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

1. Introduction

Currently the genus Japalura (sensu lato) comprises 31 recognized species (Manthey et al., 2012; Uetz and Hošek, 2016; Wang et al., 2015, 2016). Several recent intergeneric changes modify generic arrangements: J. kaulbacki Smith, 1937 was transferred into the genus Pseudocalotes (Mahony 2010); revalidation of the genus Oriotiaris made by Käs tle and Schleich (1998) for Japalura species with a naked tympanum was not confirmed by results of molecular research (Macey et al., 2000; Schulte et al., 2004) and latest revisions (Mahony 2009, 2010; Manthey, 2010). The lizards of genus Japalura are distributed in southeastern and eastern Asia as well as in Himalaya region (Figure 1). Two species among known species: J. chapaensis Bourret, 1937 and J. fasciata Mertens, 1926 were registered in the fauna of Vietnam (Ota, 1989b, 2000; Ota and Weidenhöfer, 1992; Bobrov, 1995; Ananjeva et al., 2007; Bobrov, Semenov, 2008; Nguyen et al., 2009; Mahony, 2010; Manthey et al., 2012). During a survey in 2004, three specimens of

agamid lizard referred to Japalura genus but different from the other congeners were collected in the south of Central Vietnam. Comparison of morphometric and meristic characters revealed that these specimens differ from all recognized species of this genus. This new species is described below.

2. Materials and Methods

Specimens from the south of Central Vietnam (ZISP 25230–25231, VNMN3110) were compared with specimens of the other congeners. A set of characters was selected to ensure a maximum comparability of published data for all currently recognized species of the genus Japalura (Ota, 1989a, 1989b, 1991, 2000, 2003; Ota and Weidenhöfer, 1992; Ota et al., 1998; Zhao et al., 1999; Mahony, 2009, 2010; Manthey et al., 2012).

The set of 10 measurements is as follows, SVL, snout to vent length (distance from the tip of snout to anus); TL, tail length, HL, head length (distance from tip of snout to rear border of right angle of jaw); HW, head width at its widest point, HD, maximum head depth taken at the rear axis of the jaw; IN, internarial distance, IOD, interorbital distance (distance between outer edges of orbits), SEL, distance from tip of snout to anterior margin of right eye; FLL, forelimb length (distance from axilla to tip or finger

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

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

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

* Corresponding author: Prof. Natalia B. ANANJEVA, from the Zoological Institute, Russian Academy of Sciences, with her research mainly focusing on taxonomy, ecology, phylogeny and biogeography of Eurasian amphibians and reptiles.E-mail: Natalia.Ananjeva@zin.ruReceived: 24 April 2016 Accepted: 14 July 2016

ORIGINAL ARTICLE

1 Department of Herpetology, Zoological Institute, Russian Academy of Sciences 199034, St. Petersburg, Russia2 Vietnam National Museum of Nature, Vietnam Academy of Science and Technology, 18 Hoang Quoc Viet Road,

Cau Giay, Hanoi, Vietnam

Natalia B. ANANJEVA et al. A New Species of Japalura from Central VietnamNo. 1 15

IV, excluding claw, on right side); HLL, hindlimb length (distance from groin to tip of toe IV, excluding claw, on right side). Measurements were taken in millimeters (mm) to a precision of 0.1 mm with dial calipers. The following seven ratios TL/SVL, HL/SVL, HW/SVL, HD/SVL, HLL/SVL, HLL/SVL, HW/HL were calculated to compare them with other species of Japalura.

The following meristic data were recorded: SL, supralabials (number of scales bordering right margin of upper lip including the most posterior one); IL, intralabials (number of scales bordering right margin of lower lip including the most posterior one); NSL, number of scales between nasal and supralabials; SER, number of scales contacting rostral; SEN, number of scales contacting nasal; SEIP, number of scales contacting interparietal; DC, dorsal crest, number of mid-dorsal crest-like slightly enlarged serrated scales from neck to anterior margin of vent; F4S, finger IV subdigital scales number, T4S, toe IV subdigital scales number; MD, number of enlarged mid-dorsals from the nape to above the vent.

Comparative material was examined in the holdings of Zoological Institute, Russian Academy of Sciences, St. Petersburg (ZISP), Vietnam National Museum of Nature, Vietnam Academy of Science and Technology (VNMN) and Muséum National d’Histoire Naturelle, Paris (MNHN); additionally the well documented photographic material by Manthey (2010) and comprehensive summarized tables with data on all the species of genus including mainland species with concealed tym panum

(Ota, 1989a, 1991; Mahony, 2009, 2010; Manthey et al., 2012) were used.

3. Results

A description of a new species (Figures 2, 3, 4 and 5) Japalura ngoclinensis n. sp.Holotype ZISP 26230 (adult female), above Mang Xang Village, Ngoc Linh Mountain, Dac Glei District, Kon Tum Province, Vietnam (15°05' N 107°57' E, elevation 1650 m), on 25 March 2004 by Alexei V. Abramov (Figures 2 and 5).Paratypes ZISP 26231 (adult female) and VNMH 3110 (subadult female) from above Mang Xang Village, Ngoc Linh Mountain, Dac Glei District, Kon Tum Province, Vietnam (15°05' N 107°57' E, elevation 1650 m), on 25 March 2004 by Alexei V. Abramov (Figures 3–5).Diagnosis Medium-sized, long-tailed Japalura species with relatively long limbs, that differs from all other species of this genus by the following combination of characters: tympanum hidden, transverse gular fold absent, enlarged subocular scale row present. Third and fourth fingers have equal length. Very low dorsal crest from 5–6 slightly enlarged serrated scales. Description of holotype SVL 67.9 mm, TL/SVL 239%, FLL/SVL 41%, HLL/SVL 72%, HW/SVL 18%, HW/SVL 18%, HD/SVL 13%, HW/HL 60%, SL 7/8, IL 8/9, 54 MD, 21 F4S, 26 T4S. Body shape in cross section laterally slightly compressed; nuchal and dorsal crest from 6 very short serrated scales; head is separated from

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

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the body by the neck.Quadrangular rostral approximately 2.5 times broader

than high in contact with six scales including SL; head dorsally covered with heterogeneous strongly keeled

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

Figure 3 Ventral (A) and dorsal (B) views of paratype (ZISP 26231) of Japalura ngoclinensis sp. nov.

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

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

Natalia B. ANANJEVA et al. A New Species of Japalura from Central VietnamNo. 1 17

scales, snout covered by relatively flat scales as well as orbital and interorbital areas of head. Nasal scale separated from rostral by one scale and by one scale from the first supralabial scale; two scale rows between the orbit and supralabials; a group of enlarged, keeled scales at the poste rior edge of the orbit. Mental large, triangular. Genials divided by convex scales of irregular shape.

Backwards of parietale and orbital region the head is covered with granular scales of different size. There are 12–13 scales in interorbital distance; 5 scales in internasal distance; 8 large supralabiale scales are separated from the orbital area by two rows of large scales.

No gular pouch visible.Tympanum hidden, covered with scales; supraciliary

scales widely overlapping; gular scales homogeneous, keeled; transverse gular fold absent, numerous head scales with hair-like sense organs; dorsals heterogeneous in shape, keeled, directed caudally; some groups of enlarged scales on each side of the back paral lel to the MDs separated by smaller scales; limbs with keeled scales.

Dorsal crest starts from the neck, separated from parietale by 4 scales and mostly developed in between the first and seventh rows, posteriorly enlarged scales go to the tail. Crest scales with visible keels. The tail is flattened laterally and like a body covered with heterogeneous keeled scales above and more ordered keeled scales below. From the crest to the belly there are rows of large keeled scales, interspersed with smaller scales of irregular shape. The belly is covered with small keeled scales.Color in alcohol Dorsally grey-brownish, ventrally lighter uniform colored; brownish and light-grey tuberculate irregularly arranged scales on the head; ventral side of the head light-brownish; white stripe under the eye, gular region without spots; nape and dorsum until the base of the tail with light cross bands differing in shape and size; a light dorsolateral band, dark brown flanks with numerous lighter spots; limbs dorsally with dark bands.Variation The morphometric and meristic variations are compiled in Table 1. Etymology The specific epithet refers to the distribution of the new species in the Ngoc Linh Mountain, Kon Tum Province, Vietnam. Distribution and natural history The species is known only from the type locality (Figure 1). All three specimens were collected on a forested slope of Ngoc Linh Mount, elevation 1650 m. Comparison Japalura ngoclinensis has tympanum hidden what differs it from following species: J. dasi Shah et Kästle, 2002), J. major (Jerdon, 1870), J.

tricarinata (Blyth, 1853), J. kumaonensis (Annandale, 1907), J. varcoae (Boulenger, 1918) and J. dymondi (Boulenger, 1906), possessing exposed tympanum (Smith, 1935; Manthey et al., 2012). Additionally from: J. dasi, J. kumaonensis and J. varcoae by the presence of a transverse gular fold (vs. absent); from J. major by a fewer T4S ≤ 22 (vs. ≥ 23); from J. tricarinata by shorter limbs FLL/SVL ≤ 45.5% (vs. ≥ 50%) and HLL/SVL ≤ 82% (vs. ≥ 86%) and from J. major by a longer tail length TL/SVL ≥ 226% (vs. ≤ 182%) (Smith, 1935; Mahony, 2009, 2010; Manthey et al., 2012).

According to detailed data arrangement from Manthey et al., 2012 we made particular comparison within the group of species with tympanum concealed and absence or presence of transverse gular fold:

From the species with tympanum hidden and presence of transverse gular fold J. batangensis Li, Deng, Wu et Wang, 2001, J. brevicauda Manthey, Denzer, Hou et Wang, 2012, J. flaviceps Barbour et Dunn, 1919, J. hamptoni Smith, 1935, J. yulongensis Manthey, Denzer, Hou et Wang, 2012, J. splendida Barbour et Dunn, 1919, J. zhaoermii Goa et Hou, 2002, J. vela Wang, Jiang et Che, 2015, J. laeviventris Wang, Jiang, Siler, et Che, 2016 and J. iadina Wang, Jiang, Siler et Che, 2016; it differs by the absence of such a fold.

Additionally from J. batangensis by a narrower head HW ≤ 18% (vs. ≥ 21%), a lesser number of IL ≤ 9 (vs. ≥ 10) and by a higher number of MD ≥ 54 (vs. ≤ 44); from J. brevicauda J. ngoclinensis differs by the possessing longer tail ≥ 226% (vs. ≤ 145%); from J. flaviceps by a narrower head HW/SVL ≤ 18% (vs. ≥ 19%), from J. hamptoni by a higher number of MD ≥ 54 (vs.44) and fewer F4S ≥ 20 (vs. 24); from J. zhaoermii by a narrower head HW/SVL ≤ 18% (vs. ≥ 20%), higher number of MD ≥ 54 (vs. ≤ 47) and fewer F4S ≥ 20 (vs. ≥ 24); from J. yulongensis by a higher number of MD ≥ 54 (vs. 35–42); from J. vela by more narrow head HW/SVL ≤ 17% (vs. ≥ 21%), shorter forelimbs ≤ 45% (vs. ≥ 45%) and a higher number of MD ≥ 54 (vs. ≤ 51); from J. laeviventris by lesser number of MD ≤ 56 (vs. ≥ 57) and from J. iadina by a shorter relative snout length SEL/HL ≥ 41% (vs. 34.9%–40.2%).

Comparison within the group of species with tympanum concealed and absence of transverse gular fold:

Japalura ngoclinensis differs from J. andersoniana Annandale, 1905 by absence of arrangement of dorsal pholidosis with v-shaped rows of enlarged dorsals (vs. enlarged dorsal scales in v-shaped rows) and shorter hindlimbs HLL/SLV ≤ 82% (vs. ≥ 88%);

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from J. chapaensis Bourret, 1937 by fewer T4S ≤ 26 (vs. ≥ 28);

from J. fasciata Mertens, 1926, by slightly smaller adult size of females, SVL to 58.7 mm (vs. to 71 mm), higher number of MD ≥ 54 (vs. ≤ 38), and absence of clear white transverse band (vs. presence of such a band);

from J. grahami Steineger 1924 by more NSL ≥ 1 (vs. absent) and MD present (vs.absent); from J. micangshanensis Song 1987 by narrower head width HW/SVL ≤ 18% (vs. ≥ 21%);

from J. otai Mahony, 2009 by narrower head width HW/SVL ≤ 18% (vs. ≥ 21%);

from J. planidorsata Jerdon, 1870 by a larger SVL ≥ 57 mm (vs. ≤ 53 mm);

from J. sagittifera Smith, 1940 by the absence of a light stripe from the eye to the angle of the mouth (vs. present) and a little fewer MD ≤ 52 (vs. ≥ 53);

from J. variegata Gray, 1853 by a little more ≥ 54 (vs. ≤ 54) mm;

from J. yunnanensis (Anderson, 1878) by presence of NSL ≥ 1 (vs. 0) by a higher number of MD ≥ 54 (vs. ≤ 43).

From the East Asian insular species, it differs from J. brevipes Gressit, 1936, J. luei Ota, Cheng et

Shang, 1998 and J. makii Ota, 1989 by having more NSL ≥ 1 (vs.0);

from J. polygonata Hallowell, 1861 and J.swinhonis Günther, 1864 by absence (vs. presence) of a light lateral stripe and more clearly pronounced nuchal crest and by shorter forelimbs ≤ 45% (vs. ≥ 47.6% and ≥ 45% respectively).

The character of enlarged row of scales offered by Mahony (2010) as one of three scale characters which in different combinations can be used in attempt to stabilise the generic status of some systematically problematic draconin species. It is present in J. ngoclinensis as well as in the following East Asian Japalura s.l. species with this enlarged row of scales: J. brevipes,

Characters/№/sexZISP 26230 ZISP 26231 ZISP 26232 (=VNMN 3110)

Female Female Femaleholotype paratype paratype

1 SVL 68.56 67.92 57.052 TL 163.65 153.71 99.41*3 HL 21.09 20.27 16.854 HW 12.68 11.28 10.155 HD 8.79 8.83 7.326 IN 4.9 4.37 SEL 8.65 8.21 7.028 IOD 8.71 6.42 7.949 FLL 27.82 29.20 25.9310 HLL 49.47 49.83 47.0111 SER 6 5 612 SEN 5 5 513 SL 7/8 8/9 814 IL 8/9 7/7 915 SEIP 7 8 816 DC 6 5 817 MD 54 56 5418 F4S 21 20/22 2119 T4S 26 25/26 2420 NSL 1 1 121 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%

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

* autotomized tail.

Natalia B. ANANJEVA et al. A New Species of Japalura from Central VietnamNo. 1 19

J. chapaensis, J. fasciata, J. hamptoni, J. luei, J. makii, J. splendida and J. yunnanensis.

4. Discussion

The systematic status of many Asian Draconine agamids has long been in dispute; general arrangement within this subfamily is not clear (Mahony, 2010). Perhaps first noted by Schmidt (1927), the genus Japalura Gray has been recognized to be paraphyletic that received recent evidence and further support based on numerous molecular phylogenetic studies (Honda et al., 2000; Macey et al., 2000; Schulte et al., 2004; Zug et al., 2006). This genus is presented at least by two evolutionary lineages (Himalayan lineage and Indochina lineage). Without a comprehensive molecular phylogeny on this genus it is difficult to hypothesize the eastern range limits of Japalura sensu stricto (s.s.) due to overall morphological similarities between these two undefined groups (Mahony, 2010). Schleich and Kästle (1998) revalidated genus Oriocalotes Günther, 1864 and combined all species with a visible tympanum e.g. tricarinata in this genus, while placing species with a hidden tympanum, e.g. variegata into the genus Japalura. In contrast Macey et al., 2000 and Schulte et al., 2004 showed in their phylogenetic analysis that particularly J. variegata and J. tricarinata fall into the same group of related species which shows affinity to Draco, while J. flaviceps and J. splendida form a different groups that is closely related to the genus of Pseudocalotes. These molecular studies show strongly supported monophyly of the Southeast Asian line of Japalura, Acanthosaura, and Pseudocalotes.

Genus includes 31 species; more than a half (18) live in continental China (Ananjeva et al., 2011; Uetz and Hošek, 2016), among them 2 new species were discovered in historical collections (Manthey et al., 2012) and 3 more new species are recently described from Eastern Tibet (Wang et al., 2015, 2016). Japalura genus occurs from tropical to warm temperate region in the eastern half of Asia, ranging from northern India, Nepal and China to the northern Indochina and subtropical islands of East Asia: Taiwan, China and islands in Southern Japan (Figure 1). Many forms are morphologically poorly diverged but chromosomal and molecular-genetic study of insular species allow to recognize a variety of distinct endemic species Japalura polygonata, J. swinhonis, J. brevipes, J. makii, and J. luei inhabiting Taiwan, China, with partial syntopy and polytypic J. polygonata occuring also the Ryukyu Archipelago of Japan (Ota and Honda, 2010).

This genus is one of five recognized agamid genera (Draco, Japalura, Stellio, Calotes and Uromastyx) crossed the boundaries of the six geographical regions of endemism (Moody, 1980); it is only agamid genus which penetrates northern boundary of proposed transition zone between the Palearctic and Indomalayan kingdoms (Bobrov, 1997).

Phylogenetic studies of the genus demonstrated its polyphyly with using only 4 species. This phylogeny showed 2 clades – J. flaviceps and J. splendida from Southeast Asia with Calotes and Acanthosaura capra and A. lepidogaster, Pseudocalotes flavigula and brevipes (Macey et al., 2000) and another clade: J. tricarinata and J.variegata with Draco blanfordii and 2 species of genus Ptyctolaemus (Macey et al., 2000; Schulte et al., 2004; Zug et al., 2006). Phylogenetic position of other known species, including a new one, remains unknown. Knowing the patterns of distribution and morphological similarities we can suggest that it refers to the Southeast Asian evolutionary line. However, as has been shown above, the lizards of this genus are morphologically poorly diverged. Thus for polyphyletic Japalura lizards like for other draconin agamid we meet difficulties of establishing and evaluation of morphological characters (Mahony, 2010).

The record of a new species significantly changes existing ideas about the southern border of genus distribution, which had previously been considered in Tonkin. On the territory of Vietnam earlier two species were known (Bobrov, 1995; Ananjeva et al., 2007; Bobrov, Semenov, 2008; Nguyen et al., 2009). One of them was described as J. swinhoinis chapaensis Bourret, 1937 from Sa Pa (Chapa), Lao Cai Province, Vietnam and afterwards (Ota, 1989) its status was determined as distinct species, J. chapaensis Bourret 1937. Its distribution is limited by territory of Vietnam (northern provinces Cao Bang and Lao Cai).

Second species of Japalura from Vietnam, J. fasciata Mertens 1926, was also described from Tonkin and referred to “yunnanensis” –complex. In Vietnam it is known only known from type locality in Lang Son Province. In China this species inhabits central and southern regions and further records indicate a much wider distribution range within China.

This new species, third one for Vietnam, is known only from the type locality: Ngoc Linh Mountain, Kon Tum province (15°05' N 107°57' E, 1700–1900 m a.s.l.) that is situated in about 1000 km southwards from the earlier known and shown above distribution ranges of Japalura species (Figure 1).

This new record found in Central Highland of Vietnam

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confirms our earlier considerations about high number of endemic species of amphibians and reptiles found in Ngoc Linh Mountain. It was shown (Orlov, 2005) that 282 species of amphibians and reptiles have been recorded in Central Highland of Vietnam; among them 174 (61.7%) was found in Ngoc Linh Mountain. A record of new species of Japalura is an important supplement to the index of endemics in herpetofauna of Central Highland 26.6% (75 species) endemic of this region and index of endemics of Ngoc Linh Mountain 46.7% (35 species). It also confirms the idea about the high degree of overlapping among regions: 134 common species (47.5%) for Central Highland and north-west Tonkin (Orlov, 2005).

An identification key for Japalura species of the fauna of Vietnam is given below:Tympanum membrane and ear opening covered with scales; transverse gular fold absent…...….Japalura genus1. presence of clear white transverse band…..... J. fasciata 2. absence of clear white transverse band…………....……2a. MD ≤ 37………..................................… J. chapaensis2b. MD ≥ 54 ……………………….…..... J. ngoclinensis

Acknowledgements The study was partially supported by grants RFBR 15-04-01730, 14-04-92000 NNS, 15-29-02457ofi and 17-54-54003, and under participation of Zoological Institute (theme No. 00125-2016-0002). Many thanks to Evgeny A. Golynsky who prepared distribution map. We are especially grateful to the Joint Russian-Vietnamese Tropical Research and Technological Center under the A. N. Severtsov Institute of Ecology and Evolution RAS and Alexei V. Abramov (ZISP) who kindly provided specimens of Japalura from Ngoc Linh Mountain.

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