Otocryptis wiegmanni Wagler, 1830

Srikanthan, Achyuthan N., Adhikari, Omkar Dilip, Ganesh, Sumaithangi Rajagopalan, Deuti, Kaushik, Das, Kalpana, Kulkarni, Vidisha M., Gowande, Gaurang G. & Shanker, Kartik, 2021, A molecular and morphological study of Otocryptis Wagler, 1830 (Squamata: Agamidae) reveals a new genus from the far south of the Western Ghats, Peninsular India, Zootaxa 5016 (2), pp. 205-228 : 214-217

Otocryptis wiegmanni Wagler, 1830

( Figs. 3–4 , Tables 3 & 5)

Otocryptis bivittata Wiegmann, 1831

Otocryptis wiegmanni (nec Taylor 1953)— Wagler, 1830

Holotype. ZMB 708 , adult male specimen originally in the personal collection of M. E. Bloch the provenance of which was subsequently interpreted as Ceylon (= Sri Lanka) (also the holotype of Otocryptis bivittata ; see Bahir & de Silva 2005).

Material Examined. Adult males (n=5) SRI LANKA ‒ BNHS 278 A and BNHS 279 , collected by E. E. Green, from Paradeniya (7.26˚ N 80.60 ˚E), on 8 May 1912 ; BNHS 280 , from unknown precise locality ; BNHS 281 , col- lected by J. Woodmason, Colombo (6.92˚ N 79.90 ˚E), in November 1906 ; BNHS 282 , collected by E. E. Green, Kandy (7.28˚ N 80.63 ˚E), on 31 March 1912 . Adult female (n=1) SRI LANKA ‒ BNHS 278 B, collected by E. E. Green, from Paradeniya , on 8 May 1912 .

Diagnosis (re-defined herein). A genus of draconine agamid lizard endemic to Sri Lanka, diagnosed by the following characters of lepidosis and osteology: large dewlap appendage present in males with dewlap bone well surpassing axillary midline contact or forelimb insertion, reaching up to 1/3 of torso; dewlap brightly colored in males with a conspicuous large display spot; cephalic scales irregular, some smaller than mid-dorsal scales, smooth, feebly carinate or conical; temporal region with 4‒6 enlarged, conical scales; tympanum subdermal; dorsal crest ab- sent; a rudimentary sheath-like nuchal crest present in males; antehumeral pit absent; temporal spines absent or not developed; breeding and display coloration on the entire head including the nuchal crest; dorsal tubercles absent.

Otocryptis wiegmanni , type species of genus Otocryptis , is distinguished from its another congener Otocryptis nigristigma as follows: A distinctly maroon patch on lateral side of dewlaps in males of O. wiegmanni (vs a distinctly black patch on lateral side of dewlaps in males of O. nigristigma ); fifth toe long, 22.6‒32.3% of HL in males, and 26.2% of HL in females in O. wiegmanni (vs fifth toe short 14.3–16.0% of HL in males, and 14.6–18.5% of HL in females of O. nigristigma ); males of O. wiegmanni have first finger and first toe 21.3‒32.8 % and 16.7‒23.6 % of HL (vs males of O. nigristigma have first finger and first toe 15.7–17.8% and 14.6–17.2% of HL); females have third and fourth fingers 53.4 % of HL and 60.8 % of HL in O. wiegmanni (vs 33.9–39.5% of HL and 32.5–40.5% of HL respectively, in females of O. nigristigma ).

Osteological characters summarized for Otocryptis wiegmanni as follows (some in comparison with Agasthyagama beddomii comb. nov., also see Table 6): Two comparatively elongate, second ceratobranchials, elements responsible for erecting the dewlap, extend backwards on to belly; premaxilla forms pointed anterior margin of upper jaw; nasal process of premaxilla well developed; 14 teeth; comparatively larger maxilla; deeper and perforated by “V” shaped foramina at anterior end; pre-frontals roughly rectangular; jugal comparatively smaller with a distinct foramen located at its centre; temporal fossa small, triangular; palatine bone longer and comparatively broader at anterior end; cultriform process long and slender; sternal plate is well developed with six posterolateral sternal ribs and two posterior sternal ribs; femur, humerus, radius, ulna and vertebral column comparatively longer.

Description and variation. Based on referred material, 5 males, and 1 female —data in brackets; also see Table 3 for measurements given as % of HL: Snout to vent length 64.0−74.0 (43.0). Head longer than broad, HL 18.0−24.6 (11.6), HW 11.2−13.7 (8.2); triangular in dorsal aspect, DHW 8.0−10.7 (6.0); HH 9.8−11.4 (7.7); interorbital area concave, IOD: 7.0−9.6 (5.0) between anterior end, 10.5−12.8 (6.8) between posterior end; SNL 7.7−9.6 (5.6) area between snout-tip and interorbital region flat, 3−4 enlarged conical scales surrounded by irregular carinate scales. Eyes small, HED 3.4−5.5 (3.7); orbital rim prominent, supraorbital region with 5–7 (6) rows of large, carinate scales, the inner row extending forward to form a Y-shaped ridge on snout; superciliary scales carinate; second row of scales from inner margins of both upper and lower eyelid large, feebly carinate. Nostril oval, laterally orientated, E-N 5.9−7.6 (3.6); IND 3.3−3.6 (3.0); canthus rostralis with 10–11 (10) strongly carinate scales; nasal scales large, undivided, horizontally oval, in contact with 1 st supralabial on both sides, separated from 2 nd supralabial by row of scales; rostral wide, separated from nasal by a single scale, in contact with 1 st supralabial on both sides. Supralabials: 8−11 (11) on left side and 9−11 (11) on right side. Temporal region with 4–6 (5) enlarged, conical scales; upper temporal fossae slightly concave. Cephalic scales irregular, some smaller than middorsal scales, smooth, feebly carinate or conical. Tympanum subdermal. A rudimentary sheath-like nuchal crest present in males only, comprising 5–6 (5) triangular scales. Infralabials: 9−10 (10) on left side, 10−11 (10) on right side. Mental scale pentagonal or hexagonal, longer than wide, longer than length of rostral scale, in contact with 1 st infralabial on both sides; one pair of postmentals, smaller than mental, contacting mental and 1 st infralabial; contacting each other on posterior side; gular scales strongly carinate, larger than mid-ventrals, rhombic, set in regular series in males. SA 29.4−35.8 (17.6); dewlap long, well developed in adult males (absent in females); DL 23.2−33.3; DW 14.3−18.9.

Body slightly depressed dorsolaterally, MBW 5.8−8.9 (5.5); subtriangular in section (sub-triangular or sub-oval in females). Dorsal scales smaller than larger lateral scales, feebly to strongly carinate, unequal, irregular imbricate. Some larger dorsal scales in pelvic area form short, feebly defined transverse rows with scale angles pointing backwards. Larger lateral scales irregularly interspaced among smaller scales, strongly to feebly carinate, imbricate; scales on dorsal surface point backwards; dorsolateral scales (in mid-dorsolateral region) point upwards and backwards; lower lateral scales point backwards and downwards. Ventral scales uniform, strongly carinate, larger than lateral scales but smaller than larger gular scales in males. In females, ventral scales uniform, strongly carinate, and larger than lateral and gular scales. Mid ventral scales 95−98 (89).

Forelimbs much smaller than hindlimbs: UAL 11.2−12.8 (8.1), LAL 12.7−13.9 (7.9); FEL 23.7−26.7 (15.7), TBL 26.8−28.9 (17.2). Scales on forelimbs strongly carinate, sub-equal to ventrals; subdigitals bicarinate; distal end of keels on palm and foot elevated, acuminate; digits covered dorsally and laterally with carinate, elongate scales. Scales on hind limbs strongly carinate, larger than ventrals; subdigitals of hind limbs bicarinate; digits and claws laterally compressed; claws curved, pointed, each between two scales, one above and one below. Sub-digital squamation on left manus—I: 8−9, II: 11−14, III: 13−17, IV: 15−17, V: 9−11 in males (I: 8, II: 14, III: 15, IV: 16, V: 10 in female); on left pes—I: 9−15, II: 11−14, III: 12−17, IV: 21−28, V: 7−8 in males (I: 10, II: 13, III: 13, IV: 25, V: 8 in female). Finger lengths (left manus)—IV: 9.1−10.9> III: 8.1−9.7> II: 7.2−8.0> I: 5.2−5.9> V: 5.1−5.8 in males (IV: 7.0> III: 6.2> II: 5.6> I: 3.3> V: 3.1 in female); toe lengths (left pes)—IV: 15.1−27.4> III: 13.8−16.8> II: 7.5−8.4> V: 4.9−6.3> I: 3.4−4.6 in males (IV: 18.1> III: 12.8> II: 5.6> V: 3.0> I: 2.9 in female); tail incomplete, badly damaged.

Osteology of O. wiegmanni ( Fig. 4 ). In comparison with Agasthyagama beddomii comb. nov., based on the scan of UMMZ 65556, male—data in brackets—also see Table 6. for measurements given as % of skull length. Strongly ossified skull with slightly enlarged orbit (vertical diameter 37.6% & horizontal diameter 47%) surrounded by bone and temporal fossa, bridged over by an arch formed mainly of squamosal and a well-developed paired jugal bones. Upper jaw comprises two rami which are fused anteriorly in the midline. Premaxilla smaller (6%) and single bone at the anterior end of ramus forming pointed anterior margin of upper jaw consisting an alveolar and a nasal process. Nasal process of premaxilla well developed, posterodorsally oriented between nasals; the alveolar process of premaxilla slightly depressed and broad. Premaxilla articulating with maxillae posterolaterally, vomer posteroventrally, and nasals posteriorly. The maxilla is more elongate (53%), forming lateral margin of the upper jaw. Alveolar processes of both maxillae and premaxillae bear 14 (3 premaxillary + 11 maxillary) small, pleurodont type teeth along its ventral edge. Nasals paired, articulate anteriorly with the premaxilla, posteriorly with the frontal, anterolaterally with the maxillae and posterolaterally with the prefrontal bones. Frontal single (32.9%), forming roof of the skull, sharply concave inside, deeper and perforated by a single “V” shaped foramen at the anterior end. Prefrontal bones roughly rectangular, forming anterior end of the orbital cavity. Postfrontal bones forming posterior region of orbit, slightly shorter (22.2%) and slender. Parietal comparatively larger (33.3%) and flatter, fused to form single bone which forms the roof of the cranium in the parietal region. It has a slightly smaller oval parietal foramen (2.1%). This bone posteriorly bears supra-temporal processes, each of which articulates with the quadrates, squamosal, supra-temporal and paraoccipital process of the exoccipitals of its side. Jugal bone is entire comparatively smaller (9.8%), present below postorbitals and articulating below with posterior end of maxillae. It has an enlarged, distinct, foramen located at its centre. Temporal fossa smaller (24.8%) and roughly triangular in dorsal aspect. In ventral view, the palatine bones larger (22.6%) are separated by pyriform space, elongated, anteriorly much border, articulating anteriorly with vomer via vomerine and maxillae with maxillary process; and posteriorly articulate with pterygoids via pterygoid process. Parabasisphenoid larger (21.8%), is a single element lying between the pterygoid anteriorly and the basioccipital posteriorly and it forms the anterior floor of the braincase. The anteromedial part of the parabasisphenoid bears a long and slender cultriform process that articulates anteriorly with pterygoid bones. Supraoccipital smaller (6.8%), forming posterior roof of basioccipitals, present anterior to occipital condyle and posterior to parabasisphenoid. In dorsal view, supraoccipital located between parietal and occipital condyle. Anteriorly, bone is separated from posterior margin of parietal but the process ascendens provides a connection between parietal and supraoccipital. Quadrates are slightly longer (18.4%) articulating with the lower jaw and occupy the posterolateral corner of the skull. In the ventral view of the skull, two comparatively elongated, second ceratobranchials (67.1%) extending backwards onto belly. Vertebral column comparatively longer (257.2%) consisting of 4 cervical vertebrae, 16 thoracic vertebrae and 4 lumbar vertebrae. Atlas to 4 th vertebrae do not bear any ribs; 5 th and 6 th thoracic vertebrae have short ribs and next 7 th to 14 th vertebrae have long ribs; 7 th thoracic vertebrae bear longest ribs which are closely connected to almost posterolateral edges of the sternal plate. Pectoral girdle situated at the anterior end of trunk composed of two halves one situated on either side of interclavicle and sternum. Interclavicle lodged in a groove on the mid-ventral surface of the sternum. Sternum in form of a roughly rhomboidal plate of cartilage present embedded in ventral thoracic wall. Anterolateral edges of sternum articulating with coracoids and epi-coracoids of pectoral girdles; postero-lateral borders bear two small facets for articulation with the sternal ribs; and posterior end of sternum bears two sternal ribs. The sternal plate is well developed with six posterolateral sternal ribs and two posterior sternal ribs. Humerus (72.2%), radius (64.1%) and ulna (64.8%) longer and slender. Digits long and slightly enlarged. Pelvic girdle situated at the posterior end of the trunk, comparatively enlarged in size, consists of two halves meeting in the midline by a vertical ligament. Femur comparatively longer (137.2%) with tibia (128.6%) and fibula (125.5%) in size.

Distribution and Natural History. Otocryptis wiegmanni occurs in the wet zone of Sri Lanka, i.e., the southwestern quarter of the island, from near sea level to up to 1300 m asl. The species has been recorded from Kandy, Kegalle, Kithulgala, Ratnapura and Sinharaja regions of Sri Lanka. Diurnal, largely terrestrial, secondarily arboreal, chiefly insectivorous and oviparous laying up to 3–7 ellipsoid eggs ( Somaweera & Somaweera 2009).

Figures

FIGURE 3. Otocryptis wiegmanni in life from Kandy, Central Sri Lanka; (A) head and forebody lateral view, (B) head dorsal view, (C) gular region lateral view showing extended dewlap and maroon spot, (D) gular region ventral view, (E) entire with extended dewlap, (F) general view of an adult male showing the weak ‘nuchal crest’.

FIGURE 4. Otocryptis wiegmanni (UMMZ 65556, adult male), osteology showing general skeleton, (A) entire ventral view, (B) entire lateral view, (C) entire dorsal view, and skull (D) ventral view, (E) side view, (F) dorsal view