Cyrtodactylus muangfuangensis, Sitthivong & Luu & Ha & Nguyen & Le & Ziegler, 2019

Sitthivong, Saly, Luu, Vinh Quang, Ha, Ngoan Van, Nguyen, Truong Quang, Le, Minh Duc & Ziegler, Thomas, 2019, A new species of Cyrtodactylus (Squamata: Gekkonidae) from Vientiane Province, northern Laos, Zootaxa 4701 (3), pp. 257-275 : 260-272

Cyrtodactylus muangfuangensis sp. nov.

( Figs. 2–4 )

Holotype. VNUF R.2018.32 (Field number: SL.18.32), adult male, on a karst cliff (18°30’N, 101°57’E, 289 m elevation), nearby Phaluoang Village , Muangfuang District, Vientiane Province, Central Laos, collected on 31 December 2018 by Saly Sitthivong and Ha Van Ngoan.

Paratypes. NUOL R.2018.33 (Field number: SL.18.33), adult female , ZFMK 102701 (Field number: SL.18.40), subadult female , VNUF R.2018.41 (Field number: SL.18.41), subadult female, NUOL R.2018.39 (Field number: SL.18.39), subadult female, same collection data as the holotype .

Diagnosis. The new Cyrtodactylus species can be distinguished from remaining congeners by the following combination of characters: maximum SVL reaching 83.9 mm; dorsal pattern consisting of dark continuous nuchal loop, nape band and five dark transversal bands between limb insertions; intersupranasals two; dorsal tubercles present on occiput, body, hind limbs and tail base; 15 or 16 irregular dorsal tubercle rows at midbody; lateral folds clearly defined, without interspersed tubercles; 31–37 ventral scales between ventrolateral folds; 6 precloacal pores separated by a diastema of 6–8 poreless scales from a series of 7 (left) and 8 (right) femoral pores in enlarged femoral scales; 6 precloacal pores plus in total 10–15 femoral pores in females, which are separated by 6–8 poreless scales; enlarged precloacal and femoral scales present; one to four postcloacal tubercles on each side; median subcaudal scales transversely enlarged.

Description of holotype. Adult male, SVL 79.6 mm; body elongate (TrunkL/SVL 0.51); head elongate (HL/ SVL 0.28), relatively wide (HW/HL 0.67), depressed (HH/HL 0.41), distinguished from neck; loreal region concave; snout long (SE/HL 0.45), obtuse anteriorly, longer than diameter of orbit (OD/SE 0.55); scales on snout small, round or oval, granular, bigger than those in frontal and parietal regions; orbit large (OD/HL 0.25), pupils vertical; supraciliaries with spinous scales posteriorly; ear oval-shaped, small (EarL/HL 0.07); rostral wider than high with a straight medial suture, bordered with first supralabial and nostril on each side; supranasals in contact with each other; nostril opening oval, surrounded by supranasal, rostral, first supralabial, and two enlarged postnasals; mental triangular, slightly wider than long (ML/MW 0.88); two enlarged postmentals; 9/9 supralabials; 10/10 infralabials; dorsal scales granular; dorsal tubercles round, conical, present from occipital region to dorsum and half of tail, tubercles in 15 irregular rows at midbody, larger at midbody, each surrounded by 10 granular scales; ventrolateral body folds well defined, without interspersed tubercles; ventral scales smooth, round, midventral scales three times larger than the granules on dorsum, 34 longitudinal ventral scale rows at midbody; gular region with homogeneous, small and smooth scales; 178 ventral scales from mental to cloacal slit; precloacal groove absent; enlarged precloacal scales present; six contiguous precloacal pores, separated by 6/8 poreless scales and 7/8 femoral pores (FPl+PP+FPr: 7+6+8); enlarged precloacal and femoral scales present.

Fore- and hind-limbs moderately slender (ForeL/SVL 0.18, CrusL/SVL 0.18); enlarged tubercles on dorsal surface of forelimbs absent, but covered with slightly enlarged granules; hindlimbs with distinctly developed tubercles dorsally; interdigital webbing absent; 18/18 lamellae under fourth finger; 21/22 lamellae under fourth toe.

Tail partly regenerated, longer than SVL (TaL 104.1 mm, one-third of its posterior portion regenerated); 3/3 postcloacal tubercles; dorsal tail bearing distinct tubercles at base only; median subcaudal scales transversely enlarged, flat, smooth.

Coloration in life. Background coloration light brownish grey, with dark brown dorsal pattern; dorsal surface of head with irregular dark brown spots; dark canthal stripe, stretching from nostril to orbit; an irregularly shaped dark nuchal loop present, edged in black, in V-shape; greyish brown nuchal band present; five dark transversal bands between limb insertions at midbody, somewhat irregularly shaped, each outlined by darker brown borders anteriorly and posteriorly; tubercles at midbody greyish brown; dorsal surface of limbs with dark spots; dorsal surface of original tail (2/ 3 in the anterior part of the tail length) light brownish grey with dark bands, dark bands slightly wider than light bands; dorsal surface of regenerated tail (1/ 3 in the posterior part of the tail length) grey with dark markings; ventral surface of head, body and limbs cream; lower surface of tail greyish cream (see Fig. 2 ).

Sexual dimorphism and variation. The females differ from males by the absence of hemipenial swellings at the tail base. The subadult female paratype (ZFMK 102701) has the original tail with 11 dark bands. For further morphological characters of the paratypes see Table 2 and Fig. 4 .

Comparisons. We compared Cyrtodactylus muangfuangensis sp. nov. with other congeners from Laos and neighboring countries in the mainland Indochina region, including Vietnam, Cambodia, and Thailand based on examination of specimens (see Appendix) and data from literature ( 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 a,b; Luu et al. 2015; Nguyen et al. 2015b; Sumontha et al. 2014, 2015; Pauwels et al. 2016; Le et al. 2016; Luu et al. 2016 a,b,c; Luu et al. 2017; Nguyen et al. 2017; Pauwels et al. 2018; Nazarov et al. 2018; Murdoch et al. 2019; Pham et al. 2019). The new species can be differentiated from other known species of the genus Cyrtodactylus by morphological characters (see Table 3). Below we compared the new species with the morphologically similar species from Thailand, Vietnam, and remaining species from Laos.

Cyrtodactylus muangfuangensis sp. nov. is morphologically similar to C. dumnuii Bauer, Kunya, Sumontha, Niyomwan, Pauwels, Chanhome & Kunya ; C. tigroides Bauer, Sumontha & Pauwels , C. khelangensis Pauwels, Sumontha, Panitvong & Varaguttanonda , C. brevipalmatus (Smith) , and C. huongsonensis Luu, Nguyen, Do & Ziegler in arrangement and the number of femoral and precloacal pores. However, the new species can be distinguished from C. dumnuii by having fewer ventral scales (31–37 versus 40), the presence of pitted scaless in females (versus absent), nuchal band enlarged posteriorly, butterfly-shaped (versus interrupted posteriorly), fewer body bands (5 versus 6–7), fewer dorsal tubercles at midbody (15–16 versus 18–20), and fewer dark and light caudal bands on tail (8–10 versus 12; 9–10 versus 13, respectively); from C. tigroides by having fewer precloacal pores (6 versus 8 or 9), a heterogeneous mixture of body scalation (versus homogeneous), more numbers of conical, keeled tubercles (15–16 versus 13 small tubercle rows); from C. khelangensis by its smaller size (maximum SVL 83.9 mm versus 95.3 mm), having more pitted scales in females (5–9+5–8 versus 2+1 femoral pores in left and right side, respectively), and more precloacal pores in males (6 versus 2–5); from C. brevipalmatus by its larger size (maximum SVL 83.9 mm versus 72 mm), having fewer ventral scales (31–37 versus 44), having fewer femoral pores in males (6 versus 9), dorsal pattern banded (versus blotched); from C. huongsonensis by having fewer ventral scales (31–37 versus 48), the presence of pitted scales in females (versus absent), and fewer pitted scales in females (6 versus 8). For more details see Table 4 .

Cyrtodactylus muangfuangensis sp. nov. has transversely enlarged median subcaudal scales and thus differs from the following species which lack this structure: C. buchardi David, Teynié & Ohler , C. cryptus Heidrich, Rösler, Vu, Böhme & Ziegler , C. interdigitalis Ulber , C. pseudoquadrivirgatus Rösler, Vu, Nguyen, Ngo & Ziegler , C. vilaphongi Schneider, Nguyen, Le, Nophaseud, Bonkowski & Ziegler , and C. ziegleri Nazarov, Orlov, Nguyen & Ho.

Cyrtodactylus muangfuangensis sp. nov. has femoral pores in the male and thus differs from the following species which lack this structure: C. pageli Schneider, Nguyen, Schmitz, Kingsada, Auer & Ziegler , C. spelaeus Nazarov, Poyarkov, Orlov, Nguyen, Milto, Martynov, Konstantinov & Chulisov , C. teyniei David, Nguyen, Schneider & Ziegler , C. thathomensis Nazarov, Pauwels, Konstantinov, Chulisov, Orlov & Poyarkov. In addition, the new species differs from C. pageli by having fewer ventral scales (31–37 versus 41–44), and the presence of enlarged femoral scales (versus absent); from C. spelaeus by the presence of enlarged femoral scales and pitted scales in females (versus absent); from C. teyniei by having body pattern banded (versus blotched); and from C. thathomensis by the presence of enlarged femoral scales (versus absent), and having 16-21 pitted scales in females (versus 12 pores in females).

Cyrtodactylus muangfuangensis sp. nov. has six precloacal pores plus in total 21 femoral pores in the male, which are separated by poreless scales and thus differs from the following species which have a contiguous series of precloacal-femoral pores: C. bansocensis Luu, Nguyen, Le, Bonkowski & Ziegler (34), C. calamei Luu, Bonkowski, Nguyen, Le, Schneider, Ngo & Ziegler (35–39), C. darevskii Nazarov, Poyarkov, Orlov, Nguyen, Milto, Martynov, Konstantinov & Chulisov (38–44), C. hinnamnoensis Luu, Bonkowski, Nguyen, Le, Schneider, Ngo & Ziegler (36–44), C. jaegeri Luu, Calame, Bonkowski, Nguyen & Ziegler (44), C. jarujini Ulber (52–54), C. khammouanensis Nazarov, Poyarkov, Orlov, Nguyen, Milto, Martynov, Konstantinov & Chulisov (40–44), C. lomyenensis Ngo & Pauwels (39–40), C. multiporus Nazarov, Poyarkov, Orlov, Nguyen, Milto, Martynov, Konstantinov & Chulisov (58–60), C. rufford Luu, Calame, Nguyen, Le, Bonkowski & Ziegler (42–43), and C. soudthichaki Luu, Calame, Nguyen, Bonkowski & Ziegler (29).

Cyrtodactylus muangfuangensis sp. nov. differs from C. sommerladi Luu, Bonkowski, Nguyen, Le, Schneider, Ngo & Ziegler by having precloacal-femoral pores interrupted by poreless scales (versus in a continuous row), and dorsal tubercles being arranged in 15–16 rows (versus undefined dorsal tubercles, 0–5).

Distribution. Cyrtodactylus muangfuangensis sp. nov. is currently known only from the type locality: Muangfuang District, Vientiane Province, Central Laos ( Fig. 5 ).

Etymology. The specific epithet of the new species refers to the type locality of the new species in Muangfuang District, Vientiane Province, Central Laos. Suggested common names: Muangfuang Bent-toed Gecko (English), Ki Chiem Muang Fuang (Laotian).

Natural history. The type series of the new species was collected between 19:00 and 21:00h, on karst cliffs, at elevations of 255–289 m a.s.l. The surrounding habitat was karst forest ( Fig. 6 ).

Figures

FIGURE 2. Dorsal view of the adult male holotype of Cyrtodactylus muangfuangensis sp. nov. (VNUF R.2018.32) in life. Photo: S. Sitthivong

FIGURE 3. Cloacal region (precloacal and femoral pores marked with black dots) of the holotype of Cyrtodactylus muangfuangensis sp. nov. in preservative. Photo: V. Q. Luu

FIGURE 4. Type series of Cyrtodactylus muangfuangensis sp. nov. from right to left: adult male holotype (VNUF R.2018.32); adult female paratype (NUOL R.2018.33); subadult female paratype (ZFMK 102701); subadult female paratype (VNUF R.2018.41); adult female paratype (NUOL R.2018.39). Photo: V. Q. Luu

FIGURE 5. Distribution map of the Cyrtodactylus phongnhakebangensis species group and other closely related Cyrtodactylus species inhabiting Laos and adjacent areas: dark brown, C. khammouanensis; pastel pink, C. rufford; gold, C. bansocensis; pastel green, C. southichacki; silver, C. jaegeri; beige, C. sommerladi; teal, C. hinnamnoensis; magenta, C. dareveskii; green, C. calamei; grass green, C. lomyenensis; red, C. multiporus; indigo, C. thathomensis; dark grey, C. pageli; black, Cyrtodactylus muangfuangensis sp. nov.; dark green, C. teyniei; charcoal, C. roesleri; red orange, C. phongnhakebangensis; purple, C. huongsonensis; yellow, C. jarujini; orange, C. damnuii; white, C. tigroides; blue, C. khelangensis; pastel yellow, C. brevipalmatus.

FIGURE 6. Habitat of Cyrtodactylus muangfuangensis sp. nov. Photo: S. Sitthivong.

Tables

TABLE 2. Measurements (in mm) and morphological characters of the type series of Cyrtodactylus muangfuangensis sp. nov. (for other abbreviations see material and methods; VNUF = Vietnam National University of Forestry, Hanoi, Vietnam; National University of Laos (NUOL) ZFMK = Zoologisches Forschungsmuseum Alexander Koenig, Bonn, Germany; * = tail regenerated).

Character	VNUF R.2018.32	NUOL R.2018.33	ZFMK 102701	VNUF R.2018.41	NUOL R.2018.39
	(Holotype)	(Paratype)	(Paratype)	(Paratype)	(Paratype)
Sex	adult male	adult female	subadult female	subadult female	subadult female
SVL	79.6	83.9	58.5	60.8	60.1
TaL	104.1	81.9*	80.1	69.1	78.9
HH	9.1	10.5	7.6	7.2	7.5
HL	22.0	21.2	15.9	15.9	15.6
HW	14.7	16.4	11.4	11.2	11.6
OD	5.5	5.2	3.8	4.1	4.2
SE	10	10	7.1	7.1	7.1
EyeEar	5.7	5.6	4.3	4.3	4.6
EarL	1.5	1.9	1.1	1.2	1.5
TrunkL	40.5	37.2	24.4	24.6	25.5
ForeL	14.3	13.2	9.5	9.6	10.1
FemurL	17.6	18.2	12.6	12.5	13.3
CrusL	14.4	13.6	12.3	12.2	12.3
LD4A	7.3	7.4	5.1	5.9	5.5
LD4P	9.4	9.3	6.2	6.3	6.3
RW	3.4	3.6	2.7	2.7	2.6
RH	1.9	1.9	1.5	1.2	1.4
MW	3.2	3.4	2.3	2.4	2.5
ML	2.8	2.7	2.1	2.1	2.1
SL	9/9	11/11	11/10	11/10	11/10
IL	10/10	10/11	9/9	10/10	9/11
N	3/3	2/3	2/2	2/2	2/2
IN	0	0	0	0	0
PM	2	2	2	2	2
GP	8	9	10	9	9
DTR	15	16	15	15	15
GST	10	10	10	10	10
V	34	36	37	31	34
SLB	178	176	177	180	182
SR	102	104	112	103	110
FPl+FPr	7+8	9+6 (pitted scales)	5+5 (pitted scales)	7+7 (pitted scales)	7+8 (pitted scales)
PP	6	6 (pitted scales)	6 (pitted scales)	6 (pitted scales)	6 (pitted scales)
PAT	3/3	4/4	3/3	4/4	1/1
LD4	18/18	19/18	17/16	18/18	18/19
LT4	21/22	23/22	18/19	20/20	21/22

TABLE 3. Morphological comparisons between Cyrtodactylus muangfuangensis sp. nov. and its congeners from Laos and neighboring countries in the Indochina region (com- piled 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. 2014, 2015; Nguyen et al. 2015b; Luu et al. 2015; Luu et al. 2016a,b; Le et al. 2016; Luu et al. 2017; Nguyen et al. 2017; Pauwels et al. 2018; Nazarov et al. 2018; Murdoch et al. 2019; Pham et al. 2019). Abbreviations are as follows: – = characters unobtainable from literature; * = tail regenerated; max. = maximal; SVL = snout–vent length; TaL = tail length; V = ventral scales; EFS = enlarged femoral scales; FP = femoral pores; PP = precloacal pores; LD4 = subdigital lamellae on fourth finger; TL4 = subdigital lamellae on fourth toe; FPl = femoral pores on the left side; FPr = femoral pores on the right side.

Taxa	SVL	TaL	V	EFS	FP	PP	PP	LD4	LT4	Color pattern	Enlarged
	(mm)	(mm)				(in males)	(in females)			of dorsum	subcaudals
Cyrtodactylus muang-	79.6–83.9	81.9 *– 102.6	31–37	present	present	7+6+8	5–9+6+5–8	16–19	18–23	banded	present
fuangensis sp. nov.						(FPl+PP+FPr)	(FPl+PP+FPr
C. angularis	80.0–92.0	92–95.2	40–45	present	absent	3	(pitted scales) 3	18–19	18–19	banded	present
C. astrum	46.4–108.3	99.0*–109.0*	31–46	–	present	31–38	–	–	20–24	banded	present
C. auralensis	75.9–84.3	96	37–44	present	absent	(FP+PP) 7–9		–	19–21		present
C. auribalteatus	82.8–98.1	106.5–138.7	38–40	5–7	4–5 (in	6	8–9 absent	–	18–21	banded banded	present
C. badenensis	59.3–74.1	58.6–82.4	25–29	absent	males) absent	0	0	–	18–22	banded	present
C. bansocensis	71.0–74.0	98.5–103.5	34–35	present	present	34	unknown	16–19	18–21	banded	present
C. bichnganae	95.3–99.9	96.3–115.6	30–31	11–13	18	(FP+PP) 10	8	18–20	16–20	banded	present
C. bidoupimontis	74.0–86.3	75.0–86.0	38–43	6–8	absent	4–6	0	15–20	18–23	banded	absent
C. bobrovi	75.2–96.4	80.8–90.3	40–45	0	0	5	0	19–21	21–22	banded	absent
C. bokorensis C. brevipalmatus	77.5–93 64.0–72.0	105–120 77.0	42–48 35–44	present present	absent present	7–9 6+9+7	0 6+9+7	– –	18–20 –	banded blotched	present present
C. buchardi	60.0–65.0	46.0–54.0	30	absent	absent	(FPl+PP+FPr) 9	(FPl+PP+FPr) 0	14	12	blotched	absent
C. bugiamapensis	58.6–76.8	65.3–83.0	36–46	6–10	absent	7–8	0–7	15–17	17–20	blotched	absent
C. calamei	75.0–89.3	86.1–107.5	39–42	present	present	35–39	38	16–18	18–20	banded	present
C. caovansungi	90.4–94.0	120.0	38–44	8	6	(FP+PP) 9	(FP+PP) 0	22	23–25	banded	present
C. cardamomensis	72–84.1	78–107.8	36–43	present	absent	9–10	0	–	17–19	banded	present
C. cattienensis	43.5– 69.0	51.0–64.7	28–42	3–8	absent	6–8	0	12–16	14–19	banded	absent

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