Morphological and molecular differences in two closely related Calotes Cuvier 1817 (Squamata: Agamidae: Draconinae) with the first record of Calotes medogensis Zhao & Li, 1984 from India Author Boruah, Bitupan 0000-0001-8829-6069 Wildlife Institute of India, Chandrabani, Dehradun, Uttarakhand, 248001, India. & bitupan. kaz @ gmail. com; https: // orcid. org / 0000 - 0001 - 8829 - 6069 bitupan.kaz@gmail.com Author Narayanan, Surya SM Sehgal Foundation Centre for Biodiversity and Conservation, Ashoka Trust for Research in Ecology and the Environment (ATREE), Royal Enclave, Srirampura, Bangalore, Karnataka, 560064, India. Author Deepak, V. Senckenberg Dresden, Königsbrücker Landstrasse 159, 01109 Dresden, Germany. Author Das, Abhijit Wildlife Institute of India, Chandrabani, Dehradun, Uttarakhand, 248001, India. text Zootaxa 2022 2022-12-13 5219 5 433 455 journal article 208662 10.11646/zootaxa.5219.5.3 e4f02d98-62b3-4535-905f-31f9febb8d8b 1175-5326 7431376 F5233380-1CB1-4DF4-87D8-674C87672CBA Calotes medogensis Zhao & Li, 1984 Material examined: WII-ADR1211, a female ( Figure 3A,B )collected from Ramsing( 28°39ˊ22.71˝ N , 94°58ˊ46.22˝ E , Elevation: 601 m asl .), Mouling National Park , Upper Siang , Arunachal Pradesh on 28 October , 2021 ( Figure 1 ); WII-ADR452, a female ( Figure 3 C, D ) collected from Poba Reserve Forest , Arunachal Pradesh , on 30 September , 2018 ( Figure 1 ) . FIGURE 1. Map showing type localities and collection localities of Calotes medogensis (red) and Calotes jerdoni (blue). Type localities are represented by star. 1. Medog, China 2. Ramsing, Mouling National Park, Arunachal Pradesh, India 3. Poba Reserve Forest, Arunachal Pradesh, India, 4. Malki forest, East Khasi Hills, Meghalaya, 5. Dzuleke, Nagaland, India. Diagnosis: A moderate-sized agamid lizard, SVL 76–95.9 mm ( n= 3), TailL 212–271 mm ( n= 3). Nuchal and dorsal crests continuous, consisting of erect compressed scales, distinctly pronounced at nuchal crests than the dorsal crests, smaller posteriorly. Two series of supratympanic spines present. Body laterally compressed, scales at the trunk heterogeneous in size, larger than ventrals, larger at the mid-body towards dorsal side, dorsally and posteriorly oriented, 40–42 dorsal scales, 53–58 rows of scales at midbody, 88–91 ventrals. Mental smaller than chin shields, first pair of chin shields are narrow and elongated, separated from each other by two small scales, the gular scales at the mid-line are distinctly smaller than the adjacent gular scales away from the mid-line. Dorsally and ventrally green in colour. Morphological description based on newly collected materials: Morphometric and meristic characters are given in the Table 2 . Moderate-sized lizard, SVL 82.8–95.9 mm ( n= 2), head distinct from neck ( Figure 3 ), moderately large, longer than width (HeadW/HeadL= 0.67–0.68), slightly concave in front, snout rounded in dorsal view, longer than orbital diameter (OrbD/SnEye= 0.65–0.74), loreal slightly concave, supraciliary and canthal ridge sharp, nostril elliptical, nasal large, orbit three times larger than tympanum, pupil rounded, tympanum oval, rostral wider than mental, dorsal snout and head scales heterogenous in shape and size ( Figure 6 A, D ), scales at interorbital region, parietal region and occipital region have spinules forming irregular keel, scales at the loreal region, below and behind the orbit keeled, heterogenous in shape and size, scales surrounding the eye are granular, feebly conical, scales surrounding tympanum are smaller than that of mandibular region and nape, supralabials nearly rectangular, elongated, smooth, rostral wide and narrow, interparietal scale with a distinct pit, two series of ridge on temporal region above tympanum ( Figure 6 C, F ), upper row is consists of 9–10 scales, continuous with supraciliary scale to neck, posterior two or three scales of the upper row more pronounced or spinous, the lower row just above tympanum consisting of 9–10 scales and separated by two to three scales from tympanum, scales posteriorly more pronounced forming spine. Mental subtriangular or nearly hexagonal, smaller than chin shields, infralabials smooth, narrow and elongated, first pair of chin shields are narrow and elongated, separated by two small scales, posterior chin shields are separated from infralabials by a series of narrow scales, scales between the chin shields are smaller than the rest of the gular scales, irregular in shape and size ( Figure 7 B, E ), feebly keeled, gular scales posteriorly pointed, keeled, subimbricate, gular scales (except the middle scales) larger than that of the belly, the gular scales at the mid-line are distinctly smaller than the adjacent gular scales away from the mid-line ( Figure 7 C, F ). Scales on neck are similar shape, posteriorly pointed; upwardly and posteriorly directed, size increases dorsally and posteriorly, 8–9 nuchal spine, size increases posteriorly ( Figure 6 C, F ), length of the longest nuchal spine was more than twice the width of it (LongN/WidthN = 2.19–2.71, n= 2). Body laterally compressed; triangular in cross section. Dorsal crest reduced to mere ridge posteriorly. Dorsal and lateral scales of the trunk larger than ventrals, heterogenous in size; larger at the mid-body towards dorsal side, feebly keeled; pointed, dorsally and posteriorly oriented, 40–42 dorsal scales, 53–58 rows of scales around trunk at midbody, ventral scales smaller than that of dorsal and lateral, strongly keeled; heterogenous in size, pointed, 88–91 ventrals, scales at the axilla, groin and insertion of hind limb are smaller, a fold consisting of very small scales present in front of the shoulder. Limbs moderate and thin. Forefoot length nearly equal to upper arm length (UparmL/ForefL= 0.99–1.03). Crus length and thigh length almost equal (CrusL/UplegL= 1.02–1.09). All scales on limbs are keeled, smaller at the insertion, scales on ventral side of thigh are smaller than that of crus. Digits slender; with strong and curved claw, lamellae entire, bicarinate (more than two keels at the base). Tail long, two to three times of SVL, rounded, gradually tapering posteriorly, dorsal scales slightly smaller than that of ventral, scales at the base near cloaca smaller than the rest, ventral scales strongly keeled, forming continuous ridge. Colouration in preservative: WII-ADR1211, head, anterior part of back, forelimb, crus and foot dorsally bluish grey, large black patch from axilla to middle of the trunk, longer on left side, lower half of the trunk yellowish green, vertebral scales bluish grey all along the body, thigh scales dorsally green with black edge, almost anterior one third of the tail bluish grey, posteriorly greyish brown, three series of whitish scales on back arranged in “˄” shape. Fold on shoulder black. Head laterally bluish grey. Anterior gular scales reddish, rest gular scales intermixed with grey and yellowish green, chest scales yellowish, anterior part of belly intermixed with grey and yellowish scales, posterior part of the belly light yellow, hind limbs ventrally light yellow, forelimbs greyish, tail at the base and some extent of anterior part greenish yellow, posteriorly turn into grey. Small light brown patch on elbow and knee, a large light brown patch on heel to the base of 5 th toe. WII-ADR452, dorsally bluish grey, slightly dark patches on head, neck, above axilla and thigh, a large black patch from axilla to groin covering dorsolateral and lateral side of the trunk on both sides, ventrally entire body grey. Light cream coloured patch on elbow, knee and heel, larger on heel. Colouration in life: Dorsal, lateral and ventral side of head green ( Figure 9 ), dark stripes present on orbit radiating from eye, sometimes faint. Dorsum and tail dorsally green, series of discontinuous white scales on back arranged in “˄” shape. A dark brown fold/pit on shoulder, ventrally belly and tail light green, light brown patch on elbow, knee and heel. Morphological comparison: Calotes medogensis closely similar to Calotes jerdoni , but it can be differentiated from the latter by more pronounced nuchal spines, i.e. nuchal spines distinctly longer than that of the dorsum (vs. nuchal spines not distinctly longer than those of the dorsum), length of the longest nuchal spine more than twice the width (vs. length of the longest nuchal spine less than twice the width), scales on trunk oriented obliquely upward (vs. scales on trunk oriented backward) ( Figure 8 ), posterior 2–3 scales of the upper row of ridge above tympanum are spinous (vs. all the scales are similar), mental smaller than the first two pairs of chin shields (vs. mental larger than the chin shields) ( Figure 7 ), the first two pairs of chin shield are longer than wide (vs. chin shields are as long as wide) ( Figure 7 ), first pair of chin shield are separated by two small scales below mental (vs. first pair of chin shield are separated by small single scale below mental) ( Figure 7 ), the scales between the chin shields are irregular in shape (vs. scales between the chin shields are almost similar shaped), the gular scales at the mid-line are distinctly smaller than the adjacent gular scales away from the mid-line (vs. gular scales at the mid-line are nearly equal to the adjacent gular scales away from the mid-line) ( Figure 7 ), number ventrals is more in Calotes medogensis (88–91) than that of Calotes jerdoni (77–78). Calotes medogensis can be differentiated from Calotes maria by the presence of dark brown fold on shoulder (vs. absent) (Smith 1935), black stripes on orbit originating from eyes present (vs. absent) ( Günther 1870 ). Natural history: We encountered the individuals on bushes, commonly observed roosting on shrubs and ferns along roadside and forest edges at night. We observed individuals changing body colour from green to brown ( Figure 9B ). FIGURE 2. Phylogenetic relationship of Calotes spp based on ND2 gene. A. BI phylogeny with Bayesian posterior probability support values, B. ML phylogeny with bootstrap support values. Scalebar = substitutions per site. Outgroups except P. blanfordanus are not shown in the figure. Labelled in blue text are sequences generated in this study. FIGURE 3. Newly collected specimens of Calotes medogensis . A. & B. WII-ADR1211, C. & D. WII-ADR452. Scale bar = 10 mm. FIGURE 4: Newly collected specimens of Calotes jerdoni . A. & B. WII-ADR999, C. & D. WII-ADR966. Scale bar = 10 mm.” TABLE 2. Morphometric and meristic data of newly collected specimen of Calotes medogensis and Calotes jerdoni . All the measurements are given in millimetre. Bilateral characters are given in right, left order separated by comma. ‘-‘ indicates no data recorded.

Species	C. medogensis		C. jerdoni
	WII- ADR1211	WII- ADR452	WII- ADR966	WII- ADR999	BMNH 1946.8.11.54 (Lectotype)	BMNH 1946.8.11.49 (Paralectotype)	BMNH 1946.8.11.50 (Paralectotype)	BMNH 1946.8.11.51 (Paralectotype)	BMNH 1946.8.11.53 (Paralectotype)	BMNH 1946.8.11.56 (Paralectotype)
Location	Ramsing, Mouling NP, Arunachal Pradesh	Poba Reserve Forest, Arunachal Pradesh	Dzuleke, Kohima, Nagaland	Malki forest, East Khasi Hills, Meghalaya	Khasi Hills	Khasi Hills	Khasi Hills	Khasi Hills	Khasi Hills	Khasi Hills
Sex	Female	Female	Female	Male	Male	Male	Male	Female	Female	Female
SVL	95.9	82.8	93	63.2	76.6	76.7	75.3	88.8	102.2	74.1
HeadL	25.4	22.3	22.8	18.2	20.8	20.4	21.7	25	26.2	20.4
HeadW	17	15.2	15.6	11.7	13.8	12.4	13.3	15.2	17.7	14
HeadH	15.9	13.4	13.3	10.4	11.5	12.7	12.9	13.9	13.7	11.7
JawW	16	15	14.8	11.1	-	-	-	-	-	-
EyeEar	7.5	6.4	6.6	4.8	4.9	-	4.9	6	6.6	4.9
NarEye	6.6	5.4	5.5	3.6	5.8	-	4.9	6.1	5.8	5.1
InterOrb	11.5	9.4	9.4	7.4	9.1	-	8.5	10.8	11.2	8.8
SnEye	12	9.9	10.5	6.8	8.9	-	7.9	10.1	9.2	7.8
OrbD	7.8	7.3	7.6	5.4	6.3	-	5.8	6.7	7.1	6
SnW	6.9	6.1	6.7	4.5	4.9	-	5	5.5	6.1	5
TD	2.7	2.4	2.3	2.2	1.9	-	2.2	3.1	3.1	2
4FingLng	13.3	12.7	14.6	11.6	13.5	-	13	14	9.9	12.2
4ToeLng	18.7	18.9	20.2	15.1	17.6	-	16.5	18.8	17.1	15.3
ForefL	21.1	19	20.1	15.1	-	-	-	-	-	-
UparmL	21.7	18.9	20.3	14.8	13.8	-	13.7	15.9	17.2	13.1
LoArmL	17.3	16	16.6	11.4	13.7	-	13.4	15.9	16.6	11.7
HindfL	32.4	31.5	32.3	23.2	27.7	-	26.4	33.6	31	26
UplegL	22.1	22	21.4	15.3	18.2	19	17.9	20.2	22.6	18.9
CrusL	24.1	22.5	21.7	15.2	19	-	17	21.2	21.6	17.3
PectW	14.8	13.2	15.7	11.1	-	-	-	-	-	-
PelvW	10.9	10.5	12.3	9.1	-	-	-	-	-	-
SnForel	34.4	32.4	36.1	20.1	-	-	-	-	-	-
TailL	216	271	300	196	213	216	240	287	277	204
TailH	9.8	7.1	8.2	5.19	6.1	-	7.4	6.1	7.9	6.3
TailW	9.8	7.8	8	5.1	5.7	-	8.6	6.4	8.3	6.2
TrunkL	45	34.1	40.1	29.2	39.8	-	41.6	44.4	55.4	37
LongN	4.6	4.6	3.4	2.1	2.3	2.7	1.8	2.9	2.8	2.2
WidthN	2.1	1.7	2.1	1.2	1.2	1.4	1	2.1	1.6	1.7
CanthR	9, 9	10, 9	10, 10	9, 9	-	-	-	-	-	-
InfLab	9, 9	12, 12	10, 11	10, 10	11, 10	10, 9	10, 10	11, 11	-	-
Sns	9	7	11	10	-	-	-	-	-	-
HeadSLn	19	19	18	17	-	-	-	-	-	-
HeadSTr	15	19	14	18	-	-	-	-	-	-
SupLab	9,10	12, 11	11, 10	10, 10	10, 11	10, 10	10, 10	10, 9	-	10, 10
NC	8	9	9	9	-	-	-	-	-	-
4FingLm	28, 27	27, 27	28, 26	24, 23	-	-	-	-	-	-
4ToeLm	33, 32	32, 32	29, 34	29, 30	-	-	-	-	-	-
Dorsal	40	42	39	41	37	-	37	37	37	34
Midbody	58	53	58	54	46	-	46	53	47	49
Ventrals	91	88	77	78	71	-	73	73	68	75

Distribution: Calotes medogensis was known only from Yarang village, Beibeng Township of Medog County, China ( Che et al. 2020 ). We recorded the species in Poba reserve forest, Jengging, Tuting, Yubuk, Ramsing in Arunachal Pradesh . Earlier records of Calotes jerdoni from Komsing, Yembung, Balek ( Annandale 1912 ), Nameri National Park, Pake Tiger Reserve, Mouling National Park and Namdapha National Park ( Pawar & Birand 2001 ), Pasighat ( Sanyal & Gayen 2006 ), Zoological Survey of India campus, Itanagar ( Sinha et al. 2021 ) could be referred to Calotes medogensis based on the proximity to the current records. Also, a record of Calotes jerdoni from Bhutan by Wangyal (2011) needs further verification.