Asian Herpetological Research 2017, 8(1): 14–21 ORIGINAL ARTICLE DOI: 10.16373/j.cnki.ahr.160031

A New Species of Japalura (Agamidae: Lacertilia: Reptilia) from Central Highland, Vietnam

Natalia B. ANANJEVA1*, Nikolay L. ORLOV1 and Tao Thien NGUYEN2

1 Department of Herpetology, Zoological Institute, Russian Academy of Sciences 199034, St. Petersburg, Russia 2 Vietnam National Museum of Nature, Vietnam Academy of Science and Technology, 18 Hoang Quoc Viet Road, Cau Giay, Hanoi, Vietnam

Abstract A new species of the agamid genus Japalura is described based on three specimens from southern part of Central Vietnam. It is distinguished from remaining congeners by the following combination of characters: adult size (SVL females 68–69 mm), tail length/SVL ratio 226%–239%, HW/SVL ratio 17%–18%; FLL/SVL ratio 41%–43%; HLL/SVL ratio 72%–73%; 7–9 supralabials, 7–9 infralabials, 54–56 middorsal scales, 20–22 lamellae under finger IV, 24–26 lamellae under toe IV, 1 scale between nasal and supralabials; tympanum concealed; absence of transverse gular fold. The geographical distribution of Japalura genus in general and of a new species in particular is discussed.

Keywords Agamidae, Japalura, taxonomy, Vietnam, description of a new species, distribution

1. Introduction agamid lizard referred to Japalura genus but different from the other congeners were collected in the south Currently the genus Japalura (sensu lato) comprises of Central Vietnam. Comparison of morphometric and 31 recognized species (Manthey et al., 2012; Uetz and meristic characters revealed that these specimens differ Hošek, 2016; Wang et al., 2015, 2016). Several recent from all recognized species of this genus. This new intergeneric changes modify generic arrangements: J. species is described below. kaulbacki Smith, 1937 was transferred into the genus Pseudocalotes (Mahony 2010); revalidation of the 2. Materials and Methods genus Oriotiaris made by Käs­tle and Schleich (1998) for Japalura species with a naked tympanum was not Specimens from the south of Central Vietnam (ZISP confirmed by results of molecular research (Macey et al., 25230–25231, VNMN3110) were compared with 2000; Schulte et al., 2004) and latest revisions (Mahony specimens of the other congeners. A set of characters was 2009, 2010; Manthey, 2010). The lizards of genus selected to ensure a maximum comparability of published Japalura are distributed in southeastern and eastern Asia data for all currently recognized species of the genus as well as in Himalaya region (Figure 1). Two species Japalura (Ota, 1989a, 1989b, 1991, 2000, 2003; Ota and among known species: J. chapaensis Bourret, 1937 and Weidenhöfer, 1992; Ota et al., 1998; Zhao et al., 1999; J. fasciata Mertens, 1926 were registered in the fauna of Mahony, 2009, 2010; Manthey et al., 2012). Vietnam (Ota, 1989b, 2000; Ota and Weidenhöfer, 1992; The set of 10 measurements is as follows, SVL, snout Bobrov, 1995; Ananjeva et al., 2007; Bobrov, Semenov, to vent length (distance from the tip of snout to anus); TL, 2008; Nguyen et al., 2009; Mahony, 2010; Manthey et tail length, HL, head length (distance from tip of snout to al., 2012). During a survey in 2004, three specimens of rear border of right angle of jaw); HW, head width at its widest point, HD, maximum head depth taken at the rear * Corresponding author: Prof. Natalia B. ANANJEVA, from the Zoological Institute, Russian Academy of Sciences, with her research axis of the jaw; IN, internarial distance, IOD, interorbital mainly focusing on taxonomy, ecology, phylogeny and biogeography of distance (distance between outer edges of orbits), SEL, Eurasian amphibians and reptiles. E-mail: [email protected] distance from tip of snout to anterior margin of right eye; Received: 24 April 2016 Accepted: 14 July 2016 FLL, forelimb length (distance from axilla to tip or finger No. 1 Natalia B. ANANJEVA et al. A New Species of Japalura from Central Vietnam 15

Figure 1 Distribution range of agamid lizards of Japalura genus. Type locality of Japalura sp. nov. is marked by black triangle.

IV, excluding claw, on right side); HLL, hindlimb length (Ota, 1989a, 1991; Mahony, 2009, 2010; Manthey et al., (distance from groin to tip of toe IV, excluding claw, 2012) were used. on right side). Measurements were taken in millimeters (mm) to a precision of 0.1 mm with dial calipers. The 3. Results following seven ratios TL/SVL, HL/SVL, HW/SVL, HD/ SVL, HLL/SVL, HLL/SVL, HW/HL were calculated to A description of a new species (Figures 2, 3, 4 and 5) compare them with other species of Japalura. Japalura ngoclinensis n. sp. The following meristic data were recorded: SL, Holotype ZISP 26230 (adult female), above Mang Xang supralabials (number of scales bordering right margin Village, Ngoc Linh Mountain, Dac Glei District, Kon of upper lip including the most posterior one); IL, Tum Province, Vietnam (15°05' N 107°57' E, elevation intralabials (number of scales bordering right margin 1650 m), on 25 March 2004 by Alexei V. Abramov of lower lip including the most posterior one); NSL, (Figures 2 and 5). number of scales between nasal and supralabials; SER, Paratypes ZISP 26231 (adult female) and VNMH 3110 number of scales contacting rostral; SEN, number of (subadult female) from above Mang Xang Village, Ngoc scales contacting nasal; SEIP, number of scales contacting Linh Mountain, Dac Glei District, Kon Tum Province, interparietal; DC, dorsal crest, number of mid-dorsal Vietnam (15°05' N 107°57' E, elevation 1650 m), on 25 crest-like slightly enlarged serrated scales from neck to March 2004 by Alexei V. Abramov (Figures 3–5). anterior margin of vent; F4S, finger IV subdigital scales Diagnosis Medium-sized, long-tailed Japalura species number, T4S, toe IV subdigital scales number; MD, with relatively long limbs, that differs from all other number of enlarged mid-dorsals from the nape to above species of this genus by the following combination of the vent. characters: tympanum hidden, transverse gular fold Comparative material was examined in the holdings of absent, enlarged subocular scale row present. Third and Zoological Institute, Russian Academy of Sciences, St. fourth fingers have equal length. Very low dorsal crest Petersburg (ZISP), Vietnam National Museum of Nature, from 5–6 slightly enlarged serrated scales. Vietnam Academy of Science and Technology (VNMN) Description of holotype SVL 67.9 mm, TL/SVL 239%, and Muséum National d’Histoire Naturelle, Paris FLL/SVL 41%, HLL/SVL 72%, HW/SVL 18%, HW/ (MNHN); additionally the well documented photographic SVL 18%, HD/SVL 13%, HW/HL 60%, SL 7/8, IL 8/9, material by Manthey (2010) and comprehensive 54 MD, 21 F4S, 26 T4S. Body shape in cross section summarized tables with data on all the species of genus laterally slightly compressed; nuchal and dorsal crest including mainland species with concealed tym­panum from 6 very short serrated scales; head is separated from 16 Asian Herpetological Research Vol. 8

Figure 4 Ventral (A) and dorsal (B) views of paratype (VNMH 3110) of Japalura ngoclinensis sp. nov.

Figure 2 Ventral (A) and dorsal (B) views of holotype (ZISP 26230) of Japalura ngoclinensis sp. nov.

Figure 5 Lateral head views of Japalura ngoclinensis sp. nov.: A. Paratype ZISP 26231. B. Holotype ZISP 26230.

the body by the neck. Quadrangular rostral approximately 2.5 times broader Figure 3 Ventral (A) and dorsal (B) views of paratype (ZISP than high in contact with six scales including SL; head 26231) of Japalura ngoclinensis sp. nov. dorsally covered with heterogeneous strongly keeled No. 1 Natalia B. ANANJEVA et al. A New Species of Japalura from Central Vietnam 17 scales, snout covered by relatively flat scales as well tricarinata (Blyth, 1853), J. kumaonensis (Annandale, as orbital and interorbital areas of head. Nasal scale 1907), J. varcoae (Boulenger, 1918) and J. dymondi separated from rostral by one scale and by one scale from (Boulenger, 1906), possessing exposed tympanum the first supralabial scale; two scale rows between the (Smith, 1935; Manthey et al., 2012). Additionally from: orbit and supralabials; a group of enlarged, keeled scales J. dasi, J. kumaonensis and J. varcoae by the presence of at the posterior­ edge of the orbit. Mental large, triangular. a transverse gular fold (vs. absent); from J. major by a Genials divided by convex scales of irregular shape. fewer T4S ≤ 22 (vs. ≥ 23); from J. tricarinata by shorter Backwards of parietale and orbital region the head is limbs FLL/SVL ≤ 45.5% (vs. ≥ 50%) and HLL/SVL ≤ covered with granular scales of different size. There are 82% (vs. ≥ 86%) and from J. major by a longer tail length 12–13 scales in interorbital distance; 5 scales in internasal TL/SVL ≥ 226% (vs. ≤ 182%) (Smith, 1935; Mahony, distance; 8 large supralabiale scales are separated from 2009, 2010; Manthey et al., 2012). the orbital area by two rows of large scales. According to detailed data arrangement from Manthey No gular pouch visible. et al., 2012 we made particular comparison within the Tympanum hidden, covered with scales; supraciliary group of species with tympanum concealed and absence scales widely overlapping; gular scales homogeneous, or presence of transverse gular fold: keeled; transverse gular fold absent, numerous head scales From the species with tympanum hidden and presence with hair-like sense organs; dorsals heterogeneous in of transverse gular fold J. batangensis Li, Deng, Wu shape, keeled, directed caudally; some groups of enlarged et Wang, 2001, J. brevicauda Manthey, Denzer, Hou scales on each side of the back parallel­ to the MDs et Wang, 2012, J. flaviceps Barbour et Dunn, 1919, J. separated by smaller scales; limbs with keeled scales. hamptoni Smith, 1935, J. yulongensis Manthey, Denzer, Dorsal crest starts from the neck, separated from Hou et Wang, 2012, J. splendida Barbour et Dunn, 1919, parietale by 4 scales and mostly developed in between the J. zhaoermii Goa et Hou, 2002, J. vela Wang, Jiang et first and seventh rows, posteriorly enlarged scales go to Che, 2015, J. laeviventris Wang, Jiang, Siler, et Che, 2016 the tail. Crest scales with visible keels. The tail is flattened and J. iadina Wang, Jiang, Siler et Che, 2016; it differs by laterally and like a body covered with heterogeneous the absence of such a fold. keeled scales above and more ordered keeled scales below. Additionally from J. batangensis by a narrower head From the crest to the belly there are rows of large keeled HW ≤ 18% (vs. ≥ 21%), a lesser number of IL ≤ 9 (vs. ≥ scales, interspersed with smaller scales of irregular shape. 10) and by a higher number of MD ≥ 54 (vs. ≤ 44); from The belly is covered with small keeled scales. J. brevicauda J. ngoclinensis differs by the possessing Color in alcohol Dorsally grey-brownish, ventrally longer tail ≥ 226% (vs. ≤ 145%); from J. flaviceps by lighter uniform colored; brownish and light-grey a narrower head HW/SVL ≤ 18% (vs. ≥ 19%), from J. tuberculate irregularly arranged scales on the head; hamptoni by a higher number of MD ≥ 54 (vs.44) and ventral side of the head light-brownish; white stripe under fewer F4S ≥ 20 (vs. 24); from J. zhaoermii by a narrower the eye, gular region without spots; nape and dorsum head HW/SVL ≤ 18% (vs. ≥ 20%), higher number of MD until the base of the tail with light cross bands differing ≥ 54 (vs. ≤ 47) and fewer F4S ≥ 20 (vs. ≥ 24); from J. in shape and size; a light dorsolateral band, dark brown yulongensis by a higher number of MD ≥ 54 (vs. 35–42); flanks with numerous lighter spots; limbs dorsally with from J. vela by more narrow head HW/SVL ≤ 17% (vs. ≥ dark bands. 21%), shorter forelimbs ≤ 45% (vs. ≥ 45%) and a higher Variation The morphometric and meristic variations are number of MD ≥ 54 (vs. ≤ 51); from J. laeviventris by compiled in Table 1. lesser number of MD ≤ 56 (vs. ≥ 57) and from J. iadina Etymology The specific epithet refers to the distribution by a shorter relative snout length SEL/HL ≥ 41% (vs. of the new species in the Ngoc Linh Mountain, Kon Tum 34.9%–40.2%). Province, Vietnam. Comparison within the group of species with Distribution and natural history The species is known tympanum concealed and absence of transverse gular only from the type locality (Figure 1). All three specimens fold: were collected on a forested slope of Ngoc Linh Mount, Japalura ngoclinensis differs from J. andersoniana elevation 1650 m. Annandale, 1905 by absence of arrangement of dorsal Comparison Japalura ngoclinensis has tympanum pholidosis with v-shaped rows of enlarged dorsals (vs. hidden what differs it from following species: J. dasi enlarged dorsal scales in v-shaped rows) and shorter Shah et Kästle, 2002), J. major (Jerdon, 1870), J. hindlimbs HLL/SLV ≤ 82% (vs. ≥ 88%); 18 Asian Herpetological Research Vol. 8

Table 1 Morphometric and meristic characters of Japalura ngoclinensis sp. nov. (for abbreviations see Materials and Methods).

ZISP 26230 ZISP 26231 ZISP 26232 (=VNMN 3110) Characters/№/sex Female Female Female holotype paratype paratype 1 SVL 68.56 67.92 57.05 2 TL 163.65 153.71 99.41* 3 HL 21.09 20.27 16.85 4 HW 12.68 11.28 10.15 5 HD 8.79 8.83 7.32 6 IN 4.9 4.3 7 SEL 8.65 8.21 7.02 8 IOD 8.71 6.42 7.94 9 FLL 27.82 29.20 25.93 10 HLL 49.47 49.83 47.01 11 SER 6 5 6 12 SEN 5 5 5 13 SL 7/8 8/9 8 14 IL 8/9 7/7 9 15 SEIP 7 8 8 16 DC 6 5 8 17 MD 54 56 54 18 F4S 21 20/22 21 19 T4S 26 25/26 24 20 NSL 1 1 1 21 TL/SVL 239% 226% 174% 22 HL/SVL 31% 29% 29% 23 HW/SVL 18% 17% 18% 24 HD/SVL 13% 13% 13% 25 HW/HL 60% 56% 60% 26 FLL/SVL 41% 43% 45% 27 HLL/SVL 72% 73% 82% * autotomized tail.

from J. chapaensis Bourret, 1937 by fewer T4S ≤ 26 from J. yunnanensis (Anderson, 1878) by presence of (vs. ≥ 28); NSL ≥ 1 (vs. 0) by a higher number of MD ≥ 54 (vs. ≤ from J. fasciata Mertens, 1926, by slightly smaller 43). adult size of females, SVL to 58.7 mm (vs. to 71 mm), From the East Asian insular species, it differs higher number of MD ≥ 54 (vs. ≤ 38), and absence of from J. brevipes Gressit, 1936, J. luei Ota, Cheng et clear white transverse band (vs. presence of such a band); Shang, 1998 and J. makii Ota, 1989 by having more NSL from J. grahami Steineger 1924 by more NSL ≥ 1 ≥ 1 (vs.0); (vs. absent) and MD present (vs.absent); from J. from J. polygonata Hallowell, 1861 and J.swinhonis micangshanensis Song 1987 by narrower head width Günther, 1864 by absence (vs. presence) of a light lateral HW/SVL ≤ 18% (vs. ≥ 21%); stripe and more clearly pronounced nuchal crest and from J. otai Mahony, 2009 by narrower head width by shorter forelimbs ≤ 45% (vs. ≥ 47.6% and ≥ 45% HW/SVL ≤ 18% (vs. ≥ 21%); respectively). from J. planidorsata Jerdon, 1870 by a larger SVL ≥ The character of enlarged row of scales offered by 57 mm (vs. ≤ 53 mm); Mahony (2010) as one of three scale characters which in from J. sagittifera Smith, 1940 by the absence of a different combinations can be used in attempt to stabilise light stripe from the eye to the angle of the mouth (vs. the generic status of some systematically problematic present) and a little fewer MD ≤ 52 (vs. ≥ 53); draconin species. It is present in J. ngoclinensis as from J. variegata Gray, 1853 by a little more ≥ 54 (vs. well as in the following East Asian Japalura s.l. ≤ 54) mm; species with this enlarged row of scales: J. brevipes, No. 1 Natalia B. ANANJEVA et al. A New Species of Japalura from Central Vietnam 19

J. chapaensis, J. fasciata, J. hamptoni, J. luei, J. makii, J. This genus is one of five recognized agamid genera splendida and J. yunnanensis. (Draco, Japalura, Stellio, Calotes and Uromastyx) crossed the boundaries of the six geographical regions of 4. Discussion endemism (Moody, 1980); it is only agamid genus which penetrates northern boundary of proposed transition The systematic status of many Asian Draconine agamids zone between the Palearctic and Indomalayan kingdoms has long been in dispute; general arrangement within (Bobrov, 1997). this subfamily is not clear (Mahony, 2010). Perhaps Phylogenetic studies of the genus demonstrated its first noted by Schmidt (1927), the genus Japalura polyphyly with using only 4 species. This phylogeny Gray has been recognized to be paraphyletic that showed 2 clades – J. flaviceps and J. splendida from received recent evidence and further support based Southeast Asia with Calotes and Acanthosaura capra and on numerous molecular phylogenetic studies (Honda A. lepidogaster, Pseudocalotes flavigula and brevipes et al., 2000; Macey et al., 2000; Schulte et al., 2004; (Macey et al., 2000) and another clade: J. tricarinata and Zug et al., 2006). This genus is presented at least by J.variegata with Draco blanfordii and 2 species of genus two evolutionary lineages (Himalayan lineage and Ptyctolaemus (Macey et al., 2000; Schulte et al., 2004; Indochina lineage). Without a comprehensive molecular Zug et al., 2006). Phylogenetic position of other known phylogeny on this genus it is difficult to hypothesize the species, including a new one, remains unknown. Knowing eastern range limits of Japalura sensu stricto (s.s.) due the patterns of distribution and morphological similarities to overall morphological similarities between these two we can suggest that it refers to the Southeast Asian undefined groups (Mahony, 2010). Schleich and Kästle evolutionary line. However, as has been shown above, the (1998) revalidated genus Oriocalotes Günther, 1864 lizards of this genus are morphologically poorly diverged. and combined all species with a visible tympanum e.g. Thus for polyphyletic Japalura lizards like for other tricarinata in this genus, while placing species with a draconin agamid we meet difficulties of establishing and hidden tympanum, e.g. variegata into the genus Japalura. evaluation of morphological characters (Mahony, 2010). In contrast Macey et al., 2000 and Schulte et al., 2004 The record of a new species significantly changes showed in their phylogenetic analysis that particularly existing ideas about the southern border of genus J. variegata and J. tricarinata fall into the same group distribution, which had previously been considered in of related species which shows affinity to Draco, while Tonkin. On the territory of Vietnam earlier two species J. flaviceps and J. splendida form a different groups that were known (Bobrov, 1995; Ananjeva et al., 2007; Bobrov, is closely related to the genus of Pseudocalotes. These Semenov, 2008; Nguyen et al., 2009). One of them was molecular studies show strongly supported monophyly of described as J. swinhoinis chapaensis Bourret, 1937 from the Southeast Asian line of Japalura, Acanthosaura, and Sa Pa (Chapa), Lao Cai Province, Vietnam and afterwards Pseudocalotes. (Ota, 1989) its status was determined as distinct species, Genus includes 31 species; more than a half (18) live J. chapaensis Bourret 1937. Its distribution is limited by in continental China (Ananjeva et al., 2011; Uetz and territory of Vietnam (northern provinces Cao Bang and Hošek, 2016), among them 2 new species were discovered Lao Cai). in historical collections (Manthey et al., 2012) and 3 more Second species of Japalura from Vietnam, J. fasciata new species are recently described from Eastern Tibet Mertens 1926, was also described from Tonkin and (Wang et al., 2015, 2016). Japalura genus occurs from referred to “yunnanensis” –complex. In Vietnam it is tropical to warm temperate region in the eastern half of known only known from type locality in Lang Son Asia, ranging from northern India, Nepal and China to the Province. In China this species inhabits central and northern Indochina and subtropical islands of East Asia: southern regions and further records indicate a much Taiwan, China and islands in Southern Japan (Figure 1). wider distribution range within China. Many forms are morphologically poorly diverged but This new species, third one for Vietnam, is known only chromosomal and molecular-genetic study of insular from the type locality: Ngoc Linh Mountain, Kon Tum species allow to recognize a variety of distinct endemic province (15°05' N 107°57' E, 1700–1900 m a.s.l.) that species Japalura polygonata, J. swinhonis, J. brevipes, J. is situated in about 1000 km southwards from the earlier makii, and J. luei inhabiting Taiwan, China, with partial known and shown above distribution ranges of Japalura syntopy and polytypic J. polygonata occuring also the species (Figure 1). Ryukyu Archipelago of Japan (Ota and Honda, 2010). This new record found in Central Highland of Vietnam 20 Asian Herpetological Research Vol. 8 confirms our earlier considerations about high number Publication, 226 pp (In Russian) of endemic species of amphibians and reptiles found Honda M., Ota H., Kobayashi M., Nabhitanhata J., Hoi-Sen in Ngoc Linh Mountain. It was shown (Orlov, 2005) Yong, Sengoku S., Hikida T. 2000. Phylogenetic relationships of the family Agamidae (Reptilia: Iguania) inferred from that 282 species of amphibians and reptiles have been mitochondrial DNA sequences. Zool Sci, 17: 527–537 recorded in Central Highland of Vietnam; among them Macey J. R., Schulte II J. A., Larson A., Ananjeva N. B., Wang 174 (61.7%) was found in Ngoc Linh Mountain. A record Y., Pethiyagoda R., Rastegar-Pouyani N., Papenfuss T. J. of new species of Japalura is an important supplement 2000. Evaluating trans-Tethys migration: An example using to the index of endemics in herpetofauna of Central acrodont lizard phylogenetics. Syst Biol, 49(2): 233–256 Highland 26.6% (75 species) endemic of this region and Mahony S. 2009. A new species of Japalura (Reptilia: Agamidae) index of endemics of Ngoc Linh Mountain 46.7% (35 from northeast India with a discussion of the similar species Japalura sagittifera Smith, 1940 and Japalura planidorsata species). It also confirms the idea about the high degree Jerdon, 1870. Zootaxa, 2212: 41–61 of overlapping among regions: 134 common species Mahony S. 2010. Systematic and taxonomic revaluation of four (47.5%) for Central Highland and north-west Tonkin little known Asian agamid species, Calotes kingdonwardi Smith, (Orlov, 2005). 1935, Japalura kaulbacki Smith, 1937, Salea kakhienensis An identification key for Japalura species of the fauna Anderson, 1879 and the monotypic genus Mictopholis Smith, of Vietnam is given below: 1935 (Reptilia: Agamidae). Zootaxa, 2514: 1–23 Tympanum membrane and ear opening covered with Manthey U. 2010. 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