Evolution in karst massifs: Cryptic diversity among bent-toed geckos along the Truong Son Range with descriptions of three new species and one new country record from Laos Author Luu, Vinh Quang Author Bonkowski, Michael Author Nguyen, Truong Quang Author Le, Minh Duc Author Schneider, Nicole Author Ngo, Hanh Thi Author Ziegler, Thomas text Zootaxa 2016 4107 2 101 140 journal article 39173 10.11646/zootaxa.4107.2.1 7ea63a6e-321e-42a1-a309-d50dfad93583 1175-5326 266526 D04FFE10-BF1B-4F1C-BBAF-0396D812B830 Cyrtodactylus calamei sp. nov. ( Fig. 7 ) Holotype . VNUF R.2015.28, adult male, in the karst forest, Tham Nok Aen region, Thong Xam Village ( 17°34.179’N , 105°50.329’E , elevation 210 m a.s.l.) within Hin Nam No NPA, Khammouane Province, central Laos , was collected on 25 March 2015 by V. Q. Luu, T. Calame, and K. Thanabuaosy. Paratypes . IEBR A. 2015.36, adult male; NUOL R- 2015.22, subadult male; VNUF R.2015.27, adult female, the same data as the holotype . Diagnosis. Cyrtodactylus calamei sp. nov. can be distinguished from its congeners by a unique combination of the following characters: Adult SVL 80.0 ± 8.0 mm (mean ± SD); head dorsally with grey small scattered spots; nuchal loop present with indentations, not enlarged posteriorly, extending from each postnasal cross orbit and contacting on nape; four greyish brown, wide transverse bands between limbs, sometimes irregular; dorsal surface with homogenous, low, round, weakly keeled scales; 39–42 ventral scales at midbody; ventrolateral skin folds well-defined; 183–197 ventral scale rows from mental to cloacal slit; 101–114 scale rows at midbody; 35–39 precloacal-femoral pores in males, 38 in the female; enlarged femoral and precloacal scales present; four postcloacal tubercles; subcaudal scales transversely enlarged. Description of holotype . Adult male, medium sized, SVL 75.8 mm ; body long (TrunkL/SVL 0.44); head moderate (HL/SVL 0.29), narrow (HW/HL 0.77), somewhat depressed (HH/HL 0.42), differentiated from neck; prefrontal and postnasal regions concave; snout elongate (SE/HL 0.45), obtuse; snout scales small, homogeneous, granular, about two times larger than those in frontal and parietal regions; eye large (OD/HL 0.29), pupils vertical; supraciliaries with tiny spines posteriorly; ear opening oval, obliquely directed, small in size (EarL/HL 0.10); rostral subrectangular, wider than high (RH/RW 0.61), medially deep furrow, vertical suture, bordered by nasorostral, nare and first supralabial laterally; nares oval, surrounded by rostral anteriorly, supranasal, first supralabial laterally, and posteriorly by two enlarged postnasals; intersupranasal scale single; mental subtriangular, nearly as wide as rostral (MW/RW 0.94), bordered by two postmentals and first infralabial laterally; supralabials nine; infralabials eight. Dorsal scales granular to flattened; dorsal tubercles round, weakly keeled, extending from postocciput to base of tail; ventrolateral folds distinct; ventral scales smooth, medial scales about two times larger than dorsal scales, imbricate, 40 rows at midbody between folds; midbody scale rows 104; ventral scales from mental to cloacal slit 183; enlarged femoral-precloacal scales present; precloacal-femoral pores 39, in a continued row; precloacal groove absent. Fore and hind limbs moderately long (ForeL/SVL 0.18, CrusL/SVL 0.2); tubercles on dorsum of fore limbs absent; dorsal surface of hind limbs interspersed with tubercles; interdigital webbing weakly developed; subdigital lamellae on fourth fingers 16/16 and on fourth toes 19/18. Tail tapering to a point (TaL/SVL 1.42); four postcloacal tubercles laterally; subcaudals distinctly enlarged. Coloration in life. Ground color of dorsal head, back, limbs, and tail yellowish brown; dorsal head with small spread spots and a heart-shaped marking on postocciput; nuchal loop present, in U-shape, from posterior corner of nare crossing eye and tympanum, extending to nape, dark brown, bolder from postocular to nape, irregularly edged in yellow; four greyish brown body bands between limb insertions with indentations in mid-dorsal region, edged in yellow; dorsal surface of fore and hind limbs with grey reticulations; tail with narrow light bands; ventral surface greyish cream. Sexual dimorphism. The single adult female differs from two adult males by its larger size (maximum SVL 89.3 mm versus 75.8 mm in the males) and lacking of hemipenial swellings at the base of tail (see Table 4 & Fig. 6 ). Comparisons. We compared Cyrtodactylus calamei sp. nov. with other Cyrtodactylus species known 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 . 2014a ; 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 . 2015a ; Luu et al . 2016a , b ) (see Table 3 ). The cluster and correspondence analyses indicated that Cyrtodactylus calamei sp. nov. is nested in the same clade with C. darevskii and the species to be described in the following ( Figs. 2–3 ). Molecular phylogenetic analyses also strongly supported the sister relationship between the new species and afore mentioned taxa (see Fig. 1 ). Morphologically, Cyrtodactylus calamei sp. nov. closely resembles the other karst forest species, C. darevskii and C. phongnhakebangensis , in dorsal colour pattern. However, the new species can be distinguished from C. darevskii by its smaller size (maximum SVL 89.3 mm versus 100 mm ), having fewer dorsal tubercle rows (10–16 versus 16–20), fewer femoral and precloacal pores in males (35–39 versus 38–44), more femoral and precloacal pores in females (38 versus 24–34), the presence of heart-shaped marking on postocciput ( versus absent), four greyish brown regular transverse body bands as wide as nearly two times of nuchal band ( versus four to five dark irregular transverse breaking bands as narrow as nuchal band), first body band wide, butterfly-shaped ( versus thin, U-shaped in C. darevskii ), tail with light rings ( versus banded); from C. phongnhakebangensis by its smaller size (maximum SVL 89.3 mm versus 96.3 mm ), more scale rows from mental to the front of cloacal slit (183–197 versus 161–177), nuchal loop narrow, indented, not posteriorly enlarged ( versus wide, posteriorly enlarged), four greyish brown transverse body bands, slightly narrower light bands ( versus three to five dark transverse body bands, twice wider than light bands), and tail with light rings ( versus bands) (see Table 4 ). TABLE 4. Morphometric measurements (in mm) and meristic characters of the type series of Cyrtodactylus calamei sp. nov. (* = regenerated or broken tail, for other abbreviations see material and methods).

Character	VNUF R.2015.28 holotype	IEBR A. 2015.36 paratype	NUOL R-2015.22 paratype	VNUF R.2015.27 paratype
Sex	male	male	subadult male	female
SVL	75.8	75.0	64.1	89.3
TaL	107.5	92.4	86.1	102.0*
HL	21.7	21.5	18.8	24.8
HW	14.1	13.9	11.9	16.6
HH	9.0	7.4	6.5	10.4
OD	6.2	5.8	4.9	6.5
SE	9.7	10.0	7.8	10.8
EyeEar	6.0	5.8	5.8	6.7
EarL	2.2	2.1	2.1	2.4
TrunkL	33.4	31.6	28.4	38.2
ForeL	13.6	13.4	11.2	14.2
FemurL	17.8	16.3	14.7	18.6
CrusL	15.5	16.0	13.4	17.9
LD4A	9.0	8.8	7.6	9.1
LD4P	10.7	9.6	8.9	11.3
RW	3.3	3.3	2.8	3.8
RH	2.0	1.8	1.7	2.0
MW	3.1	3.4	2.6	3.7
ML	2.3	2.2	2.0	2.7
SL	9/9	11/11	10/10	10/10
IL	8/8	10/11	8/8	9/10
N	3/3	3/3	3/3	3/3
IN	1	1	0	1
PM	2	2	2	2
DTR	16	16	10	14
GST	8	9	8	9
V	40	39	40	42
SLB	183	193	187	197
SR	104	107	101	114
FP+PP	39	38	35	38
PAT	4/4	4/4	4/4	4/4
LD4	16/16	17/18	18/18	18/17
LT4	19/18	18/19	21/20	18/18

Distribution. Cyrtodactylus calamei is currently known only from the type locality in Tham Nok En area, Hin Nam No NPA, Khammouane Province, central Laos ( Fig. 6 ). Etymology. The new species is named in honour of our friend and colleague, Mr. Thomas Calame, from WWF Greater Mekong, Vientiane, Laos , who participated in our field research in Hin Nam No NPA, Khammouane Province between 2014 and 2015 . As common names, we suggest Calame’s Bent-toed Gecko (English), Ki Chiem Calame (Laotian). FIGURE 7. A) Dorsolateral view of the holotype (VNUF R.2015.28); B) lateral view of the paratype (NUOL R.2015.22) of Cyrtodactylus calamei sp. nov. in life. Photos: V. Q. Luu. Natural history. Specimens were found at night between 19:30 and 21:08h, on limestone outcrops, at elevations between 190 and 260 m a.s.l. The surrounding habitat was karst forest. The relative humidity was 80% and the air temperature ranged from 23 to 26o C (see Table 5 ).