Calotes wangi hainanensis, Huang & Li & Wang & Li & Hou & Cai, 2023

Huang, Yong, Li, Hongyu, Wang, Yilin, Li, Maojin, Hou, Mian & Cai, Bo, 2023, Taxonomic review of the Calotes versicolor complex (Agamidae, Sauria, Squamata) in China, with description of a new species and subspecies, ZooKeys 1187, pp. 63-89 : 63

Calotes wangi hainanensis ssp. nov.

Fig. 4

Calotes versicolor Shi et al. 2011 in part; Huang et al. 2013 in part.

Type material.

Holotype. Adult male, CIB095629 (filed number HCL200908058). Collected from Chonggongbao Village, Zayun Town, Qiongzhong County, Hainan Province, China (18.9886°N, 109.55716°E, elevation 439 m) by Yong Huang and Bo Cai in August 2009.

Allotype. Adult female, CIB095630 (filed number HCL200908059), with the same locality and collector information as the holotype.

Other material examined.

Hainan Province, China: 1. Qiongzhong , Wanling Town (2♂, 1♀) , Hongmao Town (1♂, 1♀) , Zayun Town (1♂) , Hongmao Town (3♂, 1♀) , Limushan Town (1♀) . 2. Wuzhishan , Hongshan Town (3♂, 2♀) , Fanyang Town (5♀) . 3. Dingan , Donghong (1♀) . 4. Ledong , Yongming (1♀) . 5. Tunchang , Dalupo (1♀) .

Description of holotype.

Adult male, medium-sized body, SVL 88.34 mm. Trunk length 43.82 mm, head length 24.47 mm, head depth 14.41 mm, head width 16.08 mm, interorbital width 8.51 mm, snout width 5.29 mm, eye-ear length 6.30 mm, eye diameter 7.66 mm, naris-eye length 4.49 mm, jaw width 16.06 mm, snout-eye length 7.85 mm, snout-forelimb length 30.74 mm, tail length 283.89 mm, toe IV length 21.57 mm, finger IV length 13.41 mm, upper arm length 16.64 mm, lower arm length 13.59 mm, upper leg length 23.01 mm, Crus length 20.26 mm, forelimb length 44.82 mm, hindlimb length 71.77 mm, when hind limbs are adpressed forward to reach between eyes and tympanum.

Supralabial scale count 10:10, infralabial scale count 11:11, nasal-supralabial scale rows 1:1, suborbital scale rows 3:3, gular scale count 21, ventral scale count 46, finger IV subdigital lamellae count 22:22, toe IV subdigital lamellae count 27:27, post-tympanic scale count 1:1, post-occipital scale count 2:2, vertebral scales 38, mid-body scale rows 38, dorsal eyelid scales 13:13, no scales between anterior chin-shield, scales between nasal shield and orbit 5:5, state of scales on side of neck and adjacent shoulder area pointing obliquely upward, keels on these scales are weakly to strongly developed, nuchal and dorsal crest scales short, nuchal crest scales significantly larger than dorsal crest scales, dorsal crest shortening progressively before mid-body, gular fold and fold in front of the shoulder absent, no patch of granular scales in front of forelimb insertion, pre-axillary area with moderate to large scales, postorbital spine absent, scales on side of trunk pointing obliquely upward.

In ethanol, color is dark khaki or khaki with black or pale gray (105, 105, 105) markings. Tan and black mixed marking on top of head, nine brown (165, 42, 42) radial stripes around eyes, throat coloration burly wood with blank throat stripes, inner lip, oral cavity and tongue coloration is smoky white, ventral body is tan with dark stripes, presence of blank line on vent midline from throat to pelvis, six dark-khaki bands on dorsum of trunk between axilla and inguen, Seven to eight dark-khaki cross-bands on the fore- and hindlimbs, continuous ventral striping on trunk, and 22 cross-bands on tail (anterior nine bands are black, posterior 13 are brown).

Etymology.

The specific epithet of hainanensis refers to Hainan Island where the new subspecies was discovered. We suggest the English common name Hainan garden lizard and the Chinese name 中国树蜥雷公马亚种 (zhōng guó shù xī léi gōng mǎ yà zhǒng), which comes from a colloquial name for Calotes wangi hainanensis in Hainan Province, China, meaning Thor’s mount that can predict the weather.

Coloration in life.

This species is prone to color changes with different colors during the breeding season and as a result of stress. During the breeding season, adult males are uniformly dark orange (255, 140, 0) to orange on the front half (except for the fingers), with black patches on each side of the neck, darker longitudinal stripes on the chin and darker radial stripes around the eyes. The hind body and toes are uniformly khaki or dark khaki. The dark patches on the sides of the neck are approximately triangular and do not meet, extending from the front of the shoulder to the jaw angles. In the non-breeding season, animals are dark khaki or khaki with black markings.

The coloration of adult females is a relatively pale, uniform khaki over almost the entire body with dark horizontal stripes on a khaki background, and a pale yellow (255, 255, 224) or yellow (255, 255, 0) continuous or discontinuous longitudinal stripe on each side of the trunk in most individuals. They also have black or dark khaki radial stripes around the eyes and dark longitudinal stripes on the chin. Like the females, juveniles are a uniform khaki color over almost the entire body, with dark horizontal stripes and a pair of pale-yellow dorsolateral stripes. Under stress, the coloration of most individuals quickly changes to dark khaki with pale gray markings.

Variations.

The means for the measurements of morphological characters are DorsalS 39-52 (average 44.7), Mid-bodyS 36-46 (average 41.2, 2/167 specimen is 35), Eyelid 10-14 (average 12.4, 1/167 specimen is 9 or 15), NarEyeS 4-5 (average 4.9, 6/167 specimen is 6), 4ToeLm 22-27 (average 24.25, 4/167 specimen is 20, 5/167 specimen is 21 and 1/167 specimen is 30), HAF between the eyes and snout (6/167 specimen tympanum), FS absent (5/167 specimen present), SVL 66.14-109.1 mm (average 84.9 mm, 6/187 specimens longer than 100 mm). HAF between the eyes and tympanum (6/167 reaching tympanum, 21/167 reaching eye); FS absent (5/167 present). The ranges for each of these characters are given in Table 5 .

Calotes wangi wangi ssp. nov. can be separated from C. w. hainanensis ssp. nov. by the following characters: 1. Eyelid 10-13 (average 12.4, few 9 and 15) vs 13-14 (average 13, few 12. 2. Tail/SVL 1.59-3.36 (average 2.83) vs 1.77-4.01 (average 3.15). 3. Male, UpLegL/SVL 0.11-0.26 (average 0.23) vs 0.23-0.32 (average 0.25). 4. Male, CrusL/SVL 0.09-0.24 (average 0.21) vs 0.21-0.3 (average 0.24).

Diagnosis.

Calotes wangi sp. nov. can be separated from all other species of Calotes by having the following characters: 1. Medium-sized adult male, SVL <90 mm (66.86-109.1 mm, average 85.64 mm). 2. Smaller HeadW in males, 11.64-19.72 (average 14.84). 3. Larger HindLimbL/SVL 0.64-0.89 (average 0.75). 4. Larger 4ToeLng/SVL 0.08-0.3 (average 0.21). 5. Larger 4FingLng/SVL 0.11-0.21 (average 0.14). 6. Eyelid 10-14 (average 12.4). 7. NarEyeS 4-5 (average 4.9). 8. Scales on side of trunk point obliquely upward. 9. Patch of granular scales in front of forelimb insertion absent. 10. Scales on side of neck and adjacent shoulder area point obliquely upward; keels on neck scales are weakly to strongly developed. 11. Paired dark patches are approximately triangular, extending from the front of the shoulder to the jaw angles. 12. Coloration of tongue is orange. 13. Nuchal and dorsal crest scales short; dorsal crest shortening progressively before mid-body; nuchal crest distinctly larger than dorsal crest. 14. Supratympanic spines are short. 15. Fourth toe with claw can stretch between the eyes and tympanum when hind limbs are adpressed forward. 16. Fold in front of the shoulder is absent. 17. During breeding season, males are uniformly dark orange to orange on the front half (except for the fingers) with black patches on both sides of neck and darker radial stripes around eyes; Hind body and toes a uniform khaki or dark khaki.

Comparisons.

Calotes wangi sp. nov. was previously recognized as C. versicolor by previous authors but it can easily be distinguished from true C. versicolor (India) by adult males having shorter average SVL <90 mm vs> 90 mm in C. versicolor ; fewer scales between the nasal shield and the orbit 4-5 (average 5), <6 vs> 6; dark patches on neck extending to jowl vs not extending to jowl; nuchal and dorsal crest scales short and nuchal crest larger vs nuchal and dorsal crest scales well developed and nuchal crest scales only slightly larger than dorsal crest scales; pair of short supratympanic spines on each side of the head vs two well-separated supratympanic spines; coloration of tongue orange vs pale flesh color (239, 205, 197).

This species can be separated from others in the C. versicolor complex by the following characters ( Zug et al. 2006): 36-46 mid-body scale rows (average 41.2) vs 45-55 in the Nat-Ma-Taung Calotes versicolor group and 46-53 in the Moyingyi Calotes versicolor group; forearm stripe and paired nuchal spots present in adult vs absent in the Thai-east Calotes versicolor group.

The new species differs from Calotes irawadi by its smaller HeadW in males, 11.64-19.72 vs 13.08-28.3 in C. irawadi , shorter Interorb 4.59-12.89 vs 6.9-15.07; longer 4FingLng/SVL 0.11-0.21 vs 0.09-0.15; fewer mid-body scale rows 36-46 (average 41.2) vs 38-51 (average 44.9); fewer scales between the nasal shield and the orbit 4-5 (average 4.9) vs 5-7 (average 6.1); progressively shortened dorsal crest before mid-body vs progressively shortened after mid-body (especially in males), fourth toe with claw reaching between the eyes and tympanum when hind limbs adpressed forward vs reaching tympanum.

The new species can be separated from Calotes htunwini Zug & Vindum, 2006 ( Gowande et al. 2021) by the scales on the side of the neck and adjacent shoulder area pointing obliquely upward vs horizontally in C. htunwini ; the keels on these scales weakly to strongly developed vs modestly to strongly developed. The new species can be separated from from C. farooqi Auffenberg & Rehmann, 1995 by the shorter average adult male SVL, <90 mm vs> 90 mm in C. farooqi ; the smaller HeadW/SVL, 0.14-0.21 vs 0.19-0.21; more dorsal eyelid scales, 10-14(average 12.4) vs 9-11 (average 10) in C. farooqi . The new species can be separated from C. vultuosus (Harlan, 1825) by shorter average adult male SVL, <90 mm vs> 90 mm in C. vultuosus ; smaller HeadW/SVL, 0.14-0.21 vs 0.16-0.2; smaller HeadH/SVL, 0.11-0.18 vs 0.16-0.21; more scales between the nasal shield and the orbit, 4-5 (average 4.9) vs> 6; dark patches on neck not extending to 1/2 jowl vs stretching to 1/2 jowl in C. vultuosus .

This species can be separated from other members of Calotes by a combination of the following characters: 1. Crescent-shaped patch of granular scales absent at the forelimb insertion vs present in C. emma , C. grandisquamis Günther, 1875, C. jerdoni Günther, 1870, C. mystaceus Duméril & Bibron, 1837, and C. nemoricola Jerdon, 1853. 2. Keeled dorsal scales vs smooth in C. medogensis Zhao & Li, 1984. 3. Mid-body scale rows 36-46 (average 41.2) vs 49-65 in C. emma , 27-35 in C. grandisquamis , 45-57 in C. jerdoni , 58-63 in C. maria , 48-60 in C. minor , 45-58 in C. mystaceus . 4. Nuchal and dorsal crest scales short, nuchal crest distinctly larger vs nuchal spines much longer, dorsal spines reduced in C. maria and C. nemoricola . 5. Pair of short supratympanic spines on each side of the head vs a row of three or four compressed supratympanic spines in C. grandisquamis and C. nemoricola , eight or nine compressed spines above the tympanum in C. calotes (Linnaeus, 1758), two parallel rows of supratympanic scales in C. jerdoni and C. maria Gray, 1845, and a single well-developed postorbital spine in C. emma . 6. Homogeneous scalation on the dorsal region and a comparatively well-developed dorsal crest vs heterogeneous scalation and an undeveloped dorsal crest in C. paulus Smith, 1935 and C. zolaiking Giri, Chaitanya, Mahony, Lalronunga, Lakrinchhana, Das, Sarkar, Karanth & Deepak, 2019. 7. Concave orbital region and absence of row of erect scales on sides of neck vs no concave orbital region and presence of row of erect scales in C. bhutanensis Biswas, 1975. 8. Absence of fold in front of shoulder vs presence of fold in C. chincollium Vindum, 2003, C. nigriplicatus Hallermann, 2000, C. bachae Hartmann, Geissler, Poyarkov, Ihlow, Galoyan, Rödder & Böhme, 2013, C. geissleri Wagner, Ihlow, Hartmann, Flecks, Schmitz & Böhme, 2021, C. goetzi Wagner, Ihlow, Hartmann, Flecks, Schmitz & Böhme, 2021, C. mystaceus and C. vindumbarbatus Wagner, Ihlow, Hartmann, Flecks, Schmitz & Böhme, 2021. 9. Posterodorsal orientation of lateral body scales and absence of shoulder pit vs posteroventral orientation and presence of shoulder pit in C. ceylonensis Müller, 1887, C. desilvai Bahir & Maduwage, 2005, C. liocephalus Günther, 1872, C. liolepis Boulenger, 1885, C. manamendrai Amarasinghe & Karunarathna, 2014, C. nigilabris Peters, 1860, C. pethiyagodai Amarasinghe, Karunarathna & Hallermann, 2014.

Measurements and scale counts of specimens are given in Table 5 and Suppl. material 1: table S2.

Distribution and natural history.

Calotes wangi sp. nov. is a transboundary species, ranging from southern China (Fujian Province, Guangdong Province, Hainan Province and Guangxi Zhuang Autono-mous Region, southern Hunan Province, Macao, and Hong Kong) to northern Vietnam (Lang Son). The eastern Yunnan region in China is also likely to harbor this species ( Yang and Rao 2008; Shuo Liu, pers. comm. 17 April 2023). The known altitude range of C. w. wangi ssp. nov. is 15~2000 m and C. w. hainanensis is 47~1867 m. It is found in subtropical evergreen broad-leaved forests and tropical monsoon forests in southern China and northern Vietnam, mostly in mountainous areas, hills and plains on forest edges, arable land, shrub lands, and even urban green belts. It is active at the edge of the forest, and when it is in danger, it rushes into bushes or climbs tree trunks to hide. It is active from April to October every year, while in the tropics it is active from March to November or even longer. The authors investigated in Hainan and found that C. wangi sp. nov. was active from 7:30~11:30 hrs in summer and 15:00~19:00 hrs in the afternoon. Investigations in Guangxi found that the lizards lie on sloping shrub branches at night, sleeping close to the branches.

Both Calotes wangi sp. nov. and C. irawadi are oviparous. It has been reported that females of C. wangi sp. nov. contained 5-12 (17 in one specimen in Mt. Diaoluo, Hainan) mature eggs from late April to September, while females of C. irawadi contained 12 mature eggs in June ( Zhao et al. 1999). After our dissections, 13 specimens of C. wangi sp. nov. from Hainan obtained in August had 5-8 eggs (average 6), and five C. irawadi from Lushui, Yunnan obtained in late May were pregnant with 5-15 eggs (average 11).

Calotes wangi sp. nov. eats a variety of insects, spiders, and other arthropods ( Zhao et al. 1999).

Specimens examined.

Calotes irawadi (n = 18). Myanmar: Kawnglanghpu (CIB097515). China: Yunnan Province, Lushui City (Shangjiang CIB001824-26), Baoshan City (Shangjiang CIB001819-23, CIB001827-28), Dehong Dai and Jingpo Autonomous Prefecture (Hongbenghe CIB116106 and CIB116212, Tongbiguan KIZ 059191, KIZ NB20180905, KIZ HBH20200913 and KIZ HBH20200914), Xishuangbanna Dai Autonomous Prefecture (CIB001610).

Figures

Figure 4. Scatter plots of PC 1 and PC 2 of principal component analysis (PCA) based on morphometric measurements, distinguishing Calotes wangi sp. nov. from C. irawadi.