A new species of Pseudocophotis (Agamidae: Acrodonta: Lacertilia: Reptilia) from central Vietnam

Russian Journal of Herpetology

Vol. 14, No. 2, 2007, pp. 153 – 160

A NEW SPECIES OF Pseudocophotis (AGAMIDAE: ACRODONTA: LACERTILIA: REPTILIA) FROM CENTRAL VIETNAM

Natalia B. Ananjeva,1 Nikolai L. Orlov,1 Nguyen Quang Truong,2 and Roman A. Nazarov1,3 Submitted March 23, 2006. A new agamid lizard, Pseudocophotis sp. nov. from central mountain region of Vietnam (Kon Tum Province) if described. The new species strongly differs from another agamid species of Eastern Himalaya-China-Indochina region by combination of pholidosis, morphometric characters and coloration. Keywords: arboreal Agamidae, Draconinae, Pseudocophotis, Pseudocalotes, Japalura, central Vietnam, a new species.

The species composition of lizards of the agamid genera (subfamily Draconinae) is still controversial and obscure. This subfamily is one of the most diverse groups of arboreal and semi-arboreal agamids with a high percentage of endemics (Moody, 1980). Draconine agamids exhibit very high degree of diversification at the generic level. Phylogenetic studies conducted in recent decades have improved our knowledge about the composition and phylogeny of the subfamily. The problems often arise relating to newly described species in certain genera (Moody, 1980; Manthey and Grossmann, 1997; Macey et al., 2000; Ananjeva and Stuart, 2001; Hallermann and Böhme, 2000; Hallermann and McQuire, 2001; Schulte et al., 2004; Ananjeva et al., 2007). The genus Pseudocophotis Manthey and Grossmann, 1997 includes one species P. sumatrana (the former Cophotis sumatrana) distributed in Sumatra and Java. In his unpublished dissertation Moody (1980, p. 304) included Cophotis sumatrana in the list of new taxa in the process of being described and concluded that numerous major characters separate this species which can be cladistically related to other genus (p. 178). Recently Pseudocophotis sumatrana was re-

ferred to the genus Pseudocalotes (Hallermann and Böhme, 2000; Hallermann and McQuire, 2001). Because of complicated situation and preliminary conclusion of phylogenetic analysis (Moody, 1980) the generic referral of Pseudocophotis sumatrana to the genus Pseudocalotes seems to be prematural. We prefer to remain Pseudocophotis genus until more detailed phylogenetic revision. During a survey in 2005, one specimen of arboreal agamid lizard was collected by Nikolai Orlov, Sergei Ryabov, and Ho Thu Cuc in the central mountain region of Vietnam. In 2006 five more specimens were collected in the same region by Nikolai Orlov, Ho Thu Cuc, and Roman Nazarov. This agamid lizard was very different from other known agamids of Vietnam. Comparison of morphometric and meristic characters revealed that this specimen differs from all recognized species of the genera Japalura and Pseudocalotes and have the most affinities to the genus Pseudocophotis. This new species is described below.

1

Specimen from Central Vietnam (1 male, 3 females, 3 juveniles) were compared with specimens of the other S-E Asian agamids. Meristic characters and measurements were compared with those of currently recognized species of the genus Japalura (Mertens, 1926; Bourret, 1937; Ota, 1989, 1991; Ota et al., 1998; Zhao et al., 1999), Pseudocalotes (Hallermann and Böhme, 2000;

2

3

Zoological Institute, Russian Academy of Sciences, Universitetskaya nab., 1, St. Petersburg 199034, Russia; E-mail: [email protected] Institute of Ecology and Biological Resources, Vietnamese Academy of Science and Technology (VAST); 18 Hoang Quoc Viet St., Hanoi, Vietnam; E-mail: [email protected] Zoological Museum, Moscow State University, B. Nikitskaya ul. 6, Moscow 125009, Russia; E-mail: [email protected].

MATERIAL AND METHODS

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154 Hallermann and McQuire, 2001) and other draconin agamids (Boulenger, 1885; Smith, 1935; Taylor, 1963; Orlov et al., 2005). A set of characters was selected to be comparable with thorough data of Ota for the genus Japalura (Ota, 1988, 1989, 1989a, 1991; Ota and Weidenhöfer, 1992; Ota et al., 1998), Pseudocalotes (Hallermann and Böhme, 2000; Hallermann and McQuire, 2001), and other arboreal agamids (Boulenger, 1885; Smith, 1935; Taylor, 1963; Orlov et al., 2006). They are following: 1) snout to vent length (SVL), distance from the tip of snout to anus; 2) snout to eye length (SEL), distance from tip of snout to anterior margin of right eye; 3) interorbital distance (IOD), distance between outer edges of superciliary series; 4) head length (HL), distance from tip of snout to occiput; 5) head width (HW), head width at the level of tympanum; 6) head depth (HD) at the parietal region; 7) number of nuchal crest spines (NS); 8) supralabials (SL), number of scales bordering right margin of upper lip including the most posterior one; 9) infralabials (IL), number of scales bordering right margin of lower lip including the most posterior one; 10) number of scales contacting rostral (SER); 11) number of scales contacting nasal (SEN); 12) number of scales contacting interparietal (SEIP); 13) number of scales around midbody (M); 14) horizontal diameter of eye (E); 15) dorsal crest (DC), number of mid-dorsal crest-like scales from neck to anterior margin of vent; 16) number of subdigital scales under IV finger (FS4); 17) number of subdigital scales under IV toe (T4S); 18) forelimb length (FLL), distance from axilla to tip of finger IV, excluding claw, on right side; 19) hindlimb length (HLL), 20) distance from groin to tip of toe IV, excluding claw, on right side, distance from axilla to groin (AGL); 21) tail length (TL), distance from anus to tip of tail (only in specimens with undamaged tail tips). Comparative material was examined in the holdings of Zoological Institute (Russian Academy of Sciences, St. Petersburg, Russia, ZISP), Muséum national d’Histoire naturelle (Paris, France, MNHN), Zoological Museum (Berlin, Germany, ZMB), and Zoological Museum (Hamburg, Germany, ZMH). Additional data were taken from the literature (Ota, 1988, 1989, 1989a, 1991; Ota and Weidenhöfer, 1992; Manthey and Grossmann, 1997; Ota et al., 1998; Zhao et al., 1999; Hallermann and Böhme, 2000; Hallermann and McQuire, 2001). Detailed morphological characteristics of currently recognized species of Japalura and Pseudocalotes were taken from Ota (1989: Tables 1 and 2), Hallermann and McQuire (2001: Table 1), and Zhao et al. (1999).

Natalia B. Ananjeva et al. DESCRIPTION Pseudocophotis kontumensis sp. nov. (Figs. 1 – 8) Holotype (Figs. 1 – 5). ZISP 24251 (Fn 32923), an adult male, from Kon Du, Mang Canh village, Konplong district, Kon Tum Province, Vietnam (14°41¢25¢¢ N 108°19¢31¢¢ E), 1210 m elevation, collected on April 7, 2005, by Nikolai L. Orlov, Sergei Ryabov, and Ho Thu Cuc. Paratypes (Figs. 6 and 7). ZISP 24252 – 24257 (Fn 1952 – 1957) (3 adult females, 3 juveniles) from Kon Du, Mang Canh village, Konplong district, Kon Tum Province, Vietnam (14°41¢ N 108°19¢ E), 1200 – 1250 m elevation, collected on May 25 – June 5, 2006, by Nikolai Orlov, Ho Thu Cuc, and Roman Nazarov. Diagnosis. Pseudocophotis kontumensis differs from other agamid lizards known to occur in Vietnam based on morphological characters. It is a middle-sized agamid lizard with laterally compressed body, very large dorsal and lateral scales, smaller keeled ventral scales, hidden tympanum, fold across the throat and relatively short flexible prehensile tail. No gular sac. From Acanthosaura a new species can be distinguished by the absence of postorbital and nuchal spines; from Calotes — by heterogeneous unequal dorsal scales intermixed with larger irregular scales. From both genera it differs by tympanum covered with skin. From Bronchocela it is distinguished by its much shorter tail. From Gonocephalus P. kontumensis differs by hidden tympanum, much smaller adult size and smaller ventral scales in comparison with dorsal. A new species differs from 3 species of Pseudocalotes occurring in Vietnam (Ananjeva et al., 2007; Bain et al., 2007): P. brevipes, P. microlepis, and P. floweri by hidden tympanum and smaller scale number around the midbody. P. brevipes and P. microlepis have much shorter tail (Table 1; Hallermann and McQuire, 2001: Table 1). Japalura species with hidden tympanum (including Vietnamese J. chapaense and J. fasciata) have much longer tail and limbs (Table 1; Ota, 1989: Tables 1 and 2). Pseudocophotis kontumensis differs from P. sumatrana by higher number of scales around midbody. Description of holotype. An adult male with a SVL 85.6 mm; TL 134 mm; HL 29.9 mm; HW 15.3 mm; HD 14.1 mm; 38 scales around midbody. Fore limb length 33.4 mm, Hind limb length 46.6 mm. Snout to eye length 11.9 mm; interorbital distance 12.6 mm. Tympanum hidden, covered by scales; horizontal. diameter of orbit 7.4 mm.

A New Species of Pseudocophotis from Central Vietnam Head relatively large: HL/SVL 34.9; HW/SVL 17.9; HD/SVL 16.5; distinctly separated from the body. Intraorbital distance includes 12 – 13 scales and internasal distance — 5 scales. 9 supralabials and 8 infralabials. Supralabials are separated from orbital regions by two rows of large scales. 7 scales contacting rostral; 7 scales contacting nasal; 7 scales contacting interparietal. Posterior to parietal scale and orbital region head is covered by very raised granulate scales of different size. Snout with relatively flat scales. Orbital and interorbital regions of the head are also covered by relatively flat scales. Rostral shield is large, triangle-shaped. Chin shield isosceles in shape. Genials are separated by prominent

155 irregular scales. There is fold across the throat; no gular sac. Gular scales small, moderately granulated. Nuchal crest comprises six erect, compressed scales, triangular in shape, pointed, maximal length 6 mm. It starts from nuchal region and separated from parietale by five scales. Second, third and fourth nuchal crest spines are mostly developed; posterior spines are lower. Low dorsal crest present. Scales along the keel have visible keels. Body laterally compressed, covered by large heterogeneous unequal scales. From nuchal and dorsal crest laterally towards venter rows of large keeled scales are situated. They are intermixed by scales of irregular shape. Belly is covered by small keeled scales.

TABLE 1. Scale Counts, Morphometric Measurements (in mm), and Their Indexes for the Type Series of P. kontumensis Character 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30

SVL TL SEL IOD HL HW HD NS SL IL SER SEN SEIP M E DC FS4 T4S FLL HLL AGL Index TL/SVL HL/SVL HW/SVL HD/SVL SEL/SVL IOD/SVL FLL/SVL HLL/SVL AGL/SVL

Male ZISP 24251

Female ZISP 24252

Female ZISP 24253

Female ZISP 24254

Juvenile ZISP 24255

85.6 134.0 11.9 12.6 29.9 15.3 14.1 5 9 8 7 7 7 48 7.4 10? 19 24 9.5 + 23.9 14.1 + 32.5 34.8

79.4 118.9 11.7 10.6 24.4 13.1 11.1 6 8 6 7 7 9 50 6.7 5 18 23 8.3 + 22.3 14.1 + 31.1 38.4

87.8 132.3 11.3 12.0 25.4 13.0 11.5 7 8 9 7 6 8 51 6.6 7 19 25 7.4 + 22.2 12.2 + 29.9 43.4

84.4 124.8 11.9 12.4 26.6 13.9 11.2 5 8 9 7 6 8 50 5.7 5 19 23 9.5 + 24.5 14.9 + 30.4 45.3

60.4 98.9 9.0 8.8 19.3 10.7 9.5 5 8 9 7 7 8 51 3.6 5 20 24 7.7 + 18.1 12.2 + 24.9 25.7

138.9 34.9 17.9 16.5 13.9 14.7 39.0 54.4 40.6

149.7 30.7 16.5 14.0 14.7 13.4 38.5 56.9 48.4

150.7 28.9 14.8 13.1 12.8 13.7 33.7 47.9 49.4

147.9 31.5 16.5 13.3 14.1 14.7 40.3 53.7 53.7

163.7 31.9 17.7 15.7 14.9 14.6 42.7 61.4 42.5

Note. For abbreviations see Material and Methods.

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Fig. 3. Holotype ZISP 24251 of Pseudocophotis kontumensis sp. nov. Dorsal view. Fig. 1. Holotype ZISP 24251 of Pseudocophotis kontumensis sp. nov. Diurnal coloration.

Fig. 4. Holotype ZISP 24251 of Pseudocophotis kontumensis sp. nov. Ventral view.

Fig. 2. Holotype ZISP 24251 of Pseudocophotis kontumensis sp. nov. Nocturnal coloration.

Limbs robust, relatively short, covered with large strongly keeled scales, ventral part is covered by small

scales. Fourth finger with 19 subdigital lamellae; fourth toe with 24 subdigital lamellae. Tail comparatively short, strongly swollen at the first third from its base, laterally compressed and covered by strongly keeled irregular scales on back not forming regular rings. Scales of ventral surface of tail is more regular, keeled. Enlarged scales on tail look like continuation of low dorsal crest. Coloration in life. There are two kinds of physiological coloration — for day and night. Diurnal coloration is brighter with two transversal wide white bands between limbs and less developed light band in the neck region. Six diffuse bands cross tail. General background of coloration is brown with greenish free design on head. Supra-and infralabials gray. Above supralabials from tip of snout to neck interception pass light band which is light-green from snout to eye. From eye to neck this band becomes yellow-brown. Color of ventral side grayish-green. General background and pattern create favorable camouflage conditions on trunk of trees. Nocturnal coloration is in general darker. Bands as well as other pattern are developed not so clear. Coloration in alcohol. Gray-bluish coloration with the same patterns as in color in life.

A New Species of Pseudocophotis from Central Vietnam

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a

b

c

d Fig. 5. Head of holotype of Pseudocophotis kontumensis sp. nov. a, Lateral view; b, dorsal view; c, nuchal region; d, swollen tail.

Fig. 6. Paratype ZISP 24253 of Pseudocophotis kontumensis sp. nov. Dorsal view.

Etymology. The specific epithet refers to Kon Tum Province the mountainous type locality of the new species. Variation. Variation in morphological characters is presented in Table 1. The paratypes (females and juveniles) differ from the holotype in having 50 – 51 (vs. 48) scales around midbody, 8 (vs. 9) supralabials, 6 – 9 (vs. 8) infralabials, smaller head (more short and nar-

Fig. 7. Paratype ZISP 24253 of Pseudocophotis kontumensis sp. nov. Ventral view.

row) and longer tail. Coloration of paratypes (females) differs in general rusty-brown background of dorsum and lighter belly. Distribution. The species is known only from type locality (Fig. 9). Natural history. Pseudocophotis kontumensis was observed by us from 1200 – 1250 m a.s.l. in mountain-

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Fig. 8. Female ZISP 24253 of Pseudocophotis kontumensis sp. nov. using prehensive tail.

ous region of Central Vietnam (Kon Tum Province) (Fig. 9). This arboreal agamid prefers the rain polydominant forest. All individuals were met on the vegetation (bushed and small trees) in the bottomland of forest streams under closed canopy. They are very secretive and well camouflaged on the bark. They were observed in day time moving very slowly and never been noted on the ground. At night they were found sleeping on the branches of bushes and trees on the height of 2 – 4 m (Fig. 8). In captivity they eat crickets and cockroaches. Females laid clutches in June (6 – 8 eggs). DISCUSSION AND COMPARISONS The new species differs from all known agamids of the fauna of Vietnam of genera Physignathus, Leiolepis, Acanthosaura, Bronchocela, Calotes, Draco, Gonocephalus, Japalura, Pseudocalotes. From Japalura species also having tympanum covered with skin P. kontumensis distinguished by shorter tail (Table 1), laterally compressed body, very large dor-

sal and lateral scales. New species differs by concealed tympanum from J. tricarinata, J. major, J. kumaonensis, and J. daymonsi possessed exposed tympanum (Smith, 1935). The body of new species is moderately compressed whereas J. planidorsata has subquadrangular body. From short-tailed species of Japalura (J. dymondi, J. varcoae) the new species differs by concealed tympanum. A new species possess a relatively shorted tail when compared with J. andersoniana, J. dymondi, J. flaviceps, J. chapensis, J. makii, J. splendida, J. szechwanensis, and J. yunnanensis (Zhao et al., 1999). From all species of Pseudocalotes genus also can be distinguished by concealed tympanum. The latest interpretation of scope of genus Pseudocalotes (Hallermann and Böhme, 2000; Hallermann and McQuire, 2001) referring Pseudocophotis sumatrana to the genus Pseudocalotes allow to attribute the new species to genus Pseudocalotes. However because of complicated situation we prefer to reserve generic name Pseudocophotis (Manthey and Grossmann, 1997) and to refer the new species to this genus based of tympanum scaledover, scalation intermixed with very large irregular

A New Species of Pseudocophotis from Central Vietnam scales among dorsal, lateral and caudal scales, relatively short prehensile tail (TL/SVL 150.2 in P. kontumensis and 148 in P. sumatrana). In his unpublished dissertation Moody (1980, p. 304) noted Cophotis sumatrana in the list of new taxa in the process of being described and concluded that numerous major characters separate this species which can be cladistically related to other genus (p. 178). From P. sumatrana the new species differs by the number of scales around midbody (32 – 34 in P. sumatrana and 48 – 51 in P. kontumensis). P. sumatrana and P. kontumensis have disjunctive distribution in Sumatra and Java, from one hand and Central Vietnam, from another hand. Taking into account very secretive mode of life of these animals we can suppose that they could be found in other regions of southern Indochina. The common faunistic elements between southern Indochina and Sumatra are known for a number of species of frogs Rhacophoridae, Ranidae, Micrihylidae families, colubrid snakes of Boiga genus and viperid snakes of Trimeresurus and Ovophis genera, and others. Acknowledgments. We are very grateful to Vietnam Institute of Ecology and Biological Resources (IEBR) for the organization of the fieldwork in Vietnam, Director of IEBR, Prof. Le Xuan Canh, Director of Dept. of Vertebrate Zoology, Dr. Nguyen Xuan Dang and research scientists of this department Ho Thu Cuc. The opportunity to work in Vietnam was made possible by “International Foundation of Conservation of Biological Diversity “Fora” (Tula, Tatyana S. Moiseeva, President) and contribution of Sergei Ryabov. Our research was partially supported by grants NSH 4212.2006.4, RFFI 05-04-48156, 07-04-90004-VAST, and MNHN professorship in 2004 – 2005 to Natalia Ananjeva and Nikolai Orlov. We would like to thank Alain Dubois and Annemarie Ohler (MNHN), Rainer Günther (ZMB), and Jakob Hallermann (ZMH) providing us museum specimens for this study.

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159 100°

105°

110° E

CHINA Hanoi Gulf of

20° E

To n k i n Hainan

THAILAND 15°

CAMBODIA

Gulf of

Ho Chi Minh City

10°

Thailand

Fig. 9. Type locality (yellow star) of Pseudocophotis kontumensis sp. nov.: Kon Du, Mang Canh village, Konplong district, Kon Tum Province, Vietnam (14°41¢25¢¢ N 108°19¢31¢¢ E).

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