Agasthyagama beddomii ( Boulenger, 1885 ) Srikanthan & Adhikari & Ganesh & Deuti & Das & Kulkarni & Gowande & Shanker, 2021

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 : 218-225

Agasthyagama beddomii ( Boulenger, 1885) comb. nov

( Figs. 6–9 , Tables 3 & 5)

Otocryptis beddomii Boulenger, 1885

Otocryptis beddomi — Inger et al. 1984

Otocryptis beddomei — Manthey 2010

Syntype. ZSI 15733, adult female, collected by R. H. Beddome from Sivagiri Ghat (9.14˚ N 77.24 ˚E), Tamil Nadu, India. Other syntypes are in the Natural History Museum, London, UK (fide Boulenger 1885; Das et al. 1998).

Other Material Examined. Adult males (n=2) INDIA ‒ CESL 99 , collected by Saunak P. Pal, from Bonnacaud estate (8.68˚ E 77.16 ˚E), Agasthyamalai hills, Kerala, in Aug 2010 ; CESL 268 , collected by Saunak P. Pal, from Kanayar (9.12˚ N 77.17 ˚E), Devar Malai hills, Kerala in Aug 2010 . Adult female (n=1)‒ZSI 26316, collected by L. Bindu and Party, from Pandimotta, Shendurney Wildlife Sanctuary, Kerala, on 19 Nov. 2016 .

Etymology. Named after its distribution range in the Agasthyamalai, that in turns gets its name from the Vedic sage Agathya—a sage in Hindu tradition—suffixed with the nomen “ agama ”, commonly used to refer to agamid lizards, a term believed to have been brought to Dutch Guiana ( Suriname) by West African slaves, whose term for chameleons was ‘agama’ in their native Gbe language ( Arends 2017). The gender is masculine.

Diagnosis (for genus and species). A small sized genus of draconine agamid lizard (average male SVL 38.2 mm; female SVL 40.8 mm), endemic to the Western Ghats of India, diagnosed by the following characters of lepidosis and osteology: dewlap appendage and second ceratobranchials not extending beyond forelimb insertion, dewlap diminished, males with small brightly colored gular sac; scales on gular sac equal sized; nuchal and dorsal crest not developed; small, rounded antehumeral pit present; 2 or 3 median rows of elongate and more or less regular scales present between supraoculars; temporal region with 1 or 2 enlarged, conical scales; yellow to orange coloration only in gular region; presence of large trihedral scales on the dorsolateral aspect of the body.

Osteological characters summarized for Agasthyagama beddomii comb. nov. as follows: (some in comparison with Otocryptis wiegmanni , also see Table 6): Two comparatively smaller, second ceratobranchials present, extending backwards to level of anterior chest; premaxilla bone forms convex anterior margin of upper jaw; nasal process of premaxilla partially developed; 13 teeth; comparatively smaller maxilla; frontal slightly broader and perforated by somewhat oval-shaped foramina at anterior end; pre-frontals roughly triangular; postfrontals elongate and broad, slightly inflated anteriorly; jugals larger with no distinct foramen, slightly enlarged, and posteriorly concave; temporal fossa enlarged, oval; palatines anteriorly thinner; cultriform process short and roughly cylindrical; sternal plate poorly developed with only two posterior sternal ribs and posterolateral sternal ribs absent. Femur, humerus radius, ulna and vertebral column comparatively shorter.

Comparisons with regional, related genera. Agasthyagama gen. nov. differs from South Asian draconine agamid genera as follows (only opposing suite of character states listed): Otocryptis —absence of the anteheumeral pit and presence of large dewlap that extends beyond the axilla; Draco Linnaeus, 1758 —presence of patagium membrane between ipsilateral limbs; dorsoventrally depressed trunk; exposed tympanum; presence of bilateral frills along tail; Calotes Cuvier, 1817 – exposed tympanum; dorsal crest with lanceolate spines, extending on to tail; dorsolateral body scales generally distinctly larger than ventral scales; supraorbital and temporal spines present in some species; Monilesaurus , and Microauris —exposed tympanum; dorsal scales equally sized; Psammophilus Fitzinger, 1843 —exposed tympanum and dorsoventerally depressed body; Salea —exposed tympanum; dorsal crest with lanceolate spines; caudal crest well-developed in adult males; dorsolateral body scales distinctly larger than ventral scales; supralabials always in broad contact with nasal scales; Sitana and Sarada ‒exposed tympanum, presence of a large dewlap and absence of fifth toe; Ceratophora —presence of well-developed or rudimentary nasal appendage in some species; dorsolateral body scales larger than ventral scales in some species; Lyriocephalus —presence of a bulbous rostral knob; dorsolateral body scales distinctly heterogenous, with a basal layer of tiny granular scales that are smaller than ventral scales, intermixed with very large, keeled trihedral tuberculate scales; 5–7 paravertebral rows of spines along trunk; canthal ridge raised and well-defined; Cophotis —dorsal scales very large, shield-like, loosely imbricate and chaotically arranged; dorsolateral body scales larger than ventral scales; tail prehensile.

Re-description of syntype ZSI 15733. Overall the syntype specimen is in good condition. Adult female; Snout to vent length, SVL 40.6 mm. Head longer than broad, HL 9.2 mm; broad, HW 6.9 mm; roughly triangular-shaped in dorsal aspect, DHW 5.9 mm; HH 6.2 mm; interorbital area concave; IOD between anterior end, 5.9 mm; between posterior end, 6.3 mm. Snout elongated, roughly pointed; SNL 5.5 mm; area between snout-tip and interorbital region concave; temporal region with 2 enlarged, conical scales surrounded by small, unequal sized carinate scales; upper temporal fossae slightly concave; SA 20.2 mm, 2.2 times larger than HL. Eyes small, HED 2.0 mm; orbital rim prominent, supraorbital region with 6 rows of large, carinate scales, the inner row extending forward to form a Y-shaped ridge on snout; superciliary scales carinate. Nostril oval, laterally orientated; E-N 3.7 mm, 1.5 times larger than SNL; IND 2.8 mm; canthus rostralis with 11 strongly carinate scales; nasal scales large, undivided, roughly oval, separated from 1 st supralabial by a row of scales; rostral separated from nasal by a single scale, in contact with 1 st supralabial on both sides. Supralabials: 11/11 (left/right). Cephalic scales irregular, smooth, feebly carinate. Tympanum subdermal. Infralabials: 9/10 (left/right). Mental scale pentagonal or hexagonal, longer than wide, in contact with 1 st infralabial on both sides; one pair of postmentals, smaller than mental, in strong contact with 1 st infralabial and mental; posteriorly separated by almost equal-sized gular scale; gular scales strongly keeled, larger than midventrals, rhombic, dewlap absent.

Body slightly compressed dorsolaterally; MBW 11.9 mm; dorsal scales slightly larger than smaller lateral scales, feebly to strongly carinate, unequal, imbricate, large trihedral scales present on dorsolateral aspect of the body; smaller lateral scales irregularly arranged, 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 and smaller gular scales in both sexes; mid-ventral scales 89.

Forelimbs comparatively much smaller than hindlimbs, UAL 6.3 mm, LAL 6.1 mm; FEL 13.2 mm; TBL 13.1 mm; scales on forelimbs strongly carinate, sub-equal to ventrals; sub-digitals 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. Subdigital squamation on left manus—I: 8, II: 12, III: 13, IV: 15, V: 6, on left pes—I:5, II: 11, III: 15, IV: 22, V: 5; relative finger lengths of left manus—IV: 5.2> III: 4.7> V: 3.4> II: 3.3> I: 2.3, toe lengths of left pes—IV: 8.6> III: 4.1> II: 3.6> I: 2.6> V: 1.5; tail incomplete, broken.

Variation in other referred materials, male (female). measurements (in mm)—DL, 6.9‒7.2; DW, 1.1‒1.3; SVL, 37.3‒39.0 (40.9); MBW, 5.3‒5.9 (9.7); TAL, 23.7‒48.1 (64.0); TW, 1.7‒3.8 (3.3); AG, 14.8‒15.6 (15.5); HL, 12.1‒13.5 (9.6); HW, 6.7‒7.4 (8.5); DHW, 5.7‒6.3 (6.7); HH, 6.4‒6.7 (7.8); UAL, 7.2‒7.6 (6.3); LAL, 7.6‒8.6 (6.3); FEL, 12.3‒13.9 (12.3); TBL, 13.9‒14.8 (14.0); HED, 2.2‒3.0 (2.5); SA, 14.9‒15.8 (21.5); SNL, 5.6‒6.0 (5.9); E-N, 3.4‒3.8 (4.0); IND, 2.5 (2.1); IOD: anterior end, 5.4‒5.7 (6.1), posterior end, 6.1‒7.2 (6.8); finger lengths (left manus), I: 1.6−1.8 (1.8), II: 2.5−2.9 (2.6), III: 3.0−4.2 (3.7), IV: 4.4−4.6 (3.9), V: 2.3−2.7 (2.5), toe lengths (left pes), I: 2.2−2.4 (2.0), II: 3.5−3.6 (2.5), III: 6.3−7.6 (5.4), IV: 9.9−10.1 (9.2), V: 1.7−2.0 (2.0). Scalation —SL, 10−11 (10−11); IL, 10−11 (9−11); scales on canthus rostralis, 11 or 12 (11 or 12); supraorbital scale rows, 5 or 6 (6); subdigital squamation on left manus—I: 8−9 (9), II: 11−13 (12), III: 13−14 (13), IV: 14−15 (14), V: 6−7 (7), on left pes—I: 5−6 (6), II: 10−11 (12), III: 15−17 (16), IV: 22−23 (22), V: 4−5 (5).

Osteology of A. beddomii comb. nov. In comparison with Otocryptis weigmanni , based on the scan of CESL 099, male ( Fig. 9 )—data in brackets—also see Table 6 for measurements given as % of skull length. Skull strongly ossified with smaller orbit (36.4% vertical diameter and 39.7% horizontal diameter) surrounded by bone and temporal fossa, bridged over by an arch formed mainly of squamosal and well-developed paired jugals. Upper jaw consists of two halves or rami which are fused anteriorly in midline. Premaxilla slightly larger (6.6%) and single bone at the anterior end of ramus forming convex anterior margin of upper jaw consisting an alveolar and a nasal process. Nasal process of premaxilla partially developed, posterodorsally oriented between nasals; alveolar process of premaxilla slightly depressed and broad. Premaxilla articulating with maxillae posterolaterally, vomer posteroventrally, and nasals posteriorly. Maxilla smaller (46.2%) forming lateral margin of upper jaw. Alveolar processes of both maxillae and premaxillae bearing 13 (3 premaxillary + 10 maxillary) small, pleurodont type teeth along its ventral edge. Nasal bones paired, articulating with premaxilla anteriorly, with frontal on posterior side, maxillae on anterolateral side and prefrontal bones on posterolateral side. Frontal slightly larger (33.9%) single forming the roof of the skull, sharply concave inside and perforated by vaguely oval-shaped foramen at the anterior end. Pre-frontal bones approximately triangular, forms anterior of the orbital cavity. Postfrontals larger (24.8%) forming the posterior region of orbits, slightly elongated and broad, slightly inflated anteriorly. Parietal smaller (26.4%), flat, broad anteriorly and narrow posteriorly, fused to form single bone, forms the roof of the cranium in the parietal region, with a comparatively smaller, oval parietal foramen. This bone posteriorly forms supra-temporal processes, each of which articulating with quadrates, squamosal, supra-temporal and paraoccipital process of exoccipital of its side. Jugal much larger (17.4%), entire, flatter, and posteriorly concave, lies below postorbitals and articulating below with posterior end of maxillae and lacks any visible foramen. Temporal fossa larger (27.3%) and roughly oval in dorsal aspect. In ventral view, palatine bone much smaller (14.4%) separated by the pyriform space, anteriorly slightly thinner, articulating anteriorly with vomer via vomerine and maxillae with maxillary process; and posteriorly articulating with pterygoids via pterygoid process.A smaller parabasisphenoid (18.2%) forms the anterior floor of braincase, whereas basioccipital constitutes posterior part, present between pterygoid anteriorly and basioccipital posteriorly. Bone bears short and somewhat cylindrical cultriform process located at anteromedial part of parabasisphenoid articulating anteriorly with pterygoid bones. Basioccipital becoming posterior floor of braincase, whereas a comparatively larger supraoccipital (11.6%) forming posterior roof of basioccipital, situated anterior to occipital condyl and posterior to parabasisphenoid. In dorsal view, supraoccipital located between parietal and occipital condyl. Anteriorly, this bone separated from posterior margin of parietal but process ascendens provide connection between parietal and supraoccipital. Quadrates slightly smaller (17.4%), articulating with lower jaw, and occupying posterolateral corner of skull. In the ventral view of skull, two comparatively small, elongated, second ceratobranchials (52.1%) extending backwards onto chest. Vertebral column comparatively shorter (227.3%) with 4 cervical vertebrae, 13 thoracic vertebrae and 4 lumbar vertebrae. Atlas to 4 th cranial vertebrae do not bear any ribs; 5 th thoracic vertebrae have short ribs and next 6 th to 13 th thoracic vertebrae have long ribs. Pectoral girdle situated at anterior end of trunk composed of two halves one present on either side of interclavicle and sternum. Interclavicle lodged in a groove on mid-ventral surface of sternum. Sternum roughly rhomboidal, cartilaginous, present embedded in ventral thoracic wall. Anterolateral edges of sternum articulating with coracoids and epi-coracoids of pectoral girdles; postero-lateral borders bears two small facets for articulation with sternal ribs; and posterior end of sternum bears two sternal ribs. Sternum plate poorly developed with only two posterior sternum ribs and posterolateral sternum ribs absent. Humerus (65.3%), radius (53.7%) and ulna (57%) comparatively shorter. Digits comparatively short. Pelvic girdle situated at posterior end of trunk, smaller in overall size, consisting two halves meeting in midline by a vertical ligament. Femur comparatively shorter (117.4%) with tibia and fibula unrenderable from the scan.

Distribution and Natural History. Agasthyagama beddomii comb. nov. is a strictly terrestrial and diurnal lizard that is insectivorous and breeds by oviparous mode ( Smith, 1935). Inger et al. (1984) observed nearly 75 individuals of this species in the Ponmudi hills in lower elevation (110‒650 m asl) western slopes of Agasthyama-

lai. The bulk of the sightings were of individuals “scampering over leaf litter” with a few sightings on vegetation such as shrubs and trees up to 1.5 m perching height. They even recorded gravid females with a clutch size of 3‒5 near-term ova and remarked that the breeding season appears restricted. Daniels (1991) reported this species from the Balamore and Maramalai hills further south and in the Ashambu ranges at 400‒500 m asl, on leaf-litter strewn forest paths. Its body coloration and patterns also resembling dried leaves on the forest floor, most likely used for camouflage. Subsequently, Jose et al. (2007) elaborated on the ethology and spatio-seasonal dynamics in the Myristica swamp ecosystem in Kulathupuzha-Shendurney areas further north. The authors also remarked on the apparent variations in sightings of this species in monsoon and other seasons, across riparian ( Myristica swamp) vs. non-riparian habitats. Later Chandramouli (2009) worked on the spatial distribution and microhabitat associations of Agasthyagama beddomii comb. nov.

Figures

FIGURE 6. Agasthyagama beddomii comb. nov. syntype ZSI 15733, adult female (A) close-up of the specimen, entire view, (B) with jar label visible.

FIGURE 7. Agasthyagama beddomii comb. nov. live uncollected individuals from the Agasthyamalai Hills, Kerala, India, showing (A) head dorsal view, (B) head ventral view, (C) head lateral view, (D) entire, lateral view, (E) head lateral view with extended gular sac, (F) entire dorsal view, (G) gular profile, showing differential colouration.

FIGURE 8. Agasthyagama beddomii comb. nov. showing (A) ventral and (B) dorsal sides of referred materials, adult males, CESL 099 and CESL 268; (C) profile photo and (D) drawing of hemipenis.

FIGURE 9. Agasthyagama beddomii comb. nov. (CESL 099, 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.