Cyrtodactylus pageli Schneider, Nguyen, Schmitz, Kingsada, Auer & Ziegler, 2011, sp. n.

Cyrtodactylus pageli sp. n.

Holotype. IEBR A.2010.36, adult male from Phoukham Cave (18°55.606´N 102°23.273´E, elevation 270 m a.s.l.), near Ban Na Thong Village, Vang Vieng District, Vientiane Province, northwestern Laos, collected on 5 May 2010 by Truong Quang Nguyen and Phouthone Kingsada (Figs. 2–5).

Paratypes. ZFMK 91827, adult female, MHNG 2723.91, adult male, NUOL 2010.3, adult female, and NUOL 2010.4, subadult male, NUOL 2010.5, subadult male, from Pouna Cave (18°55.547'N 102°22.943'E, elevation 260 m a.s.l.), near Ban Na Thong, Vang Vieng District, Vientiane Province, northwestern Laos, collected on 5 May 2010 by Truong Quang Nguyen and Phouthone Kingsada.

NUOL 2010.6, subadult male, NUOL 2010.7, adult female, MTD 48025, adult male, and IEBR A.2010.37, adult male, from Phouthong Cave (18°54.997'N, 102°24.701'E, elevation 300 m a.s.l.), near Ban Phongeuan, Vang Vieng District, Vientiane Province, northwestern Laos, collected on 5 May 2010 by Truong Quang Nguyen and Phouthone Kingsada (Figs. 5–6).

Diagnosis. A medium sized Cyrtodactylus with maximum SVL 81.8 mm, head distinct from slender body, belly flat. Cyrtodactylus pageli sp. n. can be distinguished from all congeners on the basis of the following combination of characters: 9–12 supralabials; 9 infralabials; 9–14 rows of dorsal tubercles; no distinct lateral body fold; 41–46 midbody ventrals; precloacal pores present in both sexes (males 4, females 5 or 6); no precloacal groove; femoral scales not distinctly enlarged; no femoral pores; subdigital lamellae under the fourth toe 19–26; subcaudals transversally enlarged; dark nuchal band, followed by a transversal row of dark neck blotches (rarely forming an additional transversal neck band) in-between nuchal band and first body band; five in part irregular dark dorsal bands between limb insertions plus transversal band between hind limbs; tail with banded pattern.

Description of holotype. Adult male with total length of 178.4 mm (SVL 76.2 mm, TL 102.2 mm. Rostral Yshaped, wider than high, medially with a straight, vertical rostral suture, in contact with nasorostral, nare, and first supralabial on each side; rostral as wide as mental; 11 supralabials; 5 or 6 scale rows between supralabials and orbit; nares in contact with rostral, nasorostral, supranasal, two postnasals, and first supralabial; no internasals; snout scales distinctly larger than head scales, largest snout scales medially up to rostral; loreal region only slightly upraised; 53 scales between fifth supralabials across the dorsal head surface; 39 scales between anterior corners of eyes; interorbital region with small round, convex scales, outer ones more oval; scales in postorbital region distinctly smaller (ca. half the size) than snout scales, irregular in shape; head without enlarged tubercles; pupil vertical; 16 spinous ciliaria, posterior ones more developed; ear opening vertical, oval; mental triangular, in contact with two postmentals and the first infralabial on each side; nine infralabials; postmentals surrounded by seven granular scales, of which the two outer ones are enlarged, first infralabial on each side and by second infralabial (on right side only); gular scales granular.

Dorsal scales small, twice the size of head scales between and behind the eyes; dorsal tubercles arranged in 11 longitudinal rows at midbody; sides of body without tubercles; dorsal tubercles surrounded by 8 or 9 dorsal scales; dorsal tubercles separated by 2–4 dorsals from each other; no lateral body folds; ventral scales round, slightly arched, imbricated, 2 or 3 times larger than gular and throat scales, and twice the size of dorsals; 41 ventrals plus 115 dorsals around midbody; 239 scales between mental and cloacal slit; dorsal surface of limbs without tubercles; no distinctly enlarged femoral scales; fingers and toes free of webbing; relative finger length formula I <II <V <III <IV, relative toe length formula I <II <V <III <IV; claw bordered by two scales; finger I with 16–17, finger II with 16–17, finger III with 20–21, finger IV with 20–21, and finger V with 18–19 subdigital lamellae, of which finger I has 6–7 basally broadened lamellae, finger II 7, finger III 7–8, finger IV 8 and finger V 7; toe I with 17, toe II with 19, toe III with 21–22, toe IV with 21, toe V with 21–23 subdigital scales, of which toe I has 7–8 basally broadened lamellae, toe II 7, toe III 9, toe IV 7–8, and toe V 7–8; no precloacal depression; 4 well discernible precloacal pores; pore bearing scales posteriorly surrounded by five enlarged scales; adjoining scales continuously decreasing in size; 4/4 well developed postcloacal tubercles; hemipenes only partially everted; original tail lacking distinct whorls, dorsally with tubercles at base only; median row of subcaudals transversally enlarged.

Coloration in life (Fig. 2): Top of head light brown with dark pattern; each side with a dark stripe stretching from the snout tip towards the anterior corner of the eye; indistinct dark blotches between eyes and on occiput; eyelids with green cast; ciliaria bright yellow; iris of eyes metallic-yellowish, with dark texture and with orange margin around pupil slit; a dark, posteriorly widened nuchal loop with thin light margins stretches from the posterior corners of the eyes along the neck; the basic dorsal color is light brown with transverse dark brown body bands; four dark elongate blotches between arm insertions, with a median dark spot just behind the neck; five partly irregular dark brown bands between limb insertions; posterior margin of these dark, more or less jagged body bands bordered with a thin bright stripe of light brown; laterally the body bands merge partially, but being interspersed with irregular light brown spots; dorsal pattern of front legs consisting of a patchy mixture of light and dark brown; dorsum of hind limbs mostly dark brown with some small light brown spots; tail with nine dark brown continuous bands on a light brown ground, however, these bands are much clearer and straighter than body bands, except for anterior bands; ventral side of body solid light beige, with ventral side of tail being slightly darker.

Color in preservative (70 % ethanol) (Figs. 3–5): Color in alcohol differs only slightly from the coloration in life. The overall color scheme is somewhat less pronounced, because the color slightly fades in alcohol. Main structures are still clearly visible, the slight green cast of the eyelids and the bright yellow color of the ciliaria are no longer visible, but have the same beige-brown color compared to remaining parts of the head.

Variation. The paratype series largely corresponded with the description of the holotype. For measurements, scalation, and colour pattern variation see Figs. 5–6 and Tables 3–5. Noteworthy is the higher precloacal pore count in females (5 or 6) versus 4 in males. However, pores in females are only slightly developed by minute openings.

Sexual dimorphism was also discernible in terms of distinct hemipenial swellings at the lower tail base in adult males, and the postcloacal tubercles of the males are somewhat larger than the female postcloacal tubercles. Transversal body bands individually vary in intensity (darker bands versus lighter bands) and shape (continuously developed body bands versus interrupted or irregularly shaped body bands). In the subadult males NUOL 2010.4, NUOL 2010.5, and NUOL 2010.6, the posterior region of the tail was very light to white, which thus may indicate juvenile to subadult stages, besides smaller size. In some of the paratypes lateral body fold like structures were discernible at least in parts of the flanks; however, this also may be due to the preservation state of the specimens.

TABLE 3 a. Selected measurements of the male holotype and male paratypes of Cyrtodactylus pageli sp. n.; measurements in mm, 1 = broken tail, m = mean, max. = maximum; s = standard deviation.

Further specimens. Beyond the type series there have been further specimens seen in March 2008 and May 2010, which have not been collected (Fig. 9). These individuals largely agreed in colour pattern and morphology with the type series. In March 2008, a male specimen of 81.8 mm snout-vent length was captured around Pouna Cave, Vang Vieng District, Vientiane Province, northwestern Laos. In this specimen the hemipenes could be partially everted (Fig. 7). The hemipenes appear stout, apically thickened, heart-shaped; sulcus spermaticus basally with bulging lips; on the sulcal side, between the not fully everted lobes, medially there is a protruding tissue lobe (well discernible from Fig. 7 B); upper truncus and apex with calyces, in part elongated at the base, getting smaller towards the lobes, with denticulated margins.

Comparisons. Comparisons are based on the original descriptions or descriptions provided in broader faunal and taxonomic publications (e.g., Smith 1920, 1921, 1935; Ulber & Grossmann 1991; Das 1993, 1996; Darevsky & Szczerbak 1997; Manthey & Grossmann 1997; Bauer 2002, 2003; Bauer et al. 2002, 2003; David et al. 2004; Pauwels et al. 2004; Batuwita & Bahir 2005; Grismer 2005; Grismer & Leong 2005; Nguyen et al. 2006; Youmans & Grismer 2006; Hoang et al. 2007; Orlov et al. 2007; Rösler et al. 2007; Grismer & Ahmad 2008; Grismer et al. 2008; Hayden et al. 2008; Kraus 2008; Linkem et al. 2008; Nazarov et al. 2008; Ngo 2008; Ngo & Bauer 2008; Ngo et al. 2008; Oliver et al. 2008; Rösler & Glaw 2008; Rösler et al. 2008; Bauer et al. 2009; Geissler et al. 2009; Mahony 2009; Oliver et al. 2009; Welton et al. 2009; Bauer et al. 2010; Grismer et al. 2010; Chan & Ahmad 2010; Ngo & Chan 2010; Ngo & Grismer 2010; Ngo & Pauwels 2010; Ngo et al. 2010; Nguyen et al. 2010; Shi & Zhao 2010; Sumontha et al. 2010; Welton et al. 2010; Ziegler et al. 2010).

Cyrtodactylus pageli sp. n. has 4–6 precloacal pores only (in both sexes). The new species thus differs from the following Cyrtodactylus species, which have distinctly lower or higher precloacal pore counts: C. aaroni Günther & Rösler, 2003 (9–12 precloacal pores), C. aequalis Bauer, 2003 (9), C. agusanensis (Taylor, 1915) (8–11), C. annandalei Bauer, 2003 (11–12), C. ayeyarwadyensis Bauer, 2003 (10–28), C. baluensis (Mocquard, 1890) (9–11), C. bichnganae Ngo, 2010 (10), C. biordinis Brown & McCoy, 1980 (11–14), C. brevidactylus Bauer, 2002 (8), C. caovansungi Orlov, Nguyen, Nazarov, Ananjeva & Nguyen, 2007 (9), C. capreoloides Rösler, Richards & Günther, 2007 (13), C. chrysopylos Bauer, 2003 (10), C. consobrinus (Peters, 1871) (9–11), C. cryptus Heidrich, Rösler, Vu, Böhme & Ziegler, 2007 (10), C. durio Grismer et al., 2010 (12), C. erythrops Bauer et al., 2008 (9), C. gansi Bauer, 2003 (16–29), C. gubernatoris (Annandale, 1913) (9), C. ingeri Hikida, 1990 (8), C. interdigitalis Ulber, 1993 (14), C. intermedius (Smith, 1917) (8–10), C. irianjayaensis Rösler, 2000 (9–17), C. khasiensis (Jerdon, 1870) (12–14), C. lateralis (Werner, 1896) (13), C. louisiadensis (De Vis, 1892) (18–20), C. malayanus (De Rooij, 1915) (8–10), C. malcolmsmithi (Constable, 1949) (12), C. marmoratus Gray, 1831 (16), C. nigriocularis Nguyen, Orlov & Darevsky, 2006 (0–2), C. pantiensis Grismer, Chan, Grismer, Wood & Belabut, 2008 (8–9), C. philippinicus (Steindachner, 1867) (8–12), C. pulchellus Gray, 1828 (8), C. russelli Bauer, 2003 (15), C. sadleiri Wells & Wellington, 1985 (8–13), C. slowinskii Bauer, 2002 (9–11), C. spinosus Linkem, McGuire, Hayden, Setiadi, Bickford & Brown (12–13), C. stresemanni Rösler & Glaw, 2008 (10), C. sumonthai Bauer, Pauwels & Chanhome, 2002 (2), C. teyniei David, Nguyen, Schneider & Ziegler, 2011 (14 in the single known specimen, an adult female), C. tigroides Bauer, Sumontha & Pauwels, 2003 (8–9), C. tuberculatus (Lucas & Frost, 1900) (13), C. wakeorum Bauer 2003 (12), C. wetariensis (Dunn, 1927) (11), and C. yoshii Hikida, 1990 (8–12).

The following Cyrtodactylus species have a series of precloacal-femoral or precloacal and femoral pores, which are lacking in Cyrtodactylus pageli sp. n.: C. auribalteatus Sumontha, Panitvong & Deein, 2010 (having 6 precloacal pores + 4–5 femoral pores), C. brevipalmatus (Smith, 1923) (7–10+6-7), C. chanhomeae Bauer, Sumontha & Pauwels, 2003 (having 32–34 precloacal-femoral pores), C. consobrinoides (Annandale, 1905) (26), C. deveti (Brongersma, 1948) (18–23), C. dumnuii Bauer, Kunya, Sumontha, Niyomwan, Pauwels, Chanhome & Kunya, 2010 (5–6+6), C. epiroticus Kraus, 2008 (60–82), C. feae (Boulenger, 1893) (32), C. fumosus (Müller, 1895) (42–52), C. jarujini Ulber, 1993 (42–54), C. klugei Kraus, 2008 (66–76), C. lomyenensis Ngo & Pauwels, 2010 (39–40), C. loriae (Boulenger, 1898) (30–81), C. novaeguineae (Schlegel, 1837) (24–43), C. phongnhakebangensis Ziegler, Rösler, Herrmann & Vu, 2003 (32–42), C. redimiculus King, 1962 (5–8+8–9), C. robustus Kraus, 2008 (75–85), C. roesleri Ziegler, Nazarov, Orlov, Ngyuen, Vu, Dang, Dinh & Schmitz, 2010 (20–28), C. salomonensis Rösler, Richards & Günther, 2007 (71–72), C. seribuatensis Youmans & Grismer, 2006 (40–44), C. serratus Kraus, 2007 (87), C. tamaiensis Mahony, 2009 (40), C. tiomanensis Das & Lim, 2000 (19), C. tripartitus Kraus, 2008 (64–78) and C. variegatus (Blyth, 1859) (32).

The following Cyrtodactylus species differ from Cyrtodactylus pageli sp. n. by the absence of precloacal and femoral pores in both sexes: C. badenensis Nguyen, Orlov & Darevsky, 2006, C. darmandvillei (Weber, 1890), C. eisenmanae Ngo, 2008, C. gordongekkoi (Das, 1993) (see Biswas 2007), C. grismeri Ngo, 2008, C. jarakensis Grismer, Chan, Grismer, Wood & Belabut, 2008, C. jellesmae (Boulenger, 1897), C. laevigatus (Darevsky, 1964), C. semenanjungensis Grismer & Leong, 2005, C. sermowaiensis (de Rooij, 1915), C. thirakhupti Pauwels, Bauer, Sumontha & Chanhome 2004, and C. wallacei Hayden, Brown, Gillespie, Setiadi, Linkem, Iskander, Umilaela, Bickford, Riyanto, Mumpuni & McGuire, 2008.

Cyrtodactylus pageli sp. n. has transversally enlarged subcaudals and thus differs from C. adleri Das, 1997, C. annulatus (Taylor, 1915), C. batucolus Grismer, Chan, Grismer, Wood & Belabut, 2008, C. buchardi David, Teynié & Ohler, 2004, C. cattienensis Geissler, Nazarov, Orlov, Böhme, Phung, Nguyen & Ziegler, 2009, C. cavernicolus Inger & King, 1961, C. derongo Brown & Parker, 1973, C. halmahericus Mertens, 1929, C. huynhi Ngo & Bauer, 2008, C. irregularis (Smith, 1921), C. jambangan Welton, Siler, Diesmos & Brown, C. mandalayensis Mahony, 2009, C. matsuii Hikida, 1990, C. murua Kraus & Allison, 2006, C. nuaulu Oliver, Edgar, Mumpuni, Iskandar & Lilley, 2009, C. papuensis (Brongersma, 1934), C. pseudoquadrivirgatus Rösler, Vu, Nguyen, Ngo & Ziegler, 2008, C. pubisulcus Inger, 1957, C. quadrivirgatus Taylor, 1962, 2010, C. sworderi (Smith, 1925), C. tautbatorum Welton, Siler, Diesmos & Brown, 2009, C. zhaoermii Shi & Zhao, 2010, C. ziegleri Nazarov, Orlov, Nguyen & Ho, 2008, and C. zugi Oliver, Tjarhan, Mumpuni, Krey & Richards, 2008.

Cyrtodactylus pageli sp. n. lacks distinctly enlarged femoral scales and thus differs from C. condorensis (Smith, 1921), C. phuquocensis Ngo, Grismer & Grismer, 2010 and C. takouensis Ngo & Bauer, 2008.

Cyrtodactylus pageli sp. n. has 41–46 ventral scales at midbody and thus differs from C. agamensis (Bleeker, 1860) (having 67 ventral scales), C. cracens Batuwita & Bahir (31), 2005, C. edwardtaylori Batuwita & Bahir, 2005 (29–30), C. fraenatus (Günther, 1864) (27–35), C. leegrismeri Chan & Norhayati, 2010 (27–35), C. macrotuberculatus Grismer & Norhayati, 2008 (19–22), C. papilionoides Ulber & Grossmann, 1991 (30–34), C. paradoxus (Darevsky & Szczerbak, 1997) (26–36; which in addition has different back pattern and different dorsal tubercle arrangement: 16 versus 9–14 longitudinal midbody rows in Cyrtodactylus pageli sp. n.), C. ramboda Batuwita & Bahir, 2005 (28–32), C. soba Batuwita & Bahir, 2005 (30–34), C. subsolanus Batuwita & Bahir, 2005 (30–32), and C. wayakonei Nguyen, Kingsada, Rösler, Auer & Ziegler, 2010 (31–35).

Cyrtodactylus pageli sp. n. has 9–14 rows of dorsal tubercles and thus differs from C. mimikanus (Boulenger, 1914) (having 19–25), C. oldhami (Theobald, 1876) (19–24), C. peguensis (Boulenger, 1893) (20) and C. yangbayensis Ngo & Chan, 2010 (20-23).

Cyrtodactylus pageli sp. n. lacks a precloacal groove and thus differs from C. aurensis Grismer, 2005, and C. rubidus (Blyth, 1860). Cyrtodactylus pageli sp. n. has 19–26 subdigital lamellae under the fourth toe and thus differs from C. angularis (Smith, 1921) (having 15–16), and in part from C. elok Dring, 1979 (18–19), but both of which in addition have distinctly different dorsal pattern.

Cyrtodactylus pageli sp. n. lacks lateral body folds, which are present in C. chauquangensis Hoang, Orlov, Ananjeva, Johns, Hoang & Dau, 2007; this latter species furthermore differs amongst others by the presence of 128–130 rows of scales around the midbody (versus 113–122 midbody scales in Cyrtodactylus pageli sp. n.) and by a differing dorsal pattern.

Cyrtodactylus pageli sp. n. has five dorsal body bands and thus differs from C. hontreensis Ngo, Grismer & Grismer, 2008, which has three broad dorsal body bands.

In addition, Cyrtodactylus pageli sp. n. differs from the representatives of the subgenus Geckoella by the presence of transversally enlarged subcaudals and by having precloacal pores in both sexes: C. (Geckoella) albofasciatus (Boulenger, 1885), C. (Geckoella) collegalensis (Beddome, 1870), C. (Geckoella) deccanensis (Günther, 1864), C. (Geckoella) jeyporensis (Beddome, 1877), C. (Geckoella) nebulosus (Beddome, 1870), C. (Geckoella) triedrus (Günther, 1864), and C. (Geckoella) yakhuna (Deraniyagala, 1945).

Etymology. The new Cyrtodactylus species is named after Theodor Bernhard Pagel, Director of the Cologne Zoological Garden, in recognition of his long-term support of biodiversity research and conservation in the Indochinese Subregion of Southeast Asia, in particular in Vietnam and Laos.

Distribution. The species is currently known only from Vang Vieng District, Vientiane Province, northwestern Laos (Fig. 8).

Natural history. The type series was collected at night between 19:00–23:00 in the karst forest at elevations of 220– 300 m. Specimens were found in limestone crevices, about 1–1.5 m above the forest floor and some other geckos were observed hanging on the cliff outside of the cave entrance or hiding in a small hole at the ceiling of the high entrance hall of the cave ca. 50 m away from entrance (Fig. 9). Largest eggs were found inside the female ZFMK 91827, with maximum diameter of 2.9 mm.

comparison; * = left/right side of body, m = mean, min. = minimum, max. = maximum, s = standard deviation. continued next page