Cyrtodactylus martini, another new karst-dwelling Cyrtodactylus Gray, 1827 (Squamata: Gekkonidae) from Northwestern Vietnam Author Tri, Ngo Van text Zootaxa 2011 2834 33 46 journal article 10.5281/zenodo.277341 eff30799-13bb-4407-9d8b-87f65577e062 1175-5326 277341 Cyrtodactylus martini sp. nov. Plates 1–3 Holotype . Adult male (University of Natural Sciences [ UNS ] 0471) collected on 6 May 2009 by Ngo Van Tri from the karst forest around Lai Chau Town, Lai Chau province, northwestern Vietnam ( 22°23’N , 103°24’E ; ca. 1000 m a.s.l.). Paratypes . All paratypes were collected from the same locality as the holotype . UNS 0467–0469 were collected on 5 May 2009 . UNS 0470–0472 were collected on 6 May 2009 . Diagnosis. Cyrtodactylus martini differs from all other congeners by the following combination of characters: maximum SVL 96.2 mm ; original tail long (TL/SVL: 1.07); symmetrical or subsymmetrical reticulations on top of head; no nuchal loop; dorsal pattern consisting of four to six irregular, yellowish–white bands between limb insertions; six or seven white rings on original tail; 1–2 intersupranasal scales; 9–11 supralabials; 8–10 infralabials; 15– 16 interorbital scales on the frontal bone; 17–21 scales between eye and nostril; 39–43 rows of ventral scales between ventrolateral folds; 16–19 irregular, longitudinal rows of smooth or conical tubercles at midbody between the lateral folds; 24–27 paravertebral tubercles; four precloacal pores separated medially by one poreless scale in males, no pores in females; 14–18 enlarged scales beneath thighs which are continuous with precloacal pores; 15– 18 subdigital lamellae on first toe; 22–24 subdigital lamellae on fourth toe; no enlarged scales on heel; two medial longitudinal rows of enlarged, irregularly shaped subcaudal scales. PLATE 1. A: Male holotype (UNS 0472) of Cyrtodactylus martini sp. nov . ; B: Male paratype (UNS 0471) of Cyrtodactylus martini sp. nov . in its natural habitat. Description of holotype . Adult male, SVL: 88.0 mm ( Plate 1A ). Head moderately long (HeadL/SVL: 0.26), relatively narrow (HeadW/HeadL: 0.68), depressed (HeadH/HeadL: 0.37), distinct from neck. Lores and interorbital region inflated, canthus rostralis indistinct, frontonasal region concave. Snout elongate (SnEye/HeadL: 0.39), pointed; longer than eye diameter (OrbD/SnEye: 0.60); scales on snout and forehead small, rounded, granular, homogeneous; scales on snout larger than those on occipital region. Eye large (OrbD/HeadL: 0.23); eye red, pupil with crenulated margins when constricted and round when dilated; supraciliaries short, bearing tiny conical spines posteriorly. Ear opening oval, oblique, small (EarL/HeadL: 0.08); eye to ear distance greater than diameter of eye (EyeEar/OrbD: 1.26). Rostral incompletely divided by a shallow dorsal groove; two enlarged supranasals in broad contact with one smaller intersupranasal ( Plate 2A ), rostral in contact with first supralabial and nostril; nostril oval, surrounded by supranasal, rostral, first supralabial, and three enlarged postnasals; 3–4 rows of small scales separating orbit from supralabials. Mental triangular, wider ( 3.2 mm ) than deep ( 2.1 mm ); one pair of enlarged postmentals, in broad contact medially, bordered anteromedially by mental, anterolaterally by first infralabial, posterolaterally by five enlarged lateral chinshields ( Plate 2B ). 10 (R), 11(L) supralabials; 9 (R, L) infralabials; 15 interorbital scale rows on the frontal bone; 18 scales between eye and nostril in straight line. PLATE 2. A: Rostral scales of holotype Cyrtodactylus martini sp. nov . ; B : Mental scales of holotype Cyrtodactylus martini sp. nov . ; C : Dorsal tubercle rows of holotype Cyrtodactylus martini sp. nov . ; D : Tail base of Cyrtodactylus martini sp. nov . PLATE 3.A: Precloacal region of holotype Cyrtodactylus martini sp. nov . showing four precloacal pores separated by one poreless scale; B: subcaudal scales of holotype Cyrtodactylus martini sp. nov . showing two medial rows of enlarged scales scattered with slightly enlarged scales. Body slender, elongate (TrunkL/SVL: 0.43), ventrolateral folds with small, smooth tubercles. Dorsal scales granular to weakly conical; regularly distributed tubercles (4–7 times size of adjacent scales) extending from occipital region to base of tail; each tubercle weakly keeled; tubercles in 16 rows at midbody between ventrolateral folds ( Plate 2C ), smallest on flanks and occipital region; 24 paravertebral tubercles. Ventral scales larger than those of dorsum, smooth, relatively round, subimbricate, largest posteriorly; 43 scale rows across belly between ventrolateral folds; gular region with relatively homogeneous, smooth scales. No precloacal groove, about 40 smooth enlarged scales in the precloacal area, four of which bear pores, separated medially by one poreless scale ( Plate 3A ); 16 (R) and 14 (L) enlarged femoral scales beneath thighs continuous with enlarged precloacal scales ( Plate 3A ). Scales on palm and hind limbs smooth, granular, scattered with weakly keeled tubercles, tubercles smaller than those on dorsum, decreasing in size on dorsal region of manus and pes. Fore- and hindlimbs moderately slender (ForeL/SVL: 0.15; CrusL/SVL: 0.17); digits moderately robust, strongly inflected at basal interphalangeal joints, all bearing slightly curved claws; basal subdigital lamellae nearly as broad as digit, without scansorial surface: 7–8–8–10 –8 manus; 6–9–10–11–10 pes; narrow lamellae distal to digital inflection: 9–11–12–12–11 manus; 9–10–13–13–13 pes; one or three rows of small, non lamellar granules between basal and distal lamellar series; interdigital webbing present but weakly developed. Relative length of digits in mm: (manus) III (8.4)> IV (8.2)> II (7.1)>V (6.9)> I (5.2); (pes): V (10.0)> IV (9.7)> III (8.5)> II (7.1)>I (5.0). Tail length 94.2 mm , slender, tapering to a point; longer than SVL (TailL/SVL: 1.15); two whitish tubercles at base; 6–7 rows of scales in each segment and one weakly keeled or smooth tubercle on both sides of first three segments, except for basal-most segment where six parasagittal rows of enlarged, weakly keeled tubercles continuous from dorsum ( Plate 2D ); subcaudal scales smooth, juxtaposed, two medial longitudinal rows of enlarged, irregularly shaped extending length of tail; dorsal caudal scales flat, smooth, heterogeneous. Coloration in life (Plate 1A). Head chocolate brown with yellowish or yellowish–white symmetrical reticulations, no nuchal loop, four or six yellowish–white narrow bands on dorsum between limb insertions, one anterior to forelimbs forming a V–shape, one immediately posterior to hind limbs; flanks scattered with yellowish or cream tubercles. Original tail chocolate brown with seven incomplete, white rings, regenerated portion reddish or dark brown scattered with small, yellowish–white spots; dorsal surface of limbs lighter in color, scattered with yellowish–white tubercles. It is suspected that dorsal color can change depending on color of substrate ( Plate 1B ). Variation. Color pattern variations of Cyrtodactylus martini are shown in Plate 1C . Mensural differences are presented in Table 1 . TABLE 1. Mensural and meristic data for the type series of Cyrtodactylus martini sp. nov. Abbreviations are stated in materials and methods; all measurements in mm; (F)=female, (M)=male; Reg=regenerated tail.

Sex	Holotype Paratype UNS 0 471 UNS0467 M F	Paratype UNS 0 468 M	Paratype UNS 0 469 M	Paratype UNS 0 470 F	Paratype UNS 0 472 M	min-max x ±S.D. n = 6
SVL	88.0 96.2	90.1	85.4	64.4	88.1	64.4–96.2 85.4 ± 10.9
HeadL	22.8 24.8	24.7	22.3	18.1	22.8	18.1–24.8 22.6 ± 2.4
HeadW	15.4 16.6	16.0	14.4	11.3	15.4	11.3–16.6 14.9 ± 1.9
HeadH	8.5 9.3	9.3	8.6	5.7	8.5	5.7–9.3 8.3 ± 1.3
EarL	1.8 2.3	2.0	2.0	1.1	1.9	1.1–2.3 1.9 ± 0.4
OrbD	5.3 6.7	6.2	6.0	4.5	6.1	4.5–6.7 5.8 ± 0.8
NarEye	6.4 7.0	6.9	6.8	5.4	6.7	5.4–7.0 6.5 ± 0.6
SnEye	8.8 9.6	10.0	9.1	7.3	9.1	7.3–10.0 9.0 ± 0.9

continued next page Holotype Paratype Paratype Paratype Paratype Paratype min-max TABLE 1. (continued) UNS 0 471 UNS0467 UNS 0 468 UNS 0 469 UNS 0 470 UNS 0 472 x ±S.D. EyeEar 6.7 7.6 6.0 6.1 5.3 6.8 5.3–7.6 6.4 ± 0.8 Etymology. The specific epithet honors Mr. Shaun Martin, the Director of Education Program for Nature— WWF in the U.S.A. who sponsored a small grant in 2009 for gecko expeditions in Vietnam . Suggested English name: Martin’s Bent–toed Gecko; Vietnamese name: “Thằn lằn chân ngón Martin”.

InterNar 2.0 2.5	2.4	2.4	1.8	2.0	1.8–2.5 2.2 ± 0.3
InterOrb 7.0 7.1	7.2	6.8	5.2	6.8	5.2–7.2 6.7 ± 0.7
TrunkL 37.8 40.3	38.1	36.4	26.1	39.3	26.1–40.3 36.3 ± 5.2
BW 13.7 20.9	12.6	13.0	9.7	12.4	9.7–20.9 13.7 ± 3.8
ForeaL 13.0 14.4	13.9	14.0	10.0	13.7	10.0–14.4 13.2 ± 1.6
CrusL 14.8 18.0	17.0	16.6	12.5	16.2	12.5–18.0 15.9 ± 2.0
TL 94.2 67.5 (R)	79.2(R)	80.7(R)	76.0	101.2	76.0–101.2 79.8 ± 16.6
TW 6.6 6.9	7.7	6.9	4.6	7.8	4.6–7.8 6.8 ± 1.2
SupL 10/11 10/9	9/10	11/11	10/10	11/10	9–11/9–11
Infra 9/9 8/9	8/9	10/10	10/10	10/9	8–10/9–10
TubR 16 18	16	19	16	16	16–19
Paravert 24 26	25	26	24	27	24–27
VenS 43 42	41	43	40	39	39–43
Pre–pores 2 + 2 0	2 + 2	2 + 2	0	2 + 2	0 (females) or 4 (males)
EnlfemS 16/14 14/16	15/16	17/18	17/17	16/15	14–17/14–18
PCT 2/2 2/2	2/2	2/2	2/1	2/2	2/1–2
NSF I: 7+9 I: 7+10 II: 8+11 II: 8+9 III: 8+12 III: 8+11 IV: 10+12 IV: 8+11 V: 8+11 V: 9+9	I: 7+9 II: 7+10 III: 7+12 IV: 8+12 V: 9+11	I: 7+9 II: 7+11 III: 8+11 IV: 10+12 V: 8+11	I: 8+9 II: 9+10 III: 8+12 IV: 7+12 V: 9+10	I: 8+9 II: 9+10 III: 9+13 IV: 11+12 V: 10+11	I: 16–17 II: 17–19 III: 19–22 IV: 19–23 V: 18–21
NST I: 6+9 I: 7+9 II: 9+10 II: 9+10 III: 10+13 III: 10+11 IV: 11+13 IV: 10+12 V: 10+13 V: 9+12	I: 7+8 II: 8+9 III: 9+12 IV: 10+13 V: 10+13	I: 8+10 II: 8+9 III: 8+12 IV: 10+13 V: 8+13	I: 7+9 II: 7+10 III: 9+11 IV: 10+12 V: 11+12	I: 8+10 II: 8+11 III: 11+12 IV: 11+13 V: 9+13	I: 15–18 II: 17–19 III: 20–23 IV: 22–24 V: 21–23
FronS 15 15	16	16	15	16	15–16
Orb–nosS 18 19	18	21	19	17	17–21
IntersupS 1 2	1	1	1	1	1–2
Enlar–chin 5 5	5	5	6	5	5–6

Distribution and natural history. The karst forest surrounding Lai Châu Town has long been degraded and converted into agricultural lands and farms, rendering the remaining karst forest as secondary forest, with a number of peaks rising to approximately 1200–1300 m . a.s.l ( Plate 4A ). The karst outcroppings are isolated from Hoang Lien Son Mountain and are formed by underground water movement inside deep caves. Cyrtodactylus martini were found at night between 1900–2030h, mostly within crevices at the entrance of a small, dry cave ( Plate 4B ) but some individuals were also found on the karst surface outside the cave at night. Paratype UNS 0 467, an adult female, was gravid with two eggs. PLATE 4. A: Karst forest, macrohabitat of Cyrtodactylus martini sp. nov . in Lai Chau province; B : Karst crevices, microhabitat of Cyrtodactylus martini sp. nov. in Lai Chau province. Comparison with other species. Cyrtodactylus martini sp. nov. differs from C. badenensis Nguyen et al . , C . darmandvillei (Weber) , C. eisenmanae Ngo, C . grismeri Ngo , C. jarakensis Grismer et al . , C . jellesmae (Boulenger) , C . laevigatus (Darevsky, C . semenanjungensis Grismer & Leong , C . sermowaiensis (De Rooij) , C . thirakhupti Pauwels et al . , and C . wallacei Hayden et al . by the presence (versus absence) of precloacal pores in males. It further differs from Cyrtodactylus aaroni Günther & Rösler , C . aequalis Bauer , C . agusanensis (Taylor) , C . annandalei Bauer , C. auribalteatus Sumontha et al . , C . baluensis Mocquard , C. batucolus Grismer et al . , Cyrtodactylus bichnganae Ngo & Grismer , C. biordinis Brown & McCoy , C . brevipalmatus (Smith) , C . caovansungi Orlov et al . , C . capreoloides Rösler et al . , C . chanhomeae Bauer et al . , C . consobrinoides (Annandale) , C . consobrinus Malkmus , C . deveti (Brongersma) , C. epiroticus Kraus , C . erythrops Bauer et al . , C . feae (Boulenger) , C . fumosus (Müller) , C . gubernatoris (Annandale) , C. halmahericus (Mertens) , C . huynhi Ngo & Bauer , C . interdigitalis Ulber , C . jarujini Ulber , C. klugei Kraus , C . loriae (Boulenger) , C . louisiadensis (De Vis) , C . macrotuberculatus Grismer & Ahmad , C . marmoratus (Gray) , C . mimikanus (Boulenger) , C . novaeguineae (Schlegel) , C . phongnhakebangensis Ziegler et al . , C . pulchellus Gray , C . redimiculus King , C. robustus Kraus , C . roesleri Ziegler et al . , C . russelli Bauer , C . sadleiri Wells & Wellington , C . salomonensis Rösler et al . , C. seribuatensis Youmans & Grismer , C . serratus Kraus , C . slowinskii Bauer , C . takouensis Ngo & Bauer, C. tamaiensis (Smith) , C . tigroides Bauer et al . , C . tiomanensis Das & Lim , C. tripartitus Kraus , C . tuberculatus (Lucas & Frost) , C . variegatus (Blyth) , C . wetariensis (Dunn) , C. yangbayensis Ngo & Chan , C. ziegleri Nazarov et al . and C. zugi Oliver et al . by the absence of femoral pores. Cyrtodactylus martini sp. nov. can be distinguished from C . aurensis Grismer , C. cavernicolous Inger & King , C. macrotuberculatus Grismer & Norhayati , C. marmoratus Gray , C. pubisulcus Inger , C. pulchellus Gray , C. semenanjungensis Grismer & Leong , C. stresemanni Rösler & Glaw , and C . tiomanensis Das & Lim by lacking a precloacal groove. Cyrtodactylus martini sp. nov. is similar to C. zhaoermii Lei & Hui in having precloacal pores (4 vs. 3–4), but these pores are smaller and separated by one poreless scale, it has a larger size ( 64.4–96.2mm vs. 48.1–59.8mm ), and lacks distinct tail segments, bearing large, posteriorly directed tubercles in whorls, numbering nine to ten per caudal annulus. Cyrtodactylus martini sp. nov. is superficially similar to C . consobrinus (Peters) in having reticulations on the head but differs from the latter by being significantly smaller in size ( 64.4–96.2mm vs. 125mm SVL), having fewer precloacal pores (4 vs. 9–11), fewer tubercle rows (16–19 vs. 18–20), and fewer lamellae under the fourth toe (16– 19 vs. 22–28), lacking tubercles on the lateral skin folds, and having the subcaudal scales consisting of two medial longitudinal rows of enlarged, irregularly shaped scales as opposed to a single medial row of enlarged scales. Cyrtodactylus martini sp. nov. also resembles C. wayakonei Nguyen et al. in having reticulations on top of the head but differs by being larger in size (maximum 96.2mm vs. maximum 86.8 mm ), having fewer precloacal pores in males (four separated by one poreless scale vs. 6–8 in a chevron), higher number of ventral scales (39–43 vs. 31– 35), intersupranasals (1–2 vs. 0), number of subdigital lamellae on fourth toe (22–24 vs. 19–20), and lower number of paravertebral tubercles (24–27 vs.> 30 as counted from the published figure of the holotype of C. wayakonei , IEBR A.2020.01). Other differences between the two species are summarized in Table 2. Cyrtodactylus martini sp. nov. differs from other Sunda shelf Cyrtodactylus including C . leegrismeri Chan & Norhayati , C. baluensis (Mocquard) , C. batucolus Grismer et al. , C. brevipalmatus (Smith) , C. elok Dring , C . fumosus (Müller) , C . ingeri Hikida , C . jarakensis Grismer et al. , C. lateralis (Werner) , C . malayanus (De Rooij) , C . matsuii Hikida , C. nuaulu Oliver et al. , C. oldhami (Theobald) , C . pantiensis Grismer et al. , C. peguensis (Boulenger) , C . quadrivirgatus Taylor , C. seribuatensis , C. sworderi (Smith) , C. tautbatorum Welton et al . , C . yoshii Hikida by the presence of reticulations on top of head and the absence of caudal tubercles. Comparison among other species of Cyrtodactylus with precloacal pores in Indochina is presented in Table 2.