Cyrtodactylus rufford, Luu, Vinh Quang, Calame, Thomas, Nguyen, Truong Quang, Le, Minh Duc, Bonkowski, Michael & Ziegler, Thomas, 2016

Luu, Vinh Quang, Calame, Thomas, Nguyen, Truong Quang, Le, Minh Duc, Bonkowski, Michael & Ziegler, Thomas, 2016, Cyrtodactylus rufford, a new cave-dwelling bent-toed gecko (Squamata: Gekkonidae) from Khammouane Province, central Laos, Zootaxa 4067 (2), pp. 185-199 : 188-196

Cyrtodactylus rufford sp. nov.

( Figs. 2–3 )

Holotype. VFU R.2015.14, adult male, on a karst cliff, near the entrance of Nang Log cave (17°30.282’N, 105°23.107’E, elevation 160 m a.s.l.), Gnommalath District, Khammouane Province, central Laos, was collected on 29 March 2015 by V. Q. Luu, T. Calame and K. Thanabuaosy.

Paratypes. IEBR R.2015.35, adult male, and NUOL R.2015.15, subadult male; same collection data as was provided for the holotype.

Diagnosis. The new species differs from other congeners of the genus Cyrtodactylus by the following combination of characters: medium size, SVL reaching 72.5 mm; dorsal pattern consisting of three or four light transverse bands between limb insertions; one intersupranasal; 14–16 irregular dorsal tubercle rows at midbody, weakly developed in the paravertebral region; 27–29 ventral scale rows between ventrolateral folds; 42–43 precloacal and femoral pores in a continuous row in males, enlarged femoral and precloacal scales present; 4 or 5 postcloacal tubercles on each side; dorsal tubercles present at base of tail; subcaudal scales medially enlarged.

Description of the holotype. Adult male, SVL 68.3 mm; body slender, elongate (TrunkL/SVL ratio 0.47); head elongate (HL/SVL ratio 0.29), relatively narrow (HW/HL ratio 0.57), depressed (HH/HL ratio 0.36), distinct from neck; loreal region inflated, posterior nasal region concave; snout long (SE/HL ratio 0.47), obtuse, longer than orbit diameter (OD/SE ratio 0.57); scales on snout small, rounded, homogeneous, granular, larger than those on frontal and parietal regions; eye large (OD/HL ratio 0.27), pupils vertical; supraciliaries with tiny spines posteriorly; ear opening oval, small (EarL/HL ratio 0.10); rostral wider than high with a shallow inverted Ϙ-shaped medial suture; supranasals separated from each other by one small scale; rostral in contact with first supralabial and nostril on each side; nostrils oval, each surrounded by supranasal, rostral, first supralabial, and two enlarged postnasals; mental triangular, as wide as rostral (RW 2.8, MW 2.7); one pair of enlarged postmentals, in broad contact posteriorly, bordered laterally by first infralabials and triangular mental anteriorly; supralabials 11/12; infralabials 10/11. Dorsal scales granular to weakly conical; intermixed with conical, round dorsal tubercles (2–3 times larger than adjoining scales) present from occipital region to dorsum and base of tail, tubercles in 16 rows at midbody, larger on flanks, each surrounded by 9 granular scales; ventrals larger than lateral scales, smooth, round, largest posteriorly, in 28 longitudinal scale rows between ventrolateral folds; gular region with homogeneous, smooth scales; ventral scales from mental to cloacal slit 153; precloacal groove lacking; enlarged femoral and cloacal scales present; femoral and precloacal pores 43, in a continuous row.

Fore and hind limbs moderately slender (ForeL/SVL ratio 0.17, CrusL/SVL 0.20); dorsum of forelimbs with weakly developed tubercles; hind limbs dorsally with distinct tubercles; interdigital webbing slightly developed; lamellae under fourth finger 19/19; lamellae under fourth toe 19/18.

Tail partly regenerated, longer than SVL (TaL 94.5 mm, TaL/SVL ratio 1.38); 5/4 postcloacal tubercles; tubercles present at base of tail dorsum; subcaudals transversely enlarged in the middle, flat, smooth.

Coloration in life. Head dorsally greyish brown with dark blotches; nuchal loop brown, uninterrupted, in U– shape; labials grey; three light transverse bands between limb insertions, edged in black anteriorly, somewhat irregular in the posterior parts of the dorsal bands; tubercles at midbody brown; tubercles on body bands grey; dorsal surface of fore- and hind- limbs with dark reticulated markings; dorsal surface of tail brown with light rings (2–3 times narrower than dark bands), ventral surface of head, body and limbs greyish cream; venter of tail grey.

Variation: The adult male paratype (IEBR R.2015.35) has wider light bands on the midbody with dark spots within each band compared to those of the holotype and subadult male paratype (NUOL R.2015.15). Further morphological characters of the paratypes are provided in Table 3 .

Comparisons. We compared Cyrtodactylus rufford sp. nov. to other species of Cyrtodactylus from Laos and neighbouring countries in the mainland Indochina region, including Vietnam, Cambodia, and Thailand based on examination of specimens (see Appendix) and data provided from taxonomic publications ( Luu et al. 2014; Nazarov et al. 2014; Nguyen et al. 2014; Panitvong et al. 2014; Pauwels et al. 2014; Pauwels & Sumontha 2014; Schneider et al. 2014; Sumontha et al. 2015; Nguyen et al. 2015; Luu et al. 2015) (see Table 4).

Cyrtodactylus rufford sp. nov. has distinctly enlarged median subcaudals and thus differs from the following species which are lacking enlarged median subcaudals: C. bidoupimontis Nazarov, Poyarkov, Orlov, Phung, Nguyen, Hoang & Ziegler , C. bobrovi Nguyen, Le, Pham, Ngo, Hoang, Pham & Ziegler , C. buchardi David, Teynié & Ohler , C. bugiamapensis Nazarov, Poyarkov, Orlov, Phung, Nguyen, Hoang & Ziegler , C. cattienensis Geissler, Nazarov, Orlov, Böhme, Phung, Nguyen & Ziegler , C. cucdongensis Schneider, Phung, Le, Nguyen & Ziegler , C. cryptus Heidrich, Rösler, Vu, Böhme & 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, Vu, Nguyen, 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 rufford sp. nov. has femoral and precloacal pores in males, both of which are lacking in the following species: C. angularis (Smith) , C. badenensis Nguyen, Orlov & Darevsky , C. chauquangensis Hoang, Orlov, Ananjeva, Johns, Hoang & Dau , C. cucphuongensis Ngo & Chan , C. eisenmanae Ngo , C. grismeri Ngo , C. martini Ngo , C. nigriocularis Nguyen, Orlov & Darevsky , C. oldhami (Theobald) , C. pageli Schneider, Nguyen, Schmitz, Kingsada, Auer & Ziegler , C. paradoxus (Darevsky & Szczerbak) , C. puhuensis Nguyen, Yang, Le, Nguyen, Orlov, Hoang, Nguyen, Jin, Rao, Hoang, Che, Murphy & Zhang , C. saiyok Panitvong, Sumontha, Tunprasert & Pauwels , C. samroiyot Pauwels & Sumontha , C. sanook Pauwels, Sumontha, Latinne & Grismer , TABLE 4. Morphological comparisons between Cyrtodactylus rufford sp. nov. and its congeners from Laos and neighbouring countries in the Indochina region (compiled after Luu et al. 2014;

Nazarov et al. 2014; Nguyen et al. 2014; Panitvong et al. 2014; Pauwels et al. 2014; Pauwels & Sumontha 2014; Schneider et al. 2014; Sumontha et al. 2015; Nguyen et al. 2015; Luu et al.

2015). Abbreviations are as follows: FPl = femoral pores in the left side; FPr = femoral pores in the right side; – = characters unobtainable from literature; * = tail regenerated, for other abbrevi-

ations see material and methods.

. darevskii 84.6–100.0 95.0–113.0 38–46 present present 38–44 24–34 17–20 18–22 banded present

(FP+PP)

. dumnuii 76.2–84.2 100.2* 40 present present in 6+5–6+6–7 0– 7 16 19 banded present

males/abse (FPl+PP+FPr)

nt in

females

……continued on the next page TABLE 4. (Continued) TABLE 4. (Continued) C. spelaeus Nazarov, Poyarkov, Orlov, Nguyen, Milto, Martynov, Konstantinov & Chulisov , C. sumonthai Bauer, Pauwels & Chanhome , C. wayakonei Nguyen, Kingsada, Rösler, Auer & Ziegler , C. takouensis Ngo & Bauer , C. thirakhupti Pauwels, Bauer, Sumontha & Chanhome , and C. yangbayensis Ngo & Chan.

Cyrtodactylus rufford sp. nov. has a contiguous series of femoral and precloacal pores in males and thus differs from the following species which have poreless scales separating femoral from precloacal pores: C. bichnganae Ngo & Grismer , C. brevipalmatus (Smith) , C. dumnuii Bauer, Kunya, Sumontha, Niyomwan, Pauwels, Chanhome & Kunya , C. erythrops Bauer, Kunya, Sumontha, Niyomwan, Panitvong, Pauwels, Chanhome & Kunya , C. huongsonensis Luu, Nguyen, Do & Ziegler , C. interdigitalis , C. khelangensis Pauwels, Sumontha, Panitvong & Varaguttanonda , and C. tigroides Bauer, Sumontha & Pauwels.

Cyrtodactylus rufford sp. nov. has 42–43 femoral and precloacal pores in males and thus differs from the following species which have distinctly fewer femoral and precloacal pores: C. auribalteatus Sumontha, Panitvong & Deein (10–11), C. caovansungi Orlov, Nguyen, Nazarov, Ananjeva & Nguyen (15), C. chanhomeae Bauer, Sumontha & Pauwels , C. intermedius (Smith) (8–10), C. kingsadai Ziegler, Phung, Le & Nguyen (7–16), C. roesleri (20–28), and C. soudthichaki (29).

Cyrtodactylus rufford sp. nov. can be distinguished from C. astrum Grismer, Wood, Quah, Anuar, Muin, Sumontha, Ahmad, Bauer, Wangkulangkul, Grismer & Pauwels by its smaller size (maximum SVL 72.5 mm versus 108.3 mm), having fewer ventral scale rows (27–29 versus 31–46), fewer lamellae under the fourth toe (18– 19 versus 20–24), and more femoral and precloacal pores in males (42–43 versus 31–38); from C. jarujini Ulber by its smaller size (maximum SVL 72.5 mm versus 90.0 mm), having fewer ventral scale rows (27–29 versus 32–38), and fewer femoral and precloacal pores in males (42–43 versus 52–54); from C. lekaguli Grismer, Wood, Quah, Anuar, Muin, Sumontha, Ahmad, Bauer, Wangkulangkul, Grismer & Pauwels by its smaller size (SVL reaching 72.5 mm versus 103.5 mm), having fewer ventral scales (27–29 versus 31–43), and more femoral and precloacal pores in males (42–43 versus 30–36); from C. multiporus by its smaller size (maximum SVL 72.5 mm versus 98.0 mm), having fewer ventral scale rows (27–29 versus 30–38), and fewer femoral and precloacal pores in males (42– 43 versus 58–60); from C. teyniei David, Nguyen, Schneider & Ziegler by its smaller size (maximum SVL 72.5 mm versus 89.9 mm), having fewer ventral scale rows (27–29 versus 38), the presence of a nuchal band (versus being absent), and having a banded dorsal pattern (versus blotched).

Cyrtodactylus rufford sp. nov. is similar to the members of the C. phongnhakebangensis group ( Nazarov et al. 2014) including C. khammouanensis , C. lomyenensis , C. jaegeri , C. darevskii , and C. phongnhakebangensis Ziegler, Rösler, Herrmann & Vu in having a high number of femoral and precloacal pores and in the dorsal pattern. However, the new species can be distinguished from C. khammouanensis by having fewer ventral scale rows (27– 29 versus 32–38), fewer dorsal tubercles at midbody (14–16 versus 16–21), fewer lamellae under the fourth toe (18–19 versus 20–23), dorsum of head with dark blotched markings (versus absent), having light transverse bands between limb insertions with wavy margins (versus transverse bands with smooth margins), and tail with light rings (versus light bands); from C. lomyenensis by having fewer ventral scale rows (27–29 versus 35–36), fewer supralabials (10–12 versus 13–14), and more femoral and precloacal pores in males (42–43 versus 39–40); from C. darevskii by its smaller size (maximum SVL 72.5 mm versus 100.0 mm), having fewer ventral scale rows (27–29 versus 38–46), and fewer ventral scales between mental and cloacal slit (153–167 versus 180–208); from C. jaegeri by having fewer ventral scale rows (27–29 versus 31–32), fewer femoral and precloacal pores in males (42–43 versus 44), fewer lamellae under the fourth toe (18–19 versus 20–23), and having irregular shaped light transverse bands between limb insertions (versus regular shaped light transverse bands); and from C. phongnhakebangensis by its smaller size (maximum SVL 72.5 mm versus 96.3 mm), having fewer ventral scale rows (27–29 versus 32– 42), generally more femoral and precloacal pores in males (42–43 versus 32–42), and fewer ventral scales between mental and cloacal slit (153–167 versus 180–208). For more details see Table 5 .

Distribution. Cyrtodactylus rufford sp. nov. is currently known only from the type locality in the karst forest near Daen Village, Gnommalath District, Khammouane Province, central Laos ( Fig. 4 ).

Etymology. The new species is named in honour of the Rufford Foundation ( UK) for its support to herpetofaunal research and conservation in Laos. The species epithet is to be treated as a noun in apposition, invariable. As common names, we suggest Ki Chiem Rufford (Laotian) and Rufford Bent-toed Gecko (English) .

Natural history. The type series of the new species was found between 20:00 and 21:00, on a karst cliff near the entrance of Nang Log Cave, from 0.3 m to 2 m height above the forest floor, at elevations between 160 and 180 m a.s.l. Nang Log Cave is situated within an isolated karst mountain surrounded by secondary forest, plantations, and some huts for tourists. The cave is a tourism site, only 50 m distant from Road 12 and 64 km from the border of Vietnam. The humidity was approximately 80% and the air temperature ranged from 24 to 26o C ( Fig. 5 ). Other species found on the same karst wall within a 300 meters range of the type locality were the lizard Gekko gecko (Linnaeus) , and the tree frog species Rhacophorus spelaeus Orlov, Gnophanxay, Phimminith & Phomphoumy , as well as the Huntsman Spider Heteropoda maxima Jaeger , and the long-legged cave centipede Thereuopoda longicornis (Fabricius) .

Transverse dorsal bands 4 light bands 3-4 sometimes irregular shaped 2–3 light bands between limbs light bands

Tail pattern light rings light bands light bands

Figures

FIGURE 2. Adult male holotype of Cyrtodactylus rufford sp. nov. (VFU R. 2015.14) in life from Khammouane Province, central Laos: A) dorsal view, and B) lateral view. Photos: V. Q. Luu and T. Calame.

FIGURE 3. Dorsal view of the paratype of Cyrtodactylus rufford sp. nov. (IEBR R. 2015.34) in life from Khammouane Province, central Laos. Photo: V. Q. Luu.

FIGURE 4. Map showing the type locality of Cyrtodactylus rufford sp. nov. (red circle) and the type locality of Cyrtodactylus khammouanensis (white circle) in Khammouane Province, central Laos.

FIGURE 5. Habitat of Cyrtodactylus rufford sp. nov. at its type locality. Photo: V. Q. Luu.

Tables

TABLE 3. Measurements (in mm) and morphological characters of the type series of Cyrtodactylus rufford sp. nov. (* = partly regenerated, for other abbreviations see material and methods).

Character Sex	VFU R.2015.14 Holotype male	IEBR R.2015.34 Paratype male	NUOL R.2015.15 Paratype subadult male
SVL	68.3	72.5	56.5
TaL HH HL	94.5* 7.1 19.5	96.8 6.7 20.6	84.6 5.3 16.3
HW OD SE	11.2 5.2 9.1	12.9 5.6 8.6	9.8 4.8 7.0
EyeEar EarL TrunkL	4.9 2.0 32.2	5.3 2.3 31.3	4.3 1.8 27.4
ForeL FemurL CrusL	11.5 15.2 13.9	10.9 15.0 14.2	9.5 11.4 10.6
LD4A LD4P RW	6.5 8.3 2.8	6.2 8.1 2.8	5.9 7.2 2.3
RH MW ML	1.4 2.7 2.2	1.8 2.7 2.2	1.3 2.4 1.4
SL IL N	11/12 10/11 3/3	12/12 9/9 3/3	11/12 11/11 3/3
IN PM DTR	1 2 16	1 2 14	1 2 15
GST V SBL	9 28 153	9 27 167	9 29 156
SR FP+PP PAT	78 43 5/4	74 42 5/5	79 43 5/5
LD4 LT4	19/19 19/18	19/19 19/18	20/19 18/18

TABLE 5. Comparison of Cyrtodactylus rufford sp. nov. with other members of the C. phongnhakebangensis group (data obtained from Ziegler et al. 2002; Ngo & Pauwels 2010; Luu et al. 2014, Nazarov et al. 2014; and own data based on specimens examined, see Appendix).

Character	Cyrtodactylus rufford sp. nov.	C. khammouanensis C. lomyenensis
V	27–29	32–38 35–36
DTR	14–16	16–21 20–24
PP + FP (in males)	42–43	40–44 39–40
SLB	153–167	155–172 –
PAT	4–5	5–6 5
Rostral suture	Ϙ	Y Y
LT4	18–19	20–23 19–23
Transverse dorsal bands between limbs	3 or 4 light sometimes irregular shaped bands	4 light bands 4 light bands
Tail pattern	light rings	light bands light rings
continued.
Character	C. jaegeri C. darevskii	C. phongnhakebangensis
V	31–32 38–46	32–42
DTR	15–17 16–20	11–20
PP + FP (in males)	44 38–44	32–42
SLB	156–164 180–208	–
PAT	3–6 4–5	4–5
Rostral suture	I Y	I
LT4	20–23 18–22	18–26