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 hinnamnoensis sp. nov. ( Fig. 8 ) Cyrtodactylus phongnhakebangensis— Luu, Nguyen, Calame, Hoang, Soudthichack, Bonkowski, Ziegler, 2013 . Biodiversity Data Journal , e1015: 4. Holotype . IEBR A.2013.90, adult male, from Ban Dou Village ( 17°30.385’N , 105°49.160’E , elevation 183 m a.s.l.) within Hin Nam No NPA, Khammouane Province, central Laos , was collected on 11 June 2013 by V. Q. Luu and N. V. Ha. Paratypes . IEBR A.2013.89, adult male, 7 May 2013 , from Hang Toi region, Noong Ma Village ( 17°17.766’N , 106°08.803’E , elevation 580 m a.s.l.); VNUF R.2013.1 and NUOL R- 2013.2, adult males, 9 June 2013 , from Vang Ma No Village ( 17°30.778’N , 105°49.259’E , elevation 180 m a.s.l.); VNUF R.2014.99, adult male, 27 May 2014 , from Cha Lou Village ( 17°19.504’N , 105°57.630’E , elevation ca. 300 m a.s.l.); ZFMK 95235, adult female, 8 May 2013 , from Hang Toi region, Noong Ma Village ( 17°17.763’N , 106°08.778’E , elevation 555 m a.s.l.); ZFMK 95236, adult female, 30 May 2013 , from Noong Choong Region, Cha Lou Village ( 17°20.248’N , 105°56.693’E , elevation 252 m a.s.l.); NUOL R- 2013.3, adult female, 11 June 2013 , from Ban Dou Village ( 17°31.545’N , 105°49.086’E , elevation 197 m a.s.l.); VNUF R.2015.3, female, 13 March 2015 , from Xebangfai cave, Noong Ping Village ( 17°22.459’N , 105°49.626’E , elevation 182 m a.s.l.); NUOL R- 2015.9, female, 13 March 2015 , from Xebangfai cave, Noong Ping Village ( 17°22.648’N , 105°52.931’E , elevation 182 m a.s.l.); VNUF R.2015.11, female, 14 March 2015 , from Xebangfai cave, Noong Ping Village ( 17°22.759’N , 105°52.931’E , elevation 285 m a.s.l.). The paratypes ( VNUF R.2015.3, NUOL R- 2015.9, and VNUF R.2015.11) were collected by V. Q. Luu and K. Thanabuaosy in March 2015 ; the paratype ( VNUF R.2014.99) was collected by V. Q. Luu, N. V. Ha, T. Calame, D. V. Phan and K. Thanabuaosy in May 2014 , the remaining type series was collected by V. Q. Luu, N. V. Ha, and K. Thanabuaosy in May and June 2013 (V. Q. Luu et al. ). Diagnosis. Cyrtodactylus hinnamnoensis sp. nov. is characterized by: Adult SVL 84.1 ± 11.7 mm (mean ± SD); dorsal head with dark blotches; nuchal loop wide, distinct, posteriorly enlarged; dorsal body with four to six blackish brown bands between limb insertions; 13–19 irregular, weakly keeled dorsal tubercle rows; 35–48 ventral scale rows; ventral scale rows from mental to cloacal slit 179–201; scale rows at midbody 93–112; ventrolateral folds present, without tubercles; 36–44 precloacal-femoral pores in the males; 0–28 pores in females; enlarged femoral and precloacal scales present; 4–6 postcloacal tubercles; subcaudals enlarged. FIGURE 8. A) Dorsal view of the holotype (IEBR A.2013.90); and dorsal views of the paratypes of Cyrtodactylus hinnamnoensis sp. nov. in life B) VNUF R.2014.99; and C) VNUF R.2013.3. Photos: V. Q. Luu. Description of holotype . Adult male, snout-vent length (SVL) 83.6 mm ; body elongate (TrunkL/SVL 0.42); head elongate (HL/SVL 0.27), relatively wide (HW/HL 0.68), depressed (HH/HL 0.40), distinguished from neck; loreal region concave; snout long (SE/HL 0.43), obtuse, longer than diameter of orbit (OD/SE 0.60); snout scales small, homogeneous, granular, larger than those on frontal and parietal regions; eye large (OD/HL 0.26), pupils vertical; eyelid fringe with tiny spines posteriorly; ear oval-shaped, small (EarL/HL 0.08); rostral wider than high (RH/RW 0.60), with a median suture; supranasals in contact anteriorly and separated from each other by a small scale posteriorly; rostral in contact with first supralabial and nostril scales on each side; nares oval, surrounded by supranasal, rostral, first supralabial, and two enlarged postnasals; mental triangular, wider than high (ML/MW 0.65); two enlarged postmentals in broad contact posteriorly, bordered by mental anteriorly, first two infralabials laterally, and eight small scales posteriorly; supralabials 12/10; infralabials 10/9. Dorsal scales granular to flattened; dorsal tubercles round, weakly keeled, present on occiput, back and tail base, each surrounded by 8 granular scales, in 15 irregular longitudinal rows at midbody; ventral scales smooth, medial scales 2–3 times larger than dorsal scales, round, subimbricate, largest posteriorly, in 35 longitudinal rows between lateral folds at midbody; ventrolateral folds present, without tubercles; gular region with homogeneous smooth scales; ventral scales in a line from mental to cloacal slit 186; precloacal groove absent; enlarged femoral scales present; femoral and precloacal pores 42. Fore and hind limbs moderately slender (ForeL/SVL 0.17, CrusL/SVL 0.21); dorsal fore limbs with slightly developed tubercles; dorsal hind limbs covered by distinctly developed tubercles; fingers and toes free of webbing; lamellae under fourth fingers 19/19, under fourth toes 20/20. Tail regenerated, postcloacal tubercles 5/5; dorsal tail bearing tubercles at base; subcaudals distinctly enlarged, flat, smooth. Coloration in life. Ground coloration of dorsal head greyish brown with dark blotches; nuchal loop black, in U-shape, from posterior corner of eye through tympanum to the neck, dark brown, edged in yellow; body bands between limb insertions five, somewhat irregular, dark brown, edged in white; dorsal surface of fore and hind limbs with dark bars; tail brown dorsally with light brown rings, edged by yellowish white; chin, throat, and belly cream; upper and lower lips with dark brown bars; tail ventrally grey with light dots. Sexual dimorphism. The females differ from the males by lacking or having fewer precloacal-femoral pores (0–28 versus 36–44 in the males) and the absence of hemipenial swellings at the tail base (see Table 6 ). FIGURE 9. Dorsal pattern of three Cyrtodactylus sibling species: A) Cyrtodactylus phongnhakebangensis from Vietnam (Phong Nha-Ke Bang National Park); B) Cyrtodactylus calamei sp. nov. ; and C) Cyrtodactylus hinnamnoensis sp. nov. (VNUF R.2013.4), from Laos. Photos: T. Ziegler & V. Q. Luu. Comparisons. We compared the new species with its congeners from Laos and neighbouring countries in the mainland Indochina region, including Vietnam , Cambodia , and Thailand based on examination of specimens (see Appendix) and data integrated from the literature (compiled after 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 Tables 3 ). The cluster and correspondence analyses of morphological characters supported Cyrtodactylus hinnamnoensis sp. nov. as a sister taxon to C. darevskii ( Figs. 2–3 ). Molecular phylogenetic analyses also demonstrated the close relationships between these species (see Fig. 1 ). Morphologically, Cyrtodactylus hinnamnoensis sp. nov. is closely related to the C. phongnhakebangensis group including C. darevskii , C. phongnhakebangensis , C. calamei by dorsal colour pattern and the number of cloacal and femoral pores in males. However, the new species can be distinguished from C. darevskii by having fewer cloacal and femoral pores in females (maximum 0–28 versus 24–34), four to six blackish brown transverse body bands, as wide as light bands ( versus four to five dark irregular transverse breaking body bands, 0.5 times narrower than light band), first body band wide, butterfly-shaped ( versus thin, U-shaped in C. darevskii ), the presence of tubercles on fore limbs ( versus absent), and tail consisting of light rings ( versus banded); Cyrtodactylus hinnamnoensis sp. nov. differs from C. phongnhakebangensis by its slightly larger size (SVL reaching 100.6 mm versus 96.3 mm ), having fewer cloacal and femoral pores in females (0–28 versus 0–41), having more scale rows from mental to the front of cloacal slit (179–201 versus 161–177), the presence of tubercles on fore limbs ( versus absent), a narrower nuchal loop, not enlarged posteriorly (wide, enlarged posteriorly), four to six blackish brown transverse body bands as wide as light bands ( versus three to five dark transverse body bands as wide as double light bands, light transverse bands with small spots ( versus with big black blotches), and tail pattern consisting of light rings ( versus banded); Cyrtodactylus hinnamnoensis sp. nov. differs from C. calamei by its larger size (SVL reaching 100.6 mm versus 89.3 mm ), fewer cloacal and femoral pores in females (0–28 versus 38), more postcloacal tubercles (4–6 versus 4), dorsal head marking with distinctly dark spots and blotches ( versus indistinct dots), the absence of heart-shaped marking on postocciput ( versus present), four to six blackish brown body = standard deviatiοn, fοr οther abbreviatiοns see material and methοds). TABLE 6. Mοrphοmetric measurements (in mm) and meristic characters οf Cyrtodactylus hinnamnoensis sp. nov. (* = regenerated οr brοken tail, m = mean. max. = maximum,

Character	IEBR A.2013.90 Hοlοtype	IEBR A.2013.89 paratype	vNUF R.2013.1 paratype	NUΟL R-2013.2 paratype	vNUF R.2014.99 paratype	m±SD	min‾max
Sex	Male	male	male	male	male
SVL	83.6	91.9	83.7	92.5	85.0	87.3±4.5	83.6‾92.5
TaL	70.6*	100.9*	71.4*	101.6	lοss	101.6 (n=1)	101.6 (max)
HL	22.7	26.3	23.6	26.2	23.8	24.5±1.6	22.7‾26.3
HW	15.4	18.5	17.0	16.5	15.4	16.6±1.3	15.4‾18.5
HH	9.0	10.5	9.6	9.1	8.2	9.3±0.8	8.2‾10.5
OD	5.9	6.2	5.9	6.1	5.5	5.9±0.3	5.5‾6.2
SE	9.8	11.0	10.3	10.9	9.4	10.3±0.7	9.4‾11.0
EyeEar	6.2	7.2	7.7	7.3	6.6	7.0±0.6	6.2‾7.7
EarL	1.8	3.0	1.9	2.1	2.1	2.2±0.5	1.8‾3.0
TrunkL	34.7	39.0	36.6	38.4	38.1	37.4±1.7	34.7‾39.0
ForeL	14.5	15.2	15.1	16.2	14.9	15.2±0.7	14.5‾16.2
FemurL	18.1	19.4	18.9	20.4	17.3	18.8±1.2	17.3‾20.4
CrusL	17.5	19.4	17.9	19.8	16.2	18.2±1.5	16.2‾19.8
LD4A	8.0	9.3	9.0	8.5	8.4	8.6±0.5	8.0‾9.3
LD4P	9.6	11.3	10.1	10.6	10.9	10.7±0.5	10.1‾11.3
RW	3.5	4.0	3.7	4.4	3.7	3.8±0.4	3.5‾4.4
RH	2.1	2.5	2.3	2.5	2.3	2.3±0.2	2.1‾2.5
MW	3.4	4.0	3.7	3.7	3.7	3.7±0.2	3.4‾4.0
ML	2.2	3.0	2.7	2.7	2.6	2.6±0.3	2.2‾3.0
SL	12⁄10	10⁄10	10⁄10	11⁄9	9⁄11	10.2±0.9	9‾12
IL	10⁄9	9⁄9	11⁄8	10⁄10	10⁄10	9.6±0.8	8‾11
N	3	3	3	3	3	3.0±0.0	3‾3
IN	0	0	0	0	1	0.2±0.4	0‾1
PM	2	2	2	2	2	2.0±0.0	2‾2
DTR	15	18	17	17	16	16.6±1.1	15‾18
GSΤ	8	7	9	9	7	8.0±1.0	7‾9
V	35	44	37	36	36	37.6±3.6	35‾44
SLB	186	183	189	201	188	189.4±6.9	183‾201
SR	103	110	99	101	98	102.2±4.8	98‾110
PP+FP	42	38	36	44	38	39.6±3.3	36‾44
PAT	5⁄5	4⁄4	4⁄4	5⁄5	5⁄5	4.6±0.5	4‾5
LD4	19⁄19	18⁄19	19⁄17	18⁄18	19⁄18	18.0±0.7	17‾19
LT4	20⁄20	19⁄19	19⁄19	21⁄21	19⁄21	20.0±0.9	19‾21

……continued on the next page TABLE 6 . (Cοntinued)

Character	ΖFMK.95235 paratype	NUΟL R‾2013.3 paratype	ΖFMK 95236 paratype	vNUF R.2015.3 paratype	NUΟL R- 2015. 9 paratype	vNUF R. 2015.11 paratype	m±SD	min‾max
Sex	female	female	female	female	female	female
SVL	100.6	95.0	88.7	70.0	62.1	71.8	81.4±15.5	62.1‾100.6
TaL	108.3*	92.8*	82.9*	86.2	76.1	100.0	91.1±11.8	76.1‾108.3
HL	29.0	26.7	25.6	19.5	19.3	20.4	23.4±4.2	19.3‾29
HW	19.6	18.0	16.3	12.4	12.0	13.2	15.3±3.2	12‾19.6
HH	9.2	9.4	7.8	6.0	5.5	5.5	7.2±1.8	5.5‾9.4
OD	6.2	5.7	5.5	5.3	5.3	5.4	5.6±0.3	5.3‾6.2
SE	11.9	11.9	10.8	8.0	8.6	8.7	10.0±1.8	8.0‾11.9
EyeEar	8.4	7.8	6.7	5.3	4.6	4.6	6.2±1.6	4.6‾8.4
EarL	3.6	2.8	3.1	2.3	1.8	2.0	2.6±0.7	1.8‾3.6
TrunkL	43.0	38.4	37.9	32.5	25.5	30.6	34.7±6.3	25.5‾43.0
ForeL	16.0	15.9	15.2	11.7	10.9	12.3	13.7±2.3	10.9‾16.0
FemurL	21.3	19.6	19.6	15.3	14.2	15.9	17.7±2.9	14.2‾21.3
CrusL	20.3	17.9	18.1	14.2	12.7	15.0	16.4±2.9	12.7‾20.3
LD 4A	9.8	8.9	9.2	7.7	7.5	8.1	8.5±0.9	7.5‾9.8
LD 4P	11.0	10.9	11.2	8.8	8.6	9.8	10.1±1.2	8.6‾11.2
RW	4.6	4.1	3.8	2.8	2.9	3.1	3.6±0.7	2.8‾4.2
RH	3.0	2.7	2.5	1.6	1.6	1.7	2.2±0.6	1.6‾3.0
MW	4.2	3.7	3.8	2.9	2.8	2.9	3.4±0.6	2.8‾4.2
ML	3.3	2.5	3.1	2.3	2.0	2.2	2.6±0.5	2.0‾3.3
SL	10⁄10	10⁄10	9⁄11	9⁄9	9⁄10	11⁄10	9.8±0.7	9.0‾11.0
IL	9⁄9.	9⁄9.	9⁄9.	7⁄7.	7⁄8.	9⁄8.	8.3±0.9	7‾9
N	3	3	3	3	3	3	3.0±0.0	3‾3
IN	1	1	1	0	1	0	0.7±0.5	0‾1
PM	2	2	2	2	2	2	2.0±0.0	2‾ 2
DTR	19	17	16	13	16	15	16.0±2.0	13‾19
GSΤ	10	9	10	9	9	8	9.2±0.8	8‾10
V	43	48	38	42	39	40	41.7±3.6	38‾48
SLB	179	197	199	188	182	199	190.7±8.9	179‾199
SR	112	105	110	93	108	106	105.7±6.7	93‾112
PP +FP	28	0	24	13	19	14	16.3±9.9	0‾28
PAT	6⁄6	4⁄4	5⁄5	4⁄4	4⁄4	4⁄4	4.5±0.8	4‾6
LD 4	16⁄19	18⁄18	20⁄20	19⁄19	18⁄17	21⁄20	18.8±1.4	16‾21
LT 4	20⁄20	21⁄19	22⁄22	20⁄21	17⁄19	21⁄21	20.3±1.4	17‾22

TABLE 7. Cοmparisοn οf Cyrtodactylus calamei sp. nov. and Cyrtodactylus hinnamnoensis sp. nov. with οther members οf the C. phongnhakebangensis grοup (data οbtained frοm Ζiegler et al . 2002; Nazarοv et al. 2014; and οwn data). SLB = number οf scales alοng the midbοdy frοm mental tο anteriοr edge οf clοaca οpening.

Character	Cyrtodactylus calamei sp. nov.	Cyrtodactylus hinnamnoensis sp. nov.	C. darevskii	C. phongnhakebangensis
Number οf specimens	4	11	7	11
Maximal SvL (mm)	89.3	100.6	100	96.3
ventral scales	39‾42	35‾48	38‾46	32‾42
Dοrsal tubercle rοws	10‾16	13‾19	16‾20	11‾20
Femοral and preclοacal pοres (in males)	35‾39	36‾44	38‾44	32‾42
Femοral and preclοacal pοres (in females)	38	0‾28	24‾34	0‾41
SLB	183‾197	179‾201	180‾208	161‾177
Τubercle οn fοre limbs	present	present	absent	absent
Pοstclοacal tubercles	4	4‾6	4‾5	3‾5
Nuchal lοοp	narrοw, nοt enlarged pοsteriοrly	narrοw, nοt enlarged pοsteriοrly	thin, nοt enlarged pοsteriοrly	wide, enlarged pοsteriοrly
Dοrsal pattern between limb insertiοns	4 greyish brοwn transverse bands, narrοwer light bands	4‾6 blackish brοwn transverse bands, as wide as light bands	4‾5 dark transverse breaking bands, as narrοw as nearly half light bands	3‾5 dark transverse bands, as wide as dοuble light bands
Τail pattern	light rings	light rings	light bands	light bands

TABLE 8. Ecοlοgical details fοr the type series οf Cyrtodactylus hinnamnoensis sp. nov. frοm Hin Nam Nο NPA, central Laοs. Abbreviatiοns are as fοllοws∶ m.= male; f.= female; Τemp. = temperature).

Nr.	Museum Nο.	Τype	Date	Lοcality	Sex	Τime	Τemp.	Humidity	Elevatiοn	Micrοhabitat
1	IEBR A.2013.90	Hοlοtype	11 June 2013	Ban Dοu	m.	20∶50	28.3ο C	81%	193 m	karst cliff, in 0.5 m height
2	IEBR A.2013.89	Paratype	7 May 2013	Nοοng Ma	m.	20∶52	24.9ο C	81%	580 m	karst cliff, in 3 m height
3	vNUF R.2013.1	Paratype	9 June 2013	vangmanο	m.	22∶03	29.9ο C	90%	175 m	karst cliff, in 5 m height
4	NUΟL R- 2013.2	Paratype	9 June 2013	vangmanο	m.	21∶59	30.7ο C	87%	180 m	karst cliff, in 3 m height
5	vNUF R.2014.99	Paratype	27 May 2014	Chalοu	m.	21∶59	30.6ο C	81%	ca. 300 m	tree trunk, in 1 m height
6	ΖFMK95235	Paratype	8 May 2013	Nοοng Ma	f.	21∶35	29.8ο C	90%	555 m	cave entrance, karst cliff, in 2 m height
7	NUΟL R- 2013.3	Paratype	11 June 2013	Ban Dοu	f.	20∶40	28.8ο C	81%	197 m	karst cliff, in 2 m height
8	ΖFMK 95236	Paratype	30 May 2013	Cha Lοu	f.	20∶36	27.1ο C	81%	252 m	in the slit οf karst cliff, in 0.5 m height
9	vNUF R.2015.3	Paratype	13 March 2015	Xebangfai cave	f.	21∶15	28.3ο C	78%	182 m	tree trunk, near karst cliff, in 0.3 m height
10	NUΟL R- 2015. 9	Paratype	14 March 2015	Xebangfai cave	f.	19∶41	26ο C	80%	200 m	οn the karst cliff, in 0.3 m height
11	vNUF R.2015.11	Paratype	13 March 2015	Xebangfai cave	f.	20∶57	26.3ο C	81%	285 m	οn the karst cliff, in 0.3 m height

transverse bands, as wide as light bands ( versus four greyish brown transverse bands, narrower than light bands) (for more details see Table 7 , Fig. 9 ). The results of the correspondence analysis comparing all adult male morphological maesurements of Cyrtodactylus hinnamnoensis sp. nov. and the latter species indicated four distinct groups between these species (see Fig. 4 ). Distribution. Cyrtodactylus hinnamnoensis sp. nov. is currently known only from the type locality in the karst forest of Hin Nam No NPA, Khammouane Province, central Laos ( Fig. 6 ). Etymology. We name this species after its type locality, Hin Nam No NPA where the new Cyrtodactylus species was discovered and propose the following common names: Hinnamno Bent-toed Gecko (English), Ki Chiem Hin Nam No (Laotian). Natural history. Specimens were found at night between 19:41 and 22:03h on karst walls, ca. 0.3–5 m above the ground, near cave entrances in the limestone forest, at elevations between 175 and 580 m a.s.l. Only one male specimen VNUF R.2014.99 was collected on a tree trunk, about 1 m from the forest floor. The surrounding habitat was karst forest, dominated by species of Ebenaceae , Dracaenaceae, Arecaeae , Poaceae , Meliaceae , and Moraceae . The relative humidity ranged from 78% to 90%, and temperatures were from 24.9 to 30.7o C (see Table 8 ). When capturing individuals of the species, we observed an increased rate of tail autotomy and many individuals had regenerated tail, for example, seven of 11 specimens of Cyrtodactylus hinnamnoensis sp. nov. had dropped or/and regenerated tails. This suggests that these populations might be under the stress of predators (see also Grismer et al . 2016 ).