Calotes bachae Hartmann, Geissler, Poyarkov, Ihlow, Galoyan, Rödder & Böhme 2013

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 : 143-166

Calotes bachae Hartmann, Geissler, Poyarkov, Ihlow, Galoyan, Rödder & Böhme 2013 : 252

( Fig. 2 , Clade E)

Hartmann, T., Geissler, P., Poyarkov, N. A. J., Ihlow, F., Galoyan, E. A., Rödder, D. & W. Böhme (2013). Anew species of the genus Calotes Cuvier, 1817 (Squamata: Agamidae) from southern Vietnam. Zootaxa 3599 (3): 246–260.

Holotype. ZFMK 88935 (adult male, Fig. 5 A-B, Clade E) from “ Vietnam, Dong Nai Province, Cat Tien National Park (11.6344444° N 107.456667° E), 104 m elevation,” collected by Peter Geissler on May 10 th 2009 .

Original Diagnosis. Amedium-sized Calotes with a maximum SVL of 97 mm. It can be distinguished from all taxa of the C. mystaceus complex by the combination of the following characters: 1) head and body robust; 2) body scales homogeneous, relatively small, feebly keeled and arranged in regular rows; 3) 44–50 midbody scale rows; 4) upper dorsolateral scales pointing backwards and upwards; 5) two short and well separated spines, surrounded by three to four scales on either side of the upper head above the tympanum; 6) nuchal and dorsal crest continuous, composed of erected compressed scales, directed posteriorly; 7) vertebral spines and scales in males 35–42 and in females 43–46; 8) oblique fold of skin in front of fore limb insertion distinct, covered with small granular dark scales; 9) extremities and tail relatively long and slender (see comparison with C. mystaceus for details); 10) bluish to turquoise head and anterior body part, this coloration not well exceeding front limb insertion; 11) yellowish light stripe at upper lip reaching from below anterior corner of eye to posterior end of head; 12) no dorsolateral brownish blotches, sometimes faint medial brownish blotches across the vertebral crest.

Male coloration. The brilliant coloration of adult males is characterized by a brightly colored bluish to turquoise head, with bluish coloration continuing posteriorly to fore limb insertion. There is a relatively faded light (bright yellowish, when under acute distress, a character typical for C. bachae only) stripe at the upper lip crossing the tympanum from beneath the eye to end of head. Gular pouch colored in darker blue, interscale skin black. Atriangular to crescent-shaped patch of small black scales is present in front of shoulder. Very faint brownish blotches extending mid-dorsally over the vertebral crest, from above fore limb insertion on to tail; posterior to fore limb insertion brownish orange in color on trunk and tail; hind limb in a slightly darker brown; venter cream.

Distribution. Calotes bachae is known from southern Vietnam, with two specimens reported from the Vietnam-China border in northern Vietnam, and from eastern Cambodia ( Fig. 1 ).

Ecology. Calotes bachae is a diurnal, semi-arboreal lizard, often observed climbing on tree trunks at a height of 5–10 meters above the ground. The species mainly inhabits dipterocarp lowland forests and cultural landscapes up to 700 m a.s.l. The species seems to be quite heliophilic, preferring more open habitats without closed canopies ( Hartmann et al. 2013), but was also observed in dense tropical monsoon forests with closed canopy, in open gallery forest, and anthropogenic habitats like roadsides within the forest or open park landscapes within the headquarters of the park. In these anthropogenic habitats C. bachae occurs in syntopy with Calotes versicolor ( Hartmann et al. 2013). According to Hartmann et al. (2013) C. bachae feeds onnumerous arthropods including Formicidae , Coleoptera , Orthoptera , and Myriapoda. Males acquire their breeding coloration at the end of February, while gravid females have been found mid-April and egg-laying of a clutch of five eggs was observed in April and May. Juveniles hatched after 56 days while incubated at 22 to 25° Cin captivity.

Calotes bachae (n = 17): BMNH 1927.5.20.30 , BMNH 1927.5.20.31 , BMNH 1927.5.20.32 , MNHN 1927.44 , MNHN 1927.46 , ZFMK 88935 , ZFMK 88936 (now IEBR A.2012.23 ), ZFMK 88937 , ZFMK 92028 , ZFMK 92029 , ZFMK 94395 , ZFMK 94396 , ZFMK 94397 , ZMMU NAP 01509 , ZMMU NAP 01512 , ZMMU NAP 02910 , ZMMU NAP 02911 .

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. 5. Calotes bachae Hartmann et al., 2013. A. Holotype (ZFMK 88935, adult male), general view from above. B. Holotype, lateral view.C. Living holotype from Cat Tien National Park, Vietnam.

Fig. 1. Geographic distribution of examined specimens of the Calotes mystaceus complex. Colors correspond to the identified OTUs. Diamonds mark the type localities of the species described herein. Records with a bold margin were also included in the phylogenetic analyses. Imprecise (i.e., country-level) records are marked with a question mark. Detailed localities are: Calotes bachae: Vietnam: Dong Nai Prov.: Cat Tien National Park (1); Dong Nai Nature Reserve (2); Bu Gia Map National Park (3); Kon Tum (4); Trung Khanh (5); Cambodia: Banlung (7); Calotes goetzi sp. n.: Cambodia: Phnom Kulen National Park (8); Kulen Prom- tep Wildlife Sanctuary (9); Laos: Muang Phon Hong (10); Luang Prabang (11); Muang Pak Lay (12); Thailand: Sakon Nakhon Prov. (13); Lam Dom Noi River (14); Khon Buri (15); Nakhon Ratchasima (16); Dilang (17); Ban Nam Len (18); Lom Sak (19); Ban Bueng (20); Khlong Luang (21); Cha-am (22); Ban Phai (23); Kwai River bridge (24); Ban Dong Noi (25); Phitsanulok Prov. (26); Nan Prov. (27); Chiang Mai Prov. (28); Fang (33); Myanmar: Kawkareik (30); Mudon (31); Karen Hills (32); Parsa Wildlife Sanctuary (29); Kyaitong Township (34, 35); Inle Lake Wetland Sanctuary (36, 37); Taunggyi (38); Pindaya (39); Panlaung and Padalin Cave Wildlife Sanctuary (40); Mandalay-Yangon road (41); Minsontaung Wildlife Sanctuary (42); Popa Mountain Park (43); Mandalay (44); Shwebo (45); Bhamo (46); Indawgyi Lake (47); “Pegu” (54, see discussion in text); China: Baihualing (48); Longyang (49, 50); Liuku-Longling road (51); Liuku (52); Liuku-Fugong road (53); Calotes mystaceus: Myanmar: “Pegu” (54; see discussion in text); Kyaiktiyo Pagoda (56, 57); Taungoo (58); Letpein village (59); Hlawga National Park (60); Ngapudaw township (61); Ngayokekaung village (62); Calotes vindumbarbatus sp. n.: Myanmar: Gat Shang Yang village (63); Hepu village (64); Linpha village (65); Swekawngaw (66); Calotes geissleri sp. n.: Myanmar:Alaungdaw Kathapa National Park (67, 68); Mauk village (69, 70); Natzang village (71); Simggial village (72); India: Nagaland: Kohima (73).

Fig. 3. Principal component analyses results for the morphologically examined specimens. Details of Eigenvalues and explained variance are given in Table 3.

Fig. 4. Principal component analysis results for the morphologically examined specimens, colored according to to the genetic clades. Details of Eigenvalues and explained variance are given in Table 3.