The new species can be distinguished from the remaining bent-toed geckos by a combination of the following characters: maximum SVL 65.9 mm; 16–19 dorsal tubercle rows; 41–44 ventral scal
Trang 1ZOOTAXA ISSN 1175-5326 (print edition)
ISSN 1175-5334 (online edition)
Copyright © 2014 Magnolia Press
Zootaxa 3785 (4): 518–532
www.mapress.com / zootaxa / Article
http://dx.doi.org/10.11646/zootaxa.3785.4.2 http://zoobank.org/urn:lsid:zoobank.org:pub:E03B1389-367A-485C-B16C-B6E0AE2A7286
A new Cyrtodactylus (Squamata: Gekkonidae) from Khanh Hoa Province,
southern Vietnam
1 Rheinische Friedrich-Wilhelms-Universität Bonn, Germany E-mail: schneider.nicole87@googlemail.com
2 Cologne Zoo, Riehler Straße 173, 50735 Cologne, Germany E-mail: ziegler@koelnerzoo.de
3 Dong Khoi 9A, Tam Hiep, Bien Hoa, Dong Nai Province, Vietnam; E-mail: pmytrung@yahoo.com
4 Faculty of Environmental Sciences, Hanoi University of Science, Vietnam National University, 334 Nguyen Trai Road, Hanoi, Vietnam E-mail: le.duc.minh@hus.edu.vn
5 Centre for Natural Resources and Environmental Studies, Hanoi National University, 19 Le Thanh Tong, Hanoi, Vietnam
6 Department of Herpetology, American Museum of Natural History, Central Park West at 79th Street, New York 10024-5192
7 Institute of Ecology and Biological Resources, Vietnam Academy of Science and Technology, 18 Hoang Quoc Viet Road, Hanoi, Vietnam E-mail: nqt2@yahoo.com
8 Department of Terrestrial Ecology, Zoological Institute, University of Cologne, Zülpicher
Strasse 47b, D-50674 Cologne, Germany
9 Corresponding author
Abstract
We describe a new species of the genus Cyrtodactylus from southern Vietnam, based on morphological and molecular datasets Cyrtodactylus cucdongensis sp nov is described on the basis of seven specimens collected from Cuc Dong Cape, Khanh Hoa Province The new species can be distinguished from the remaining bent-toed geckos by a combination
of the following characters: maximum SVL 65.9 mm; 16–19 dorsal tubercle rows; 41–44 ventral scales at midbody; 5 or
6 precloacal pores in males, 4–6 pitted precloacal scales in females; no femoral pores; 6–13 enlarged precloacal scales; 5–
9 enlarged femoral scales; no transversally enlarged subcaudals; dorsal pattern consisting of irregular dark bands This is the 33rd species of Cyrtodactylus known from Vietnam
Key words: Gekkonidae; Cyrtodactylus cucdongensis sp nov., new species, morphology, taxonomy, molecular phylog-eny
Introduction
In the past decades, Vietnam has shown to be a hotspot of Cyrtodactylus diversity Since 1997, when only three species had been known from the country, 29 additional species have been described from Vietnam (Nguyen et al 2013; Ziegler et al 2013; Phung et al 2014) Recent field research in southern Vietnam’s coastal region led to the discovery of another population of Cyrtodactylus from Cuc Dong Cape, Khanh Hoa Province Both morphological examination and molecular analyses revealed that this population is a member of the Cyrtodactylus irregularis species complex, the most species-rich group of bent-toed geckos According to Nazarov et al (2012) and Nguyen
et al (2013) the C irregularis group currently consists of ten species, namely C bidoupimontis Nazarov, Poyarkov, Orlov, Phung, Nguyen, Hoang & Ziegler, 2012, C bugiamapensis Nazarov, Poyarkov, Orlov, Phung, Nguyen, Hoang & Ziegler, 2012, C cattienensis Geissler, Nazarov, Orlov, Böhme, Phung, Nguyen & Ziegler,
2009, C cryptus Heidrich, Rösler, Vu, Böhme & Ziegler, 2007, C huynhi Ngo & Bauer, 2008, C irregularis (Smith, 1921), C phuocbinhensis Nguyen, Le, Tran, Orlov, Lathrop, Macculloch, Le, Jin, Nguyen, Nguyen, Hoang, Che, Murphy & Zhang, 2013, C pseudoquadrivirgatus Rösler, Vu, Nguyen, Ngo & Ziegler, 2008, C taynguyenensis Nguyen, Le, Tran, Orlov, Lathrop, Macculloch, Le, Jin, Nguyen, Nguyen, Hoang, Che, Murphy &
Trang 2Zhang, 2013, and C ziegleri Nazarov, Orlov, Nguyen & Ho, 2008 Based on morphological examination and molecular analyses, we herein describe the recently discovered population as a distinct species belonging to the taxonomically complicated C irregularis group (Nazarov et al 2012; Nguyen et al 2013)
Material and methods
Molecular data and phylogenetic analyses We included a newly collected sample of Cyrtodactylus yangbayensis Ngo & Chan, 2010 (VNMN 03373) from Nha Trang, Khanh Hoa, Vietnam and all species of the Cyrtodactylus irregularis group in our analyses, except for C irregularis since sequences of this species were not specified by Nazarov et al (2012) and Nguyen et al (2013) A member of the C pulchellus Gray, 1828 complex was used as an outgroup
We used the protocols of Le et al (2006) for DNA extraction, amplification, and sequencing A fragment of the mitochondrial gene, cytochrome c oxidase subunit 1 (COI), was amplified using the primer pair VF1-d and VR1-d (Ivanova et al 2006) After sequences were aligned by Clustal X v2 (Thompson et al 1997), data were analyzed using maximum parsimony (MP) and Bayesian analysis, as implemented in PAUP*4.0b10 (Swofford 2001) and MrBayes v3.2 (Ronquist et al 2012), respectively Settings for these analyses followed Le et al (2006), except that
evolution was set to TrN+I+G as selected by Modeltest v3.7 (Posada & Crandall 1998) The cutoff point for the burn-in function was set to 5 in the Bayesian analysis, as -lnL scores reached stationarity after 5,000 generations in both runs Nodal support was evaluated using Bootstrap replication (BP) as calculated in PAUP and posterior probability (PP) in MrBayes v3.2 Uncorrected pairwise divergences were calculated in PAUP*4.0b10 (Table 1) Morphological characters Measurements were taken with a slide-caliper to the nearest 0.1 mm Abbreviations are as follows: snout-vent length (SVL), from tip of snout to vent; tail length (TL), from vent to tip
of tail; maximum head height (HH), from occiput to underside of jaws; head length (HL), from tip of snout to posterior margin of ear; maximum head width (HW); greatest diameter of orbit (OD); snout to eye distance (SE), from tip of snout to anteriormost point of eye; eye to ear distance (EE), from posterior margin of eye to anterior margin of ear opening; internarial distance (IND), measured between inner border of nostrils; interorbital distance (IOD), measured across narrowest point between orbits; ear diameter (ED), greatest diameter of ear; axilla to groin distance (AG); forearm length (ForeaL), from the base of the palm to the elbow; shank length (SL), from the base
of heel to the knee; length of first finger (LeF1) and of fourth finger (LeF4), from the base to the tip of finger; length of first toe (LeT1) and of fourth toe (LeT4), from the base to the tip of toe; rostral width (RW); rostral height (RH); mental width (MW); mental length (ML)
Scale counts were taken as follows: supralabials (SPL); infralabials (IFL); nasal scales surrounding nare, from rostral to labial (not including rostral and labial), i.e nasorostral, supranasal, postnasals (N); postrostrals or internasals (I); scale rows between fifth supralabials across the dorsal head surface (SC5SPL); interorbitals (IO), scales between the anterior corners of the eyes; spinous ciliaries (CS); postmentals, i.e scales bordering mental shield, except infralabials (PM); gulars bordering the postmentals (GP); granular scales surrounding dorsal midbody tubercles (GSDT); dorsal tubercle rows at midbody (DTR); ventral scales in longitudinal rows at midbody (V); number of scales along the midbody from mental shield to anterior edge of cloaca (SMC); scales around midbody (MS); precloacal pores (PP); femoral pores (FP); total number of postcloacal tubercles (PAT); subdigital lamellae under the first finger (LF1); subdigital lamellae under the fourth finger (LF4); subdigital lamellae under the first toe (LT1); subdigital lamellae under the fourth toe (LT4) All bilateral scale counts were given as left/right
Specimens were deposited in the collections of the Institute of Ecology and Biological Resources (IEBR), Hanoi, Vietnam; the Vietnam National Museum of Nature (VNMN), Hanoi, Vietnam; and the Zoologisches Forschungsmuseum Alexander Koenig (ZFMK), Bonn, Germany
Trang 4Molecular phylogeny The combined matrix contained 657 aligned characters MP analysis of the dataset recovered eight most parsimonious trees with 1007 steps (CI = 0.44; RI = 0.66) The Bayesian topology is similar
to the one produced by the MP analysis in the shallow nodes, except for the sister relationships between C kingsadai Ziegler, Phung, Le & Nguyen, 2013 and C cryptus, C pseudoquadrivirgatus and C taynguyenensis, and
C pubisulcus Inger, 1957 and C quadrivirgatus Taylor, 1962 However, many basal nodes are either poorly corroborated or unresolved in both analyses (Fig 1) The new species is strongly supported as a sister taxon to C yangbayensis (BP = 97, PP = 100) It is also significantly divergent from others in terms of genetic distance with a minimum pairwise divergence of approximately 9% in the mitochondrial fragment of COI (Table 1)
FIGURE 1 Bayesian consensus tree based on the partial COI gene Number above and below branches are Bayesian posterior probabilities and bootstrap values (>50%), respectively Cyrtodactylus pulchellus = C pulchellus sensu lato (before the revision of Grismer et al 2012) The bracket and number indicate the sister relationship between two taxa and the bootstrap value supported by the MP analysis, respectively
Cyrtodactylus cucdongensis sp nov
(Figs 2, 3)
Holotype IEBR A.2013.104, adult male, collected by T M Phung on 12 June, 2011, from Cuc Dong Cape, Ninh Hoa District, Khanh Hoa Province, Vietnam
Paratypes VNMN A.2013.18, adult male, ZFMK 95513, adult female, ZFMK 95514 subadult male and ZFMK 95515, subadult female, collected on 12 June, 2011, IEBR A.2013.105, adult female and VNMN A.2013.19, adult female, collected on 2 September, 2011, the same data as the holotype
Trang 5Diagnosis Cyrtodactylus cucdongensis sp nov is distinguished from the remaining bent-toed geckos by a combination of the following characters: maximum SVL 65.9 mm; 16–19 dorsal tubercle rows; 41–44 ventral scales at midbody; 5 or 6 precloacal pores in males, 4–6 pitted precloacal scales in females; no femoral pores; 6–13 enlarged precloacal scales; 5–9 enlarged femoral scales; no transversally enlarged subcaudals; dorsal pattern consists of irregular dark bands
Description of the holotype Adult male with a total length of 147.2 mm (SVL 65.9 mm, tail broken, TL 81.3 mm); rostral Y-shaped, wider than high (RW 2.3 mm, RH 1.7 mm, RW/RH 1.35), medially with a straight, vertical rostral suture, in contact with nasorostral, nare and first supralabial on each side, medially in contact with internasal; mental wider than rostral (MW 2.9 mm); supralabials 9; infralabials 8; supralabials separated from orbit
by 3 or 4 rows of granular scales; nares in contact with rostral, nasorostral, supranasal, two postnasals, and first supralabial; internasal single; snout scales larger than head scales; scales between fifth supralabials 53; scales between anterior corner of eyes 57; interorbital region with small round, convex scales; scales in postorbital region distinctly smaller (ca half the size) than snout scales, posteriorly increasing in size, irregular in shape; postorbital region with enlarged tubercles in 5 or 6 rows on each side; pupil vertical; spinous ciliaria 9/10, posterior ones more developed; ear opening vertical, oval; mental triangular, in contact with two postmentals and the first infralabial on each side; postmentals surrounded by four granular scales, of which two outer ones distinctly enlarged, and first infralabial on each side; gular scales granular
FIGURE 2 Male holotype (IEBR A.2013.104) of Cyrtodactylus cucdongensis sp nov Photos T M Phung
Dorsal scales small, twice the size of head scales between the eyes; dorsal tubercles in 18 longitudinal rows at midbody; dorsal surface of body with tubercles; dorsal tubercles surrounded by 9–12 dorsal scales, separated from each other by 2–4 scales; lateral body folds slightly developed, without tubercles; ventral scales round, slightly
Trang 6arched, imbricated, 3 or 4 times larger than gular and throat scales, three times larger than dorsals; ventral scales in
42 rows; midbody scales rows 116; scales between mental and cloacal slit 183; dorsal surface of limbs with tubercles, more developed on hind limbs; enlarged femoral scales 6 on each side, two series of three enlarged scales separated by two granular scales on right hind limb, outer enlarged femoral scale separated by six granular scales from other enlarged femoral scales on left hind limb; fingers and toes free of webbing; relative finger lengths I < II
< V < IV < III, relative toe lengths I < II < III < V < IV; claw sheathed by two scales; subdigital lamellae: finger I
10 or 11 (including 3 basally broadened lamellae), finger II 13 (4), finger III 15 (4), finger IV 17 (4–5), finger V 12
or 13 (3–4), toe I 10 or 11 (3–4), toe II 14 or 15 (4), toe III 17 (5), toe IV 18–20 (7), toe V 17–18 (6); precloacal depression absent; precloacal pores 6; pore-bearing scales posteriorly surrounded by nine enlarged scales, arranged
in a diamond shape; adjoining scales continuously decreasing in size; two postcloacal tubercles on each side, well developed; hemipenis not everted; original tail without distinct whorls; dorsal surface covered by 14 rows of tubercles, each row with 4 or 5 tubercles, tubercles absent in distal part of tail; median row of subcaudals not transversally enlarged
FIGURE 3 A) Adult male paratype (VNMN A.2013.18) and B) adult female paratype (ZFMK 95513) of Cyrtodactylus cucdongensis sp nov Photos T M Phung
Coloration in life: Top of head light brown; each side with a dark stripe, from between tip of snout and eye, running below the eyes and connecting with the other band at the neck, stripe on right side not clear, blurred
Trang 7between eye and neck; irregular dark blotches on dorsal surface of the head indistinct; eyelids with green cast; ciliaria bright yellow; iris marbled in black and metallic-yellow; dorsal surface of body light brown with irregular transverse dark brown bands; dark bands between limb insertions four, two inner ones in irregular shape, consisting
of blotches and small bands; a median dark spot just behind the neck band; five or six blotches in one row on each side from neck to hind limbs; flanks without dark bands or blotches; dorsal pattern of limbs consisting of a mixture
of light and dark brown; dorsal tail with seven dark brown bands or broad blotches on a light brown ground; ventral surface of body solid light beige, with ventral side of tail being slightly darker
Color in preservative (70 % ethanol): The overall color scheme is somewhat less pronounced, slightly fades in alcohol Main characteristics are still clearly visible, the slight green cast of the eyelids and the bright yellow color
of the ciliaria are not visible, but have the same bright 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 color pattern variation see Tables 2, 3 and Fig 3 ZFMK 95515 has, in contrast to the holotype and the other paratypes, distinct whorls on the dorsal surface of tail Transverse body bands individually vary in intensity (darker bands versus lighter bands) and shape (continuously developed body bands versus interrupted or irregularly shaped body bands) Sexual dimorphism was also discernible in terms of hemipenial swellings at the under tail base in adult males, the enlarged precloacal scales are distinctly larger in males (even in the subadult male) and the precloacal pores in females are indistinct, like pitted scales
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; Darevsky & Szczerbak 1997; Bauer 2002, 2003; Bauer et al 2002, 2003, 2009; David et al 2004, 2011; Pauwels et al 2004, 2013, 2014; Nguyen S.N et al 2006, 2013; Hoang et al 2007; Orlov et al 2007; Nazarov et al 2008; Ngo 2008; Ngo & Bauer 2008; Rösler & Glaw 2008; Rösler et al 2008; Geissler et al 2009; Mahony 2009; Ngo & Chan 2010; Ngo & Pauwels 2010; Ngo et al 2008, 2010, 2011; Nguyen T.Q et al 2010; Sumontha et al 2010; Schneider et al 2011; Grismer et al 2012; Nazarov et al 2012; Ziegler et al 2010, 2013; Luu et al 2014; Pauwels & Sumontha 2014) Cyrtodactylus cucdongensis sp nov can be distinguished from all the Vietnamese congeners as follows: The new species has no transversally enlarged subcaudal scales and thus differs from the following species: C badenensis Nguyen, Orlov & Darevsky, 2006, C bichnganae Ngo & Grismer, 2010, C caovansungi Orlov, Nguyen, Nazarov, Ananjeva & Nguyen, 2007, C chauquangensis Hoang, Orlov, Ananjeva, Johns, Hoang & Dau,
2007, C condorensis (Smith, 1920), C cucphuongensis Ngo & Chan, 2011, C eisenmanae Ngo, 2008, C grismeri Ngo, 2008, C hontreensis Ngo, Grismer & Grismer, 2008, C huongsonensis Luu, Nguyen, Do & Ziegler, 2011, C intermedius (Smith, 1917), C kingsadai, C martini Ngo, 2011, C nigriocularis Nguyen, Orlov & Darevsky, 2006,
C paradoxus (Darevsky & Szczerbak, 1997), C phongnhakebangensis Ziegler, Rösler, Herrmann & Vu, 2002, C phuquocensis Ngo, Grismer & Grismer, 2010, C roesleri Ziegler, Nazarov, Orlov, Nguyen, Vu, Dang, Dinh & Schmitz, 2010, C takouensis Ngo & Bauer, 2008, C thochuensis Ngo & Grismer, 2012, and C yangbayensis The following species have femoral pores, which are lacking in Cyrtodactylus cucdongensis sp nov.: C huynhi (3–8) and C dati Ngo, 2013 (3–4) Cyrtodactylus cucdongensis sp nov has enlarged femoral scales which are absent in
C cryptus Cyrtodactylus cucdongensis sp nov can be distinguished from C thuongae Phung, van Schingen, Ziegler & Nguyen, 2014 by having more precloacal pores in males (5–6 vs 0–1)
From the representatives of the C irregularis complex, Cyrtodactylus cucdongensis sp nov differs as follows (only characters are mentioned, which are not listed in Table 4): from C bidoupimontis by lacking a dark neckband and tail bands which are distinctly broader; from C bugiamapensis by having fewer dorsal tubercle rows (16–19
vs 20–24), additionally, the dorsal pattern of Cyrtodactylus cucdongensis sp nov consists of irregular dark bands
vs transverse bands formed by dark spots in C bugiamapensis; from C cattienensis by having more enlarged precloacal scales in most specimens (8–21 vs 6–13); from C irregularis by lacking a thickened tail base that shows very large triple-edged knobs and forms 3–5 pronounced semi-rings of tail segments, additionally, the dorsal pattern of Cyrtodactylus cucdongensis sp nov differs from the pattern of C irregularis (banded vs blotched); from C phuocbinhensis by having irregular dark bands vs two dark brown stripes or blotches on dorsum; from C pseudoquadrivirgatus by the absence of enlarged lateral tubercles (vs present), the presence of enlarged femoral scales (vs absent), and by having uniformly bright colored limbs versus striped or mottled limbs in C pseudoquadrivirgatus; from C taynguyenensis by the presence of enlarged femoral scales (vs absent) and by having irregular dark bands (vs irregular blotches bordered by light brown edges); from C ziegleri by having fewer dorsal tubercle rows (16–19 vs 20–24)
Trang 10FIGURE 4 Map showing the type locality (red circle) of Cyrtodactylus cucdongensis in Khanh Hoa Province, southern Vietnam