NHMUK 1946.8.11.54, adult male from Khasi Hill, India.

NHMUK 1946.8.50., adult male; NHMUK 1946.8.11.49, NHMUK 1946.8.11.51, NHMUK 1946.8.11.53, NHMUK 1946.8.11.56, adult females. Same collection information as the lectotype.

Calotes jerdoni can be diagnosed from members of the species complex by a combination of following morphological characters: (1) body size large, SVL 65.7–102.2 mm; (2) tail slender, long, TAL 271.0–324.3% SVL; (3) posterior-most conical scales of parallel ridges of head not elongated into spine shape, relatively low; (4) inferior row of conical scales 1 or 2 scales away from superior tympanum, TRD 38.4–52.5% TD; (5) scales posterolateral of parietal bearing longitudinal keels; (6) gular scales large, 16–19 along central midline, homogeneous in posteromedial region, much larger than ventrals; (7) gular scales strongly keeled, mucronate with distinct, elongated tips; (8) distinct gular pouch absent, transverse gular fold absent; (9) body scales large, ABR 46–53; (10) shoulder fold present, short, covered with granular scales underneath; (11) neck scales oriented straight posteriorly; (12) axillary scales oriented posteriorly or posterosuperiorly at an angle less than 60°; (13) nuchal crests relatively short, triangular shaped, differentiated from dorsal crest, TNC 8.1–13.2% HL; (14) middorsal crest scale count 34–40; (15) scale rows around midbody 46–53; (16) ventral scale count 62–75, (17) F4S 19–26, T4S 25–31; (18) dorsal and ventral background coloration Yellow Green (Color 103) to Grass Green (Color 110) in normal condition, can change drastically to dark brown to blackish red under stress; (19) Medium Chrome Orange (Color 75) patches present on elbows, knees, and ankles; and (20) Pale Pinkish Buff (Color 3) to Medium Chrome Orange (Color 75) dorsolateral stripes from neck to basal tail present in some individuals.

Calotes platyceps Blyth, 1852: 354

NHMUK 1946.8.11.24, adult male from “Afghanistan” (see remarks on its type locality below); designated by Boulenger (1885).

Figure 5. 

Lectotype of Calotes maria (NHMUK 1946.8.11.24), showing dorsal (A), ventral (B), lateral head (C), ventral head (D), dorsal head (E), and axillary region (F). Photos by V. Deepak.

NHMUK 1870.11.29.30a, adult male; 1870.11.29.30b, adult female; both from Khasia (Khasi Hill), India; NHMUK 1860.3.19.1020, adult female from “Afghanistan”.

Figure 6. 

Non-vouchered individuals of Calotes maria in life, showing an adult (A) from Royal Manas National Park and a juvenile (B) from Garo Hill, Meghalaya. Photos by Abhijit Das.

On its original description, Gray (1845) described the species based on four specimens: three individuals (a juvenile, a subadult, and an adult) from “Affghanistan and a single half-grown individual from Khassia Hill”. Later Boulenger (1885) designated a male from Affghanistan as the “type” (= lectotype) of C. maria (NHMUK 1946.8.11.24), and he identified two of the young male syntypes of C. maria as C. jerdoni.

Based on Blyth (1853), the initial syntypes of C. maria, along with other species of the genus Calotes, were collected by Dr. Griffith and later handled by the East India Company before donating them to the Natural History Museum. Blyth (1853) suspected that those specimens from “Affghanistan” (= Afghanistan) and “Khassia Hills” (= Khasi Hill) were mixed during the handling by the East India Company. As the tropical species C. maria certainly cannot survive in desert or cold highlands of Afghanistan, and given Afghanistan is zoogeographically distinct from northeast India, the type locality of C. maria was certainly mistaken as Afghanistan and should be replaced by “Khassia Hills”, which refers to the Khasi Hill in Meghalaya State, Northeast India.

First described in 1852, C. platyceps was described in a short, rather simple description without a clear statement regarding its type specimen. Later, based on specimens collected from Khasi hills, Jerdon (1870) identified his specimens as C. platyceps and added a diagnosis of C. platyceps against C. maria, including a smaller body size, larger and fewer gular and body scales, and a narrower distance between supratympanic crest and the upper edge of tympanum. While the diagnosis of “C. platyceps” described by Jerdon (1870) matches the diagnosis of C. jerdoni, they were not based on the type specimen of C. platyceps. Subsequently, Boulenger (1885) without explanation, considered C. platyceps a junior synonym of both C. maria and C. jerdoni—a taxonomic treatment followed by subsequent authors (Boulenger 1890; Smith 1935; Che et al. 2020; Uetz et al. 2023).

Although the presence of a supratympanic crest on each side of the head aligns C. platyceps with the C. jerdoni complex, based on the limited information in the original morphological description, it is impossible to confirm whether C. platyceps represents a junior synonym of C. maria or C. jerdoni. As the type specimen of C. platyceps cannot be located, and since the original description of the species provides little useful information on its taxonomic identity, we treat the name C. platyceps as a nomen dubium. As such, the name is nomenclaturally still available, but it has no bearing on the taxonomic and nomenclatural decisions made here.

Calotes maria can be diagnosed from its congeners by a combination of following morphological characters: (1) body size large, SVL 101–117.5 mm; (2) tail slender, long, TAL 318.3–323.9% SVL; (3) posterior-most conical scales of parallel ridges of head distinctively elongated into spine shape, about same length as longest nuchal crest; (4) inferior row of conical/spinous scales 4 scales away from superior tympanum; (5) dorsal head scales posterolateral of parietal bearing longitudinal keels; (6) mental smaller than first pair of chin shields; (7) gular scales count 27–31, homogeneous in posteromedial region, much larger than ventrals, mucronate with distinct, moderate-lengthed tips; (8) body scales small, ABR 58–67; (9) distinct gular pouch absent, transverse gular fold absent; (10) shoulder fold absent; (11) neck scales oriented posterosuperiorly; (12) axillary scales oriented posterosuperiorly at an angle less than 60°; (13) nuchal crests long, elongated, lanceolate-shaped, differentiated from dorsal crest, TNC 17.6–24.0% HL; (14) middorsal crest scale count 47–52; (15) ventral scale count 63–76, (17) F4S 23–28, T4S 28–32; (18) dorsal background coloration Pratt’s Payne’s Gray (Color 293) to Sky Blue (Color 192) under long-term preservation, ventral Paris White (Color 139); and (19) short, Pale Pinkish Buff (Color 3) to Yellow Ocher (Color 14), oblique stripes present on dorsum, similar colored patches present on tail base.

Large sized Calotes, SVL 101.0 mm; body not compressed dorsally, lachrymiform in cross-section; tail swollen at base, long, TAL 322 mm, 318.8% SVL. Limbs slender, forelimb 52.7 mm, 52.2% SVL; hindlimb 69.8 mm, 69.1% SVL. Rostral rectangular, bordering four small scales excluding supralabials; supralabials 10/11, feebly keeled posteriorly; nasal elongated oval, bordering second supralabial, 6/7 scales away from orbit circle; loreals distinctively keeled, each bearing single keel; canthal ridge distinct, canthus rostralis scales 10/10, elongated except anterior most two, supraciliaries only slightly overlapping, overlapping length less than half of scale length; suborbital scale rows 3/3, about equal size, each scale bearing single, distinct, inferior-located lateral keel. Distinctively protruding, enlarged keeled scales each bearing multiple keels aligned in two parallel lateral ridges: superior row continuing from posterior most supraciliary to superior squamosal head, consisting 8 raised scales, anterior-most 2 and 4 scales conical on left and right, respectively, remaining 6 and 4 much more elongated into spikes; inferior row two small scales away from posterior mid orbit to inferior squamosal head, consisting 12/10 scales, anterior-most 5 sub-pyramidal or conical, remaining posterior scales elongated into spikes. Inferior one of parallel ridges four small scale rows away from anterior tympanum. Tympanum exposed, round, TD 10.5% HL; no post orbital spikes, posterosuperior tympanic and temporal spikes present, which are located on parallel ridges. Dorsal head scales heterogeneous in size and shape, slightly larger above eyes, distinctively smaller on temporal, moderately keeled, each bearing single keel; enlarged scales arranged in Y-shape three scales posterior to rostral along longitudinal midline, with posterior most scale on stem of “Y” figure largest; 17 scales transversely across dorsal head between and excluding supraciliary at widest point; parietal scale elongated, narrow, parietal eye present; scales posterolateral of parietal bearing longitudinal keels.

Except scales anterior to shoulder, all dorsal body scales broad triangular shaped, regularly arranged, imbricate, mostly homogeneous, weakly keeled; neck scales oriented posterosuperiorly; axillary scales oriented posterosuperiorly; remaining lateral and superior body scales oriented posterosuperiorly, inferolateral body scales more distinctively keeled, oriented straight posteriorly or inferoposteriorly; midbody scale row around body 67. Nuchal crest 13, distinctively elongated into spikes, much taller than remaining dorsal crests, tallest one 5.6 mm in height, 19.5% HL; dorsal crests much lower, curving posteriorly; mid-dorsal crest scale count 52 anterior to cloaca; shoulder fold absent. Scales of dorsal limbs more or less homogeneous, keeled. Dorsal tail scales homogeneous, feebly keeled or smooth anteriorly, more distinctively keeled posteriorly.

Mental spear-shaped, enclosed by and smaller than first pair of chin shields; chin shields feebly keeled, larger than nearby gulars, 5/6, anterior most 3/2 in contact with infralabials on each side, remaining ones separated by one or two rows of small scales; infralabials 10/10, more distinctively keeled than supralabials; gular scales all distinctively keeled; anterior gulars smaller, juxtaposed, various in shape and size; remaining posterior gular scales larger, similar size to dorsals, rhomboid shaped, imbricated, mucronate with moderate-length tip on posterior end of each gular; post mental gular scales 30 along medial line. Ventral body cut open from chest to pelvis, ventral scales imbricated, slightly smaller than larger gulars, distinctively keeled, carinated, ventral scale count 76 along medial axis from neck to cloaca. Ventral limb scales homogeneous in size, more distinctively keeled than dorsal limb scales; subdigital lamellae 27/28 under finger IV, 32/32 under toe IV. Tail scales distinctively keeled, carinated (ventral tail cut open at base).

CIB 1477/8380177, adult male from Yarang, Medog County, Nyingchi Prefecture, Xizang Autonomous Region, China.

Figure 7. 

Holotype of Calotes medogensis (CIB 1477/8380177), showing dorsal (A), ventral (B), lateral head (C), ventral head (D), dorsal head (E), and axillary region (F). Photos by Kai Wang.

KIZ 014003, 019546, adult females; KIZ 013861, subadult female; all from Beibeng, Medog County, Nyingchi Prefecture, Xizang Autonomous Region, China.

Figure 8. 

Topotypic individuals of Calotes medogensis in life. KIZ 07372 (A), KIZ 014003 (B), unvouchered individuals (C–G). Photos by Kai Wang and Jian Wang.

WII-ADR 1211, WII-ADR 452, both adult females from the East Himalaya (Boruah et al. 2022).

Calotes medogensis is diagnosed from congeners by a combination of the following characters: (1) body size large SVL 75.7–95.9 mm; (2) tail slender, long, TAL 286.7–326.1% SVL; (3) conical scales forming two parallel ridges from posterior eye to temporal head, posterior most 3–5 scales of superior ridge distinctively raised and conical, but not elongated into spikes; (4) inferior ridge of conical scales 2–3 scale-rows away from superior tympanum, TRD 36.9–63.2% TD; (5) dorsal head scales posterolateral of parietal bearing transverse keels; (6) mental smaller than first pair of chin shields; (7) gular scale count 20–25 along central medial line, larger than ventrals, mucronate with feeble tips; (8) gular scales keeled, but keel often extending less than ½ of scale length on posterolateral region; (9) gular pouch present in life, weakly developed, but absent after preservation; (10) transverse gular fold absent; (12) shoulder fold present, short, covered with granular scales underneath; (13) nuchal crests relatively short, lanceolate shape, dorsal crests moderately developed, TNC 5.2–7.3% HL in males, 10.3–15.7% in females; (14) neck scales oriented posterosuperiorly, larger than ventrals; (15) axillary scales oriented near vertically; (16) middorsal crest scale count 37–42, (17) scale rows around midbody 53–60; (18) ventral scale count 57–65; (19) F4S 25–29, T4S 30–34; (20) dorsal and ventral background coloration Yellow Green (Color 103) to Grass Green (Color 110) in normal condition, can change drastically to dark brown to blackish under stress; (21) region underneath shoulder fold Dark Carmine (Color 61) to Poppy Red (Color 63); (22) white to Medium Paris White (Color 140), narrow, oblique streaks present on lateral dorsum, running anterosuperiorly to posteroinferiorly, each streak formed by a single row of scale.

Medium sized Calotes, SVL 75.7 mm, body not compressed dorsally, lachrymiform in cross-section; tail complete, swollen at base, long, TAL 217 mm, 286.7% SVL. Limbs slender, forelimb 42.0 mm, 55.4% SVL; hindlimb 59.2 mm, 78.2% SVL. Rostral rectangular, bordering six small scales excluding supralabials; supralabials 11/11, feebly keeled posteriorly; nasal elongated oval, bordering second supralabial, six scales away from orbit circle; loreals distinctively keeled, each bearing single longitudinal keel; canthal ridge distinct, canthus rostralis scales 10/11, elongated except posterior most two, supraciliaries only slightly overlapping, overlapping length one third of scale length; suborbital scale rows 4/4, scales about equal size, each bearing single distinct lateral keel. Distinctively protruding, enlarged keeled scales each bearing single lateral keel aligned laterally in two parallel ridges: superior row continuing from posterior most supraciliary to superior squamosal head, consisting of 9/10 conical scales; inferior row from posterior mid orbit to inferior squamosal head, consisting of 10/10 scales, anterior five sub-pyramidal, posterior most four conical. Inferior ridge two small scale rows away from anterior tympanum. Tympanum exposed, round; no post orbital, posterosuperior tympanic, or temporal spikes. Dorsal head scales heterogeneous in size and shape, distinctively keeled, each bearing single longitudinal keel except ones on temporal, which bearing single transverse keel; enlarged scales arranged in Y-shape four scales posterior to rostral along longitudinal midline, with posterior most scale on each opening branch of “Y” figure largest; 17 scales transversely across dorsal head between supraciliary at widest point; parietal scale enlarged, spear-shaped pointing posteriorly, parietal eye distinct; scales posterolateral of parietal bearing transverse keels.

Except scales beneath shoulder fold, all dorsal body scales broad, triangularly shaped, regularly arranged, imbricate, homogeneous, weakly keeled; neck scales oriented posterosuperiorly; axillary scales oriented vertically upward; remaining lateral and superior body scales oriented posterosuperiorly, inferolateral body scales more distinctively keeled, oriented straight posteriorly; midbody scale row around body 46. Nuchal crest low triangular shaped, slightly damaged, ninth nuchal crest tallest, 1.2 mm in height, 1.0 mm in longitudinal width at base; nuchal crest gradually transitioned to lower, serrated dorsal crest posteriorly; mid-dorsal crest scale count 40 anterior to cloaca; distinct shoulder fold present, short, with fine granular scales underneath. Dorsal scales of dorsal limbs homogeneous, slightly larger on distal appendages, keeled. Tail scales homogeneous, feebly keeled, or smooth anteriorly, more distinctively keeled posteriorly.

Mental pentagonal, not enclosed by and smaller than first pair of chin shields; chin shields smooth anteriorly, feebly keeled posteriorly, slightly enlarged than nearby gulars, 6/6, anterior most 2 in contact with infralabials on each side, remaining ones separated by one or two rows of small scales; infralabials 11/12, feebly keeled; gular scales all distinctively keeled; anterior gulars small, juxtaposed, various in shape and size; remaining posterior gular scales larger, similar size to dorsals, rhomboid shaped, more homogeneous, imbricated, keel of individual scale only extending less than half of scale length; posterior gular scales slightly mucronate with feeble tips; post mental gular scales 23 along medial line. Ventral body scales imbricated, smaller than gulars and dorsals, distinctively keeled, carinated, ventral scale count 58 along medial axis from neck to cloaca. Ventral limb scales homogeneous in size, more distinctively keeled than dorsal limb scales; subdigital lamellae 29/27 under finger IV, 34/31 under toe IV. Ventral tail scales distinctively keeled, carinated.

The lateral sides of head are uniformly Yellow Green (Color 103). The dorsal surface of the head is darker than lateral sides, which are Light Grass Green (Color 103). Lateral sides of the head gradually transition to Pistachio (Color 102) or Light Grass Green (Color 103) posteriorly to the dorsal and lateral dorsum, and further inferiorly to Pale Cyan (Color 156) on ventrolateral dorsum. The region underneath the shoulder fold is Dark Carmine (Color 61) to Poppy Red (Color 63). Most individuals have white to Medium Paris White (Color 140) oblique streaks that run anterosuperiorly to posteroinferiorly, and each streak is formed by a single row of scale. The ventral surfaces of the head and body are uniform Yellow Green (Color 103). Some individuals have a small patch on the knee and ankle, respectively, which is Orange-Rufous (Color 56) in color.

Calotes medogensis is arboreal, inhabiting tropical forests at mid to low elevations, and individuals were observed sleeping on twigs and vines less than 2 m above the ground at night. Calotes medogensis is an insectivore, feeding on various beetles and other insects. Currently, the species is known from Medog County, Xizang Autonomous Region, China and the adjacent region in northeastern India (Boruah et al. 2022). In Medog, C. medogensis is sympatric with other agamids such as C. paulus, Japalura andersoniana, and Mictopholis austeniana. Possible predation may come from lizard-feeding snakes such as Lycodon cf. fasciatus, Oligodon spp., and Ptyas nigromarginata (Che et al. 2020). One of the voucher females (KIZ 014003) is gravid with 11 eggs. With no pressing threats and protection coverage by nature reserves in both China and India, we propose to list the species as Least Concern (LC) for China’s Red List Assessment and IUCN Assessment.

Calotes jerdoni – Zug et al. (2006: 60)

CAS 219992, adult male from Baw Khue Plantation, Nat Ma Taung National Park, Min Dat Township, Min Dat District, Chin State, Myanmar (21.38375°N, 98.89886°E, elevation 1787 m).

Figure 9. 

Holotype of Calotes iadina sp. nov. (CAS 219992), showing dorsal (A), ventral (B), lateral head (C), ventral head (D), dorsal head (E), and axillary region (F). Photos by Kai Wang.

CAS 219993, adult male, same collection information as the holotype; CAS 233207, adult female, from Chun Kyone, Haka Township, Chin State, Myanmar (22.77231°N, 93.56411°E, elevation 1705 m) on July 15, 2003; CAS 233229, 233233, 233235, adult males; CAS 233232, 233234, adult females, all from Falam Township, Chin State, Myanmar (22.81439°N, 93.55752°E, elevation 1630 m), collected on July 18, 2003.

Figure 10. 

Type series of Calotes iadina sp. nov. in life, including the holotype (CAS 219992) (A) and paratypes [CAS 233207 (B), CAS 224511 (C), and CAS 219993 (D)]. Photos by Jens V. Vindum.

The species name iadina is derived from Greek, which means “emerald” and refers to the bright green coloration of the new species.

Calotes iadina sp. nov. can be diagnosed by a combination of the following morphological characters: (1) body size large, SVL 63.2–102.5 mm; (2) tail slender, long, TAL 293.9–367.9% SVL; (3) conical scales of two parallel ridges triangular shaped, relatively low, not in spine shape; (4) inferior row of conical scales 2–3 scale rows away from superior tympanum, TRD 14.8–28.8% TD; (5) mental larger than first pair of chin shields; (6) gular scale count 16–19, much larger than ventrals, strongly keeled, mucronate, each bearding a distinctively elongated tip on posterior end; (7) gular pouch feeble or absent, transverse gular fold absent; (8) shoulder fold present, covered with granular scales underneath; (9) nuchal crests relatively short, lanceolate shape, TNC 8.27–13.3% HL, dorsal crests moderately developed; (10) neck scales oriented posteriorly or posterosuperiorly; (11) axillary scales oriented vertically or near vertically; (12) dorsal body scales keeled, oriented posterosuperiorly, larger than ventrals; (13) middorsal crest scale count 39–43, (14) scale rows around midbody 48–53; (15) F4S 22–27, (16) T4S 27–33; (16) dorsal and ventral background coloration Yellow Green (Color 103) to Grass Green (Color 110) in normal condition, can change drastically to brownish to blackish under stress; (17) shoulder fold Pale Neutral Gray (Color 296) to Jet Black (Color 300); (18) white to Pale Pinkish Buff (Color 3) patches sometimes present on elbows, knees, and ankles, and same colored dorsolateral stripes sometimes present from neck to base of tail.

The new species can be diagnosed from all congeners other than members of the C. jerdoni complex by having two parallel rows of conical or spinous scales on temporal region of the head (vs. absence). Within the C. jerdoni complex, C. iadina sp. nov. is most similar to C. jerdoni, but it differs from the latter by having vertical or near vertical orientation of axillary scales (vs. mostly oriented posteriorly or slightly upward, with an angle less than 60°; Fig. 4) and larger and fewer ventral scales (48–54 vs. 62–75). Moreover, the ANOVA analyses corroborated this statistical difference, recovering varying numbers of significantly different mean values between C. iadina sp. nov. and C. jerdoni, in which females of the new species differ from the females of C. jerdoni in having significantly different mean values head width (HD), interorbital distance (IOD), middorsal crest scale length (MD), tympanum diameter (TD), finger IV length (F4L), and orbital diameter (OD) (Appendix V). Males of C. iadina sp. nov. differ significantly from males of C. jerdoni in head length (HD), interorbital distance (IOD), tympanum diameter (TD), toe IV length (T4L), femur length (FEL), ventral scale count (VN), and orbital diameter (OD) (Appendix V).

For remaining species of the C. jerdoni complex, C. iadina sp. nov. differs from C. yunnanensis and C. medogensis by having larger and fewer gular scales along medial central axis (16–19 vs. 23–26 for C. medogensis, 20–23 for C. yunnanensis), presence of a distinct, elongated posterior tip on each gular scale (vs. absent or much shorter), larger and fewer ventral body scales (VEN 48–53 vs. 57–61 in C. medogensis, 54–62 in C. yunnanensis), and greatly overlapping supraciliaries (overlapping half of scale length vs. less than one third). Additionally, C. iadina sp. nov. differs from C. yunnanensis by having shorter nuchal crest scales (TNC 8.3–13.3% HL vs. 14.1–19.1%); from C. medogensis by having greatly overlapping supraciliaries (overlapping more than half of scale length vs. less than half); and from C. maria by having fewer scale rows between supratympanic ridge and superior edge of tympanae (1 or 2 rows vs. >3 rows), larger and fewer gular scales (16–19 vs. 27–31), much shorter nuchal crests (TNC 8.3–13.3% HL vs. >17.6%), the presence of shoulder fold (vs. absence), and the absence of spike-shaped scales on posterior end of the parallel ridges of the head (vs. presence). For the nomen dubium, C. platyceps, C. iadina sp. nov. differs by having more conical scales on the supratympanic ridge (6 or 7 vs. 8 or 9) and by the presence of a black stripe running through the eyes (vs. absence).

Adult male, body size medium, SVL 84.5 mm, body not compressed, cross-section lachrymiform shaped; tail slender, long, TAL 364.0% SVL. Head narrow, somewhat elongate, HW 58.9% HL, HD 53.0% HL; jaw muscular, slightly swollen, HWJ 111.8% HW; snout length moderate, SEL 38.6% HL. Rostral rectangular, length four times height, bordering five small scales excluding supralabials, single scale away from nasal; nasal oval, in contact with first two supralabials; supralabials 11/10, 3–5 feebly keeled; loreal slightly concave, loreal scales medium in size, all keeled, 5–7 scales away from orbit; eye surrounded by fine ciliary scales; suborbital scale rows 3/3, medial row much enlarged; canthal ridge distinct, canthus rostralis 10/11, supraciliaries overlapping about half scale length of consecutive ones. Two distinct ridges on temporal head, parallel anteriorly but joining posteriorly on squamosal region; superior ridge continuous from last canthal rostralis, consisting 7/6 conical scale that each bearing single lateral keel; inferior ridge starting post orbit, consisting 11/10 enlarged scales, anterior most two on each side smooth, convex, remaining ones conical, with posterior-most 2 largest and tallest, but none of them elongated into spikes; inferior ridges 1/2 scale rows away from tympana; region between two ridges slight concave; tympana exposed, oval, vertically oriented, TD 13.3% HL, OTD 22.0% HL; jaw scales enlarged, distinctively keeled, mucronate, imbricate. Dorsal head scales weakly keeled, heterogeneous in size and shape; enlarged scales arranged in Y-shape, single scales posterior to rostral along longitudinal midline, all about equal size; 14 scales transversely across dorsal head between supraciliary at widest point; parietal scale enlarged, narrow, elongated, parietal eye distinct; scales posterolateral of parietal bearing oblique keels.

Mental triangular, separating and much larger than first pair of chin shields; chin shield 8/7, first three on each side in direct contact with infralabials, remaining ones separated by single, small scale-row; first four chin shields smooth, remaining one each bearing single indistinct lateral keel; remaining gular scales all distinctively keeled, imbricate, mucronate with long tips, gradually increasing in size posteromedially, gular scales 16 along central-medial line.

Dorsal body scales feebly keeled, more distinct anteriorly on neck and inferiorly close to ventrolateral flank, mostly homogeneous in size; scales on neck oriented horizontally backward or obliquely upward at less than 45°; axillary scales smaller, oriented near vertically, which then gradually change to a more oblique orientation superiorly and posteriorly; shoulder fold present, distinct, with granular fine scales underneath; flank scales large, with minimal lateral keel on posterior tip only, imbricate in regular oblique rows, 45 scale rows at mid body. Crest distinct, nuchal crest not significantly differentiated from dorsal crests, lanceolate shaped, relatively short, TNC 11.9% HL; middorsal crest scale 41, gradually reducing height posteriorly, crest scale height equal width from anterior one third of body, remaining dorsal crests longer than height. Dorsal scales on limbs mostly homogeneous in size and shape, smooth or feebly keeled, keel status slightly more distinct proximally on lower appendages. Tail swollen at base, feebly keeled on anterior one sixth of length, gradually increasing in keel status posteriorly and forming carinate rows.

Ventral body scales smaller than gulars and dorsals, distinctively keeled, mucronate, imbricate, forming carinate rows; ventral scale row 51; F4S 27/25, T4S 32/33; ventral limb scales more distinctively keeled; ventral tail scales more distinctively keeled, carinate at base.

In life, background coloration is light grass green. Distinct black radial stripes are present around eyes, forming a transverse oriented Y-shape across eyes, with its opening facing posteriorly. The shoulder fold is black. Numerous indistinct, black, oblique transverse stripes are present on flank. The gular and ventral body are lighter and more yellowish. Two parallel rows of single white scales are present on ventrolateral body between axillary and groin. Under preservation, the dorsal background green body coloration turns into dark bluish gray, and the ventral color turns into light blue.

Morphometric and pholidosis variation are summarized in Table 4. Additionally, all paratypes have more distinctly keeled scales on the dorsal surface of the limbs than the holotype. Two preserved female paratypes (CAS 233232 and 233234) possess light-gray dorsolateral stripes, which extend from the neck to the base of tail. Some individuals also have indistinct oblique transverse stripes (CAS 233233 and 233234).

Based on collection notes, specimens were found along a road during morning hours, although the species is likely to be arboreal. Some of the females were gravid, suggesting that breeding occurs during July. Currently, the species is only known from western Myanmar along the Chin Hills, and no imminent conservation threat is known. The species is known to be capable of changing its ground color, from blackish brown to its original green (Fig. 9). Given the lack of data on habitat quality and population trend, we propose to list the new species as Data Deficient (DD) for its IUCN assessment.

Calotes maria – Anderson (1879: 806)

Calotes jerdoni – Pope (1935: 468–469), Smith (1935: 194–195), Zhao and Adler (1993: 188), Zhao and Yang (1997: 156–157), Zhao et al. (1999: 90–91), Yang and Rao (2008: 189), Wang et al. (2020: appendix), Wang et al. (2022: appendix)

ZSI 6571, adult female from Teng-yue, western Yunnan, China.

Figure 11. 

Holotype of Calotes yunnanensis (ZSI 6571), showing dorsal (A), ventral (B), lateral head (C), ventral head (D), dorsal head (E), and axillary region (F). Photos by V. Deepak and Abhijit Das.

Most studies have considered Teng-yue as interchangeable with the modern Tengchong County in Yunnan Province (Zhao and Adler, 1993), including the museum label of the holotype at ZSI. Although Teng-yue includes the modern Tengchong County, they are not interchangeable: Teng-yue was the administrative unit of the Qing Dynasty and encompassed a much larger area west of the Gaoligong Mountains, which also includes modern-day Dehong Prefecture. Therefore, the type locality of the species should be revised as both modern-day Dehong Prefecture and Baoshan (which include Tengchong County).

Figure 12. 

Topotypic individuals of Calotes yunnanensis in life, including KIZ 014146 (A), KIZ 014148 (B), KIZ 014147 (C), and a non-vouchered individual (D). Photos by Yu-Fan Wang and Zheng-Pan Duan.

Annandale (1905) noted that the holotype of C. yunnanensis is a male. However, our examination shows it to be a female (e.g., no distinct bulge at base of tail as in male specimens such as KIZ 014148, 029968, and 039747). Also, in the original description, Annandale (1905) measured the head length of the holotype as 34 mm and body length as 65 mm, which differs greatly from our measurement of head length (27.2 mm) and SVL (94.8 mm). As Annandale (1905) did not specify his measurement method, these differences could be explained by different measurement methods.

Topotypes: KIZ 014148, 029968, 039747, adult males; KIZ 015993, 014146, 014147, 030060, 50692, 034358, adult females; all from Yingjiang County, Dehong Prefecture, Yunnan, China. CAS 224511, adult male, from Nagmung Township, Putao, Kachin State, Myanmar (27.440861°N, 97.8950278°E, 573 m); CAS 221514, adult female, from Naung Mon Township, Putao, Kachin State, Myanmar (27.44122°N, 97.91875°E, 535 m); CAS 221551, adult female, from Machanbaw Township, Putao District, Kachin State, Myanmar (27.28994°N, 97.858472°E, 708 m); CAS 245278, adult female from Lahe township, Khandi District, Sagaing Division, Myanmar (26.324194°N, 95.44875°E, 900 m); CAS 245364, adult female, from Laung Nguk Village, Lahe Township, Sagaing Division, Myanmar (26.1590278°N, 95.529167°E, 833m).

Calotes yunnanensis is diagnosed from congeners by a combination of the following characters: (1) body size large, SVL 73–120 mm; (2) tail slender, long, TAL 311.8–353.7% SVL; (3) conical scales forming two parallel ridges from posterior eye to temporal head, posterior most 3–5 scales of superior ridge distinctively differentiated but relatively low; (4) inferior ridge of conical scales 2–3 scale-rows away from superior tympanum, TRD 38.4–52.5% TD; (5) dorsal head scales posterolateral to parietal bearing oblique or lateral keels; (6) mental larger or about equal to first pair of chin shields; (7) gular scale count 21–25 along central-medial line, equal or larger than ventrals, strongly keeled, slightly mucronate, keel extending across entirety of each scale and elongated on posterior end into very short tips; (8) gular pouch present in life, weakly developed, but absent after preservation; (9) transverse gular fold absent; (10) shoulder fold present, short, covered with granular scales underneath; (11) nuchal crests long, lanceolate shape, dorsal crests moderately developed, TNC 14.1–21.2% HL; (12) neck scales and dorsal body scales weekly keeled, oriented upwards and backwards, larger than ventrals; (14) axillary scales oriented near vertically; (13) middorsal crest scale count 38–45, (14) scale rows around midbody 51–60; (15) ventral scale count 54–68; (16) F4S 26–29, T4S 31–34; (17) dorsal and ventral background coloration Yellow Green (Color 103) to Grass Green (Color 110) in normal condition, can change drastically to dark brown to blackish under stress; (18) shoulder fold Pale Neutral Gray (Color 296) to Brick Red (Color 36); (19) white to Pale Pinkish Buff (Color 3) patches present on elbows, knees, and ankles; and (20) Pale Pinkish Buff (Color 3) dorsolateral stripes from neck to basal tail present in some individuals.

Calotes yunnanensis can be diagnosed from all congeners other than members of the C. jerdoni complex by having two parallel rows of conical or spinous scales on temporal region of the head (vs. absence) and by the absence of post orbital spikes (vs. presence).

For members of the C. jerdoni complex, C. yunnanensis was confused with C. maria and C. jerdoni, but it can be diagnosed from C. maria by the presence of a shoulder fold (vs. absence), a shorter distance between supratympanic ridge and tympanum (1–3 scales away vs. four or five scales); and from C. jerdoni by having a different shape of gular scales (heterogeneous in size, smaller in posteromedial region, without elongated tips or tips very short vs. homogeneous in size, with distinct, elongated tips), smaller gular scales and higher scale count (22–25 vs. 16–19), and by having longer nuchal crest scales (TNC 14.1–19.1% HL vs. 8.1–13.2%).

Morphologically, C. yunnanensis is most similar to C. medogensis, but it differs from the latter by having obliquely or longitudinally oriented keels on scales posterolateral to the parietal (vs. transversely oriented keels), larger mental scale relative to first pair of chin shields (vs. smaller), more distinctly keeled gular scales, particularly on posterolateral region (keel extending across the entirety of each scale vs. only extending across posterior half or less), and by the presence of white dorsolateral stripes on some individuals (vs. always absent) (Fig. 4).

Adult female, body size medium, SVL 94.8 mm, body not dorsally compressed, cross-section triangular shaped; tail slender, broken. Head somewhat elongate, HW 63.6% HL, HD 60.4 % HL; jaw not distinctively swollen; snout length moderate, SEL 44.4% HL. Rostral rectangular, length four times height, two scales away from nasal; nasal oval, separated from first supralabials by single scale; supralabials 11/10, all smooth; loreal region slightly concave, loreal scales medium in size, all keeled, 6 scales away from orbit; eye surrounded by fine ciliary scales; suborbital scale rows 3/3, medial row much enlarged; canthal ridge distinct, canthus rostralis 10/10, supraciliaries overlapping about one third length. Two distinct ridges on temporal head, parallel anteriorly but joining posteriorly on squamosal region; anterior ridge continuous from last canthal rostralis, consisting 10/10 conical scale that each bearing single lateral keel; inferior ridge starting post orbit, consisting 9/10 enlarged scales, each bearing single lateral keel; inferior ridges 3 scale rows away from tympana; region between two ridges slight concave; tympana exposed, oval, vertically oriented, TD 12.7% HL; jaw scales enlarged, moderately keeled, slightly mucronate, imbricate. Dorsal head scales weakly keeled, heterogeneous in size and shape; slightly enlarged scales arranged in Y-shape, single scales posterior to rostral along longitudinal midline, all about equal size; 16 scales transversely across dorsal head between supraciliary at widest point; parietal scale enlarged, rectangular, parietal eye distinct; scales posterolateral of parietal bearing oblique keels.

Mental triangular, smaller than and separating first pair of chin shields; chin shields 6/7, first two on each side in direct contact with infralabials, remaining ones separated by one or two small scale-row; first four chin shields smooth, remaining one each bearing single weak lateral keel; remaining gular scales all distinctively keeled, imbricate, mucronate, some bearing single short tip, gradually increasing in size posteromedially, gular scale 23.

Dorsal body scales feebly keeled, mostly homogeneous in size; scales on neck oriented obliquely upward at about 45°; axillary scales smaller, oriented near vertically, which then gradually changing to a more oblique orientation superiorly and posteriorly; shoulder fold present, distinct, with granular fine scales underneath; flank scales large, mostly smooth, some with minimal lateral keel on posterior tip only, imbricate in regular oblique rows, 56 scale rows at mid body. Crest distinct, nuchal crest not significantly differentiated from adjacent dorsal crests, lanceolate shaped, relatively long; middorsal crest scale 45, gradually reducing height posteriorly. Dorsal scales on limbs mostly homogeneous in size and shape, smooth or feebly keeled, keel status slightly more distinct proximally on lower appendages. Tail not swollen at base, broken in four sections, scales feebly keeled on anterior part, gradually increasing in keel status posteriorly and forming carinate rows.

Ventral body scales smaller than gular and dorsals, distinctively keeled, mucronate, imbricate, forming carinate rows; ventral scale row 58; ventral scales of limbs more distinctively keeled. Limbs slender, partially broken at elbow and knee joints.

The holotype is nearly uniform light-gray, with the temporal region slightly darker, and the ventral surface paler. For newly collected specimens, the coloration in life varies drastically based on the mood and status of the lizards. The background coloration of dorsal and lateral surfaces ranges from Cinnamon (Color 255) to bright Pistachio (Color 102). Some individuals possess Light Buff (Color 2) dorsolateral stripes from the neck to the anterior one third of tail, some Light Buff (Color 2) or Clay Color (Color 18) oblique stripes on dorsum, and others are uniformly colored with no ornamentation patterns. All individuals possess Light Buff (Color 2) patches on limb joints, either on elbows, knees, ankles, or any combination of thereof. The palate and buccal mucosa are Jet Black (Color 300), but the gums and the tongue are Pink (Color 242).

Currently, C. yunnanensis has been recorded from the southwestern Yunnan Province of China and Sagaing and Kachin States of Myanmar. It likely inhabits northern Shan State in Myanmar as well. Anderson (1879) speculated that C. yunnanensis (identified as C. maria at the time) is less arboreal than C. versicolor and C. emma, because the collecting site was mostly deforested. However, based on our field observations, we found this is not true: C. yunnanensis is highly arboreal, and individuals were found inhabiting well-forested tropical and subtropical regions in Southwest Yunnan.

In China, although the habitat of C. yunnanensis is well-protected by nature reserves, it has very restricted range, and has been targeted by domestic illegal pet traders (yingjiang.gov.cn, 2019). On the other hand, globally, C. yunnanensis has a considerable distribution range that consists of well-preserved habitats. Therefore, we assess the species as Near Threatened (NT) for China’s domestic Red List Assessment, and as Least Concern (LC) globally based on IUCN criteria.