Cyrtodactylus roesleri, Ziegler, Thomas, Nazarov, Roman, Orlov, Nikolai, Vu, Truong Quang Nguyen Thanh Ngoc, Dang, Kien Ngoc, Dinh, Tri Huy & Schmitz, Andreas, 2010

Ziegler, Thomas, Nazarov, Roman, Orlov, Nikolai, Vu, Truong Quang Nguyen Thanh Ngoc, Dang, Kien Ngoc, Dinh, Tri Huy & Schmitz, Andreas, 2010, A third new Cyrtodactylus (Squamata: Gekkonidae) from Phong Nha-Ke Bang National Park, Truong Son Range, Vietnam, Zootaxa 2413, pp. 20-36 : 24-33

Cyrtodactylus roesleri sp. nov.

Holotype. Adult male ( ZFMK 89377) collected on 22 May 2009 by Thomas Ziegler and Kien Ngoc Dang nearby Phong Nha – Ke Bang National Park, Minh Hoa District, Quang Binh Province, Vietnam.

Paratypes. Three adult males ( IEBR A.0932, MHNG 2713.79, VNUH 220509) and one juvenile ( ZFMK 89378) from the type locality, collected by Thomas Ziegler and Kien Ngoc Dang on 22 May 2009; one adult female ( ZFMK 86433) collected by Thomas Ziegler and Astrid Heidrich in June 2006 within Phong Nha – Ke Bang National Park, Quang Binh Province, Vietnam; ten adult males ( ZISP 25777-25782, ZMMU R- 13061-1, ZMMU R- 13061-2, ZMMU R- 13061-3, ZMMU R- 13061-4) and four adult females ( ZISP 25783, ZISP 25784, ZMMU R- 13061-5, ZMMU R- 13051-6) collected by Nikolai Orlov in July 2003 within Phong Nha – Ke Bang National Park, Quang Binh Province, Vietnam.

Diagnosis. A small, slender Cyrtodactylus with a maximum SVL of 75.3 mm, distinguished from all congeners by the combination of the following characters: 1) dorsal pattern consisting of dark nuchal band and 4–5 sometimes irregular shaped dark transversal bands between limbs; 2) dorsal tubercles present on the back of head, body, hind limbs and tail base; 3) ventrals in 34–40 longitudinal rows at midbody; 4) lateral folds absent; 5) continuous series of 20–28 precloacal and femoral pores in males; 6) enlarged femoral scales present; 7) enlarged precloacal scales present; 8) precloacal groove absent; 9) 5–8 postcloacal spurs in both sexes; 10) subcaudal scales transversally enlarged; 11) fourth toe with 17–21 subdigital lamellae.

Description of holotype. Size small (SVL 65.7 mm, TaL 77.5 mm), distance from posterior corner of eye to anterior margin of ear including ciliaria 5.0 mm, maximum horizontal ear diameter 1.0 mm; for further measurements see Table 1.

Rostral wider than high (RW 3.3 mm, RH 1.9 mm, RW/RH 1.7) with an inverse Y-shaped median suture; supralabials 11(r)/11(l); scales between orbit and the eighth supralabial 3/3, small; infralabials 9(r)/8(l); nares bordered by rostral anteriorly, first supralabial laterally and 3(r)/4(l) nasals posteriorly; supranasals separated from each other by two nasorostrals and an internasal; medial snout scales granular, those in contact with and nearby the supralabials flattened and approximately 2–4 times larger than the medial scales; upper anterior ciliaries 2–3 times larger than posterior cilaries; head scales somewhat granular, one-half as large as median snout scales; back of dorsal head and temporal region with rounded, conical tubercles, 2–3 times larger than the surrounding scales; mental triangular, slightly wider than rostral; 1 pair of enlarged postmentals, longer than wide, bordering with mental anteriorly, first two labials on the right side, first labial on the left side, and followed by 2 pairs of enlarged gular scales, and some merged smaller gular scales in between; dorsal scales granular, as large as or somewhat larger than medial snout scales; dorsal tubercles round, conical, weakly keeled posteriorly, surrounded by 8–9 granular scales, tubercles forming about 13 ill-defined (irregular) longitudinal rows at midbody; ventral scales smooth, medial scales 2–3 times larger than the dorsal granules, 39 longitudinal rows at midbody; lateral folds absent; upper and lower arm lacking enlarged tubercles; dorsal hind limb covered with scattered, smooth, flat to conical tubercles; 1–2 outermost femoral pores bearing scales enlarged, attached to nine enlarged, poreless femoral scales on each hind limb; fingers and toes lacking distinct webbing; lamellae numbering 14 under first finger (5 basal and 9 distal lamellae), 17 under fourth finger (6 basal and 11 distal lamellae), 14–15 under first toe (5 basal and 9–10 distal lamellae), and 19–20 (7– 8 basal and 12 distal lamellae) under fourth toe; claws surrounded by a small scale on upper and a large scale on lower sides; 28 contiguous precloacal and femoral pores in an angular series; precloacal region covered by a patch of about 24 enlarged scales; precloacal groove absent; 5(r)/6(l) enlarged postcloacal tubercles on lateral surface of hemipenial swelling; dorsal tail surface only at early base with distinct tubercles; subcaudals distinctly enlarged, flat, smooth. The apex of the hemipenis is thickened and deeply bifurcated. Sulcus spermaticus thin, deep, bordered by bulging lips; the sulcus bifurcates in the upper truncus region. Apical lobes with spinose calyces.

Coloration in ethanol. Dorsal head light brownish grey with some ill-defined dark blotches especially in the back; nuchal band not distinctly broadened, dark brown and with white margins, extending from the neck to the posterior margins of eyes; between anterior margin of eye and nostril some ill-defined and in part connected dark blotches; labials greyish-brown without distinct dots. Dorsum brownish-grey with 4 dark brown transverse bands in between limbs; dark bands anteriorly somewhat lighter, with light margins; anterior three dorsal bands distinctly widened towards vertebral region; the last, not distinctly widened dorsal band is located in the sacral region; more or less distinct dark lateral blotches are discernible between dorsal bands; flanks greyish-brown marbled, somewhat lighter than dorsal surface and darker than cream to grey venter. Lower side of toes and fingers grey; dorsal limbs and digits greyish-brown with dark marbling or blotches. Tail dorsum with about 12 light (whitish-grey) transversal bands; tail tip light; in front of the anterior 4–5 light bands there are black transversal bands; the wide interspaces between these black and white or only white transversal bands are grey in colour. Ventral side of tail anterior dark grey with white spots, posterior paler.

For coloration in life see Figs. 2 and 4–6 . Especially conspicuous are yellow portions in the pattern, such as the light borders of the dorsal bands including the nuchal and neck band. Here, especially the dorsal tubercles are brightly yellow in colour. Also parts of the ciliaries, the posterior supralabials, and the overall surface colour of limbs, head and body are yellow to yellowish-beige (which seemingly fades to brownish grey in preservative).

Variation of paratypes. For the variation of paratypes see Tables 1–2. Males are somewhat smaller than the females: SVL males 51.1–73.5 mm (64.8± 6.5 mm), n = 14; females 63.5–75.3 mm (71.7± 4.9 mm), n = 5. Paratypes largely correspond with the description of the holotype. Noteworthy differences are up to about 19 ill-defined dorsal tubercle rows at midbody, 7–10 enlarged poreless femoral scales on each hind limb in males, and minimally 10 enlarged scales in the precloacal region. Concerning the colour pattern, most of the male and female paratypes show a more or less distinct dark stripe stretching from the anterior corner of the eye towards the nostril. Several paratypes also have up to five dark dorsal bands between limbs, and some of them are irregular shaped (such as Y-shaped), medially interrupted or developed only at one side of the dorsum.

Dark lateral blotches between dorsal bands are only discernible in some paratypes and are also lacking in the single juvenile paratype. The male paratype IEBR A.0932 is especially brightly coloured (see Fig. 6 ), with intensive black blotches on the head, deep black transversal dorsal bands and bright yellow tubercles in lighter interspaces. In this specimen, the gular region is dark grey with light blotches in preservative, which are yellow in live. Such a throat pattern is more or less also discernible in some other adult paratypes as well as in the juvenile paratype. The latter specimen also has a white posterior half of the tail with only faint dark transversal bands, which differs from the adult condition. Original tails of adults may show thin, bright yellow borders of dark transversal bands, with light interspaces of last third of tail being white.

Comparisons. Comparisons are based on the original descriptions or descriptions provided in broader faunal and taxonomic publications (e.g., Smith 1920, 1921, 1935; Taylor 1963; Ulber & Grossmann 1991; Ulber 1993; Darevsky & Szczerbak 1997; Manthey & Grossmann 1997; Ziegler et al. 2002; Bauer 2002, 2003; Bauer et al. 2002, 2003; David et al. 2004; Pauwels et al. 2004; Grismer 2005; Grismer & Leong 2005; Ngo & Grismer 2006; Nguyen et al. 2006; Grismer et al. 2007; Heidrich et al. 2007; Hoang et al. 2007; Orlov et al. 2007; Grismer & Norhayati 2008; Grismer et al. 2008; Nazarov et al. 2008; Ngo 2008; Ngo & Bauer 2008; Ngo et al. 2008; Rösler & Glaw 2008; Rösler et al. 2008; Bauer et al. 2009; Geissler et al. 2009).

5 6 - - 13061 13061

86433 25783 25784 R- R- max. s) 89378 ZFMK ZISP ZISP ZMMU ZMMU min -. (m ± ZFMK

Sex f f f f f Juv. SVL 75.3 71.3 63.5 74.6 74 max. 75.3 38.3 TaL 74.2 86.0 * 74.0* 65.0* max. 86.0 44.4

In comparison with Vietnamese congeners, Cyrtodactylus roesleri sp. nov. is distinguishable from C. cattienensis , C. cryptus , C. huynhi , C. irregularis , C. pseudoquadrivirgatus , and C. ziegleri by the presence of transversally enlarged subcaudals which are lacking in the latter species. Cyrtodactylus roesleri sp. nov. furthermore has series of 20–28 precloacal and femoral pores in males, and 17–22 in females, whereas precloacal pores are completely lacking in males of C. badenensis , C. eisenmanae and C. grismeri , or are lacking or only rudimentarily developed in males of C. nigriocularis (0–2) and C. paradoxus (0–4); in addition, Cyrtodactylus roesleri sp. nov. differs from C. phongnhakebangensis in a lower number of precloacal and femoral pores in males (20–28 versus 32–42), and the new species differs from the remaining species by distinctly higher pore counts: C. caovansungi (9 PP + 6 FP), C. chauquangensis (6–7), C. condorensis (4–7), C. hontreensis (7–8), C. intermedius (8–10), and C. takouensis (3–4 PP + 1 FP). For detailed comparisons see Table 3.

C. cryptus and Cyrtodactylus roesleri sp. nov. (after Ziegler et al. 2002, Heidrich et al. 2007, and own data).

Cyrtodactylus Cyrtodactylus cryptus Cyrtodactylus roesleri sp. nov. phongnhakebangensis

SVL max. 104.4 mm max. 90.8 max. 75.3 mm

nuchal band enlarged in neck region enlarged in neck region (triangular) not enlarged in neck region Compared with the sympatric species Cyrtodactylus cryptus and C. phongnhakebangensis , Cyrtodactylus roesleri sp. nov. at first glance mostly resembles the latter species. Here, closer examination allows for proper separation (see Table 4). In addition, C. phongnhakebangensis is a relatively large species and we examined a subadult C. phongnhakebangensis with a SVL of 79.4 mm (which is beyond the currently known maximum SVL of adult Cyrtodactylus roesleri sp. nov.), that still showed the juvenile colour pattern on its tail, i.e. light dorsal tail bands that are completely yellow at base, and becoming lighter towards the tail tip which is completely white.

When comparing Cyrtodactylus roesleri sp. nov. with remaining congeners from the neighbouring countries in mainland Southeast Asia, it differs from C. angularis , C. ayeyarwadyensis , C. brevidactylus , C. buchardi , C. gansi , C. pantiensis , C. quadrivirgatus , C. semenanjungensis , C. stresemanni , C. sworderi , and C. wakeorum by having enlarged subcaudal scales (versus lacking in latter species). Cyrtodactylus roesleri sp. nov. has a higher precloacal (precloacal-femoral) pore count in males (20–28) compared with C. consobrinoides (4), C. feae (8–10), C. khasiensis (12–14), C. nebulosus (0), C. oldhami (1–4), C. papilionoides (4–6), C. peguensis (7–9), C. rubidus (12–14), C. sumonthai (1–2), and C. thirakhupti (0). In contrast, Cyrtodactylus roesleri sp. nov. has a lower precloacal (precloacal-femoral) pore count in males (20– 28) compared with C. chanhomeae (32–34) and C. jarujini (52–54). Cyrtodactylus roesleri sp. nov. has a continuous series of precloacal and femoral pores, whereas in C. pulchellus , precloacal and femoral pores are well distinguishable, and in the species C. aequalis , C. annandalei , C. brevipalmatus , C. erythrops , C. gubernatoris , C. interdigitalis , C. russeli , C. slowinskii , and C. tigroides there is a diastema separating femoral from precloacal pores. Cyrtodactylus roesleri sp. nov. differs from C. consobrinus by fewer ventrals in longitudinal rows at midbody (34– 40 versus 65–70); the new species differs from C. elok , C. macrotuberculatus and C. variegatus by a higher number of ventrals at midbody (34–40 versus 6–10 in C. elok , 19–22 in C. macrotuberculatus , and 22 in C. variegatus ). Finally, Cyrtodactylus roesleri sp. nov. differs by fewer (4–5) transversal dark dorsal bands between limbs from C. chrysopylos who has a dorsal pattern consisting of nine bands between occiput and sacrum.

Distribution. Cyrtodactylus roesleri sp. nov. is currently known only from Phong Nha – Ke Bang National Park and surroundings in Quang Binh Province, Vietnam (see Fig. 7 ).

Etymology. We name this species after our friend and colleague Herbert Rösler to honour his long-term engagement in breeding and studying geckonid lizards. Herbert Rösler published several books dealing with geckos and is specialized in geckonid morphology and taxonomy, which resulted among others in the description of diverse new gecko species.

Ecological notes. Most of the observations took place at night. The specimens were usually found on karst outcrops in limestone forest, at a height of about 1–1.5 m (sometimes also at 2 m height) above ground (even during rain).

Figures

FIGURE 2. Male holotype of Cyrtodactylus roesleri sp. nov. in life (ZFMK 89377). Photo T. Ziegler

FIGURE 4. Male paratype of Cyrtodactylus roesleri sp. nov. in life (VNUH 220509) with five dorsal bands between limbs. Photo T. Ziegler

FIGURE 5. Adult male paratype of Cyrtodactylus roesleri sp. nov. in life with asymmetrical dorsal banded pattern and with original tail. Photo N. Orlov

FIGURE 6. Portrait of an especially brightly coloured male paratype of Cyrtodactylus roesleri sp. nov. in life (IEBR A. 0932). Photo T. Ziegler

FIGURE 7. Map showing the type locality of Cyrtodactylus roesleri sp. nov. in Vietnam.