Description of a new diminutive, rupicolous species of day-gecko (Squamata: Gekkonidae: Cnemaspis) from southern Sri Lanka Author Batuwita, Sudesh Author Agarwal, Ishan Author Bauer, Aaron M. text Zootaxa 2019 2019-03-08 4565 2 223 234 journal article 28393 10.11646/zootaxa.4565.2.6 57166eb2-9e55-45a1-bfb9-0282c1516b94 1175-5326 2589921 F79A0800-1463-40D8-AF1A-AC0A508C49C2 Cnemaspis ingerorum sp. nov. Ingers’ dwarf gecko ( Figs. 1A, B ; Table 1–3 ) Cnemaspis sp. 2 Agarwal, Biswas, Bauer, Greenbaum, Jackman, De Silva & Batuwita, 2017 Cnemaspis kumarasinghei [non Cnemaspis kumarasinghei Wickramasinghe & Munindradasa, 2007 ] Manamendra-Arachchi, Batuwita & Pethiyagoda, 2007 Holotype . Male , NMSL WHT 7332 , 26.9 mm SVL , collected from Sandagala near Tissamaharama , Hambantota District , Southern Sri Lanka , 6.3416°N , 81.2666° E , ~ 100 m . , by S. Batuwita and J. R. De Lile , 29 July 2005 . Paratypes . Females, NMSL WHT 7330 , 22.2 mm SVL , NMSL WHT 7331 , 24.1 mm SVL , same collection data as holotype . Diagnosis. Small-sized gecko, maximum SVL 30.0 mm; pupil rounded; paired postmentals separated by a medial scale; nostril not in contact with first supralabial; 17–21 ventral scales across mid - body; dorsal scales heterogeneous, isolated scattered spinous tubercles on dorsum; gular, pectoral and abdomen scales smooth; two precloacal pores, five femoral pores on each side in male; subcaudals smooth, median row of enlarged subcaudals semicircular, consisting of a series with a single enlarged scale alternating with a pair of narrow scales; supralabials to angle of jaws seven; digits not dilated towards distal end; proximal lamellae wider than distal lamellae; all digits bearing recurved well-developed claws; subdigital lamellae under 4th digit of pes 16–18; dorsum olive green in life with about five black bands on dorsum; nape with a black spot and occipital area with two dark markings; venter white ( Fig. 1A, B ). TABLE 1. Cnemaspis species of the kandiana clade sampled in this study with locality, voucher number and GenBank accession numbers. Abbreviations for voucher collections are as follows: AA, Rohan Pethiyagoda field series; ADS, Anslem de Silva field series; AMB, Aaron M. Bauer field series (now in NMSL); JB, Jon Boone private collection; MVZ, Museum of Vertebrate Zoology (University of California, Berkeley); SB, Sayantan Biswas field series; WHT, Wildlife Heritage Trust (Sri Lanka).

Species	Voucher	Locality	GenBank Accession Numbers
Number	ND2	CYTB
Cnemaspis ingerorum sp. nov.	WHT 7331	Sri Lanka, Hambantota District, Sandagala	KY037990	KY037945
Cnemaspis kallima	AA 82	Sri Lanka, Matale District, Rattota, Gammaduwa	KY037970	KY037925
Cnemaspis kandiana 1	AA 57	Sri Lanka, Kandy District, Gannnoruwa	KY037971	KY037926
Cnemaspis kandiana 2	AMB 7487	Sri Lanka, Kandy District, Gampola	KY037972	KY037927
Cnemaspis kandiana 3	AA 0 1	Sri Lanka, Kandy District, Loolecondera Tea Estate	KY037973	KY037928
Cnemaspis kumarasinghei	AMB 7431	Sri Lanka, Moneragala District, Rathataakanda (Buttala)	KY037974	KY037929
Cnemaspis cf. kumarasinghei	AA 13	Sri Lanka, Budulla District, Tonacombe Estate (Namunukula)	KY037975	KY037930
Cnemaspis latha	WHT 7214	Sri Lanka, Nuwara Eliya District, Bandarawela	KY037976	KY037931
Cnemaspis modigliani	MVZ 239314	Sumatra, Kecematan Enggano, Pulau Enggano, near Malakoni	KY037977	KY037932
Cnemaspis pava	AA 19	Sri Lanka, Nuwara Eliya District, Ambegamuwa	KY037981	KY037936
Cnemaspis retigalensis	AMB 7448	Sri Lanka, Anuradahapura District, Mihintale	KY037982	KY037937
Cnemaspis samanalensis	AMB 7505	Sri Lanka, Nuwara Eliya District, Upcot tea estate	KY037983	KY037938
Cnemaspis silvula	AA 88	Sri Lanka, Galle District, Hiyare forest reserve	KY037984	KY037939
Cnemaspis upendrai	AA 12	Sri Lanka, Nuwara Eliya District, near Dimbula Junction	KY037987	KY037942
Cnemaspis sp. 1	AA 17	Sri Lanka, Matara District, Naotunna (near Thalalla)	KY037989	KY037944
Cnemaspis sp. 3	AMB 7508	Sri Lanka, Ratnapura District, Masimbula, Godakawela	KY037991	KY037946
Cnemaspis sp. 4	AMB 7529	Sri Lanka, Galle District, Haycock	KY037992	KY037947
Cnemaspis sp. 5	AA 87	Sri Lanka, Budulla District, Haputale	KY037993	KY037948
Cnemaspis sp. 6 India	SB 0 48	India, Karnataka, Kodagu District, Kumarahalli	KY037995	-
Cnemaspis sp. 7 India	JB 239	India, Karnataka (Pet Trade)	KY037996	KY037950
Cnemaspis gemunu	WHT 7348	Sri Lanka, Nuwara Eliya District, near Hakgala	KY037999	KY037953
Cnemaspis phillipsi	AA 81	Sri Lanka, Matale District, Rattota, Gammaduwa	KY038001	KY037955

Cnemaspis ingerorum sp. nov. is morphologically most similar to C. kumarasinghei Wickramasinghe & Munindradasa ( Fig. 1C ). The new species differs from this species by the following suite of characters: 17–21 ventral scales across midbody ( versus 24–26; not 34–36 as given by Wickramasinghe & Muindradasa, 2007 ), heterogeneous ( versus homogeneous) dorsal scales, ventral sides of forelimb and hind-limb with smooth ( versus keeled) imbricate scales, caudal scales on dorsal side unkeeled ( versus keeled) and distinct banded ( versus diffuse dark markings) dorsum colouration ( Table 2 ). The new species also differs genetically from C. kumarasinghei by a 4.5% genetic difference in ND2 ( Fig. 2 ). Cnemaspis ingerorum sp. nov. is most closely related to C. silvula Manamendra-Arachchi, Batuwita & Pethiyagoda ( Fig. 1D ) with 4.5% uncorrected ND2 sequence divergence. The new species differs from C. silvula by the absence ( versus presence) of keeled pectoral scales and dorsal scales, in having 8 ( versus 10–15) pairs of irregular spine-like tubercles on flank, and by the presence of unkeeled ( versus keeled) subcaudal scales. FIGURE 1. Holotype of Cnemaspis ingerorum sp. nov. , from Sandagala, Tissamaharama: A , dorsolateral view; B , ventral view; C, dorsolateral view of Cnemaspis kumarasinghei , not preserved, from Buttala; and D, dorsal view of Cnemaspis silvula , not preserved, from Kanneliya Forest Reserve. FIGURE 2. Maximum likelihood phylogeny of the kandiana clade of South Asian Cnemaspis . Numbers at nodes depict bootstrap support, values <50 and outgroups not shown. TABLE 2. Comparison of morphological and meristic data of Cnemaspis ingerorum sp. nov. , C. kumarasinghei and C. silvula .

Cnemaspis ingerorum sp. nov. ( n = 3)	C. kumarasinghei ( n = 4)	C. silvula ( n = 5)
Dorsal scales	Heterogeneous	Homogeneous	Heterogeneous
Scales across midbody	17–21	24–26	20–22
Ventral scales	Smooth	Smooth	Keeled
Caudal scales	Smooth	Keeled	Keeled
Subcaudals	Smooth	Keeled	Keeled
Scales on ventral sides of forelimb	Smooth	Keeled	Keeled
Scales on ventral sides of hind-limb	Smooth	Keeled	Keeled

Description of holotype (For mensural and meristic data, see Table 3 ). Adult male, 26.9 mm SVL. Head relatively long (HL 26.3% of SVL), moderately broad (HW 17.4% of SVL, HW 66.1% of HL), distinct from neck. Snout length greater than eye diameter (ED 66.6% of ES), interorbital distance narrow (IO 4.2% of HL), pupil rounded. Granules on snout unkeeled, larger than those on occipital region; interorbital scales and those above eye and on ventral side of head granular, unkeeled. Rostral scale partially divided by a medial groove, contacts first supralabial. Nostril openings separated by two enlarged supranasals and enlarged internasal scale. Nostrils rounded, dorsally oriented. Two (on left side) or three (on right side) postnasals, the lower one in broad contact with first supralabial. Mental sub-pentagonal, wider than long, in contact posterolaterally with two enlarged postmentals and medially with a single elongate postmental scale; postmentals bordered posteriorly by four scales. Supralabials eight, infralabials seven, decreasing in size towards gape. Ear opening small, semicircular (EL 36.6% of HL). Scales on ventral side of neck smooth, imbricate. Dorsal and lateral areas of neck covered by unkeeled granules and intermixed with enlarged blunt tubercles. Body robust, short (AG 39.4 % SVL). Middorsal granules heterogeneous; blunt tubercles on ventrolateral region of body and; a series of eight blunt tubercles on flank; scales on pectoral and abdomen smooth, imbricate, semicircular; 20 ventral scales across midbody; scales around vent and at base of tail smooth, imbricate; two precloacal and five femoral pores on each side separated by nine (on left side) or eight (on right side) unpored scales. Forelimbs moderately short, slender (LAL 14.1 % SVL, UAL 14.8 % SVL); hind limbs relatively long (TBL 18.5 % SVL, FEL 19.3% SVL); tibia short. Dorsum of upper and lower arms with granules and keeled imbricate scales; ventral scales on upper arm granular, unkeeled; ventral scales on lower arm with smooth imbricate scales. Scales on anterior border and dorsal side of thigh carinate, scales smooth and imbricate ventrally, posterior border of thigh granular. Scales on anterior border and dorsal side of shank intermixed with granules, smooth and keeled imbricate scales. Scales on ventral side of shank smooth. Scales on dorsum of manus carinate, ventral scales smooth; scales both on dorsal and ventral sides of pes smooth. Digits elongate, slender, all bearing slightly recurved claws; subdigital lamellae entire, unnotched; widened lamellae near base of digits, more than twice width of other scansors; subdigital lamellae on manus, 11-12-16-16-14 and pes, 10-14-16-18 -16; interdigital webbing absent; length of digits (fingers): III ( 2.4 mm )> IV ( 2.2 mm ) ~ V ( 2.2 mm )> II (2.0 mm)> I ( 1.4 mm ); (toes) IV ( 3.1 mm )> III ( 2.9 mm )> V ( 2.5 mm )> II ( 2.4 mm )> I ( 1.4 mm ). Tail tapering (broken in holotype , 13.5 mm ); tail base distinctly swollen. Six large, conical tubercles on base of tail, about as large as granules on mid-dorsum. Tail with eight rows of pointed enlarged tubercles forming whorls; caudal scales granular, unkeeled; one pair of conical postcloacal spurs; subcaudals smooth; median row of enlarged semicircular subcaudals comprising single enlarged scale alternating with two consecutive narrow scales. FIGURE 3. Distribution of Cnemaspis ingerorum sp. nov. (circle), C. silvula (squares) and C. kumarasinghei (triangles); type localities in white, other locations in black. FIGURE 4. Type locality of Cnemaspis ingerorum sp. nov. , Sandagala, near Tissamaharama. Colour in life ( Fig. 1A ). Dorsum mottled beige (anteriorly) to whitish (posteriorly) in vertebral region, becoming more pale greenish-brown towards flanks. Dorsum bears a series of darker brown, predominantly transverse markings with irregular reddish-brown margins. Anteriormost marking on occiput confluent with narrow stripe extending from posterodorsal corner of eye; interrupted in vertebral line by a bright, cream-colored longitudinal marking. A second marking, on neck just anterior to forelimb insertion, comprises three small dark oval markings, one middorsal and the others ventrolateral. Six additional transverse markings run from the shoulder to sacrum, four of these relatively bold, the second and last incomplete mid-dorsally. Each band is expanded or forked on flanks. Head mottled beige, darker brown markings across snout continuing through anterior margin of eye forming postorbital streaks from centre of posterior rim of eye to neck. Frontoparietal boundary has an incomplete transverse band and posterior parietal bears a pair of transversely-oriented dashes. Supralabial scale row with alternating dark markings on a yellowish background. Limbs similar to trunk with irregular banding, most clearly evident on the shanks and on the digits of both manus and pes. Tail dirty whitish with a series of 11 narrow brown bands, the more proximal of which have distinctive orange-to-red highlights. The most proximal marking is bold and bears a yellow spot that partially divides the cross band. Flank tubercles and tubercles on ventrolateral margins of tail whitish to pale-yellowish. Venter whitish with some scattered dark flecks and a pale yellowish wash along the margins of the jaws and throat; semi-translucent trunk, allowing internal organ positions to be seen. Ventral surfaces of forelimbs and anteroventral margins of thighs with continuation of dorsal pattern; palms, soles, and undersides of digits with dark markings. Tail venter white with narrow dark free margins on most subcaudal scales. Colour in preservative. Dorsally light brown with indistinct dusky - brown bands on body; limbs with dark bands; ventral side dusky white.Tail dusky brown; with distinct bands. Variation. Paratypes have pointed tubercles on ventrolateral region of body and eight spine-like flank tubercles. Unregenerated tail of paratype WHT 7331 is longer than its snout-vent length (104.0 % of SVL). TABLE 3. Mensural and meristic data of type series of Cnemaspis ingerorum sp. nov.

Holotype	Paratype	Paratype
WHT 7211	WHT 7331	WHT 7330
SVL	26.9	22.2	24.1
TAL	13.5 (broken)	23.1	8.8 (broken)
TBW	2.8	2.5	2.9
HL	7.1	6.3	6.6
HW	4.7	4.4	4.3
ED	2.2	2.2	2.1
IO	0.3	0.4	0.4
IN	0.9	0.9	0.9
EN	2.3	2.1	2.0
ES	3.3	2.9	2.9
EL	2.6	2.2	2.5
AG	10.6	10.0	10.0
LAL	3.8	3.5	3.5
UAL	4.0	3.2	3.1
DLM I	1.4	1.3	1.4
DLM II	2.0	1.6	1.8
DLM III	2.3	2.0	2.1
DLM IV	2.2	2.2	2.1
DLM V	2.2	1.7	1.9
TBL	5.0	3.9	4.4
FEL	5.2	4.2	4.3
DLP I	1.4	1.2	1.3
DLP II	2.4	2.1	2.1
DLP III	2.9	2.3	2.5
DLP IV	3.0	2.6	2.9
DLP V	2.5	2.5	2.4
Supralabials	7	7	7
Infralabials	7	7	7
Precloacal pores	2	-	-
Femoral pores (left, right)	5,5	-	-
scales across midbody	20	17	21
Subdigital lammellae, manus	11-12-16-16-14	11-11-13-14-13	9-10-13-13-14
Subdigital lammellae, pes	10-14-16-18-16	10-13-15-17-14	9-12-16-17-16

Ectoparasites. Numerous trombiculid mites are present on the holotype ( Fig.1A ): 10 behind neck, eight found in axillae, three to five attached to flank region and four behind the hind-limb insertions. Distribution and natural history. Cnemaspis ingerorum sp. nov. is restricted to Sandagala in the Tissamaharama area of Southern Province of Sri Lanka ( Fig. 3 ), though it may be distributed in the surrounding isolated mountains in the Dry Zone of Sri Lanka . In Sandagala, it is sympatric with following geckos: Hemidactylus frenatus Schlegel , H. hunae Deraniyagala , H. leschenaultii Duméril & Bibron , H. depressus Gray and H. lankae Deraniyagala. The new species is most active around 1600 to 1800 hrs. An adult female (30.0 mm SVL) laid two hard shelled eggs, each about 6.0 x 5.0 mm, which hatched after 53 days; hatchling were 11.9 and 12.2 mm in SVL. Cnemaspis ingerorum sp. nov. is a rupicolous gecko usually observed on rock surfaces, but close to the ground or in the litter layer in rock caves ( Fig. 4 ). When alarmed, it jumps into the litter layer and hides. Etymology. The specific epithet is an eponym in the Latin genitive plural honouring Robert Frederick Inger and Tan Fui Lian Inger of the Field Museum of Natural History (FMNH), for their enormous contributions to herpetology and in appreciation of their support for biodiversity exploration in Sri Lanka , and for their guidance to the first author. Comparisons with regional congeners. Cnemaspis ingerorum sp. nov. differs from following South Asian congeners of Cnemaspis by having heterogeneous dorsal scales and unkeeled gular, pectoral and abdominal scales ( versus having either homogeneous dorsal scales and/or keeled ventral scales): Cnemaspis alwisi Wickramasinghe & Munindradasa , C. amith Manamendra-Arachchi, Batuwita & Pethiyagoda , C. andersonii (Annandale) , C. australis Manamendra-Arachchi, Batuwita & Pethiyagoda , C. gemunu Bauer, De Silva, Greenbaum & Jackman , C. girii Mirza, Pal, Bhosale & Sanap , C. goaensis Sharma , C. gracilis (Beddome) , C. heteropholis Bauer , C. indraneildasii Bauer , C. kandambyi Batuwita & Udugampala , C. kandiana (Kelaart) , C. kumarasinghei , C. latha Manamendra-Arachchi, Batuwita & Pethiyagoda , C. littoralis (Jerdon) , C. menikay Manamendra-Arachchi, Batuwita & Pethiyagoda , C. molligodai Wickramasinghe & Munindradasa , C. monticola Manamendra-Arachchi, Batuwita & Pethiyagoda , C. nilagirica Manamendra-Arachchi, Batuwita & Pethiyagoda , C. pava Manamendra- Arachchi, Batuwita & Pethiyagoda, C. phillipsi Manamendra-Arachchi, Batuwita & Pethiyagoda , C. podihuna Deraniyagala , C. pulchra Manamendra-Arachchi, Batuwita & Pethiyagoda , C. punctata Manamendra-Arachchi, Batuwita & Pethiyagoda , C. rajakarunai Wickramasinghe, Vidanapathirana & Rathnayake , C. rammalensis Vidanapathirana, Rajeev, Wickramasinghe, Fernando & Wickramasinghe , C. retigalensis Wickramasinghe & Munindradasa , C. samanalensis Wickramasinghe & Munindradasa , C. scalpensis (Ferguson) , C. silvula , C. tropidogaster (Boulenger) , C. upendrai Manamendra-Arachchi, Batuwita & Pethiyagoda , and C. wicksii (Stoliczka) . Cnemaspis ingerorum sp. nov. has spine-like flank tubercles which differentiates it from the following species that lack these tubercles: C . indica (Gray) , C. nairi Inger, Marx & Koshy , C. otai Das & Bauer , C. yercaudensis Das & Bauer , C. sisparensis (Theobald) , C. kolhapurensis Giri, Bauer & Gaikwad , and C. wynadensis (Beddome) . The new species is distinguished from C. boiei (Gray) by having imbricate ventral scales ( versus juxtaposed in C. boiei ). The new species also differs from the following species by having both femoral and precloacal pores: Cnemaspis jerdonii (Theobald) (lacks precloacal pores), C. ornata (Beddome) (lacks femoral pores), and C. beddomei (Theobald) (lacks femoral pores). Cnemaspis ingerorum sp. nov. is distinguished from C. mysoriensis (Jerdon) by having 5 ( versus 2–3) femoral pores.