A new species of Cyrtodactylus (Squamata: Gekkonidae) from Ninh Binh Province, Vietnam Author Le, Dzung Trung Author Nguyen, Truong Quang Author Le, Minh Duc Author Ziegler, Thomas text Zootaxa 2016 4162 2 268 282 journal article 10.11646/zootaxa.4162.2.4 9340386c-6af9-4d62-8603-0b1d670db330 1175-5326 263897 A6C2B048-C5F3-4B55-B4D7-F60855E2836E Cyrtodactylus soni sp. nov. ( Figs. 2–4 ) Holotype . HNUE VL .2015.78, adult male, collected on 24 July 2015 by D. T. Le , A. M. Luong , D. T. Pham , and N. H. Nguyen , in the karst forest near Da Han Village ( 20o25.067’N , 105o51.467’E , elevation 17 m a.s.l. ), Gia Hoa Commune , within Van Long Wetland Nature Reserve , Gia Vien District , Ninh Binh Province , Vietnam . Paratypes. IEBR R.2016.4, adult female, HNUE VL.2015.94, adult female, collected on 24 July 2015 ; IEBR R.2016.5, adult male, HNUE VL. 2015.131, adult female, and HNUE VL .2015.132, adult female, collected on 18 August 2015 , the same data as the holotype . Diagnosis. The new species can be distinguished from other members of the genus Cyrtodactylus from Indochina by a combination of the following characters: medium size (SVL up to 103 mm ); internasal single; dorsal tubercles in 10–13 irregular rows; ventral scale rows 41–45; lateral skin folds present, without interspersed tubercles; precloacal pores 6 or 7 in males, 7 or 8 pitted scales in females, in a continuous row; femoral pores 6–8 on each side in males, separated by 8–11 poreless scales from precloacal pore series; enlarged femoral scales present; postcloacal spurs 2 or 3; subcaudal scales transversely enlarged; lamellae under toe IV 18–22 ; dorsal pattern consisting of a dark nuchal loop, a continuous or partly interrupted neck band, and five or six in part irregular transverse body bands between limb insertions. FIGURE 1. Phylogram based on the Bayesian analysis. Number above and below branches are MP/ML bootstrap values and Bayesian posterior probabilities (>50%), respectively. Asterisk represents 100% value. Description of holotype . Adult male, snout-vent length (SVL) 89.5 mm ; body elongate (AG/SVL 0.44); head distinct from neck, elongate, depressed (HL/SVL 0.28, HW/HL 0.71, HH/HL 0.44); loreal region concave; snout long (SE/HL 0.44), round anteriorly, longer than diameter of orbit (OD/SE 0.54); snout scales small, round, granular; eye large (OD/HL 0.24), pupils vertical, spinous ciliaries 32; ear oval shaped, small (ED/HL 0.08); rostral wider than high with a medial suture, bordered by first supralabial, nostril, and supranasal on each side; supranasals separated from each other by two small scales; nares round, surrounded by supranasal, rostral, first supralabial, and three postnasals; mental triangular, slightly wider than rostral; postmentals two, enlarged, in broad contact posteriorly, bordered by mental anteriorly, first infralabial laterally, and an enlarged chin scale posteriorly; supralabials 10/10; infralabials 9/9. Dorsal scales granular; dorsal tubercles round, conical, present on occipital region and back, each surrounded by 9 or 10 granular scales; ventral scales smooth, medial scales 2 or 3 times larger than dorsal scales, round, subimbricate, largest posteriorly, in 41 longitudinal rows at midbody; lateral skin folds distinct without tubercles; gular region with homogeneous smooth scales; 186 ventral scales between mental and cloacal slit; precloacal groove absent; enlarged femoral scales present; precloacal pores 6, femoral pores 6 on each side, in distal scale portion of enlarged femoral scales, separated by 8 poreless scales from precloacal pore series. Fore and hind limbs moderately slender (ForeaL/SVL 0.17, CrusL/SVL 0.2); forelimbs dorsally covered by few slightly developed tubercles; hind limb dorsally covered by distinctly developed tubercles; fingers and toes without distinct webbing; each claw bordered by two scales; subdigital lamellae: finger I 11 (including 2 basally broadened lamellae), finger II 14 (4), finger III 15 (4), finger IV 16 (4), finger V 16 (5), toe I 11 (2), toe II 14 (4), toe III 17 (5), toe IV 18 (6), toe V 19 (7). Tail regenerated (TaL 70.6 mm ); postcloacal spurs 2/2; dorsal tail base with tubercles; subcaudals distinctly enlarged. Coloration in life. Ground color light brownish grey; dorsal surface of head with irregular dark markings, largest at occiput; a dark postocular streak, edged in yellowish white, in contact with nuchal loop; neck with a dark transverse band; dorsum with five distinct dark transverse bands between limb insertions, edged by yellowish tubercles; upper surfaces of limbs with dark bands and reticulations; dorsal surface of the regenerated tail with indistinct transverse bands; gular region cream with indistinct grey marbling; venter cream; ventral surface of the regenerated tail dark grey with yellow marbling. Morphological comparisons. We compared the new species with its congeners from Vietnam and neighbouring countries in mainland Indochina, including Laos , Cambodia , and Thailand based on the examination of specimens (see Appendix) and data obtained from the literature ( Bauer et al. 2002 , 2003 , 2010 ; David et al. 2004 , 2011 ; Geissler et al. 2009 ; Hoang et al. 2007 ; Kunya et al. 2014 , 2015; Luu et al. 2011 , 2014 , 2015 , 2016a , 2016b , 2016c ; Nazarov et al. 2008 , 2012 , 2014 ; Ngo 2011, 2013; Ngo & Grismer 2010 ; Ngo & Chan 2011 ; Nguyen et al. 2010 , 2015; Nguyen et al . 2013 , 2014; Panitvong et al. 2014; Pauwels & Sumontha 2014; Pauwels et al. 2013 , 2014a , 2014b , 2016 ; Phung et al. 2014; Schneider et al. 2011, 2014 a, 2014b; Smith 1921; Sumontha et al. 2015; Ziegler et al. 2010 , 2013). Cyrtodactylus soni sp. nov. has distinctly enlarged subcaudals, which are only slightly or not enlarged in the following species: C. bidoupimontis Nazarov, Poyarkov, Orlov, Phung, Nguyen, Hoang & Ziegler , C. buchardi David, Teyni & Ohler , C. bugiamapensis Nazarov, Poyarkov, Orlov, Phung, Nguyen, Hoang & Ziegler , C. cattienensis Geissler, Nazarov, Orlov, Böhme, Phung, Nguyen & Ziegler , C. cryptus Heidrich, Rösler, Vu, Böhme & Ziegler , C. cucdongensis Schneider, Phung, Le, Nguyen & Ziegler , C. huynhi Ngo & Bauer , C. irregularis (Smith) , C. otai Nguyen, Le, Pham, Ngo, Hoang, Pham & Ziegler , C. phuocbinhensis Nguyen, Le, Tran, Orlov, Lathrop, Macculloch, Le, Jin, Nguyen, Nguyen, Hoang, Che, Murphy & Zhang , C. pseudoquadrivirgatus Rösler, Nguyen, Vu, Ngo & Ziegler , C. quadrivirgatus Taylor , C. ranongensis Sumontha, Pauwels, Panitvong, Kunya & Grismer , C. taynguyenensis Nguyen, Le, Tran, Orlov, Lathrop, Macculloch, Le, Jin, Nguyen, Nguyen, Hoang, Che, Murphy & Zhang , C. thuongae Phung, van Schingen, Ziegler & Nguyen , C. vilaphongi Schneider, Nguyen, Le, Nophaseud, Bonkowski & Ziegler , and C. ziegleri Nazarov, Orlov, Nguyen & Ho. Cyrtodactylus soni sp. nov. has enlarged femoral scales which are absent in C. angularis (Smith) , C. badenensis Nguyen, Orlov & Darevsky , C. bobrovi Nguyen, Le, Pham, Ngo, Hoang, Pham & Ziegler , C. buchardi , C. chauquangensis Hoang, Orlov, Ananjeva, Johns, Hoang & Dau , C. cryptus , C. grismeri Ngo , C. nigriocularis Nguyen, Orlov & Darevsky , C. otai , C. pageli Schneider, Nguyen, Schmitz, Kingsada, Auer & Ziegler , C. pseudoquadrivirgatus , C. sumonthai Bauer, Pauwels & Chanhome , C. taynguyenensis , C. vilaphongi , and C. wayakonei Nguyen, Kingsada, Rösler, Auer & Ziegler. Cyrtodactylus soni sp. nov. has more femoral and precloacal pores in males (6–7+6–7+6–8) than the following species: C. angularis (3), C. condorensis (Smith) (0–4), C. kunyai Pauwels, Sumontha, Keeratikiat & Phanamphon (5+3+6), C. martini (4), C. nigriocularis (0–2), C. oldhami (1–4), C. pageli (4), C. phetchaburiensis Pauwels, Sumontha & Bauer (0+5+0), C. quadrivirgatus (4), C. sumonthai (2), C. sanook Pauwels, Sumontha, Latinne & Grismer (3 or 4), C. sayiok Panitvong, Sumontha, Tunprasert & Pauwels (0+5+0), C. takouensis Ngo & Bauer (3 or 4), and absent in C. badenensis , C. cucphuongensis , C. eisenmanae Ngo , C. grismeri , and C. ranongensis . Cyrtodactylus soni sp. nov. has more femoral and precloacal pores (or pitted scales) in females (6–8+7–8+5–6) than the following species: C. angularis (3), C. cucdongensis (4–6), C. dumnuii Bauer, Kunya, Sumontha, Niyomwan, Pauwels, Chanhome & Kunya (0+0–7+0), C. irregularis (0–6), C. pageli (4), C. quadrivirgatus (4); femoral and precloacal pores (or pitted scales) are absent in the females of C. auribalteatus Sumontha, Panitvong & Deein , C. badenensis , C. bidoupimontis , C. bobrovi , C. buchardi , C. caovansungi Orlov, Nguyen, Nazarov, Ananjeva & Nguyen , C. cattienensis , C. condorensis , C. cryptus , C. eisenmanae , C. grismeri , C. interdigitalis Ulber , C. jarujini Ulber , C. martini , C. multiporus Nazarov, Poyarkov, Orlov, Nguyen, Milto, Martynov, Konstantinov & Chulisov , C. nigriocularis , C. otai , C. phuocbinhensis , C. sanook , C. soudthichaki Luu, Calame, Nguyen, Bonkowski & Ziegler , C. spelaeus Nazarov, Poyarkov, Orlov, Nguyen, Milto, Martynov, Konstantinov & Chulisov , C. sumonthai , C. takouensis , C. taynguyenensis , C. thuongae , C. thirakhupti , C. vilaphongi , and C. yangbayensis Ngo & Chan. FIGURE 2. The male holotype (HNUE VL.2015.78) of Cyrtodactylus soni sp. nov. in life (A: Dorsal view, B: Ventral view). Photo D. T. Le. FIGURE 3 . Cloacal (A) and femoral (B) regions of the holotype (HNUE VL.2015.78) of Cyrtodactylus soni sp. nov. in preservative. Photo D. T. Le. FIGURE 4. Female paratype (IEBR R.2016.4) of Cyrtodactylus soni sp. nov. in life: A) Dorsal view and B) Ventral view. Photo D. T. Le. TABLE 2. Measurements (in mm) and scalation of Cyrtodactylus soni sp. nov. from Ninh Binh Province, Vietnam (M = male, F = female, * = regenerated or broken tail, min = minimum, max = maximum, other abbreviations defined in the text).

HNUE VL.2015.78	IEBR IEBR A.2016.5 A.2016.4	HNUE VL.2015.94	HNUE VL.2015.131	HNUE VL.2015.132	Min–Max (n=6)
Sex	M	M F	F	F	F
SVL	89.5	89.2 103	88.7	89.5	98.5	88.7–103
TaL	70.6*	82.8* 79.8*	107.8	112	113	70.6–113
HL	25.2	25.1 27.6	25.5	24.6	28	24.6–28
HW	17.8	18.7 19	17.3	17.7	19.3	17.3–19.3
HH	11.2	11 11	10	10	11.1	10–11.2
OD	6.1	6.4 6	5.7	5.3	6.3	5.3–6.4
SE	11.2	10.7 11.6	10.9	10	11.7	10–11.7
EE	8.3	7 8.6	7.9	7.5	8.4	7–8.6
NE	8.7	8.4 8.7	8.8	8.3	9.1	8.3–9.1
ED	1.9	1.8 2.1	2.2	2	1.9	1.8–2.2
ForeaL	15	13.8 15.6	14.8	13.9	16.3	13.8–16.3
CrusL	17.5	17.2 18.8	17.2	16.7	19.5	16.7–19.5
AG	39.8	36 47.4	37.9	38.8	42.2	36–47.4
BW	17	16.2 18	16.3	16.8	19	16.2–19
IND	3.8	3.2 3.5	2.9	3.2	3.3	2.9–3.8
IOD	6.8	7.4 6	7.4	6.7	7.5	6–7.5
SPL	10/10	11/10 13/13	12/12	12/11	11/11	10–13
IFL	9/9	10/9 9/10	10/10	9/10	9/9	9–10
N	5/5	5/4 4/4	4/4	4/4	3/3	3–5
IN	1	1 1	1	1	1	1
CIL	3 2	3 2 3 3	3 4	3 1	3 3	3 1–3 4
PM	2	2 2	2	2	2	2
GST	9 or 10	9–11 9 or 10	9–11	9–10	9–11	9–11
V	4 1	4 2 4 5	4 5	4 4	4 3	4 1–4 5
SLB	186	188 203	202	209	212	186–212
FP	6/6	7/8 6/6	8/6	6/5	8/6	5–8
PP	6	7 7 (pitted scales)	8 (pitted scales)	7 (pitted scales)	7 (pitted scales)	6–8
PAT	2/2	3/3 3/2	2/2	2/3	2/3	2–3
TubR	10	13 10	10	13	11	10–13
EFS	8/8	8/9 8/8	9/9	11/11	9/10	8–11
NSF
I	2+9	3+10 3+10	3+8	2+10	2+10	11–13
II	4+10	5+12 5+12	5+9	4+13	5+11	14–17
III	4+1 1	5+1 3 5+1 3	5+1 2	5+1 4	5+1 2	1 5–1 9
IV	4+12	5+14 5+10	5+10	5+14	6+12	15–19
V	5+11	4+11 6+12	4+11	4+12	4+11	15–18
NST
I	2+9	2+9 3+1 2	2+1 0	2+1 0	2+1 0	1 1–1 5
II	4+10	5+13 5+12	5+12	5+13	5+12	14–18
III	5+1 2	6+1 4 7+1 4	5+1 4	5+1 5	7+1 2	1 9–2 1
IV	6+12	7+13 9+13	9+12	7+15	7+13	18–22
V	7+1 2	7+1 4 7+1 5	8+1 4	7+1 6	7+1 4	1 9–2 3

FIGURE 5. Map showing the type locality of Cyrtodactylus soni sp. nov. (red circle) in Van Long Wetland Nature Reserve, Ninh Binh Province, Vietnam. Cyrtodactylus soni sp. nov. differs from the following species by having fewer femoral and precloacal pores in males (6–7+6–7+6–8): C. astrum Grismer, Wood, Quah, Anuar, Muin, Sumontha, Ahmad, Bauer, Wangkulangkul, Grismer & Pauwels (31–38), C. bansocensis Luu, Nguyen, Le, Bonkowski & Ziegler (34), C. calamei Luu, Bonkowski, Nguyen, Le, Schneider, Ngo & Ziegler (35–39), C. chanhomeae Bauer, Sumontha & Pauwels (32), C. darevskii Nazarov, Poyarkov, Orlov, Nguyen, Milto, Martynov, Konstantinov & Chulisov (38–44), C. erythrops Bauer, Kunya, Sumontha, Niyomwan, Panitvong, Pauwels, Chanhome & Kunya (10+9+9), C. hinnamnoensis Luu, Bonkowski, Nguyen, Le, Schneider, Ngo & Ziegler (36–44), C. jaegeri Luu, Calame, Bonkowski, Nguyen & Ziegler (44), C. jarujini (52–54), C. khammouanensis Nazarov, Poyarkov, Orlov, Nguyen, Milto, Martynov, Konstantinov & Chulisov (40–44), C. lekaguli Grismer, Wood, Quah, Anuar, Muin, Sumontha, Ahmad, Bauer, Wangkulangkul, Grismer & Pauwels (31–41), C. lomyenensis Ngo & Pauwels (39–40), C. multiporus (58–60), C. phongnhakebangensis Ziegler, Rösler, Herrmann & Vu (32–42), C. rufford Luu, Calame, Nguyen, Le, Bonkowski & Ziegler (42–43), C. sommerladi Luu, Bonkowski, Nguyen, Le, Schneider, Ngo & Ziegler (20–26), and C. soudthichaki (29). Cyrtodactylus soni sp. nov. has a dorsum with banded color pattern, which is blotched in C. brevipalmatus (Smith) , C. buchardi , C. bugiamapensis , C. erythrops , C. irregularis , C. jarujini , C. phetchaburiensis , C. phuocbinhensis , C. pseudoquadrivirgatus , C. taynguyenensis , C. teyniei David, Nguyen, Schneider & Ziegler , and C. thuongae ; the dorsum is uniformly brown in C. nigriocularis and striped in C. oldhami , C. quadrivirgatus , and C. ranongensis . Cyrtodactylus soni sp. nov. differs from C. kingsadai Ziegler, Phung, Le & Nguyen by having fewer dorsal tubercle rows (10–13 vs. 17–23 in C. kingsadai ). Cyrtodactylus soni sp. nov. differs from the following species by having more ventral scale rows (41–45): C. badenensis (25–29), C. bichnganae (30–31), C. buchardi (30), C. chanhomeae (36–38), C. chauquangensis (36– 38), C. doisuthep Kunya, Panmongkol, Pauwels, Sumontha, Meewasana, Bunkhwamdi & Dangsri (29–35), C. erythrops (28), C. grismeri (33–38), C. inthanon Kunya, Sumontha, Panitvong, Dongkumfu, Sirisamphan & Pauwels (29–34), C. jaegeri (31–32), C. jarujini (32–38), C. khammouanensis (32–38), C. khelangensis Pauwels, Sumontha, Panitvong & Varaguttanonda (32–35), C. lomyenensis (35–36), C. multiporus (30–38), C. oldhami (34– 38), C. puhuensis Nguyen, Yang, Le, Nguyen, Orlov, Hoang, Nguyen, Jin, Rao, Hoang, Che, Murphy & Zhang (36), C. rufford (27–29), C. sanook (27–28), C. soudthichaki (32–33), C. spelaeus (36–39), C. sumonthai (33–36), C. teyniei (38), C. tigroides (34), C. vilaphongi (34–36), C. wayakonei (31–35), and C. ziegleri (33–39). FIGURE 6. Habitat of Cyrtodactylus soni sp. nov. in Van Long Wetland Nature Reserve, Ninh Binh Province, Vietnam. Photo: D. T. Le. Morphologically, Cyrtodactylus soni sp. nov. is most similar to C. huongsonensis but it differs from the latter by having fewer femoral and precloacal pores (6–7+6–7+ 6–8 in males vs. 10+6+ 7 in C. huongsonensis ) and more spinous ciliaries (31–33 vs. 26–28 in C. huongsonensis ). Sexual dimorphism and variation. The females differ from the males in the absence of hemipenial swellings at the tail base. Three female paratypes have complete tails with distinctly enlarged subcaudals. For other morphological characters see Table 2 . Distribution. Cyrtodactylus soni sp. nov. is currently known only from the type locality, Van Long Wetland Nature Reserve, in Ninh Binh Province , Vietnam ( Fig. 5 ). Ecological notes. Specimens were found at night between 18:30 and 21:00, on limestone cliffs and in rock crevices, approximately 0.3–1.5 m above the ground, at elevations between 17–28 m a.s.l. The surrounding habitat consisted of secondary karst forest of medium and small hardwoods mixed with shrubs and vines ( Fig. 6 ). Air temperature ranged from 25 to 32.1o C and relative humidity was 70–90%. Etymology. We name this new species in honor of our colleague and friend, Assoc. Prof. Dr. Son Lan Hung Nguyen, Faculty of Biology, Hanoi National University of Education, Vietnam , in recognition of his support of our research and conservation work in Vietnam . As common names we suggest Son’s Bent-toed Gecko (English) and Thạch sùng ngón sƠn (Vietnamese).