A new species of the genus Calotes Cuvier, 1817 (Squamata: Agamidae) from southern Vietnam Author Hartmann, Timo Author Geissler, Peter Author Poyarkov, Nikolay A. Author Ihlow, Flora Author Galoyan, Eduard A. Author Rödder, Dennis Author Böhme, Wolfgang text Zootaxa 2013 3599 3 246 260 journal article 10.11646/zootaxa.3599.3.3 1f3cab5a-f723-4be3-aea0-413348526bca 1175-5326 218612 B32DC7D9-E158-4355-A02B-EA64BEDC3E63 Calotes bachae sp. nov. Fig. 4 ; Tables 4–5 Holotype : ZFMK 88935, an adult male ( Fig. 4 A) from Vietnam , Dong Nai Province, Cat Tien National Park ( 11°38'04'' N 107°27'24'' E ), 104 m elevation, collected 10 May 2009 by PG. Paratypes : An adult female ( IEBR A.2012.23; Fig. 4 B) and a juvenile ( ZFMK 88937; Fig. 4 E), same data as holotype (except collection dates: 20 April 2009 ( IEBR A.2012.23) and 14 June 2009 ( ZFMK 88937)) by PG. Two adult males ( ZMMU NAP- 02910-911; Fig. 4 D) from Phuoc Hoa environs, Bu Gia Map National Park, Dak Ka River valley, Bu Gia Map commune, Bu Gia Map District, Binh Phuoc Province, southern Vietnam ( 12°19.6’ N , 107°25.2’ E ; ca. 400 m above sea level), collected on 26 May 2011 by NAP . Two adult females ( ZMMU NAP- 0 1509, 01512) from Ma Da forestry, Dong Nai Nature Reserve (former Vinh Cuu Nature Reserve), Dong Nai Province, southern Vietnam , 80 m elevation, collected 12.06.2011 by NAP . Diagnosis: A medium-sized Calotes with a maximum SVL of 97 mm , distinguished from all congeners by the combination of the following characters: 1) head and body robust; 2) body scales relatively small, homogeneous, feebly keeled, arranged in regular rows; 3) upper dorsolateral scales pointing backwards and upwards; 4) 44–50 midbody scale rows; 5) two short and well separated spines, surrounded by three to four scales on either side of the upper head above tympanum; 6) nuchal and dorsal crest continuous, composed of erect compressed scales, directed posteriorly; 7) vertebral spines and scales 35–42 in males and 43–46 in females; 8) oblique fold of skin in front of forelimb insertion dinstict, 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) very faint or no dorsal brownish blotches, medially extending across the vertebral crest. FIGURE 4. Calotes bachae sp. nov. in life. A) Holotype, adult male (ZFMK 88935), Dong Nai Province, Vietnam. Photograph by PG. B) Paratype, adult female (IEBR A.2012.23), Dong Nai Province, Vietnam. Photograph by PG. C) Uncollected adult male during breeding season, Dong Nai Province. Photograph by PG. D) Close up of paratype, adult male (ZMMU NAP-02910), Binh Phuoc Province, Vietnam. Photograph by NAP. E) Paratype, juvenile (ZFMK 88937), Dong Nai Province, Vietnam. Photograph by PG. Description of Holotype : Moderately large male (SVL 90 mm ). Tail and extremities relatively long and slender (TailL 199 mm ). For further measurements see Table 5 . TABLE 5. Measurements and morphological characters of the type series of Calotes bachae sp. nov. ; measurements in mm. *= tail broken.

ZFMK 88935 (Holotype)	IEBR A.2012.23	ZFMK 88937	ZMMU NAP-01509	ZMMU NAP-01512	ZMMU NAP-02910	ZMMU NAP-02911
Gender	3	juv.	Ƥ	Ƥ	3	3
SVL	90	74	27.4	86	85	97	83
EyeEar	5.3	3.91	1.81	4.82	5.24	7.15	4.84
HeadH	14.88	12.19	4.9	14.09	13.83	18.22	13.88
HeadL	21.8	16.92	9.24	20.91	21.5	25.72	19.15
HeadW	17.04	14.12	5.93	14.6	15.4	20.02	13.71
Interorb	10.09	7.85	3.44	9.43	8.94	11.69	8.66
JawW	15	13.45	5.74	13.78	14.23	17.94	12.62
NarEye	6.16	3.75	1.88	6.19	5.65	6.97	5.12
SnEye	10.44	7.58	3.02	9.32	10.2	11.54	9.07
SnW	5.45	4.06	1.89	5.83	5.1	6.56	5.32
4FingL	11.79	8.88	3.69	10.93	10.42	10.94	10.24
4ToeL	15.08	11.83	5.37	14.59	12.96	14.17	13.78
CrusL	19.21	14.58	5.78	18.15	18.33	19.83	16.03
ForefL	13.74	12.11	4.97	12.79	14.17	15.59	13.28
HindfL	26.14	20.34	7.68	25.3	24.74	26.97	24.35
LoArmL	15.59	11.95	5.49	14.87	15.75	18.64	13.55
SnForeL	30	25	9.7	31	32	37	27
TailH	8.71	5.12	1.38	6.76	6.62	10.35	8.62
TailL	199	157	50.2	202	203	*	191
TailW	8.44	5.27	1.58	7.12	6.56	9.57	8.54
TrunkL	42	34	11.5	39	39	41	35
UpArmL	14.73	11.77	23.9	12.68	13.28	15.78	14.79
UpLegL	20.24	14.85	24.8	19.08	20.14	21.83	16.65
CanthR	9	8	9	9	10	8	9
Eyelid	14	14	13	13	13	13	13
HeadSLn	17	18	14	14	14	14	15
HeadSTr	16	18	14	15	17	17	15
Inflab	11	10	10	10	11	10	10
SnS	9	8	6	8	9	10	8
Suplab	10	10	10	10	11	10	10
4FingLm	21	18	20	22	20	20	20
4ToeLm	23	22	24	25	25	23	24
VertS	42	46	43	43	44	42	35
MidbS	48	48	48	46	46	44	45

Head distinct from neck; area posterior of jaw-angle swollen, snout-tip blunt; rostral small; nostril in a single scale, separated from rostral by one scale; a median longitudinal row of four bluntly ridged scales on snout; canthus rostralis sharp and straight; canthal scales continous with supraciliary scales; lateral sides of head flat; supralabials 10/10 (right/left), separated from the orbit by three rows of small scales; five feebly ridged scales from orbit to above tympanum; tympanum distinct; two spines above tympanum, surrounded by four or five slightly pointed scales; infralabials 11/11; mental small, bordered by two postmentals; separated from each other; only first pair of postmentals touching infralabials; scales on chin and throat strongly keeled; a continuous vertebral crest consisting of elevated spine-like scales from nape to well behind tailbase, spine height gradually decreasing posterior of neck; dorsal and lateral scales keeled, pointing upwards and backwards; caudal scales directed backwards and slightly upwards; scales on venter parallel keeled; subcaudal scales strongly and parallel keeled. Arms and legs relatively slender, fourth finger and toe longest. Color of Holotype in life: ( Fig. 4 A). The breeding coloration of the adult male is a brightly colored bluish to turquoise head, bluish coloration continuing posteriorly to behind the forelimb. A relatively faded light stripe at upper lip crossing the tympanum from beneath the eye to end of head. Gular pouch being colored in a darker blue, interscale skin black. Triangular to crescent-shaped patch of small black scales in front of shoulder. Very faint brownish blotches extending middorsally over the vertebral crest, from above forelimb insertion on to tail; posterior to forelimb insertion brownish orange in color on trunk and tail; hind limb in a slightly darker brown; venter cream. Color of Holotype in preservative: Head and anterior part of body bluish, although artificially faded to whitish grey at top of head and between eyes and tympanum. Coloration behind forelimb insertion brownish to grey at flanks; dorsum and tail light grey; at posterior tail faded darker blotches still discernible; venter cream. Variation: For the variation of paratypes see Table 5 . The paratypes largely correspond with the description of the holotype (see Fig. 4 B, 4D). The juvenile paratype (ZFMK 88937) differs from the other types in coloration pattern (see Fig. 4 E): speckled greyish to beige in ground color; head with two well defined dark brown bands transversely above eyes; dark brown radiating streaks around the eye; well defined dark brown blotches middorsally from neck onto tail, below and between each dark blotch an indifferently light grayish-colored mottle. Comparisons: Comparisons with all Indochinese congeners are based on examined specimens (see Appendix 1) and data obtained from the literature ( Smith 1935 , Taylor 1963, Vindum et al. 2003 , Zug et al. 2006 ). Calotes bachae sp. nov. is distinguished from C. htunwini , C. irawadi and C. versicolor by possessing a crescent to triangular-shaped patch of small granular scales in front of the shoulder. It differs from C. jerdoni by the absence of two parallel rows of compressed scales above tympanum. It can be distinguished from C. emma by the absence of a large postorbital spine. It differs from C. chincollium in having a lower number of midbody scale rows (44-50 vs. 59-74) and a bluish coloration of the head in life (vs. brownish-grey with black mask). Calotes bachae sp. nov. resembles C. mystaceus in all the above mentioned characters, but can be distinguished from the latter by a combination of the following characters (for further details see table 4): In males, C. bachae sp. nov. tends to have relatively longer fourth fingers, upper arms and upper legs than male of C. mystaceus . Furthermore, their tails tends to be relatively slimmer and flatter at base than in C. mystaceus . In females, C. bachae sp. nov. tends to have relatively longer forth toes, hind foots and upper legs than in C. mystaceus . Compared to C. mystaceus , their tails are relatively longer, slimmer and flatter at base. In both sexes, the relative distance between leg insertions was significantly shorter in C. bachae sp. nov. compared to C. mystaceus . In addition, the number of vertebral spines and scales (M: 35–42, F: 43–46) and the number of midbody scale rows (44–50) tend to be lower compared to C. mystaceus ( VertS : M: 42–51, F: 44–53; MidbS : 48–59). The color pattern of C. bachae sp. nov. is also different from C. mystaceus (see Fig. 4–5 ). The bluish to turquoise colored anterior body part of the former does not well exceed behind the forelimb insertion, whereas in the latter it often reaches posteriorly up to behind the hind limb. In C. mystaceus the eponymous whitish stripe at upper lip extend from between nostril and eye to well behind the head often reaching the end of neck (vs. a yellowish light stripe from below anterior corner of eye to posterior end of head in C. bachae sp. nov. ). C. bachae sp. nov. bears no or only faint brownish dorsal blotches, which are fused medially across the vertebral crest (vs. well defined dorsolateral dark brown blotches, which are well separated by the vertebral crest). Etymology: We dedicate this species to Rike Bach (Bonn), in gratitude for supporting the second author (PG) during several fieldtrips in Indochina. Distribution: Calotes bachae sp. nov. is currently only known from the territory of Vietnam . The type series consists of specimens from Dong Nai (Cat Tien NP and Dong Nai NR) and Binh Phuoc (Bu Gia Map NP) Provinces. We also recorded the new species from Dak Lak (Yok Don NP), Lam Dong (Cat Tien NP, Loc Bao Forestry), Ba Ria—Vung Tau (Binh Chau—Phuoc Buu NP) and Tay Ninh (Lo Go—Xa Mat NP) Provinces, as well as from Ho Chi Minh City (Saigon). However, the actual distribution of the Calotes mystaceus complex within Indochina is known to be much wider ( Bobrov & Semenov 2008 ; Nguyen et al. 2009 ). Future research has to show which documented populations of the Calotes mystaceus complex in Indochina have to be assigned to Calotes bachae sp. nov. . FIGURE 5. Calotes mystaceus in life. A) Adult male (ZFMK 92606), Siem Reap Province, Cambodia. Photograph by PG. B) Adult female (ZFMK 92607), Siem Reap Province, Cambodia. Photograph by PG. C) Subadult (ZFMK 90413), Preah Vihear Province, Cambodia. Photograph by TH. Ecology: The new species was mainly observed in dipterocarp lowland forests and cultural landscape up to 700 m above sea level (a.s.l.) (maximum altitude recorded in Loc Bao Forestry, Lam Dong Province). During observations in Cat Tien National Park the species appeared to be quite heliophile, preferring more open habitats without closed canopies, but was also observed in dense monsoon tropical forests with closed canopy in Dong Nai Reserve and Bu Gia Map National Park. Within the Cat Tien National Park the species was abundant both in open gallery forests along Dong Nai River as well as in anthropogenic habitats like roadsides within the forest or open park sceneries within the headquarters of the park. The gallery forests were dominated by deciduous Lagerstroemia species and Afzelia xylocarpa . In Ho Chi Minh City the new species is abundant within the city center in the old park of the Saigon Zoo and Botanical gardens, as well as in the park of the Palace of Reunification and other green areas. In these anthropogenic habitats C. bachae sp. nov. occurs sympatrically with Calotes versicolor . In Cat Tien and Lo Go – Xa Mat National Parks and Dong Nai Reserve these two sympatric species of Calotes seem to differ in preferable habitats and C. versicolor is found in more open and disturbed habitats and usually is not observed in primary forests, whereas C. bachae sp. nov. often penetrates to undisturbed primary forest. C. bachae sp. nov. is a diurnal, semiarboreal lizard, easily climbing on tree trunks at a height of 5–10 meters above the ground. Like many other tropical agamid lizards these animals spend night time on thin terminal branches and leaves. According to the content of feaces obtained from the population in the Cat Tien National Park, Calotes bachae sp. nov. forages on numerous insects ( Hymenoptera : Formicidae ; Coleoptera; Orthoptera) and other arthropods (Myriapoda). In the end of February to beginning of March, males acquire bright coloration, as described above (see coloration of holotype in life). Four gravid females were found in the middle of April 2012 . Oviposition by two females was observed in April and May 2009 . First, the female paratype (IEBR A.2012.23) laid four eggs on 20 April 2009 . The eggs were incubated in soil substrate at temperatures of 22 to 25° C. After 56 days of incubation four juveniles hatched (SVL 24–31 mm , TailL 56–63 mm ). One juvenile ( Paratype ZFMK 88937; Fig. 4 E) was euthanized and fixed after hatching. Second, on 25 May 2009 , an uncollected female from Cat Loc Area, Cat Tien National Park, Lam Dong Province was digging a ca. 4 cm deep hole in which it laid five eggs (see Fig. 6 B). After oviposition the hole was covered with soil and leaf litter.