Cnemaspis jayaweerai, Karunarathna & Ukuwela & De Silva & Bauer & Madawala & Poyarkov & Botejue & Gabadage & Grismer & Gorin, 2023

Karunarathna, Suranjan, Ukuwela, Kanishka D. B., De Silva, Anslem, Bauer, Aaron M., Madawala, Majintha, Poyarkov, Nikolay A., Botejue, Madhava, Gabadage, Dinesh, Grismer, L. Lee & Gorin, Vladislav A., 2023, A phylogenetic and taxonomic assessment of the Cnemaspis alwisi group (Reptilia: Gekkonidae) in Sri Lanka with a description of two new species from isolated misty-mountains, Vertebrate Zoology 73, pp. 205-236 : 205

Cnemaspis jayaweerai sp. nov.

Figs 4 , 5

Holotype.

NMSL.2019.23.01, adult male, 36.8 mmSVL, collected inside a granite cave, Ethagala mountain, Ampara District, Eastern Province, Sri Lanka (7.5092N, 81.5198E, WGS1984; elevation 220 m a.s.l., around 10.00 hrs) on 20 July 2019 by Suranjan Karunarathna.

Paratypes (n = 2).

NMSL.2019.23.02, adult female, 34.8 mmSVL and NMSL.2019.23.03, adult female, 28.5 mmSVL, collected from a granite cave close to a stream in Ethagala mountain, Ampara District, Eastern Province, Sri Lanka (7.4978N, 81.5084E, WGS1984; elevation 238 m a.s.l., around 12.00 hrs) collected on 20 July 2019 Suranjan Karunarathna.

Diagnosis.

Cnemaspis jayaweerai sp. nov., is readily distinguished from its Sri Lankan congeners by a combination of the following morphological and meristic characteristics and also color pattern: maximum SVL 36.8 mm; dorsum with homogeneous, smooth granular scales; 2/2 supranasals, 1 internasal and 1/1 postnasal present; 3 enlarged postmentals; postmentals bounded by 5 or 6 chin scales (Fig. 4 ); smooth chin and gular scales, granular, juxtaposed; pectoral and abdominal scales smooth; 6 or 7 poorly developed tubercles on posterior flank; 121-126 paravertebral granules linearly arranged; 22 or 23 belly scales across the venter; precloacal pores absent in males, 9-12 femoral pores on each side in males, separated by 15-17 unpored interfemoral scales, 5-7 unpored posterior femoral scales; 121-129 ventral scales; 70-77 midbody scales; subcaudals smooth, subhexagonal, enlarged, subequal, forming a regular median row; 7-9 supralabials; 7 or 8 infralabials; 15 or 16 total lamellae on digit IV of manus, and 17 or 18 total lamellae on digit IV of pes. It is also differentiated by its significanltly different morphospatial placement in the PCoA and the MFA except for C. gunasekarai in the latter.

Comparisons with other Sri Lankan species.

Among species of the Cnemaspis podihuna clade, Cnemaspis jayaweerai sp. nov. can be readily differentiated from C. kandambyi Batuwita & Udugampala, 2017, C. manoae Amarasinghe & Karunarathna, 2020, C. molligodai Wickramasinghe & Munindradasa, 2007 and C. podihuna Deraniyagala, 1944 by the absence (vs presence) of precloacal pores, and subhexagonal (vs hexagonal) subcaudals; from C. anslemi Karunarathna & Ukuwela, 2019 by the presence of fewer midbody scales (70-77 vs 87-91), more non-pored interfemoral scales (15-17 vs 9 or 10), greater number of flank spines (6 or 7 vs 3 or 4), fewer femoral pores (10 or 11 vs 14 or 15), and fewer lamellae under the 4th toe (17 or 18 vs 20 or 21); from C. gemunu Bauer et al., 2007 by the presence of greater number of belly scales (22 or 23 vs 13-16), greater number of paravertebral granules (121-126 vs 79-93), more non-pored interfemoral scales (15-17 vs 10-12), and greater number of ventral scales (121-129 vs 112-118); from C. godagedarai De Silva et al., 2019 by the presence of fewer ventral scales (121-129 vs 133-137), fewer midbody scales (70-77 vs 98-102), greater number of paravertebral granules (121-126 vs 101-106), more non-pored interfemoral scales (15-17 vs 7-9), and fewer lamellae under 4th toe (17 or 18 vs 20 or 21); from C. phillipsi Manamendra-Arachchi et al., 2007 by the presence of a greater number of paravertebral granules (121-126 vs 86-93), more non-pored interfemoral scales (15-17 vs 11-14), fewer femoral pores (10 or 11 vs 15 or 16), and throat color (bright yellow vs white); from C. scalpensis (Ferguson, 1877) by the presence of greater number of belly scales (22-23 vs 17-19), greater number of paravertebral granules (121-126 vs 102-112), more non-pored interfemoral scales (15-17 vs 8-12), fewer flank spines (6 or 7 vs 9-11), and fewer femoral pores (10 11 vs 13-15); from C. alwisi Wickramasinghe & Munindradasa, 2007 by the presence of fewer ventral scales (121-129 vs 145-153), fewer belly scales (22 or 23 vs 27-31), greater number of paravertebral granules (121-126 vs 89-97), and throat color (bright yellow vs dirty white); from C. gunasekarai Amarasinghe et al., 2021a by the presence of fewer midbody scales (70-77 vs 89-95), more femoral pores (10 or 11 vs 7 or 8), and ventral head color (bright yellow vs dirty white); from C. gunawardanai Amarasinghe et al., 2021a by the presence of fewer ventral scales (121-129 vs 159-162), fewer midbody scales (70-77 vs 96-98), fewer paravertebral granules (121-126 vs 148-155), fewer non-pored interfemoral scales (15-17 vs 21-23), more femoral pores (10 or 11 vs 6 or 7), fewer lamellae under 4th finger (15 or 16 vs 19-21), and fewer lamellae under 4th toe (17 or 18 vs 21-23); from C. hitihami Karunarathna et al., 2019c by the presence of fewer midbody scales (70-77 vs 96-99), fewer paravertebral granules (121-126 vs 143-149), fewer non-pored interfemoral scales (15-17 vs 24-26), fewer lamellae under 4th toe (17 or 18 vs 21 or 22), and lower SVL (36.8 mm vs 41.7 mm); from C. kohukumburai Karunarathna et al., 2019c by the presence of fewer paravertebral granules (121-126 vs 150-159), fewer non-pored interfemoral scales (15-17 vs 22-26), fewer lamellae under 4th finger (15 or 16 vs 21 or 22), and fewer lamellae under 4th toe (17 or 18 vs 23-25); from C. nilgala Karunarathna et al., 2019a by the presence of a greater number of belly scales (22 or 23 vs 17-19), fewer paravertebral granules (121-126 vs 179-187), greater flank spines (6 or 7 vs 3 or 4), and greater femoral pores (10 or 11 vs 7-9); from C. punctata Manamendra-Arachchi et al., 2007 by the presence of greater number of paravertebral granules (121-126 vs 83-91), fewer non-pored interfemoral scales (15-17 vs 25-27), fewer flank spines (6 or 7 vs 11-13), and more femoral pores (10 or 11 vs 5-7); from C. rajakarunai Wickramasinghe et al., 2016 by the presence of fewer ventral scales (121-129 vs 146-186), fewer belly scales (22 or 23 vs 26-29), greater number of paravertebral granules (121-126 vs 81-85), fewer non-pored interfemoral scales (15-17 vs 20-22), and more femoral pores (10 or 11 vs 7 or 8); from C. rammalensis Vidanapathirana et al., 2014 by the presence of fewer ventral scales (121-129 vs 186-207), fewer midbody scales (70-77 vs 119-131), greater number of paravertebral granules (121-126 vs 94-96), fewer femoral pores (10 or 11 vs 14-16), fewer lamellae under 4th finger (15 or 16 vs 22 or 23), and fewer lamellae under 4th toe (17 or 18 vs 22 or 23) respectively.

Among species of the Cnemaspis kandiana clade, Cnemaspis jayaweerai sp. nov. differs by the presence (vs absence) of clearly enlarged, subhexagonal subcaudal scales and absence (vs presence) of precloacal pores from the following species: C. amith Manamendra-Arachchi et al., 2007; C. butewai Karunarathna et al., 2019c; C. dissanayakai Karunarathna et al., 2019b; C. gotaimbarai Karunarathna et al., 2019c; C. ingerorum Batuwita et al., 2019; C. kal-lima Manamendra-Arachchi et al., 2007; C. kandia-na (Kelaart, 1852); C. kawminiae Karunarathna et al., 2019b; C. kivulegedarai Karunarathna et al., 2019c; C. ko-ta-ga-mai Karunarathna et al., 2019b; C. kumarasinghei Wickramasinghe & Munindradasa, 2007; C. latha Manamen-dra-Arachchi et al., 2007; C. lokugei Karunarathna et al., 2021; C. menikay Manamendra-Arachchi et al., 2007; C. nandimithrai Karunarathna et al., 2019c; C. pava Manamendra-Arachchi et al., 2007; C. pulchra Manamendra-Arachchi et al., 2007; C. retigalensis Wickramasinghe & Munindradasa, 2007; C. samanalensis Wickramasinghe & Munindradasa, 2007; C. silvula Manamendra-Arachchi et al., 2007; C. tropidogaster (Boulenger, 1885), and C. upendrai Manamendra-Arachchi et al., 2007.

Description of holotype.

An adult male, 36.8 mmSVL, and 41.1 mmTAL. Body slender, relatively long (TRL/SVL ratio 41.3%). Head relatively small (HL/SVL ratio 27.8% and HL/TRL ratio 67.4%), very narrow (HW/SVL ratio 16.8% and HW/HL ratio 60.4%), less depressed (HD/SVL ratio 10.3% and HD/HL ratio 36.9%) and distinct from neck. Snout relatively short (ES/HW ratio 80.6% and ES/HL ratio 48.7%), more than thrice the eye diameter (ED/ES ratio 37.6%), more than half the length of jaw (ES/JL ratio 88.5%), snout slightly concave in lateral view; eye relatively small (ED/HL ratio 18.3%), twice larger than the ear (EL/ED ratio 50.3%), pupil rounded; orbit length equal to eye to ear distance (OD/EE ratio 100.0%) and shorter than digit IV of the manus (OD/DLM IV ratio 92.4%); supraocular ridges not prominent; ear opening very small (EL/HL ratio 9.2%), deep, taller than wide, larger than nostrils; single row of scales separate orbit from supralabials; interorbital distance is narrow (IO/ES ratio 76.7%), shorter than head length (IO/HL ratio 37.3%); eye to nostril distance shorter than the eye to ear distance (EN/EE ratio 83.9%).

Dorsal surface of the trunk with small, smooth, homogeneous granules, 126 paravertebral granules; 129 smooth midventral scales; 74 midbody scales across belly; 6/7 weakly developed tubercles on the flanks; ventrolateral scales small; granules on snout smooth and flat, larger than those on interorbital and occipital regions; canthus rostralis less pronounced, 12/13 smooth oval scales from eye to nostril; scales of the interorbital region oval and smooth; ear opening vertically oval, slanting from anterodorsal to posteroventral, 23/22 scales between anterior margin of the ear opening and the posterior margin of the eye. Supralabials 8/9 and infralabials 7/8, becoming smaller towards the gape. Rostral scale wider than long, partially divided (70%) by a median groove, in contact with first supralabial. Nostrils separated by 2/2 enlarged supranasals with 1 internasal and 1/1 postnasal; no enlarged scales behind the supranasals. Nostrils oval, dorsolaterally orientated, not in contact with first supralabials.

Mental subtriangular, as wide as long, posteriorly in contact with 3 enlarged postmentals (smaller than mental, and larger than chin scales); postmentals contact and bordered posteriorly by 6 smooth chin scales (larger than nostrils), contact with the 1st infralabials; ventral scales larger than chin scales, and larger than nostrils. Smooth, rounded, juxtaposed granular scales on chin and gular region; pectoral and abdominal scales smooth, subimbricate towards precloacal region, abdominal scales larger than dorsals; 23 belly scales across venter; smooth, subimbricate scales around vent and base of tail; 10/11 femoral pores; 16 unpored interfemoral scales; 7/6 small posterior femoral scales. Original tail of holotype longer than snout-vent length (TAL/SVL ratio 111.8%); hemipenial bulge greatly swollen (TBW 3.6 mm), homogeneous scales on the dorsal aspect of the tail directed backwards, spine-like tubercles absent at base of tail, latter very smooth; tail with 3 or 4 enlarged flattened obtuse scales forming whorls; a very small, round post-cloacal spur on each side; smooth subcaudals are arranged into a median series of clearly enlarged, subhexagonal scales.

Forelimbs moderately short, slender (LAL/SVL ratio 12.7% and UAL/SVL ratio 11.3%) lower arm longer than upper arm; hind limbs moderately long, tibia little shorter than the femur (TBL/SVL ratio 18.8% and FEL/SVL ratio 21.5%). Dorsal, anterior, ventral and posterior surfaces of upper arm and lower arms with smooth scales, those on anterior surface twice as large as those on other faces of limb; scales on dorsal, anterior, ventral and posterior surfaces of the femur and tibia smooth and flat granular, scales on the ventral surface is twice the size of those of the other parts. Dorsal and ventral scales on the manus and the pes smooth, granular; dorsal surfaces of digits with granular scales. Digits elongate and slender with inflected distal phalanges, all bearing slightly recurved claws. Subdigital lamellae entire (except divided at first interphalangeal joint), unnotched; total lamellae on manus (left/right): digit I (11/11), digit II (15/14), digit III (15/15), digit IV (16/16), digit V (16/15); total lamellae on pes (left/right): digit I (10/10), digit II (14/13), digit III (16/16), digit IV (18/18), digit V (16/15); interdigital webbing absent; length order of digits of left manus: I (2.2 mm), V (2.4 mm), II (2.7 mm), III (2.9 mm), IV (3.7 mm); length order of digits of left pes: I (2.1 mm), II (3.9 mm), III (4.3 mm), V (4.3 mm), IV (5.3 mm).

Variation of the type series.

The SVL of adult specimens in the type series of Cnemaspis jayaweerai sp. nov. (n = 3) ranges from 28.5 to 36.8 mm; number of supralabials 7-9, and infralabials 7 or 8; spines on flank 6 or 7; interorbital scales 26-28; supraciliaries 14 or 15; canthal scales 12 or 13; scales from eye to tympanum 21-23; total lamellae under digits of the manus: digit I (10 or 11), digit II (14 or 15), digit III (14 or 15), digit IV (15 or 16), digit V (15 or 16); total lamellae under digits of the pes: digit I (9 or 10), digit II (13 or 14), digit III (15 or 16), digit IV (17 or 18), digit V (15 or 16); ventral scales 121-129, midbody scales 70-77; paravertebral granules 121-126; chin scales 5 or 6; belly scales 20-23;, femoral pores in males 9-12; unpored interfemoral scales in males 15-17, and unpored posterior femoral scales in males 5-7 (Tables 2 - 4 ).

Color of living specimens.

The dorsal color of the head, body and limbs generally from black-brown to ash with four or five paired black blotches; in between black blotches on the dorsum are small grey-white paravertebral blotches; tail is light-brown with 11-14 faded grey cross bands along the tail dorsum (Fig. 5 ); a small black spot is present in the occipital area; dorsal pattern on head reticulated; pupil circular and black with the surrounding scales yellowish white; limbs with black and grey band like dorsal patches; manus and pes with irregular, wide, black and white crossbands on the dorsum; supralabials and infralabials are dirty white and dusted with black; lateral body has a combination of 8-10 black and grey spots; chin, gular, and throat scales are bright yellow without dark spots in males; whole ventral side with dirty white without dark spots in females; pectoral region, arms, abdomen and cloacal scales dirty white; post-cloacal spur and subcaudal scales pale yellow, without spots.

Color of preserved specimens.

Both males and females dorsally light brown, mixed with dirty white and dark paravertebral blotches; dark and grey cross bands on the dorsum of the tail; limbs with brown and dirty white band like dorsal patches; ventral surface dirty white without markings; subcaudals with dark brown margins; ventrolateral side of the trunk dusted with black.

Etymology.

The specific epithet ( Cnemaspis jayaweerai ) is a Latinized eponym in the masculine genitive singular, honouring Mr. Shanthasiri Jayaweera (a senior member and former president of the Young Zoologists’ Association (YZA) of Sri Lanka; founder of the Junior YZA; senior instructor of the Fish study group of the YZA; renowned wildlife artist; a great educator; wildlife conservationist) for his friendship and valuable contribution to environmental conservation in Sri Lanka.

Distribution and natural history.

The type locality, Ethagala forest (7.487114 to 7.514992N and 81.489325 to 81.525203E) is an isolated mountain with large pointed granite rock outcrops supporting a forest dominated by tall shade-bearing trees and bushes (e.g. Vatica obscura ). The dominant vegetation type is dry mixed semi-evergreen forests ( Gunatileke and Gunatileke 1990), and the mean annual rainfall varies between 1000 and 1500 mm, received mainly during the northeast monsoon (November-February) ( Somasekaram et al. 1988). The land area is approximately 450 ha and situated in the Ampara District, Eastern Province (dry bioclimatic zone) of Sri Lanka. The mean annual temperature of the area is 29.8-31.2°C, and its elevational range is 100-450 m a.s.l. Based on our preliminary investigations Cnemaspis jayaweerai sp. nov. appeared to be uncommon in Ethagala. Our survey of 20 ha revealed 16 ( ± 0.3) geckos for 35 man-hours (field work done on 20 July 2019, 13-14 September 2019, and 18-19 March 2020). This species was restricted to granite caves (mainly those modified by natives 1,000 years ago) in closed canopy forest, and abandoned buildings associated with granite caves inside the forest (Fig. 6 ). Further, it prefers narrow (~6-10 mm), long (~120-390 mm) and deep (~80-260 mm) crevices as hiding sites. These microhabitats were poorly illuminated (light intensity: 0-621 Lux), relatively moist (relative humidity: 68-81% and canopy cover: 75-90%) and relatively cool (ambient temperature: 31.4-32.9°C and substrate temperature: 28.2-29.1°C). The new species was sympatric with several other gecko species: Calodactylodes illingworthorum , Cnemaspis podihuna , Gehyra mutilata (Wiegmann, 1834); Hemidactylus depressus Gray, 1842; H. frenatus Duméril & Bibron, 1836; H. hunae Deraniyagala, 1937; H. leschenaultii Duméril & Bibron, 1836; H. parvimaculatus Deraniyagala, 1953, and H. triedrus Daudin, 1802. During the study periods, eggs, hatchlings, juveniles and gravid females were not observed.

Conservation status.

Application of the IUCN Red List criteria indicates that Cnemaspis jayaweerai sp. nov. is Critically Endangered (CR) due to having an area of occupancy (AOO) <10 km2 (five locations - single forest block, 0.16 km2 in total assuming a 100 m radius around each georeferenced location) and an extent of occurrence (EOO) <100 km2 (3.19 km2) in Eastern Province of Sri Lanka [Applicable criteria B2-b (iii)] (Fig. 7 ).

Figures

Figure 4. Cnemaspis jayaweerai sp. nov. male holotype (NMSL. 2019.23.01): A dorsal aspect of head, B lateral aspect of head, C ventral aspect of head, D homogeneous scales on dorsal surface of trunk, E lateral surface of trunk, F smooth ventral scales, G cloacal characters with only femoral pores, H subdigital lamellae of manus, I subdigital lamellae of pes, J dorsal side of tail, K lateral side of tail, L large subhexagonal subcaudals. Scale bar: 2 mm. Photos: Suranjan Karunarathna.

Figure 5. Cnemaspis jayaweerai sp. nov. female paratype (NMSL. 2019.23.02): A dorsal view in life in-situ, B ventral aspect showing gular, ventral, and tail coloration, C lateral aspect of male holotype (NMSL. 2019.23.01) in life in-situ. Photos: Suranjan Karunarathna.

Figure 6. General habitat of Cnemaspis jayaweerai sp. nov. in Ethagala, Ampara District of Sri Lanka: A an isolated granite mountain with dense forest and canopy cover, B a granite cave habitat with lots of cracks and crevices, C historic granite caves with ancient inscriptions in a Buddhist monastery in Ethagala. Photos: Suranjan Karunarathna.

Figure 7. Currently known distribution of the members of the Cnemaspis alwisi and Cnemaspis rammalensis groups with new species - Cnemaspis jayaweerai sp. nov. and Cnemaspis nanayakkarai sp. nov. with the three major mountain habitats and seven elevational ranges in Sri Lanka (abbreviations: AZ Semi-arid zone, DZ dry zone, IZ intermediate zone, and WZ wet zone; Inset top to bottom: Knuckles mountain massif, Central highlands, and Rakwana mountain massif). Photos: Vimukthi Weeratunga and Ashan Geganage. Map: Buddhika Madurapperuma.