Calotes goetzi, P. & F. & T. & M. & A. & W., 2021

Wagner, Philipp, Ihlow, Flora, Hartmann, Timo, Flecks, Morris, Schmitz, Andreas & Böhme, Wolfgang, 2021, Integrative approach to resolve the Calotes mystaceus Duméril & Bibron, 1837 species complex (Squamata: Agamidae), Bonn zoological Bulletin 70 (1), pp. 141-171 : 152-166

Calotes goetzi sp. n.

( Fig. 2 , Clade A) urn:lsid:zoobank.org:act:7827D48E-E121-4904-9636-3A46AE42B369

Holotype. ZFMK 92606 (adult male, Fig. 7 A-C, Clade A) from Cambodia, Siem Reap Province, near Kbal Spean within the Phnom Kulen National Park [13.699167° N, 103.998611° E] .

Paratypes. ZFMK 88341 (adult male), ZFMK 92607 (adult female) from the same locality as the holotype .

Diagnosis. Alarge species of Calotes , with a maximum SVL of 143 mm in males and 122 mm in females. It can be distinguished from other species of the group by the combination of the following characters: 1) head, body and limbs robust, tail long but not as long as in C. mystaceus ; 2) body scales mid-sized, homogeneous, keeled, arranged in regular rows; 3) upper dorsolateral scales pointing up- and backwards; 4) body scales arranged in 45–60 rows around midbody; 5) two short and separated spines, surrounded by a ring of scales between the tympanum and the vertebral crest on both sides of the head; 6) vertebral crest continuous from above the tympanum to the hind limbs, composed of erected scales, directed posteriorly, highest slightly in front of the insertion of the front limbs, becoming gradually shorter towards the hind limbs; 7) vertebral scales, including vertebral spines, 37– 52 in males and 43–60 in females; 8) oblique skin fold in front of the fore limbs; 9) head, body, and limbs bluish in males; 19) males with a white stripe from between nostril and orbit along the upper lip and the tympanum to the front limb insertion; 11) three to five distinct dark brown dorsolateral blotches.

Description of the holotype. Large male (SVL 118 mm). Body robust, tail relatively short, 236 mm long. Fore- and hind limbs relatively slender, fourth finger and toe longest. Head distinct from the neck, posterior jaw angles heavily swollen. Tip of the snout blunt, rostral small. Nostril large, in a single scale, separated from the rostral by one elongated scale and from the first two supralabial scales by two rectangular scales. Canthus rostralis distinct, formed by nine scales from the nostril to above the orbit and continuous with supraciliary scales. Lateral sides of the head flat. Supralabial scales 9/9, infralabial scales 10/10. Supralabial scales separated from the orbit by four rows of small scales. Eight scales between the orbit and the tympanum, two of them slightly ridged. Tympanum distinct, two spines, surrounded by a ring of scales, between the tympanum and the vertebral crest. Mental scale small, bordered by two postmental scales which are separated from each other. First pair of the postmental scales in contact with the infralabial scale. Scales on the dorsal part of the chin smooth to feebly keeled, becoming strongly keeled towards the throat. Vertebral crest continuous from above the tympanum to the tail, spines highest above the insertion of the front limbs and gradually decreasing towards the tail. Dorsal and lateral scales strongly keeled, pointing up- and backwards. Ventral scales parallelly keeled. Caudal scales smooth to feebly keeled, directed backwards. Subcaudal scales parallel and strongly keeled.

Male coloration. Head, body and limbs bluish. Awhite stripe, as high as the tympanum, is present from between nostril and orbit along the upper lip and the tympanum to the insertion of the front limb. The stripe is followed by three to five large reddish-brown to dark-brown blotches, with the first above the insertion of the fore limbs and the latest at around midbody or above the insertion of the hindlimbs, sometimes continuing on the tail. Throat coloration darker than the chin and the head coloration. From Thailand, males are known to have bright blue hindlimbs and crest scales. Intermediate males with a blue head and a white stripe, but body coloration brownish-black with indistinct light brown blotches. Non-brilliant coloration in males from Thailand reddish-brown, with a white non-continuous stripe from the orbit to above the hindlimbs. Blotches brownish, darker than the body coloration.

Variation. Bodymeasurements and meristic characters for adult individuals are given in Tables 4 . Specimens ranged in size between the smallest adult with a SVL of 84 mm (MNHN 1884.548) and the largest female with a SVL of 126 mm (ZFMK 44893) to the largest male measuring 145 mm (MNHN 1884.546). In general, adult males are larger than females, because of longer SVLs, tail lengths, head lengths, and widths and slightly fewer vertebral scales including the crest scales. Heads of adult males and females are proportionally equal in length, but male heads are wider than those of females, probably because of the swollen jaw angle. Dorsal coloration differs between sexes and both sexes are able to change the coloration. Coloration characters being lighter in females; head and body blue. Lateral stripe from between the nostril and orbit along the upper lips and the tympanum to the first blotch. Sometimes the first two brownish blotches are framed by scales in the same coloration as the lateral stripe. Three to five blotches present, from above the insertion of the fore limbs to about mid-body or even hindlimb insertion. Non-display coloration of females with lateral parts of the body gray and dorsal parts brown, with three to four darker crossbands between the lateral stripes on both sides of the body. Lateral stripe white, from the mental scale along the upper lips and the tympanum to the hindlimbs. Head light blue.

Etymology. The specific epithet is a patronym formed in the genitive singular honoring Dr. Stephan Goetz, Munich, Germany, in recognition of his longtime support of species conservation efforts in Cambodia.

Distribution. Calotes goetzi sp. n. is distributed in a large area of Indochina and inhabits most of the previous range of C. mystaceus . The new species is documented by museum specimens and photo vouchers from Cambodia, China (Yunnan province), Laos, Myanmar, and Thailand.

Ecology. Calotes goetzi sp. n. is diurnal and semi-arboreal to arboreal and can easily climb on tree trunks at a height of 5–10 meters or above. The species is known from dipterocarp lowland forests, cultural landscape and secondary forest. At its type locality the species was observed in more open habitats with a closed canopy, but also within dense monsoon tropical forests and solitary trees in rather open areas. Here it occurs sympatrically with Calotes versicolor . Hawkeswood & Sommung (2018) report it from farmland with e.g., rice, coconut, banana, and durian plantations. Chan-Ard et al. (2015) report the species from a wide range of forest types in Thailand, including tree-lined agricultural lands and grasslands. Similar to other Calotes species , C. goetzi sp. n. feeds on a variety of arthropods. Chan-Ard et al. (2015) report from Thailand, that mature individuals are territorial, chasing away possible intruders. Eggs are buried in soft soil and later guarded by the males. Amber et al. (2017) recognized a possible ontogenetic shift in defense strategies of C. goetzi sp. n. ( C. mystaceus at the time of their publication), with older individuals utilizing color change, while juveniles do not.

Calotes goetzi sp. n. (n = 71): BMNH 1868.4.3.61 , BMNH 1868.4.3.63 , BMNH 1891.11.26.29 , BMNH 1914.4.25.1 , BMNH 1929.12.1.11 , BMNH 1933.3.10.1 , BMNH 1933.3.10.2 , MNHN 1884.546 , MNHN 1884.547 , MNHN 1884.548 , MNHN 1893.335 , MNHN 1893.336 , NME R0581/09 , NME R0584/09 , NME R0585/09 , NME R0686/11 , NME R0751/12 , NME R0783/13 , NME R0784/13 , NME R0785/13 , NME R0786/13 , NME R0787/13 , NME R0790/14 , ZFMK 16640 , ZFMK 16641 , ZFMK 40561 , ZFMK 43906 , ZFMK 43930 , ZFMK 43931 , ZFMK 43932 , ZFMK 44893 , ZFMK 44894 , ZFMK 45490 , ZFMK 45491 , ZFMK 45492 , ZFMK 45493 , ZFMK 45494 , ZFMK 45495 , ZFMK 45496 , ZFMK 45497 , ZFMK 45498 , ZFMK 45499 , ZFMK 45500 , ZFMK 45501 , ZFMK 45502 , ZFMK 45553 , ZFMK 45554 , ZFMK 47080 , ZFMK 47081 , ZFMK 49164 , ZFMK 49202 , ZFMK 49219 , ZFMK 49220 , ZFMK 49221 , ZFMK 49242 , ZFMK 49243 , ZFMK 49244 , ZFMK 49245 , ZFMK 49246 , ZFMK 55610 , ZFMK 55611 , ZFMK 55612 , ZFMK 84867 , ZFMK 88341 , ZFMK 92606 , ZFMK 92607 , ZMB 11603A , ZMB 30186 , ZMB 30188A , ZMB 30197 , ZMB 6034 .

Figures

Fig. 2. The Bayesian consensus tree based on 988 bp of mitochondrial DNA (12S rRNA and COI) shows six distinct lineages within Calotes mystaceus. Node support in terms of Bayesian posterior probabilities is indicated by circles at nodes (nodes with a BPP ≥ 0.90 are white, BPP ≥ 0.95 are grey, BPP ≥ 0.99 are black, values <0.90 are not marked). Outgroup (Calotes versicolor) not shown for clarity. Numbers in parentheses behind taxa refer to localities mapped in Fig. 1.

Fig. 7. Calotes goetzi sp. n. A. Holotype (ZFMK 92606, adult male), general view from above. B. Holotype, lateral view.C. Living holotype from Kbal Spean, Cambodia.