Calotes farooqi Auffenberg and Rehman, 1993

Gowande, Gaurang, Pal, Saunak, Jablonski, Daniel, Masroor, Rafaqat, Phansalkar, Pushkar U., Dsouza, Princia, Jayarajan, Aditi & Shanker, Kartik, 2021, Molecular phylogenetics and taxonomic reassessment of the widespread agamid lizard Calotes versicolor (Daudin, 1802) (Squamata, Agamidae) across South Asia, Vertebrate Zoology 71, pp. 669-696 : 669

Calotes farooqi Auffenberg and Rehman, 1993 stat. nov.

Figs 8 , 9

Calotes versicolor nigrigularis Auffenberg and Rehman, 1993

Calotes versicolor farooqi Auffenberg and Rehman, 1995 (nom. nov. for C. v. nigrigularis )

Holotype.

Pakistan • 1 ♂; Khyber Pakhtunkhwa, Mansehra, Shargal (corrected to Sarhan); 34.3°N, 73.4°E, 1077 m a.s.l; 15 Jun. 1990; PMNH field crew leg.; FLMNH/UF 79470.

Other material (morphological vouchers).

All from Pakistan • 1 ♂; Punjab, Kotli Syedan ; 32.7604°N, 73.0736°E, 834 m a. s. l.; 16 Sept. 2018; Daniel Jablonski leg.; GenBank MW901312 (16S), MZ489214 (COI); CUDZ DJ 7902; • 2 ♀; Khyber Pakhtunkhwa, Mansehra; 34.5610°N, 73.2635°E, 834 m a. s. l.; PMNH 414, PMNH 1355 .

Genetic diagnosis.

The analysis included a total of three samples (MW901312-14: 16S, MZ489214: COI) from the localities Tangora, Gulbandi Buner (localities 85 and 86 in Figure 2 , both west of the Indus River in Buner District, ca. 60 km. by airline from the type locality Mansehra) and Kotli Syedan (locality 77 in Figure 2 ) from the northern hilly regions of Pakistan, representing C. farooqi stat. nov. The maximum within species divergence (not including the divergent lineage from the southern hilly and southern plains regions) was 0.4% at 16S. The species is at least 3.5% and 17% divergent from C. versicolor , 3.6% and 16.9% divergent from C. irawadi , 3.9% and 15.5% from C. vultuosus comb. nov., and 4.9% and 20.4% from C. calotes at 16S and COI respectively (Table 2 ). The species was recovered as sister to C. calotes (Fig. 3 ).

Diagnosis and comparison.

A medium to large species of Calotes , adult males ranging from 94-99 mm in SVL, body moderately compressed; head relatively long; dorso-lateral scales posterodorsally oriented, large, weakly to strongly keeled, homogeneous; ventral scales smaller than the dorso-lateral scales, strongly keeled, mucronate; anti-humeral fold absent; two distinct spines in the supratympanic region; nuchal and dorsal crest continuous, distinct; scales of the nuchal crest large, those of dorsal crest slightly smaller, slightly recurved, ending at the top of the base of the tail, males with distinct black patches on both sides of the lower jaw, extending into the forebody, in the breeding season.

The species can be separated from all the members of the C. versicolor group by a combination of characters: absence of crescent-shaped patch of granular scales at the insertion of the forelimbs (vs. present in C. emma , C. grandisquamis , C. jerdoni , C. mystaceus , and C. nemoricola ), 41-51 Mid-body scale rows (vs. 49-65 in C. emma , 27-35 in C. grandisquamis , 58-63 in C. maria Gray, 48-60 in C. minor Hardwicke and Gray, and 45-58 in C. mystaceus ); nuchal crest scales well-developed, dorsal crest scales much smaller, more or less equal in size (vs. nuchal spines much longer, dorsal spines reduced in C. maria and C. nemoricola ; nuchal spines much longer than dorsal spines in C. calotes , C. emma , C. grandisquamis ); two well-separated supratympanic spines (vs. row of 3-4 compressed supratympanic spines in C. grandisquamis and C. nemoricola , 8-9 compressed spines above tympanum in C. calotes ; two parallel rows of supratympanic scales in C. jerdoni and C. maria , single well developed postorbital spine in C. emma ). The species differs from C. paulus and C. zolaiking primarily by the homogeneous scalation on the dorsolateral region (vs. heterogeneous) and a comparatively well-developed dorsal crest. From the dubious species C. bhutanensis , the species differs in possessing longer head, concave orbital region, and by the absence of a row of erect scales on the sides of the neck. From C. chincollium , C. nigriplicatus , and members of the C. mystaceus complex ( C. bachae , C. geissleri , C. goetzi , C. mystaceus , C. vindumbarbatus , sensu Wagner et al. (2021)) by the absence of an oblique fold of skin in front of forelimbs or shoulder (vs. present). From the Sri Lankan congeners ( C. ceylonensis , C. desilvai , C. liocephalus , C. liolepis , C. manamendrai , C. nigrilabris , C. pethiyagodai ) the species differs by its posterodorsal orientation of lateral body scales (vs. posteroventral) and absence of shoulder pit (vs. present). The species differs from C. irawadi by its larger adult male body size (average male SVL 96.5 mm in C. farooqi stat. nov., vs. 82.4 mm in C. irawadi ), lesser number of dorsal crest scales (41 in C. farooqi stat. nov. vs. 48.9 in C. irawadi ); from C. htunwini by the posterodorsal or vertical orientation of scale rows on the sides of the neck and supra-axillary area (vs. horizontal in C. htunwini ).

The species is most similar in appearance to C. versicolor and C. vultuosus comb. nov., however, can be differentiated from C. versicolor by its smaller adult male body size (average male SVL 97 mm, vs. 108 mm in C. versicolor , female SVL 82 vs. 92 in C. versicolor ), dorsal crest scales composed of comparatively shorter scales, which become shorter progressively to the base of the tail (vs. dorsal crest composed of longer scales, dorsal crest continues to the base of the tail in C. versicolor ), lesser number of eyelid scales, Eyelid 9-11 (vs. 10-15 in C. versicolor , 11-14 in C. vultuosus comb. nov.), the shape and the size of the scales between the nasal shield and the orbit (large, <6 in a row between the nasal shield and the orbit in C. farooqi stat. nov., vs. small,>6 in C. versicolor and C. vultuosus comb. nov.), by the acuteness of the region between the nostril and the orbit (more acute in C. farooqi stat. nov., less acute in C. vultuosus comb. nov. and C. versicolor ). C. farooqi stat. nov. also has slightly lower number of SnS (6), in comparison to C. versicolor and C. vultuosus comb. nov. (generally 7).

The species also differs from C. versicolor and C. vultuosus comb. nov. in terms of breeding coloration of the adult males. The head and the forebody of the males of C. farooqi stat. nov. attain grey to black colour, except the vertebral region and the parts of the head above the jaw muscles; the lower portion of the orbit turns black, the black colour extends to the outer surfaces of the forelimbs, the inner surfaces of the forelimbs and the region of the venter intervening the limbs turn greyish black. In contrast, C. versicolor males attain a yellowish colour, trunk and orbital region turn orange, forelimbs and hind limbs turn dark to black, the black patches under the throat do not extend anteriorly onto the jaw muscles; C. vultuosus comb. nov. males attain a cream to brown body colour, the head and the anterior two-thirds of the trunk attain orange colour, which may extend on to the forelimbs, the posterior parts of the body remain duller.

Description of the holotype FLMNH/UF 79470 (Fig. 8).

A large-sized male, SVL 94 mm, tail complete, 245 mm. The specimen is in a mediocre condition, the head is bent to the left, tail to the right; forelimbs adpressed to the body. A vertical incision on the ventral surface between the insertion of the forelimbs.

Head large (HL/SVL 0.27), snout tip pointed in dorsal perspective; region between the nasals and the orbit slightly concave, acute, covered by heterogeneous, juxtaposed scales; loreal region from the nasal shield triangular, the anterior border of the nasal shield at the vertex of the triangle; eight CanthR scales, elongate, with their ends overlapping; supraciliary and canthal edge sharp, giving the head a flat appearance laterally from the dorsal perspective; large supraocular scales do not form shields; become parallel to convergent at the supraocular region; dorsal edges of the forehead divergent, bordered by scales of the canthus rostralis and supraciliary region, rostral broader than high; nasals single on each side, subtriangular, pointed anteriorly, rounded posteriorly, separated from the first Suplab by one prenasal scale, from rostral by two scales, from each other by six SnS; nostrils round, in a single large nasal shield each, centrally placed; scales between rostral and SnS small, juxtaposed; SnS heterogeneous, the median SnS smallest; scales of the forehead posterior to SnS sub-imbricate, very irregular in shape and size, some rugose; HeadSLn 12, bordered anteriorly by the rostral, and posteriorly by the single, large interparietal shield; orbit large, encased in a sock of granular scales, separated from the nasal shield by rows of 4-5 scales, from Suplab by three scale rows; HeadSTr 14, between the posteriormost supraciliary scales on each, just anterior to the interparietal; eye opening bordered by two rows of non-granular scales, outer row composed of larger, square-shaped scales, inner row similar in shape, slightly smaller; eyelid scales (Eyelid) 13; region between the orbit and the tympanum covered by rows of 6-7 smooth, irregular in shape; tympanum large, round, naked; 3.1 mm at its greatest height; supratympanic scales smooth to weakly keeled; two enlarged supratympanic spines, separated from each other by 3-4 scales; anterior spine slender, shorter, posterior spine prominent; posterior region of the jaws swollen, bulges out, covered by subtriangular, imbricate, postero-ventrally directed scales, some of these mucronate; upper border of the jaw muscles conceal the lower portion of the tympanum; labial scales large, sub-rectangular; Suplab 12; Inflab 11; two parallel rows of scales border the upper margin of the Suplab, lower originates above the second Suplab, separating the nasal shield and the second Suplab, terminates near the last Suplab; the upper row originates slightly posteriorly, terminates abruptly above the ninth Suplab; interparietal large, irregularly pentagonal, bordered by 9-10 smooth, heterogeneous scales; nuchal crest starts 3-4 scales behind the interparietal; mental shield large, single, subtriangular,, mental narrower than rostral; two pairs of elongate postmentals, anterior pair narrow, separated by a single small chin scale posterior to the mental; posterior pair broader, separated by three small scales; chin scales posterior to the postmental scales small, progressively become larger towards the throat; scale rows of the posterior region of the throat large, mucronate.

First nuchal scale smallest, size of the nuchal scale increases modestly towards the median nuchal scale; beyond which, the size reduces slightly, nuchal crest composed of lanceolate, recurved spines, which continue into dorsal crest scales; dorsal crest scales slightly smaller than nuchal crest scales, composed of spines that progressively become smaller towards the middle of the back; continues to the base of the tail, beyond which the mid-dorsal crest row terminates at the keeled tail scales; paravertebral scales recurved, strongly keeled, mucronate; dorso-lateral scales sub-triangular, imbricate, distinctly keeled, posterodorsally oriented; those in the supra-axillary region slightly smaller than those of the mid-body; ventrals strongly keeled, smaller than the dorso-lateral scales, mucronate, imbricate; ventral scales between the insertion of the forelimbs weakly keeled.

Limbs long and slender, covered with keeled scales, similar to dorso-laterals in shape and size, forming parallel longitudinal rows; scales on the dorso-ventral surfaces of the thigh weakly to moderately keeled, those on the crus and sole strongly keeled; first finger shortest, third and fourth almost equal, fifth longer than first; relative length of toes 4>3>5>2>1; subdigital lamellae bicarinate, keels sharp, 20 under the fourth finger, 24 under the fourth toe; tail slender, swollen at base, TailL 245 mm.

Coloration in preservation: Anterior two-thirds of the body generally greyish-tan, dorsum distinctly black till mod-body, black coloration fades away in the posterior half, vertebral and paravertebral row of scales paler; venter greyish black at the pectoral region, slightly duller towards the vent, colour abruptly changes to greyish-cream; a dark ventral midline runs from the pectoral region to the vent; dorsal surface of the forelimbs dark, lower surfaces slightly paler; hindlimbs and tail greyish-cream; supra-axillary region greyish-black, the black patches continue anteriorly along the lower surface of the jaws and gular region, except the mid-gular region which is strikingly lighter, pinkish in colour; head generally greyish above, paler laterally and posteriorly; eyelids light grey anteriorly, a small black patch near the anterior corner of the eye, a large black patch near the posterior corner of the eye.

Original description.

Auffenberg W, Rehman H (1993). Asiatic Herpetological Research 5: 14-30.

Etymology.

The specific epithet is a patronym in genitive singular case, dedicated to Farooq Ahmed, former Director, Zoological Survey Department, Pakistan.

Variation in the material examined.

One male specimen (CUDZ DJ7902) was examined. The mensural characters in mm are: HL 28.1; HW 21.0; HH 16.9; SVL 99.0; TrunkL 49.2; UpArmL 16.6; The meristic characters are: SnS 6; HeadSTr 11; HeadSLn 13; CanthR 8; Eyelid 9; Suplab 11; Inflab 12; TempSp 2; Mid-body 41; 4FingLm 17; 4ToeLm 24. Additionally, two female specimens were examined. The means for the mensural characters in mm are: HL 20.2; HW 16.8; HH 12.4; JawW 15.7; SnEye 8.6; NarEye 3.6; EyeEar 5.6; SnW 5.1; Interorb 11.8; SVL 82; TrunkL 41.3; TailH 7.3; TailW 7; PectW 14; PelvW 10.5; SnForeL 17.6; UpLegL 17.5; CrusL 17.9; HindfL 25.3; 4ToeLng 13.4; HindLimbL 60.5. The means for meristic characters are: SnS 6; HeadSTr 11; HeadSLn 14; CanthR 8; Eyelid 11; Suplab 12; Inflab 12; TempSp 2; Dorsal 42; Mid-body 43; 4ToeLm 25. The ranges for each of these characters are given in Table 4 .

Distribution.

The species has been reported from the northern hilly regions of Pakistan, while the samples from the southern plains represent a hitherto undescribed species (localities 78, 79 in Fig. 1 , 2 ). One of us (DJ) collected samples from the northern mid-elevation, hilly regions of Pakistan. Calotes farooqi stat. nov. has not been reported from within the political boundaries of India, although surveys in the hilly regions of north India adjoining Pakistan would be necessary to further comment on the presence of the species in India. Similarly, populations probably resembling C. farooqi stat. nov. have been reported from parts of Afghanistan (provinces Kabul, Laghman, Nangarhar, Paktia) adjoining Pakistan ( Wagner et al. 2016); however, their systematic status needs further morphological and genetic investigation.

Figures

Figure 1. A map of south and Southeast Asia representing the localities from which morphological and / or primary or secondary genetic data were collected. Red circles indicate the localities from which C. versicolor sensu stricto (Clade 1) originate, yellow samples indicate those of C. irawadi sensu lato (Clade 2), blue circles represent the localities of C. vultuosus comb. nov. (Clade 3), green circles indicate Clade 4, with C. farooqi stat. nov. (77, 83, 85, 86) and C. aff. farooqi (78, 79). A single sequence of Calotes irawadi from Huanan Province, China (locality 73) is not marked on the map, since it was very distant to all other localities, as also those whose precise localities were unknown. Triangles indicate type localities of the respective species. See supplementary Table S 1 for details on molecular and / or morphological vouchers used, and their respective localities.

Figure 2. The ML phylogenetic output representing the ML + BI phylogeny based on the 1130 bp long 16 S + COI combined dataset representing the clustering pattern of the sequences of the Calotes versicolor species complex. Values at the nodes indicate ultrafast ML bootstrap values / BI posterior probabilities. Colour code follows that of Figure 1. Calotes grandisquamis and Calotes nemoricola were used as the outgroup to root the trees.

Figure 3. Maximum Likelihood 16 S gene tree based on the ~ 416 base pair sequence data revealing the position of the species of the Calotes versicolor complex within the genus Calotes. Numbers at the nodes indicate bootstrap values. The tree was rooted using the Mantheyus phuwuanensis (Manthey and Nabhitabhata, 1991) as the out group. The tree was analysed under the TIM 2 + F + I + G 4 model of sequence evolution, implementing 1000 ultrafast bootstraps on the web platform of IQ-TREE. Species delimited in this study are colour coded: Green - C. farooqi stat. nov.; Yellow - C. irawadi; Red - C. versicolor; Blue - C. vultuosus comb. nov.

Figure 8. Holotype of Calotes farooqi stat. nov. FLMNH / UF 79470. A. full body ventral, B. head dorsal, C. head ventral, D. head lateral. Photographs by Coleman Sheehy.

Figure 9. Calotes farooqi stat. nov. (CUDZ DJ 7902), from Kotli Syedan, Punjab, Pakistan, in life. Photograph by Daniel Jablonski.