformly Nilambur (11º15’N, 76º15’E, alt. 300 m, ~ 50 km from granular, without tubercles; subcaudals enlarged; the coast of Kerala) and Nellakota (11º30’N, 76º25’E, enlarged mental scale; 16 or 17 femoral pores; no alt. 1500 m, ~ 80 km from the coast of Kerala), both of preanal pores; terminal scale of proximal series of which are localities quite different to the type locality enlarged subdigital lamellae three times as large as stated by Jerdon (1853): “sea coast in Malabar”. others. In fact, Beddome was so impressed by this Boulenger’s specimens had 16–18 femoral pores last character that he observed, “Its peculiar feet and no preanal pores; hexagonal, imbricate, smooth almost incline me to constitute a new genus for it.” ventral scales; uniform, smooth dorsal scalation; and In his re-description of C. littoralis, Boulenger (1890) a row of lighter, dark-edged spots along the back, noted of the subdigital lamellae, “The distal plate is none of which characters were noted by Jerdon in the largest, longer than broad, truncate anteriorly.” his original description. Considering the above, we Unfortunately, we were unable to locate the type of conceive it likely that Jerdon (1853) and Boulenger C. planipes in the collection of the BMNH, where the (1890) were referring to different species in their types of the other three species of Cnemaspis that respective accounts. In any event, later authors Beddome described from south-western India in his (e.g., Smith, 1935), while recognizing C. littoralis as 1870 paper are deposited. Considering that one of a valid species, followed Boulenger’s definition of these, C. ornata, has an enlarged subdigital lamella the species, rather than Jerdon’s original. (see Fig. 4i), it is likely that the enlarged lamella of C. In describing C. nairi, Inger et al. (1984) distin­ planipes was yet more remarkable, and a population guished this species also from C. littoralis, specimens of this evidently interesting species remains to be of which they collected from Ponmudi (= Pon Mudi, discovered. 08º45’N, 77º05’E, alt. 600 m, ~ 40 km from the coast, Three of the new species we describe from Kerala State). While their specimens were wholly Sri Lanka are remarkable in that they are from a consistent with Boulenger’s (1890) redescription, single locality, Gammaduwa Estate, near Rattota, given the distance from the coast and difference in at the northern extremity of the Knuckles Hills. The altitude, we doubt that these were in fact examples Knuckles constitute an ‘island’ habitat within Sri of C. littoralis sensu stricto. Cnemaspis littoralis there­ Lanka, with many reptile taxa endemic to it, including

106 Zeylanica

Re v i s i o n o f Cn e m a s p i s

regular, enlarged, enlarged, regular,

caudals ­ caudals

median series of sub of series median – – – – – – – – – – – – – + + + + + + + + base keeled keeled base

– – – – – – – – – – – – –

+ + + + + + + + ventral scales on tail tail on scales ventral median subcaudals cariante subcaudals median

– – – – – – – – – – – – – – – – – + + + + caudal tubercles present tubercles caudal

– + + + + + + + + + + + + + + + + + + + + gular scales tricarinate scales gular

– – – – – – – – – – – – – – – – – – – + + scales carinate scales

– – – – – – – – – – – – + + + + + + + + + pectoral and abdominal and pectoral

body carinate body

ventrolateral scales on on scales ventrolateral – – – – – – – – – – – – + + + + + + + + + carinate

– – – – – – – – – + + + + + + + + + + + + gular and throat scales throat and gular dorsal granules spinous granules dorsal

– – – – – – – – – – – – – – – – – – – – +

dorsal granules unkeeled granules dorsal

– – – – – – – – – – – – + + + + + + + + + homogeneous

– – – – – – – – – – – – – dorsal scales scales dorsal + + + + + + + +

jaws 7 7 8 7 8 7

supralabials to angle of of angle to supralabials 7–8 7–9 7–8 7–8 8–9 7–8 6–8 7–8 8–9 7–8

8–10 7–10 8–10 8–10 8–10 body

25 20 13 19 21 ventral scales across mid- across scales ventral 16–25 17–19 22–26 16–20 20–29 13–20 24–27 18–25 22–25 15–19 20–26 17–21 18–19 20–26 13–16 27–31

111 ventral scales ventral 133

113–117 113–115 112–128 132–138 122–127 128–144 121–128 131–135 121–133 129–141 141–143 129–134 126–138 121–133 123–131 128–144 133–135 146–152

121–138 toe IV subdigital lamellae subdigital IV toe 18 18 19 17 15 18 19

17–21 19–21 16–20 17–23 18–19 17–20 17–19 18–19 19–23 16–18 18–20 17–19 18–20 17–21 number of preanal pores pores preanal of number 3 3 – 3 1 – – 5 2 2 3 – 3 –

2–3 3–4 3–4 2–4 1–2 2–4 3–4 preanal pores present present pores preanal

– – – – – + + + + + + + + + + + + + + + + number of femoral pores pores femoral of number 4 5 4 3 4 4 4–5 4–5 3–5 5–7 3–4 3–6 4–6 8–9 3–4 3–4 3–5 7–9

11–14 13–15 15–16 maximum SVL (mm) (mm) SVL maximum 35.2 31.4 28.6 36.6 37.5 36.0 30.8 24.7 34.2 36.6 32.4 29.0 30.4 28.0 34.6 31.6 33.0 40.9 34.0 35.1 39.9 (+ indicates presence; – indicates absence), based on material 6. Diagnostic characters and character states separating the species of Sri Lankan Cnemaspis (+ indicates presence; Table to males only. counts relate and femoral-pore Preanal examined in this study. C. upendrai C. tropidogaster C. silvula C. scalpensis C. samanalensis C. punctata C. retigalensis C. podihuna C. pulchra C. phillipsi C. pava C. molligodai C. latha C. menikay C. kandiana C. kumarasinghei C. amith C. clivicola C. gemunu C. kallima C. alwisi

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the skinks Chalcidoseps thwaitesi and Nessia bipes; the few species of Cnemaspis occur in the lowest agamid lizards Ceratophora tennentii and Cophotis elevations (< 200 m asl) of Sri Lanka (that account dumbarae; the gecko Cyrtodactylus soba; and the for ~60% of the land area): C. silvula, C. podihuna uropeltid snake Uropeltis phillipsi. Mausakanda Estate and C. kumarasinghei. The largest number of species (now referred to as Gammaduwa), was formerly the occurs at altitudes of between 300 and 700 m asl, with property of the mammalogist-ornithologist W. W. only two (C. gemunu and C. clivicola) occurring at an A. Phillips (1892–1981), as a result of which many altitude of 1,600 m. novelties were discovered there (Wynel-Mayow, An account of the evolutionary history of Cnem­ 2002). The new species of Cnemaspis we describe aspis, and the relationships and timing of dis­persion from there were discovered in the course of searching events with and between Asia and Africa must for examples of a gecko collected by Phillips from necessarily await the results of the molecular phylo­ Mausakanda and identified by Deraniyagala (1953) genetic analysis now in progress (Aaron Bauer, pers. as C. jerdoni scalpensis. comm.), even though such an endeavour would be Each of the three species of Cnemaspis from handicapped by the difficulty of obtaining material Gammaduwa occupies a different habitat niche, from important parts of the range, especially India. and all three occur within ~200 m of one another: Bauer et al. (2007) postulated that the African C. phillipsi is arboreal; C. punctata is found on rock and Asian members of Cnemaspis are reciprocally surfaces, crevices and edificarian habitats; and C. monophyletic while demonstrating that at least kallima is essentially terrestrial. The three are also two Indonesian species were nested within the Sri easily distinguished from each other. Cnemaspis Lankan radiation. phillipsi differs from C. punctata in having 15 or 16 femoral pores; no enlarged tubercles on mid-dorsum; Acknowledgements lacking caudal tubercles; and lacking spotted dorsal We are deeply grateful to Aaron Bauer (Villanova colaration (vs 5–7 femoral pores; enlarged tubercles University), Indraneil Das (Universti Malaysia present on mid-dorsum; caudal tubercles present; Sarawak) and Madhava Meegaskumbura (Museum and a distinct spotted dorsal colaration in C. punctata); of Compatarive Zoology, Harvard Uni­versity and from C. kallima by the absence of preanal pores; and WHT), for commenting on an earlier draft of homogeneous dorsal scalation; a subequal median the manuscript and providing useful comments series of smooth subcaudals (vs 3 preanal pores for its improvement. We thank Colin McCarthy present; heterogeneous dorsal scalation; an irregular and David Gower (BMNH), Channakesava Mur­ median series of subcaudals; and carinate subcaudals thy (ZSI), Nanda Wickramasinghe, Dharma Sri on distal half of tail in C. kallima). Cnemaspis kallima Kan­damby and Lalith Kariyawasam (NMSL) for differs from C. punctata in possessing 3 preanal pores; access to material in their care. For assistance having heterogeneous dorsal scalation; an irregular in the field, we acknowledge the support of median series of subcaudals; and presence of carinate Mohomed M. Bahir and Dinesh Gabadage. We subcaudals on distal half of tail (vs no preanal pores; are grateful to Sudath Nanayakkara (WHT) for homogeneous dorsal scalation; a median series of assistance at the Agra Arboretum; and Jeffrey De subequal subcaudals; and smooth subcaudals in C. Lile, Deleepa Karunaratne, Krishan Wewelwala,­ punctata). Anoma Alagiyawadu, Chandana Yahalmulla, Given the presence of three distinct species in a Dilan Mendis, Anjana Silva, Kalana Maduwage, single locality at Gammaduwa, a relatively dry area Prasanga Wickramathilaka, Sisitha Ranasinghe (rainfall ~ 2,000 mm yr-1), it is surprising that only and Subash Batuwita for assistance with field a single species, C. silvula, has been recorded from work. Finally, we thank the Department of Wildlife the lowlands of the island’s biodiversity-rich south- Conservation for commissioning and co-funding western ‘wet zone’ quarter. Significant tracts of the the national saurian survey in 1998, and the Forest vast expanse (~10,000 km2) of rainforest that occupied Department for permission to access forest reserves this area until the early 20th century still persist, e.g., in its care. the Kanneliya-Dediyagala-Nakiyadeniya forest- reserve complex, in addition to significant extents Literature cited of ‘good’ secondary forest and home gardens. It is surprising that only one of the 21 species of Cnemaspis Abdulali, H., 1955. Extension of range of the lizard now known from Sri Lanka occurs in this area, from Cnemaspis kandiana (Kelaart). Journal of the Bombay the northern border of which C. molligodai is known Natural History Society, 53: 134. from a small range around Ratnapura. Remarkably Annandale, N., 1905 (1904). Contributions to Oriental herpet­ology I. The lizards of the Andamans, with

108 Zeylanica Re v i s i o n o f Cn e m a s p i s

the description of a new gecko and a note on the herpetology. Zoologische Mededelingen. Rijkmuseum van reproduced tail in Ptychozoon homalocephalum. Journal of Natuurlijke Historie, Leiden, 17: 161–251, pls. 1–2. the Asiatic Society of Bengal, 73 (supplement): 12–22. Cox, M. J., Van Dijk, P. P., Nabhitabhata, J. & K. Thirakhupt, Annandale, N., 1909. Report on a small collection of lizards 1998. A photographic guide to snakes and other reptiles from Travancore. Records of the Indian Museum, 3: of Peninsular Malaysia, Singapore and Thailand. New 253–257. Holland Publishers (U.K.) Ltd., London, 144 pp. Annandale, N., 1915. Herpetological notes and descriptions. Das, I., 1993. Cnemaspis gordongekkoi, a new gecko from Records of the Indian Museum, 11: 341–347. Lombok, Indonesia, and the biogeography of Oriental Bahir, M. M. & K. P. Maduwage, 2005. Calotes desilvai, a new species of Cnemaspis (Squamata: Sauria: Gekkonidae). species of agamid lizard from Morningside Forest, Hamadryad, 18: 1–9, pl. 1. Sri Lanka. Raffles Bulletin of Zoology, Supplement 12: Das, I., 2005. Revision of the genus Cnemaspis Strauch, 381–392. 1887 (Sauria: Gekkonidae), from the Mentawai and Bahir, M. M. & A. Silva, 2005. Otocryptis nigristigma, a new adjacent archipelagos off western Sumatra, Indonesia, species of agamid lizard from Sri Lanka. Raffles Bulletin with the description of four new species. Journal of of Zoology, Supplement 12: 393–406. Herpetology, 39(2): 233–247. Batuwita, S. & M. M. Bahir, 2005. Description of five new Das, I. & A. M. Bauer, 2000. Two New Species of Cnemaspis species of Cyrtodactylus (Reptilia: Gekkonidae) from from Tamil Nadu, southern India. Russian Journal of Sri Lanka. Raffles Bulletin of Zoology, Supplement 12: Herpetology, 7(1):17–28. 351–380. Das, I. & S. Sengupta, 2000. A new species of Cnemaspis Bauer, A. M., 2002. Two new species of Cnemaspis (Sauria:Gekkonidae) from Assam, north-eastern (Reptilia:Squamata:Gekkonidae) from Gund, Uttara India. Journal of South Asian Natural History, 5 (1): Kannada, India. Mitteilungen aus dem Naturhistorischen 17–24. Museum in Hamburg, 99: 155–167. Das, I. & L. L. Grismer, 2003. Two new species of Cnemaspis Bauer, A. M. & I. Das, 1999. The systematic status of the Strauch, 1887 (Sauria: Gekkonidae) from the Seribuat endemic Indian gecko, Hemiphyllodactylus aurantiacus Archipelago, Pahang and Johor States, West Malaysia. (Beddome, 1870). Journal of South Asian Natural History, Herpetologica, 59: 546–554. 4(2): 213–218. Das, I. & T.-M. Leong, 2004. A new species of Cnemaspis Bauer, A. M. & I. Das, 2000. A review of the gekkonid (Sauria: Gekkonidae) from southern Thailand. Current genus Calodactylodes (Reptilia:Squamata) from India Herpetology, 23(2): 63–72. and Sri Lanka. Journal of South Asian Natural History, Deraniyagala, P. E. P., 1944. A new Cnemaspis gecko from 5(1): 25–35. Ceylon. Journal of the Royal Asiatic Society (Ceylon Bauer, A. M., de Silva, A., Greenbaum, E. & T. Jackman, Branch), 97: 226–227. 2007. A new species of day gecko from high elevation Deraniyagala, P. E. P., 1953. A colored atlas of some vertebrates in Sri Lanka, with a preliminary phylogeny of Sri from Ceylon: tetrapod Reptilia. National Museum, Lankan Cnemaspis (Reptilia, Squamata, Gekkonidae). Colombo. 101 pp, 46 pls. Mitteilungen aus dem Zoologischen Museum in Berlin, De Rooij, N., 1915. The reptiles of the Indo-Australian Supplement 83: 22–32. Archipelago. I: Lacertilia, Chelonia, Emydosauria. E. J. Beddome, R. H., 1870a. Descriptions of some new lizards Brill, Leiden, The Netherlands, xiv + 334 pp. from the Madras Presidency, with 2 plates. Madras Dring, J. C., McCarthy, C. J. & A. J. Whitten, 1989 (1990). Monthly Journal of Medicine and Science, 1: 30–35. The terrestrial herpetofauna of the Mentawai Islands, Beddome, R. H., 1870b. Descriptions of new reptiles from Indonesia. Indo-Malayan Zoology, 6: 119–132. the Madras Presidency. Madras Monthly Journal of Gray, J. E., 1842. Description of some new species of Medicine and Science, 2: 169–176. reptiles, chiefly from the British Museum collection. Beddome, R. H., 1871. Descriptions of new reptiles from the The Zoological Miscellany, 2: 57–59. Madras Presidency. Madras Monthly Journal of Medicine Gray, J. E., 1846. Descriptions of some new sp­ecies of and Science, 4: 401–404. Indian lizards. Annals and Magazine of Natural History, Biswas, S., 2005. Cnemaspis mysoriensis (Mysore dwarf 18: 429–430. gecko): reproduction. Herpetological Bulletin, 93: Grismer, L. L. & I. Das, 2006. A New Species of Gekkonid 21–22. Lizard of the Genus Cnemaspis Strauch 1887 from Bossuyt, F & A. Dubois, 2001. A review of the frog genus Pulau Pemanggil, Johor, West Malaysia. Herpetological Philautus Gistel, 1848 (Amphibia, Anura, Ranidae, Natural History, 10(1): 1–7. Rhacophorinae). Zeylanica, 6(1): 1–112. International Trust for Zoological Nomenclature, 1999. Boulenger, G. A., 1885. Catalogue of the lizards in the British International­ Code of Zoological Nomenclature. 4th Museum (Natural History). Vol. 1. Taylor and Francis, Edition. International Trust for Zoological Nomen­ London, xii + 436 pp. clature. London, 306 pp. Boulenger, G. A., 1890. A list of the reptiles and batrachians Inger, R. F., Marx, H. & M. Koshy, 1984. An undescribed collected by Dr. E. Modigliani on Sereinu (Sipora), species of gekkonid lizard (Cnemaspis) from India Mentawei Islands. Annali dell Museo Civico di Storia with comments on the status of C. tropidogaster. Naturale di Genova, Series 2, 14: 613–618. Herpetologica, 40(2): 149–154. Brongersma, L. D., 1934. Contributions to Indo-Australian Jerdon, T. C., 1854 (1853). Catalogue of reptiles inhabiting

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the Peninsula of India.Journal of the Asiatic Society of Strauch, A., 1887. Bermerkungen über die Geckoniden- Bengal, l, 22: 462–479. sammlung im zoologischen Museum der Kaiserlichen Kelaart, E. F., 1852. Prodromus Faunae Zeylanicae, being Akademie der Wissenschaften zu St. Petersburg. contrib­utions to the zoology of Ceylon. Volume 1. Mém­oires de l’Académie Impériale des Sciences de St. Privately Published, xxxiii + 197 + 54 + (3) pp. Pétersbourg, 35: 1–72, 1 pl. Kluge, A. G., 2001. Gekkotan lizard taxonomy. Hamadryad, Taylor, E. H., 1953. A review of the lizards of Ceylon. 26: 1–209. University of Kansas Science Bulletin, 35(2)(12): 1525– Mukherjee, D., Bhupathy, S. & A. M. A. Nixon, 2005. A 1585. new species of day gecko (Squamata, Gekkonidae, Taylor, E. H., 1963. The lizards of Thailand. University of Cnemaspis) from the Anaikatti Hills, Western Ghats, Kansas Science Bulletin, 44: 687–1077. Tamil Nadu, India. Current Science, 89(8): 1326–1328. Thakur, S., 1998. Turtles and lizards of the Sahyadri. Journal Pethiyagoda, R., 2007. The ‘new species syndrome’ in Sri of the Ecological Society, Pune, 11: 38–40. Lankan herpetology: a cautionary note. Zeylanica, Theobald, W., 1868. Catalogue of reptiles in the Museum 7(1): 1–7. of the Asiatic Society of Bengal. Journal of the Asiatic Rösler, H., 1981. Bemerkungen zur Geographischen Society of Bengal, (Extra number 88), 37: 1–88. Verbreitung der Gattung Cnemaspis (Strauch 1887); Theobald, W., 1876. Descriptive catalogue of the reptiles of Anmerkungen zur Systematik von C. kandiana (Kelaart British India. Thacker, Spink and Co., Calcutta, xiii + 1852); Allgemeine Überlegungen zu ihrer Biologie. 238 pp. Sauria, Berlin, 1981(3): 7–14. Thomas, J. & P. S. Easa, 1997. Additions to the reptile fauna Sharma, R. C., 1976 (1975). Records of the reptiles of Goa. of Silent Valley, Kerala. Cobra, 27: 31–33. Records of the Zoological Survey of India, 71: 149–167. Wickramasinghe, L. J. M., 2006. A new species of Cnemaspis Smith, M. A., 1935. The fauna of British India, including Ceylon (Sauria:Gekkonidae) from Sri Lanka. Zootaxa, 1369: and Burma; Reptilia and Amphibia. Vol. 2: Sauria. Taylor 19–33. and Francis, London. xiv + 440 pp, 1 pl. Wickramasinghe, L. J. M. & D. A. I. Munindradasa, 2007. Stoliczka, F., 1870. Observations on some Indian and Review of the genus Cnemaspis Strauch, 1887 (Sauria: Malayan Amphibia and Reptilia. Proceedings of the Gekkonidae) in Sri Lanka with the description of five Asiatic Society of Bengal, 1870 (4): 103–109. new species. Zootaxa, 1490: 1–63. Stoliczka, F., 1873. Note on some Andamanese and Wynel-Mayow, E., 2002. W. W. A. Phillips: A naturalist’s Nicobarese reptiles, with the description of three new life. WHT Publications (Private) Limited, Colombo, species of lizards. Journal of the Asiatic Society of Bengal, 84 pp. 42: 162–169.

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a b

c d

e f

g h

Plate 1. a, b, dorsal and ventral aspects, respectively, of Cnemaspis boiei, BMNH 1962.181A, lectotype, male, 35.3 mm SVL; c, d, dorso-lateral and ventro-lateral aspects, respectively, of Cnemaspis indica, BMNH 46.11.22.22b, lectotype, male, 29.3 mm SVL; e, f, dorso-lateral and ventro-lateral aspects, respectively, of Cnemaspis gracilis, BMNH 74.4.29.393, lectotype, male, 29.7 mm SVL; g, h, dorso-lateral and ventral aspects, respectively, of Cnemaspis beddomei, BMNH 1946.9.4.83, lectotype, male, 46.6 mm SVL.

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a b

c d

e f

g h

Plate 2. a, b, Cnemaspis sisparensis: dorsal and ventral aspects, respectively, of BMNH 74.4.29.383, holotype, male; c, d, Cnemaspis ornata: dorso-lateral and ventral aspects, respectively, of BMNH 74.4.29.400, lectotype, male, 42.0 mm SVL; e, f, Cnemaspis ornata: dorsal aspect of BMNH 74.4.29.401, paralectotype, male, 41.7 mm SVL and BMNH 74.4.29.407, paralectotype, juvenile, 26.7 mm SVL; g, h, Cnemaspis amith new species: dorsal and ventral aspects respectively of BMNH 63.3.19.1066A, lectotype, female, 32.0 mm SVL.

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a

b

c

d

Plate 3. a, b, Cnemaspis kandiana: dorso-lateral and ventral aspects, respectively, of BMNH 53.4.1.1, lectotype, female, 33.2 mm SVL; c, dorso-lateral aspect of C. kandiana, WHT 7272, male, 31.0 mm SVL; d, communal egg laying site of C. kandiana in a cement burrow, Polgolla, Kandy District.

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a

b

c

c

Plate 4. a, Cnemaspis scalpensis: dorsal aspect of NMSL 2004.01.1, neotype, male, 29.7 mm SVL (holotype of Cnemaspis ranwellai); b, c, dorso-lateral and ventral aspects, respectively, of Cnemaspis scalpensis,WHT 7268, male, 31.3 mm SVL.

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a

b

c

Plate 5. Cnemaspis podihuna: a, dorsal aspect of uncatalouged NMSL holotype, male, 23.6 mm SVL; b, dorsal aspect of WHT 7334, male, 23.3 mm SVL; c, dorsal aspect of Cnemaspis gemunu, not preserved from Hakgala, Central Province.

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a

b

c

Plate 6. Cnemaspis alwisi: a, dorsal aspect of NMSL 2004.9.1, holotype, male, 39.92 mm SVL; b, communal egg laying site of Cnemaspis alwisi in a rock cave, Warakawehera, near Ridigama, Kurunegala District; c, Cnemaspis kumarasinghei: dorsal aspect of NMSL 2006.13.01, male, 31.61 mm SVL.

116 Zeylanica Re v i s i o n o f Cn e m a s p i s

a

b

c

Plate 7. a, Cnemaspis molligodai: dorsal aspect of NMSL 2006.14.01, holotype, male, 26.62 mm SVL; b, Cnemaspis retigalensis: dorsal aspect of NMSL 2006.12.01, holotype, male, 28.76 mm SVL; c, Cnemaspis samanalensis: dorsal aspect of NMSL 2006.15.01, holotype, male, 36.91 mm SVL.

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a

b

c

Plate 8. Cnemaspis phillipsi: a, b, dorso-lateral and ventral aspects, respectively, of WHT 7236, holotype, male, 33.5 mm SVL; c, habitat of Cnemaspis phillipsi and C. punctata, Mausakanda-Gammaduwa.

118 Zeylanica Re v i s i o n o f Cn e m a s p i s

a

b

c

Plate 9. a, b, Cnemaspis punctata: dorso-lateral and ventro-lateral aspects, respectively, of WHT 7256, holotype, male, 37.1 mm SVL; c, Cnemaspis latha: dorso-lateral aspect of WHT 7214, holotype, male, 30.4 mm SVL.

Vol. 7, No. 1 119 Ma n a m e n d r a -Ar a c h c h i e t a l .

a

b c

d

Plate 10. Cnemaspis kallima: a, dorso-lateral aspects of WHT 7251, paratype, female, 34.9 mm SVL; b, habitat of Cnemaspis kallima, Mausakanda-Gammaduwa; c, communal egg laying site under a stone at Mausakanda-Gammaduwa; d, Cne- maspis silvula: dorsal aspect of WHT 7208, male, 28.6 mm SVL.

120 Zeylanica Re v i s i o n o f Cn e m a s p i s

a

b

c

d

Plate 11. a, Cnemaspis clivicola: dorso-lateral aspect of WHT 7204, holotype, male, 36.6 mm SVL; b, edificarian habitat of Cnemaspis clivicola, near Nanu Oya railway station; c, Cnemaspis upendrai: dorso-lateral aspect of WHT 7260, paratype, male, 28.3 mm SVL; d, communal egg laying site of Cnemaspis upendrai at Pussellawa.

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a a

b

b

c

Plate 12. a, Cnemaspis pulchra: dorsal aspect of WHT 7023, holotype, male, 34.2 mm SVL; Cnemaspis menikay: dorso- lateral aspect of WHT 7349, paratype, male, 26.9 mm SVL; c, Cnemaspis pava: WHT 7286, male, 31.8 mm SVL.

122 Zeylanica