Cnemaspis lokugei, Karunarathna & Silva & Gabadage & Botejue & Majintha Madawala & Ukuwela, 2021

Karunarathna, Suranjan, Silva, Anslem De, Gabadage, Dinesh, Botejue, Madhava, Majintha Madawala, & Ukuwela, Kanishka D. B., 2021, A new species of day gecko (Reptilia, Gekkonidae, Cnemaspis Strauch, 1887) from Sri Lanka with an updated ND 2 gene phylogeny of Sri Lankan and Indian species, Zoosystematics and Evolution 97 (1), pp. 191-209 : 191

Cnemaspis lokugei sp. nov. Figs 2 , 3 ; Tables 2, 3 Lokuge’s day gecko (English) Lokugege diva-seri hoona (Sinhala)

Cnemaspis sp. 5 Agarwal et al. 2017

Cnemaspis sp. 4 Karunarathna et al. 2019c

Holotype.

NMSL. 2021.01.01, adult male, 32.9 mm SVL, collected from a granite cave bordering a stream, Haputale , Badulla District, Uva Province, Sri Lanka (6.7753°N; 80.9667°E, WGS1984; elevation 1510 m; around 10:00 hrs) on 18 December 2019 by Suranjan Karunarathna.

Paratypes.

NMSL. 2021.01.02, adult female, 30.8 mm SVL and NMSL. 2021.01.03, adult female, 30.2 mm SVL, collected from a granite cave bordering a stream, Idalgashinna , Badulla District, Uva Province, Sri Lanka (6.7791°N; 80.8967°E, WGS1984; elevation 1565 m; around 09:00 hrs) on 19 December 2019 by Suranjan Karunarathna .

Diagnosis.

Cnemaspis lokugei sp. nov., can be readily distinguished from its Sri Lankan congeners by a combination of the following morphological and meristic characteristics: maximum SVL 32.9 mm; dorsum scalation heterogeneous, mixed with smooth and keeled large granular scales; 1/1 supranasals, 1 internasal, 1/1 postnasal; 3 enlarged postmentals; postmentals bounded by 5 enlarged chin scales; chin, gular, pectoral and abdominal scales smooth, subimbricate; 15-17 belly scales across mid-body; 5 or 6 feebly-developed tubercles on posterior flank; 122-132 paravertebral granules linearly arranged; 3 precloacal pores, 4 or 5 femoral pores in males, separated by 8 or 9 proximal femoral scales lacking pores, 7 or 8 distal femoral scales lacking pores; 143-159 ventral scales; 95-103 mid-body scales; smooth subcaudals, median row comprising an irregular series of diamond shaped, small scales; 7 or 8 supralabials; 7 or 8 infralabials; 15 or 16 total lamellae on fourth digit of manus and 17 or 18 total lamellae on fourth digit of pes.

Description of holotype.

An adult male, 32.9 mm SVL and 36.9 mm TAL (regenerated). Body slender, relatively short ( TRL / SVL ratio 39.9%). Head relatively small (HL/ SVL ratio 27.7% and HL/TRL ratio 69.5%), narrow (HW/ SVL ratio 14.2% and HW/HL ratio 51.2%), depressed (HD/ SVL ratio 10.1% and HD/HL ratio 36.4%) and distinct from neck. Snout relatively long (ES/ HW ratio 75.5% and ES/HL ratio 38.6%), less than twice the eye diameter ( ED / ES ratio 53.4%), more than half length of jaw ( ES / JL ratio 64.8%), snout slightly concave in lateral view; eye relatively small ( ED /HL ratio 20.6%), larger than the ear ( EL / ED ratio 43.6%), pupil round; orbit length greater than eye to ear distance ( OD / EE ratio 125.8%) and equal to length of digit IV of manus (OD/ DLM IV ratio 100%); supraocular ridges moderately developed; ear opening small ( EL /HL ratio 9.0%), deep, taller than wide, larger than nostrils; two rows of scales separate orbit from supralabials; interorbital distance is a little shorter than snout length ( IO / ES ratio 97.7%), shorter than head length ( IO /HL ratio 37.8%); eye to nostril distance greater than the eye to ear distance ( EN / EE ratio 102.8%).

Dorsal surface of the trunk with smooth scales intermixed with keeled heterogeneous granules, 132 paravertebral granules; 148 smooth, mid-ventral scales; 95 mid-body scales; 6/5 weakly-developed tubercles on the flanks; ventrolateral scales small, irregular; granules on snout oval, keeled and raised, larger than those on interorbital and occipital regions; canthus rostralis nearly absent, 13/13 smooth oval scales from eye to nostril; scales of the interorbital region circular and keeled; short tubercles present both on the sides of the neck and around the ear; ear opening vertically oval, slanting from anterodorsal to posteroventral, 21/20 scales between anterior margin of the ear opening and the posterior margin of the eye. Supralabials 7/7, infralabials 8/7, becoming smaller towards the posterior end of the mouth. Rostral scale wider than long, partially divided (80%) by a median groove and in contact with first supralabial. Nostrils separated by 1/1 enlarged supranasals with 1 internasal; few enlarged scales behind the supranasals. Nostrils oval, dorsolaterally orientated, not in contact with first supralabials; 1/1 postnasals, smooth, larger than nostrils, partially in contact with first supralabial (Fig. 2 ).

Mental, sub-rhomboid in shape, as wide as long, posteriorly in contact with 3 enlarged postmentals (smaller than mental and lager than chin scales); postmentals in contact and bordered posteriorly by 5 unkeeled chin scales (larger than nostrils), in contact with the 1st infralabial; ventral scales smaller than chin scales. Smooth, rounded, juxtaposed scales on the chin and the gular region; pectoral and abdominal scales smooth, subimbricate towards precloacal region, abdominal scales slightly larger than dorsals; 17 belly scales across ventre; smooth scales around vent and base of tail, subimbricate; 3 precloacal pores; 4/5 femoral pores; 8/9 proximal femoral scales lacking pores on each side; 7/8 enlarged distal femoral scales. Regenerated tail little longer than the snout-vent length ( TAL / SVL ratio 112.2%); hemipenial bulge moderately swollen (TBW 2.8 mm), heterogeneous scales on the dorsal aspect of the tail directed backwards, spine-like tubercles present at the base of tail; tail with 3 or 4 enlarged flattened obtuse scales forming whorls; a large, blunt post-cloacal spur on each side, dorsoventrally flattened and narrow; subcaudals smooth and small, subrhomboidal, arranged in a single median series (Fig. 2 ).

Forelimbs very short, slender (LAL/ SVL ratio 11.6% and UAL/ SVL ratio 13.1%); hind limbs long, tibia little longer than the femur (TBL/ SVL ratio 16.7% and FEL/ SVL ratio 15.8%). Scales on anterior, upper, posterior and ventral surfaces of upper arm with keeled granules and less imbricate scales, scales of the anterior surface twice as large as those of the other surfaces; anterior, upper, posterior and ventral surfaces of lower arm with keeled and less imbricate scales, scales of the upper surface twice as large as those of the other surfaces. Scales on anterior, upper, posterior surfaces of femur keeled, ventral surface with smooth, subimbricate scales, scales on the ventral surface twice the size of those of other aspects; anterior, upper, posterior surfaces of tibia keeled, ventral surface with smooth, imbricate scales, scales on the upper surface twice the size of those of other aspects. Dorsal and ventral surfaces of manus and pes with keeled granules; dorsal surfaces of digits with granular scales (Fig. 2H, I ). Digits elongate and slender with inflected distal phalanges, all bearing slightly recurved claws (Fig. 2 ). Subdigital lamellae entire (except divided at first interphalangeal joint), unnotched; total lamellae on manus (left/right): digit I (10/11), digit II (13/14), digit III (16/16), digit IV (16/15), digit V (15/15); total lamellae on pes (left/right): digit I (11/10), digit II (15/15), digit III (17/18), digit IV (18/18), digit V (17/17); interdigital webbing absent; relative length of left manual digits: I (1.4 mm), II (1.8 mm), V (2.3 mm), III (2.5 mm), IV (3.1 mm); relative length of left pedal digits: I (1.5 mm), II (2.9 mm), III (3.4 mm), V (3.5 mm), IV (3.9 mm).

Variation in the type series

(Tables 2 and 3 ). The SVL of adult specimens in the type series of Cnemaspis lokugei sp. nov. (n = 3) ranges from 30.2 to 32.9 mm; interorbital scales 24-26; supraciliaries above the eye 14-16; supralabials 7 or 8, infralabials 7 or 8; scales from eye to tympanum 19-21; canthal scales 12 or 13; tubercles on posterior flank 5 or 6; ventral scales 143-159 (Tables 2 - 3 ); mid-body scales 95-103; paravertebral granules 122-132; belly scales across ventre 15-17; femoral pores in males 4 or 5; proximal femoral scales lacking pores in males 8 or 9; distal femoral scales lacking pores in males 7 or 8; total lamellae on digit of the manus: digit I (10 or 11), digit II (13 or 14), digit III (15 or 16), digit IV (15 or 16), digit V (14 or 15); total lamellae on digit of the pes: digit I (10 or 11), digit II (14 or 15), digit III (17 or 18), digit IV (17 or 18), digit V (16 or 17).

Colour in life.

Dorsum of head, body and limbs generally reddish-brown; yellow spot with black outer edge on neck dorsally; broken faded, yellow vertebral stripe running from occiput to tail (Fig. 3 ); seven irregular blackish-brown, chevron shaped paravertebral markings present. Tail dark brown dorsally, with 10 faded brown irregular cross-bands; pupil circular and black with the surrounding iris yellow; two very faint postorbital stripes on each side; supralabials and infralabials yellowish with tiny black spots; chin and gular scales yellow, with dark spots; pectoral, abdominal, cloacal and subcaudal scales are cream and intermixed with irregular stippling; dorsum of limbs with faded black markings; manus and pes with alternating black and cream-white cross bands.

Colour in preservation.

Dorsally grey brown with seven distinct dark, irregular blotches; pale spot with dark outer edge on neck dorsally; supralabials and infralabials dirty white; two dark postorbital stripes on each side; chin and gular scales grey; ventral surface uniformly dirty white colour with some scales on thigh, tail base and arms with dark brown margins.

Etymology.

The specific epithet is an eponym Latinised ( Cnemaspis lokugei ) in the masculine genitive singular, honouring Mr. Ajith Nethkelum Lokuge, a pioneer ecologist, analogue forestry specialist and a senior member of Young Zoologist’s Association of Sri Lanka, for his significant contribution towards environmental conservation and research in Sri Lanka.

Distribution and natural history.

The specimens of the type series were collected from the two locations, Haputale and Idalgashinna (Badulla District, Uva Province), which are situated in the central highlands of Sri Lanka (central intermediate bioclimatic zone [annual rainfall between 2000-2500 mm ( Burt and Weerasinghe 2014)]) (Fig. 4 ). Tropical sub-montane and montane forests make up the dominant vegetation type ( Gunatileke and Gunatileke 1990) of this area. The forest acreage in both areas is approximately 1200 ha and is relatively isolated from other forests due to anthropogenic habitats and tea plantations. It is very likely that the species occurs in the intervening regions between these two locations as there are similar habitats scattered between the two locations. However, this needs to be verified through a thorough field survey. These locations lie between an elevation of 1400 and 1700 m a.s.l. (Fig. 4 ). The mean annual rainfall is received mainly during the southwest monsoon (May-September), while the mean annual temperature is 26.1-28.9 ºC. Both areas are rich in granite rock boulders with 40 identified caves. Cnemaspis lokugei sp. nov. appears to be a common species in the two locations as we recorded more than 50 individuals from both locations during a two-day survey. This species was observed in granite caves and relatively old buildings on vertical surfaces, about 2 m from ground within the forested area (Fig. 5 ). The granitic cave microhabitat of C. lokugei sp. nov. was poorly illuminated (light intensity: 392-476 Lux), relatively moist (relative humidity: 76-92%), well shaded (canopy cover: 62-78%) and relatively cool (ambient temperature: 29.5-31.2 °C and substrate temperature: 27.4-28.7 °C). The new species is sympatric with several other gecko species: Cyrtodactylus sp., Gehyra mutilata , Hemidactylus frenatus , H. parvimaculatus and Hemiphyllodactylus typus . Pure white and almost spherical shaped (mean diameter 4.9 ± 0.02 mm [n = 34]) eggs with a slightly flattened side attached to a rocky substrate were observed in cave habitats where Cnemaspis lokugei sp. nov. was observed. Since these eggs were characteristic of Cnemaspis species and as there were no other Cnemaspis species observed in these habitats, it was presumed that the eggs most likely belong to C. lokugei sp. nov.

Conservation status.

Application of the IUCN Red List Criteria indicates that C. lokugei sp. nov. is Critically Endangered (CR) due to its having an area of occupancy ( AOO) < 10 km 2 (3.84 km 2 in total assuming a 100 m radius around the seven georeferenced locations), severely fragmented habitat and a projected decline in the area, extent and the quality of habitat [Applicable criteria B2ab (iii)].

Comparisons with other Sri Lankan species.

Morphologically, Cnemaspis lokugei sp. nov. most closely resembles C. butewai , C. ingerorum , C. kivulegedarai , C. kallima , C. kandiana , C. kotagamai , C. menikay and C. retigalensis because of the presence of a dorsum with heterogeneous scales and smooth belly scales (see the species comparison and Table 4 for more details). Amongst species of the C. kandiana clade sensu Agarwal et al. (2017), Cnemaspis lokugei sp. nov. differs by having heterogeneous (vs. homogeneous) dorsal scales from C. amith , C. dissanayakai , C. gotaimbarai , C. kawminiae , C. kumarasinghei , C. latha and C. nandimithrai . It can also be distinguished from C. butewai , C. kandiana , C. menikay , C. pava , C. pulchra , C. retigalensis , C. samanalensis , C. silvula , C. tropidogaster and C. upendrai by having smooth (vs. keeled) gular scales. The new species differs from C. ingerorum and C. kivulegedarai by having more ventral scales (143-159 vs. 88-95 and 109-114) and by having more mid-body scales (95-103 vs. 62-69 and 69-76); from C. kallima by having more mid-body scales (95-103 vs. 67-74) and by having more paravertebral granules (122-132 vs. 99-107); from C. kotagamai by having more mid-body scales (95-103 vs. 79-84) and by fewer scales across belly (15-17 vs. 21-22).

Amongst species of the C. podihuna clade sensu Agarwal et al. (2017), Cnemaspis lokugei sp. nov. differs by the absence of clearly enlarged, hexagonal or subhexagonal subcaudal scales from the following species with homogeneous dorsal scales: C. alwisi , C. anslemi , C. gemunu , C. godagedarai , C. hitihamii , C. kandambyi , C. kohukumburai , C. molligodai , C. nilgala , C. phillipsi , C. podihuna , C. punctata , C. rajakarunai , C. rammalensis and C. scalpensis . The new species also differs from C. alwisi , C. anslemi , C. gemunu , C. godagedarai , C. hitihamii , C. kohukumburai , C. nilgala , C. phillipsi , C. punctata , C. rajakarunai , C. rammalensis and C. scalpensis by having precloacal pores (vs. absence).

Figures

Figure 2. Cnemaspis lokugei sp. nov. male holotype (NMSL. 2021.01.01), (a) dorsal head, (b) lateral head, (c) ventral head, (d) heterogeneous scales on dorsal surface of trunk, (e) lateral surface of trunk, (f) smooth ventral scales, (g) cloacal characters with precloacal and femoral pores (h) subdigital lamellae on manus, (i) subdigital lamellae on pes; female paratype (NMSL. 2021.01.03), (j) dorsal side of tail, (k) lateral side of tail, (l) subrhomboid-shaped small subcaudals. Scale bar: 1 mm (Photos: Suranjan Karunarathna).

Figure 3. Cnemaspis lokugei sp. nov. male holotype (NMSL. 2021.01.01) in life in-situ (a) dorsal view of the full body with typical colour pattern, (b) ventral aspect showing gular and ventral colouration, (b) lateral aspect with spines on flank and labial colouration from Haputale (Photos: Suranjan Karunarathna).

Figure 4. Currently known distribution of Cnemaspis lokugei sp. nov. (holotype- red star, paratype- red circle) and its closely-related species (C. butewai - white circl, and C. pulchra - yellow circle) in Sri Lanka.

Figure 5. General habitat of Cnemaspis lokugei sp. nov. in the Uva Province of Sri Lanka (a) wet forest highland with short trees, viewed from Idalgashinna, (b) a granite cave habitat in Idalgashinna (c) communal egg clutches in Haputale. (Photos: Suranjan Karunarathna).