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 sommerladi sp. nov. ( Fig. 10 ) Holotype . VNUF R.2013.22 adult male, in karst forest, Hang Toi region, Noong Ma Village ( 17°17.795’N , 106°08.738’E , elevation 572 m a.s.l.) within Hin Nam No NPA, Khammouane Province, central Laos , collected on 0 5 May 2013 by V. Q. Luu and N. V. Ha. Paratypes . NUOL R- 2013.23, IEBR A.2015.37, ZFMK 97196, VNUF R.2013.105, VNUF R.2014.87, NUOL R- 2013.14, IEBR A.2015.39, NUOL R- 2013.21, IEBR A.2015.38, VNUF R.2013.104, ZFMK 97197, VNUF R.2014.89, VNUF R.2013.67 the same locality as the holotype . IEBR A.2015.40 adult female, in karst forest, Cha Lou Village ( 17°18.880’N , 105°57.103’E , elevation 572 m a.s.l.) within Hin Nam No NPA, Khammouane Province, central Laos , collected on 25 May 2013 . Diagnosis. Cyrtodactylus sommerladi sp. nov. is characterized by: Adult SVL 72.3 ± 3.8 mm (mean ± SD); dorsal head greyish brown without dark blotches; nuchal loop present, narrow, not enlarged posteriorly; five or six dark transverse bands between limbs; dorsal surface with homogenous, tubercle-like scales; ventral scales at midbody 31–39; ventrolateral skin folds present; ventral scale rows from mental to cloacal slit 168–192; scale rows at midbody 76–93; precloacal-femoral pores 20–26 in males, 17–21 in females; enlarged femoral and precloacal scales present; postcloacal tubercles 4–6; subcaudal scales slightly enlarged. Description of holotype . Adult male, small sized (SVL 70.3 mm ); body elongate (TrunkL/SVL 0.45); head elongate (HL/SVL 0.25), width (HW/HL 0.77), relatively depressed (HH/HL ratio 0.41), distinct from neck; loreal area concave; snout long (SE/HL ratio 0.47), obtuse, two times longer than diameter of orbit (OD/SE 0.50); snout scales small, homogeneous, granular, about one and a half times larger than those in frontal and parietal regions; eye large (OD/HL ratio 0.23), pupils vertical; eyelid fringe with tiny spines in posterior part, posterior ones more developed; ear opening oval, obliquely orientated, small (EarL/HL 0.12); rostral square-shaped, wider than high (RH/RW ratio 0.55), medially with a straight, vertical suture, in contact with nasorostral, nare and first supralabial on each side; nares oval, surrounded by rostral anteriorly, supranasal, first supralabial laterally, and two enlarged postnasals posteriorly; supranasals in contact; mental triangular, slightly narrower than rostral (RW/MW ratio 0.94), in contact with two postmentals and the first infralabial on each side, postmentals surrounded by first infralabial on each side and seven granular scales posteriorly, two outer ones enlarged; supralabials 10/10; infralabials 8/9; supralabials separated from orbit by 3 or 4 rows of granular scales. Dorsal scales homogenous, tubercle-like; dorsal tubercles indistinct; ventrolateral folds present; ventral scales smooth, medial scales 2 or 3 times larger than dorsal scales, round, and in 37 rows at midbody; midbody scale rows 87; scales between mental and cloacal slit 192; femoral scales enlarged; precloacal scales enlarged, precloacal-femoral pores 23, in a continuous row; precloacal groove absent. FIGURE 10. A) Dorsal view of the holotype (VNUF R.2013.22); and dorsal views of the paratypes of Cyrtodactylus sommerladi sp. nov. in life B) VNUF R.2013.67; C) IEBR A.2015.39; and D) NUOL R-2013.105. Photos: V. Q. Luu. FIGURE 11. Comparison of dorsal scale structure of Cyrtodactylus sommerladi sp. nov. (A: with homogenous tubercle-like scales) and C. roesleri (B: with distinct dorsal tubercle rows). Photos: N. Schneider & T. Ziegler. Fore and hind limbs moderately slender (ForeL/SVL 0.15, CrusL/SVL 0.19); dorsal surface of hind limbs with tubercles, absent on fore limbs; fingers and toes with rudimental webbing; lamellae under fourth fingers 18/18, under fourth toes 20/22. Tail longer than snout-vent length (TaiL/SVL 1.14); postcloacal tubercles 6 on each side; subcaudals slightly enlarged. TABLE 9. Mοrphοmetric measurements (in mm) and meristic characters οf the type series οf Cyrtodactylus sommerladi sp. nov. (* = regenerated οr brοken tail, m = mean, max. = maximum, SD = standard deviatiοn, sub. = subadult, fοr οther abbreviatiοns see material and methοds).

Character	vNUF R.2013.22 Hοlοtype	IEBR A.2015.39 paratype	vNUF R.2014.87 paratype	IEBR A.2015.37 paratype	ΖFMK97196 paratype	NUΟL R- 2013.105 paratype	m±SD	min‾max
Sex	male	male	male	male	male	male
SvL	70.3	71.3	70.2	71.0	67.3	69.6	70.0±1.4	67.3‾71.3
ΤaL	79.8	67.9*	lοss	69.7*	84.8*	89.4	84.6±6.8	79.8‾89.4
HL	17.9	19.4	18.3	19.7	17.6	17.9	18.5±0.9	17.6‾19.7
HW	13.7	12.3	13.5	13.8	12.9	13.3	13.3±0.6	12.3‾13.8
HH	7.3	7.1	8.3	7.6	7.1	7.3	7.5±0.5	7.1‾8.3
ΟD	4.2	4.3	4.3	4.2	3.9	4.2	4.2±0.1	3.9‾4.3
SE	8.4	8.6	8.7	8.7	7.9	8.6	8.5±0.3	7.9‾8.7
EyeEar	5.3	5.6	6.5	5.5	5.8	5.8	5.8±0.4	5.3‾6.5
EarL	2.2	1.5	1.4	1.8	1.5	1.5	1.7±0.3	1.4‾2.2
ΤrunkL	31.7	27.3	29.9	29.4	26	26.9	29.0±2.0	26.9‾31.7
FοreaL	10.6	10.6	11.8	10.7	10.8	10.6	10.9±0.5	10.6‾11.8
FemurL	14.3	15.7	14.6	15.9	12.6	14.6	14.6±1.2	12.6‾15.9
CrusL	13.6	12.9	14.4	12.5	12.8	13.7	13.3±0.7	12.5‾14.4
LD4A	6.4	6.4	5.6	7.0	6.0	5.4	6.1±0.6	5.4‾7.0
LD4P	8.3	7.8	6.9	8.2	7.4	8.1	7.8±0.5	6.9‾8.3
RW	2.9	2.5	3.4	3.0	2.8	2.9	2.9±0.3	2.5‾3.4
RH	1.6	1.6	1.9	1.2	2.1	2.1	1.8±0.4	1.2‾2.1
MW	3.1	2.9	3.2	3.5	3.3	2.9	3.2±0.2	2.9‾3.5
ML	1.9	2.4	2.4	2.0	2.6	2.2	2.3±0.3	1.9‾2.6
SL	10⁄10	9⁄10	10⁄11	11⁄10	10⁄9	10⁄10	10.0±0.6	9‾11
IL	8⁄9	9⁄9	8⁄8	9⁄7	8⁄9	9⁄9	8.5±0.7	7‾9
N	3	2	2	3	3	3	2.7±0.5	2‾3
IN	0	0	0	0	0	0	0	0
PM	2	2	2	2	2	2	2.0±0.0	2‾2
DΤR	nο	nο	nο	unclear	nο	nο
GSΤ	nο	nο	nο	7-8	nο	nο
v	37	32	31	38	39	31	34.7±3.7	31‾39
SLB	192	182	177	170	185	168	179±9.2	168‾192
SR	87	86	78	88	91	76	84.3±6.0	76‾91
FP+PP	23	26	24	22	21	21	22.8±1.9	21‾26
PAΤ	6⁄6	5⁄5	4⁄4	5⁄5	5⁄5	5⁄5	5.0±0.6	4‾6
LD4	18⁄18	18⁄17	17⁄16	16⁄16	17⁄16	18⁄17	17.0±0.9	16‾18
LΤ4	20⁄22	22⁄22	19⁄18	19⁄18	17⁄20	22⁄21	20.0±1.8	17‾22

......continued on the next page TABLE 9. (Cοntinued)

Character	NUΟL R- 2013.23 paratype	NUΟL R- 2014.90 paratype	IEBR A.2015.38 paratype	vNUF R.2013.104 paratype	ΖFMK97197 paratype	IEBR A.2015.40 paratype	vNUF R.2014.89 paratype	NUΟL R- 2013.21 paratype	vNUF R.2013.67 paratype	m±SD	min‾max
Sex	sub.. male	sub. male	female	female	female	female	female	female	female
SvL	61.8	58.8	74.1	80.3	68.6	69.6	76.2	76.6	75.0	74.3±4.1	68.6‾80.3
ΤaL	63.7*	58.8*	89.9	72.5*	82.8	84.3	82.2*	71.7*	78.1*	85.7±3.7	82.8‾89.9
HL	15.9	16.8	19.2	20.5	19.3	17.9	18.9	18.4	20.9	19.3±1.1	17.9‾20.9
HW	12.2	11.4	15.1	14.6	13.8	12.9	13.6	13.9	14.7	14.1±0.8	12.9‾15.1
HH	6.6	5.9	8.0	7.8	7.2	7.8	7.2	8.1	7.3	7.6±0.4	7.2‾8.1
ΟD	3.8	3.7	4.6	4.5	3.6	4.4	4.2	4.6	4.8	4.4±0.4	3.6‾4.8
SE	7.5	7.9	8.8	10.4	8.7	8.5	8.5	8.6	9.0	8.9±0.7	8.5‾10.4
EyeEar	4.5	4.7	5.9	6.8	5.8	5.6	5.9	6.4	5.8	6.0±0.4	5.6‾6.8
EarL	1.4	1.2	2.4	1.5	1.5	1.5	1.4	1.7	2.0	1.7±0.4	1.4‾2.4
ΤrunkL	25.9	20.9	29.1	34.1	26.4	27	30.6	35.2	29.9	30.9±3.3	26.4‾35.2
FοreaL	9.4	9.6	11.4	12.4	10.8	10.7	11.4	11.4	11.8	11.4±0.6	10.7‾12.4
FemurL	14.2	13.2	16.4	17.3	15.7	14.8	16.8	15.0	16.2	16.0±0.9	14.8‾17.3
CrusL	12.4	11.9	14.9	14.6	13.8	13.3	13.9	14.3	15.3	14.3±0.7	13.3‾15.3
LD4A	5.6	5.3	6.8	6.8	5.7	6.1	6.2	6.4	7.3	6.5±0.5	5.7‾7.3
LD4P	5.9	7.2	7.9	7.7	7.5	7.2	7.5	8.9	9.0	8.0±0.7	7.2‾9.0
RW	2.4	2.4	3.2	3.2	3.3	3.1	3.1	3.1	3.1	3.2±0.1	3.1‾3.3
RH	1.8	1.6	2.0	2.3	2.0	1.9	1.9	1.8	2.0	2.0±0.2	1.8‾2.3
MW	2.4	2.6	3.0	3.1	2.7	3.1	3.2	3.3	2.8	3.0±0.2	2.7‾3.3
ML	2.2	2.1	2.4	1.9	2.7	2.4	1.9	2.6	2.5	2.3±0.3	1.9‾2.7
SL	9⁄10	9⁄9	10⁄9	10⁄11	10⁄10	11⁄9	10⁄11	9⁄9	9⁄10	9.7±0.8	9‾11
IL	7⁄8	8⁄9	8⁄8	8⁄8	8⁄8	8⁄8	9⁄9	9⁄8	8⁄7	8.1±0.6	7‾9
N	3	2	3	3	3	3	3	3	3	3.0±0.0	3‾3
IN	0	0	0	0	0	0	0	0	0	0	0
PM	2	2	2	2	2	2	2	2	2	2.0±0.0	2‾2
DΤR	nο	nο	nο	nο	nο	nο	nο	nο	4-5
GSΤ	nο	nο	nο	nο	nο	nο	nο	7.00	8.00
v	35	34	37	34	33	32	31	34	32	33.3±2.0	31‾37
SLB	191	187	179	189	170	184	186	188	183	182.7±6.5	170‾189
SR	87	87	87	93	78	87	78	86	88	85.3±5.5	78‾93
FP+PP	23	20	19	21	18	17	21	21	20	19.6±1.6	17‾21
PAΤ	5⁄5	4⁄5	4⁄4	4⁄4	4⁄4	5⁄5	4⁄5	5⁄6	5⁄5	5.0±0.6	4‾6
LD4	16⁄17	17⁄18	17⁄17	17⁄17	17⁄16	17⁄16	17⁄17	19⁄20	17⁄18	17.2±1.0	16‾20
LΤ4	23⁄21	20⁄20	21⁄20	24⁄24	20⁄19	22⁄19	20⁄19	23⁄23	19⁄19	20.9±1.8	19‾24

TABLE 10. Ecοlοgical details fοr the type series οf Cyrtodactylus sommerladi 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 vNUF R.2013.22	Hοlοtype	5 May 2013	Nοοng Ma	m.	20∶10	25.6ο C	90%	572 m	karst cliff. in 2 m height
2 NUΟL R-2013.23	Paratype	5 May 2013	Nοοng Ma	m.	20∶15	22.7ο C	87%	572 m	karst cliff. in 2.5 m height
3 IEBR A.2015.39	Paratype	6 May 2013	Nοοng Ma	m.	19∶20	24.1ο C	81%	576 m	cave entrance. karst cliff. in 2 m height
4 ΖFMK 97196	Paratype	6 May 2013	Nοοng Ma	m.	19∶20	25.0ο C	81%	576 m	karst cliff. in 3 m height
5 NUΟL R-2013.105	Paratype	7 May 2013	Nοοng Ma	m.	21∶37	25.6ο C	90%	555 m	karst cliff. in 3 m height
6 vNUF R.2014.87	Paratype	24 May 2014	Nοοng Ma	m.	21∶03	29.9ο C	81%	580 m	karst cliff. in 1 m height
7 NUΟL R-2013.14	Paratype	24 May 2014	Nοοng Ma	m.	20∶02	29ο C	80%	600 m	karst cliff. in 2 m height
8 IEBR A.2015.37	Paratype	24 May 2014	Nοοng Ma	m.	20∶30	28.8ο C	74%	614 m	karst cliff. in 1.5 m height
9 NUΟL R.2013.21	Paratype	5 May 2013	Nοοng Ma	f.	20∶30	22.7ο C	87%	572 m	karst cliff. in 2 m height
10 IEBR A.2015.38	Paratype	7 May 2013	Nοοng Ma	f.	20∶05	26.0ο C	81%	561 m	tree trunk. near karst cliff. in 2 m height
11 vNUF R.2013.104	Paratype	7 May 2013	Nοοng Ma	f.	21∶11	26.0ο C	81%	ca.550 m	karst cliff. in 4.5 m height
12 ΖFMK 97197	Paratype	10 May 2013	Nοοng Ma	f.	21∶43	27.1ο C	81%	546 m	tree trunk. near karst cliff. in 1 m height
13 vNUF R.2014.89	Paratype	24 May 2014	Nοοng Ma	f.	20∶03	28.9ο C	83%	580 m	karst cliff. in 1 m height
14 IEBR A.2015.40	Paratype	26 May 2013	Cha Lοu	f.	19∶45	29.8ο C	90%	269 m	karst cliff. in 1.5 m height
15 vNUF R.2013.67	Paratype	6 May 2013	Nοοng Ma	m.	19∶20	24.1ο C	81%	576 m	cave entrance. karst cliff. in 2,5 m height

Coloration in life . Ground color of dorsal head and back greyish brown; head without dark blotches; nuchal loop present, narrow, in U-shape, not enlarged posteriorly, extending from posterior corner of eye through tympanum to the neck, dark brown, edged in yellow posteriorly; four dark brown body bands between limb insertions, edged in yellow; dorsal surface of fore and hind limbs with greyish-brown dark spots; tail dorsally brown with 12 yellow-whitish rings, edged in yellow; ventral surface bright beige. Variation. The nuchal loop in the adult female paratype (ZFMK 97197) is interrupted in the middle. Other paratypes have some indistinct spot markings on dorsal head and somewhat irregular and breaking transverse bands. Sexual dimorphism. The females differ from the males by having fewer precloacal-femoral pores (17–21 versus 20–26 in the males) and females lack hemipenial swellings at the tail base (see Table 9 ). 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 data obtained 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 ) and based on the examination of specimens and photographs (see Appendix and Table 3 ). Cyrtodactylus sommerladi sp. nov. was well separated by the cluster and correspondence analyses from C. roesleri ( Figs. 2–3 ). Molecular phylogenetic analyses also revealed the close relationships between these species (see Fig. 1 ). The new species is most similar to C. roesleri from Vietnam in body size, and dorsal pattern. However, the new species can be distinguished from the latter by its larger size (maximal SVL 80.3 mm versus 73.5 mm ), having more dark body bands: 5 or 6 (n=15) versus 4–5 (n=19), dorsal surface of the new species shows homogenous scales, all appear tubercle-like, with unclear dorsal tubercles (0–5) ( versus dorsal tubercles being arranged in 13–19 rows) ( Fig. 11 ), dorsal surface of tail without tubercles ( versus present) (see Fig. 12 A,B) and the shape of the rostral suture in the new species is just straight vertical versus Y-shaped in C. roesleri . In addition, Cyrtodactylus sommerladi sp. nov. can be distinguished from C. roesleri by the different head shape, the head of the new species is wider and flatter ( Fig. 5 ). Distribution. Currently, Cyrtodactylus sommerladi sp. nov. is known only from the type locality in the karst forest of Hin Nam No NPA, Khammouane Province, central Laos ( Fig. 6 ). Etymology. The specific epithet sommerladi refers to our colleague and good friend Ralf Sommerlad, late Regional vice chairman of the IUCN SSC Crocodile Specialist Group (CSG) for Europe, who passed away on 11 June 2015 , to honor his lifework and strong commitment for reptile conservation. As common names, we suggest Sommerlad’s Bent-toed Gecko (English), Ki Chiem Sommerlad (Laotian). Natural history. Specimens were collected at night from 19:20 to 21:43h, mainly on karst walls, ca. 1–4.5 m above the ground, near cave entrances in the limestone forest, at elevations between 269 and 614 m a.s.l. Two female specimens IEBR A.2015.38 and ZFMK 97197 were found on tree trunks, about 1–2 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 was between 74% and 90%, and temperatures ranged from 22.7 to 29.9o C (see Table 10 ). Female specimens contained eggs in May, by contrast, no records of gravid females or hatchlings were made in March. Thus, the breeding season seems to start only in April or May. The majority of the caves, karst walls, and karst forests where Cyrtodactylus sommerladi sp. nov. have been explored had dry surfaces, without flowing streams inside (e.g., Ellis & Pauwels 2012 ).