A new species of bent-toed gecko, genus Cyrtodactylus Gray, 1827 (Reptilia: Squamata: Gekkonidae), from Jawa Timur Province, Java, Indonesia, with taxonomic remarks on C. fumosus (Müller, 1895) Author Hartmann, Lukas Author Mecke, Sven Author Kieckbusch, Max Author Mader, Felix Author Kaiser, Hinrich text Zootaxa 2016 4067 5 552 568 journal article 51140 10.11646/zootaxa.4067.5.2 b48eb399-7654-426c-8804-6805960b2b52 1175-5326 267812 F10E6E61-6D1B-43D9-AFFD-7996E0811EE9 Cyrtodactylus klakahensis sp. nov. English: Klakah bent-toed gecko; German : Klakah Bogenfinger-Gecko Chresonym: Cyrtodactylus fumosus — Mertens 1934 , Archiv für Hydrobiologie : 689 Figures 1 –3; Table 2 & 3 Holotype . SMF 22476 ( Figures 1 & 3A–C; Table 2 ), an adult male, collected in 1928 or 1929 by members of the Deutsche Limnologische Sunda-Expedition at Klakah, Lumajang Regency, Jawa Timur Province, Java, Indonesia . Paratypes . SMF 22477 ( Figure 2 A; Table 2 ), an adult male; SMF 22478 ( Figure 2 B; Table 2 ), an adult female; and SMF 22479 ( Figure 2 C; Table 2 ), an unsexed juvenile. All specimens have the same collection information as the holotype . TABLE 1. Metric and meristic characters with abbreviations used in this study. Character Abbreviation Definition Snout-vent length SVL from tip of snout to cloaca Axial length AxialL from axilla to groin Tail length TailL length of original tail, from cloaca to tip of tail Arm length ArmL from insertion of antebrachium with body wall to claw of longest finger Leg length LegL from insertion of femur with body wall to claw of longest toe Head length HeadL from tip of snout to articulation of quadrate bone Head width HeadW measured at level of ear openings Head height HeadH measured at level of ear opening Snout length SnoutL from tip of snout to anterior margin of orbit Orbit-ear length OrbEarL from posterior margin of orbit to anterior margin of ear opening Orbital diameter OrbD from anterior to posterior margin of orbit Ear length EarL from anterior to posterior margin of ear opening Mental length MentalL maximum length of mental shield Mental width MentalW maximum width of mental shield Dorsal tubercles DTR number of tubercle rows on dorsum at midbody, counted in one row between lateral folds Paravertebral tubercles PVT number of tubercles counted in a longitudinal row between posterior insertion of fore limb and anterior insertion of hind limb Ventral scales VS number of ventral scales at midbody, counted in one row between lateral folds Precloacofemoral scales PFS number of enlarged precloacofemoral scales, counted along lowest, pore- bearing row Precloacofemoral pores PFP number of precloacofemoral pores Postcloacal tubercles PCT number of postcloacal tubercles Subdigital lamellae under 4th LT 4 subdigital scales under 4th toe, counted from first enlarged scale (true toe lamellae) on lower side of toe to scale proximal to apical scale Supralabial scales 1 SupraLab1 labial scales of upper jaw, beginning with first enlarged scale bordering rostral shield, ending with last enlarged scale bordering labial angle Supralabial scales 2 SupraLab2 labial scales of upper jaw, beginning with first enlarged scale bordering rostral shield, ending with enlarged scale below anterior margin of orbit Infralabial scales InfraLab labial scales of lower jaw, beginning with first scale bordering mental shield, ending with last enlarged scale bordering labial angle Internasal scales InterNas number of scales between rostronasals, bordering rostral shield Supraciliar scales SC number of enlarged scales extending from anterior-ventral to posterior- medial edge of orbit Interorbital scales IOS number of scales counted in one row between medial edges of orbits across occiput Gular scales GulS number of gular scales bordering pair of 1st postmentals (excluding enlarged second 2nd postmentals) TABLE 2. Metric (in mm) and meristic data for the type series of Cyrtodactylus klakahensis sp. nov. Abbreviations are defined in Table 1.

Holotype SMF 22476	Paratype SMF 22477	Paratype SMF 22478	Paratype SMF 22479
Sex	male	male	female	unsexed juvenile
SVL	68.1	61.1	67.2	37.2
AxialL ArmL	31.9 25.9	27.3 21.7	28.7 24.1	14.0 13.6
LegL	36.3	28.9	34.0	16.8
HeadL	18.1	16.8	16.6	10.4
HeadW HeadH	10.6 7.0	10.4 7.2	10.5 8.3	6.1 4.2
SnoutL	7.6	7.0	7.9	4.2
OrbEarL	5.9	5.6	6.7	2.9
OrbD EarL	3.7 1.3	3.4 1.3	3.4 1.4	1.9 0.5
DTR	19	17	17	15
PVT	26	29	33	26
VS PFS	38 40	36 38	36 38	35 38
PFP	37	38	0	0
LT 4 (proximal) LT 4 (distal) LT 4 SupraLab1 SupraLab2 InfraLab	9 11 20 R10 L10 R6 L5 R9 L9	7 10 17 R11 L10 R5 L5 R9 L10	7 10 17 R10 L11 R5 L6 R8 L10	8 9 17 R11 L10 R5 L6 R9 L8
GulS	4	4	4	4

Definition. Cyrtodactylus klakahensis can be distinguished from all other congeners occuring in the Greater Sunda Islands (including Sulawesi) and Lesser Sunda Islands by the following combination of characters: (1) a deep precloacal groove in males, (2) three rows of enlarged precloacofemoral scales, of which the third row bears 37–38 pores in males, (3) three or four rows of posterior precloacal scales, (4) raised and strongly keeled dorsal tubercles in 15–19 rows at midbody, (5) an indistinct lateral fold, (6) 17–20 subdigital lamellae under the 4th toe, and (7) subcaudal scales which are not transversely enlarged. Comparison with other species. The new species can be readily distinguished from the Greater Sunda Island congeners Cyrtodactylus batik Iskandar et al ., 2011 , C . consobrinus ( Peters, 1871 ) , C . ingeri Hikida, 1990 , C . jellesmae ( Boulenger, 1897 ) , C . lateralis ( Werner, 1896 ) , C . malayanus (de Rooij, 1915 ), C . matsuii Hikida, 1990 , C . semiadii Riyanto et al ., 2014 , C . quadrivirgatus Taylor, 1962 , C . wallacei Hayden et al ., 2008 , and C . yoshii Hikida, 1990 , and from the Lesser Sunda Island species C . darmandvillei ( Weber, 1890 ) , C . gordongekkoi ( Das, 1993 ) , C . laevigatus Darevsky, 1964 , and C . wetariensis ( Dunn, 1927 ) by the presence of a deep precloacal groove in males, bearing five pores. Data in Table 3 allow a detailed comparison of C. klakahensis with all other Sundanese bent-toed geckos, and additional characters to distinguish the new species from taxa without a precloacal groove are listed therein. In the following comparisons with species that also possess a precloacal groove, including Cyrtodactylus agamensis ( Bleeker, 1860 ) , C . baluensis ( Mocquard, 1890 ) , C . cavernicolus Inger & King, 1961 , C . celatus Kathriner et al ., 2014 , C . fumosus , C . marmoratus , C . psarops Harvey et al . 2015 , C . pubisulcus Inger, 1958 , C . semicinctus Harvey et al . 2015 , and C . spinosus Linkem et al ., 2008 , the characters for C . klakahensis are provided in parentheses. Cyrtodactylus agamensis (data from Rösler et al . 2007 , based on the single known specimen, an adult female) possesses 67 (35–38) VS; 26 (17–20) LT4; 54 enlarged PFS (38–40); and a single enlarged scale in the precloacal groove (five scales in the precloacal groove). Cyrtodactylus baluensis (data from Hikida 1990 and obtained from specimens listed in the Appendix) possesses precloacal scales that are separated from the femoral scales (enlarged PFS in both sexes); 4–10 precloacal- and 9–11 femoral pores in males (37–38 PFP in a continuous series in males); no posterior precloacal scales (posterior precloacal scales present); and enlarged subcaudals (enlarged subcaudals absent). Cyrtodactylus cavernicolus (data from Grismer & Leong 2005 ) possesses 51–58 (35–38) VS; 22–26 (17–20) LT4; no enlarged femoral scales (enlarged PFS in three rows present in both sexes); no femoral pores (pores, including those on the femur, present in males); and dorsal bands (dorsal blotches). Cyrtodactylus celatus (data from Kathriner et al . 2014 ; Rösler & Kaiser, in press.; and obtained from specimens listed in the Appendix) is a small-sized species with adult SVL of 38.4−43.6 mm ( 61.1−68.1 mm ) that possesses no femoral pores (pores, including those on the femur, present in males). Cyrtodactylus fumosus (data obtained from specimens listed in the Appendix) possesses widely scattered, roundish, flat, and smooth dorsal tubercles in 4–7 rows at midbody (closely arranged, trihedral, raised, and strongly keeled dorsal tubercles in 15–19 rows at midbody); no or a low number of scattered tubercles on the limbs (limbs strongly tuberculated); a total number of 16 pores, ten of which are precloacal pores, separated from three femoral pores by ten enlarged pore-less scales 1 in males (37–38 PFP in a continuous series in males); and an ear opening forming a horizontal cleft2 (ear opening vertically elongated). Cyrtodactylus marmoratus (data from Rösler et al . 2007 and obtained from specimens listed in the Appendix) possesses 38–47 (35–38) VS; 48–56 3 (38–40) enlarged PFS; 45–53 4 (37–38) PFP in males; and pores present in females (no pores present in females). Based on our examinations, C . marmoratus also lacks posterior precloacal scales (posterior precloacal scales present), possesses a different arrangement of postmental and gular scales, and has differently shaped postcloacal tubercles. The reader is referred to Figure 3 for a comparison of the shape and arrangement of dorsal tubercles at midbody, the precloacal region in males, and the postmental and gular scale pattern between C . klakahensis , C . fumosus , and C . marmoratus . Cyrtodactylus psarops possesses weakly keeled dorsal tubercles in 23–26 rows at midbody (strongly keeled tubercles in 15–19 rows at midbody); tubercles often present on the brachium (tubercles on brachium absent); 38–49 (35–38) VS; 28–32 (37– 38) PFP in males; a single, greatly enlarged, pore-bearing scale at the apex of the pore-bearing scale series (apical scale of pore-series not greatly enlarged); no posterior precloacal scales (posterior precloacal scales present); and a single (2–3) PCT. Cyrtodactylus pubisulcus (data from Hikida 1990 ; Das & Jim 2000 ; and obtained from specimens listed in the Appendix) possesses 37–58 (35–38) VS; no enlarged femoral scales (enlarged PFS present); no femoral pores (pores, including those on the femur, present in males); and no posterior precloacal scales (precloacal scales present). Cyrtodactylus semicinctus possesses weakly keeled dorsal tubercles in 24–27 rows at midbody (strongly keeled tubercles in 15–19 rows at midbody); and a single, greatly enlarged, pore-bearing scale at apex of the pore-bearing scale series (apical scale of pore-series not greatly enlarged). Cyrtodactylus spinosus (data from Linkem et al . 2008 ) possesses 38–44 5 (35–38) VS; no femoral pores (pores, including those on the femur, present in males); and lateral and caudal spines (lateral and caudal spines absent). 1. Boulenger (1897) provided a count of 42 PFP for C . fumosus . We re-examined the material used by Boulenger, which is clearly conspecific with the type specimen housed in NMB, and found that his single adult male specimen (BMNH 1896.12.9.3, from Rurukan, North Sulawesi) does not possess a continuous series of PFP. De Rooij (1915) provided a count of 42–52 PFP, but included data of specimens from Sulawesi ( C . fumosus ), Halmahera (identified as C . philippinicus [ Steindachner, 1867 ] by Boettger [1900] and subsequently described as a new taxon currently known as C . halmahericus [ Mertens, 1929 ]), and Java (misidentified C . marmoratus ) in her definition of C . fumosus (see Brongersma 1934 ). De Rooij’s (1915) count of 42–52 PFP for C . fumosus is often cited in the literature (e.g., Oliver et al . 2009 ; Chan & Norhayati 2010 ; Grismer et al . 2012 ), although this count is incorrect as demonstrated by our examination and literature survey (see also Remarks on the taxonomy of C . fumosus ). 2. De Rooij (1915) and Brongersma (1934) attributed the shape of the ear-opening to the state of preservation. We examined well preserved specimens of C . fumosus sensu stricto that exhibited a horizontal, slit-shaped ear opening, indicating that this character is taxonomically informative. 3. Although Rösler et al . (2007) provided a maximum count of 52 PFS for the type series, the lectotype of C . marmoratus (RMNH.RENA 2710a.1) possesses 56 PFS. 4. Rösler et al . (2007) provided a count of 45–50 PFP for male C . marmoratus , with the lectotype stated to have 45 pores. However, the lectotype possesses 52 PFP. 5. Linkem et al . (2008 ; Table 1 ) listed 38−44 VS for C . spinosus (counts listed for individual specimens), while in their comparative table ( Table 2 ) they provided a range of 40−43 VS. TABLE 3 . Characters used to distinguish Cyrtodactylus klakahensis sp. nov. from congeneric species occurring in the Sunda Islands (including Sulawesi). The presence of a diagnostic character is coded as ʻ1ʼ, the absence of a character is coded as ʻ0ʼ. For taxa possessing precloacofemoral scales (= scales in a continuous series; column entitled ‘9’), precloacal- and femoral scales (separated each other by infrascales) are coded ‘n/a’ (columns entitled ‘7’ and ‘8’). Numbers at the head of the table correspond to characters as follows: 1 = tubercles on forelimbs, 2 = tubercles on hindlimbs, 3 = tubercles on head, 4 = number of ventral scales, 5 = enlarged subcaudals, 6 = number of subdigital lamellae under 4th toe, 7 = enlarged precloacal scales (and number of pores in parentheses if present; if pores are present in one sex only, this is indicated either by ‘♂’ or ʻ♀ʼ), 8 = enlarged femoral scales (and number of pores in parentheses if present; if pores are present in sex only, this is indicated either by ‘♂’ or ʻ♀ʼ), 9 = enlarged precloacofemoral scales (and number of pores in parentheses if present; if pores are present in males only this is indicated by ‘♂’), = pores in a continuous series, 11 = precloacal groove present (if a groove is present in males only, this is indicated by ‘♂’), 12 = pattern of dorsum (bd = banded; bl = blotched; mo = mottling; pl patternless; st = striped). If data for a character are not available, this is indicated by a ‘?’. Where derived from the literature (‘Lit.’ column), references are abbreviated by letters as follows: A = publication; B = Rösler et al . 2007 ; C = Grismer & Leong 2005 ; D = Manthey & Grossmann 1997 ; E = Iskandar et al . 2011 ; F = Inger & King 1961 ; G = Kathriner et al . 2014 ; H = Rösler & Kaiser, in press.; I = Hikida 1990 ; J = de Rooij 1915 ; K = Brongersma 1934 ; L = Boulenger 1897 ; M = Das 1993 ; N = Youmans & Grismer 2006 ; O = Auffenberg 1980 ; P = Darevsky 1964; Q = Werner 1896 ; R = Das 2010 ; S = Harvey et al . 2015 ; T = Inger 1958 ; U = Riyanto et al . 2014 ; V = Linkem et al . 2008 ; W = Taylor 1962 ; X = Hayden et al . 2008 ; Y = Dunn 1927 . Under the column heading ʻ n ʼ we provide the number of adult specimens we examined personally.

Taxon SVL 1 in adults klakahensis 61–68 1 agamensis 50 1	2 1 1	3 1 1	4 35–38 67	5 0 0	6 17–20 26	7 n/a n/a (? a )	8 n/a n/a (? a )	9 10 1 (37–38, ♂ ) 1 1 (? a )?	11 12 1 bl 1 bl	Lit. A B	n 4 -
baluensis 71–95 1	1	1	36–46	1	19–23	1 (9–11, ♂)	1 (4–10, ♂)	0 0	0 bl	A, C, D	9
batik 103–113 1	1	1	48–57	1	24–27	1	0	0 0	0 bd	E	-
cavernicolus 64–81 0	1	1	51–58	0	22–26	1 (4, ♂)	0	0 0	1 bd	C, F	-
celatus 38−44 1	1	1	34–42	0	15–18	1 (4, ♂)	0	0 0	1 (♂) bl	A, G, H	3

consobrinus 97–125	1	1	1	58–71	1	22–28	1 (8–11, ♂)	0/1 (0–6, ♂)	0	0	0 bd	A, C, D, I 2
darmandvillei 80−82	1	1	1	34–36	1	23−24	n/a	n/a	1	0	0 bl	A, J, K 2
fumosus 57–78	1	1	1	37–50	0	17–23	n/a (10, ♂)	n/a (3, ♂)	1	0	1 (♂) bl	A, L 3
gordongekkoi 71–73	1	1	1	30	0	22–23	n/a	n/a	1	0	0 bl	A, M 2

ingeri 65– 76 1 1 1 40–43 1 23–27 1 (8, only ♂♂ known) 0 0 0 0 bl I, N - ...continued on next page spinosus 70– 83 1 1 1 38−44 0 19–21 1 (12–13, ♂) d 1 (? e ) 0 0 1 (♂) bd V - Cyrtodactylus agamensis is known from a single female only (see Rösler et al . 2007 ). Hence, it is not known at present if males possess pores.

jellesmae	42–64	1	1	1	40−47	0	16–19	1	0	0	0	0	bl A, L 3
laevigatus	38–47	0	1	0	30–34	0	10–15	?	0/1	?	0	0	mo, pl A, E, O, P 5
lateralis	65−85	1	1	1	60–64	0	21–22	1 (13, ♂)	0	0	0	0	bl C, D, Q -
malayanus	70–83	1	1	1	58–62	1	21–23	1 (8–10)	0	0	0	0	bd C, I, N -

TABLE 3 (continued)

Taxon marmoratus	SVL 1 in adults 45–82 0	2 1	3 1	4 38–47	5 0	6 19–24	7 1 (16, ♀)	8 1 (6−7, ♀)	9 10 11 1 (45–53, ♂) 0 (♀)/1 1	12 bl	Lit. A, B	n 2
matsuii	105 1	1	1	48−51	0	22	0 (7−8, ♂)	0	0 0 0	bl	I, N, R	-
psarops	? b 1	1	1	38–49	0	18–22	n/a (? c ,♀)	n/a (? c ,♀)	1 (28–32, ♂) 0 (♀)/1 1 (0–29,♀)	bl	S	-

pubisulcus	51–74	0/1 1	1	37–58	0	16–23	1 (7–9, ♂)	0	0 0	1	bl	A, I, N, T 10
semiadii	40–47	? 1	0	35–36	0	14–15	0	0	0 0	0	bl	U -
semicinctus	? b	0 1	1	33–44	0	19–22	n/a	n/a	1 (36–38, ♂) 1 (0–19,♀)	1	bl	S -

quadrivirgatus 51–71 1	1	1	34–42	0	19–20	n/a (3–4, ♂)	n/a	1	0	0	bl, st D, N, W	-
wallacei 92–114 1	1	1	45–49	1	17–25	1	0	0	0	0	bd, bl X	-
wetariensis 42–67 1	1	1	36–42	0	15–22	n/a (11, ♂)	n/a (12–16, ♂)	1	0	0	bl A, Y	4
yoshi 75–96 1	1	1	50–58	0	25–30	0 (8–12, ♂, absent or indistinct in ♀♀)	0	0	0	0	bl C, I	-

Harvey et al . (2015) did not provide ranges for adult specimens of C . psarops and C . semicinctus . In female C . psarops , pores can either be arranged continuously, discontinuously, or can be absent altogether. Harvey et al . (2015) only provided counts for total pore numbers, but failed to indicate individual counts for the pore-bearing parts of separated pore series. In their diagnosis and Table 1 , Linkem et al . (2008) listed 12–13 precloacal pores for C . spinosus , whereas in their Table 2 , they listed only 8–12 precloacal pores. According to the diagnosis in Linkem et al . (2008) , C . spinosus lacks femoral pores; it was listed as possessing 4–7 femoral pores in their Table 2 . FIGURE 1. Morphological features of the holotype of Cyrtodactylus klakahensis sp. nov. (SMF 22476). (A) Dorsal and ventral view of the body. (B) Precloacal region showing precloacofemoral scales (bearing precloacofemoral pores) and posterior precloacal scales. (C) Lateral view of the left side of the head. (D) Ventral view of the head, showing pattern of postmental and gular scales. Photos by Sven Mecke. FIGURE 2. Paratype series of Cyrtodactylus klakahensis sp. nov. in dorsal and ventral view. (A) SMF 22477, an adult male. (B) SMF 22478, an adult female. (C) SMF 22479, an unsexed juvenile specimen. Photos by Sven Mecke. FIGURE 3. Comparison of the shape and arrangement of dorsal tubercles at midbody, the precloacal region in males, and the postmental and gular scale pattern between Cyrtodactylus klakahensis sp. nov. , C . fumosus , and C . marmoratus . Cyrtodacytlus klakahensis sp. nov. : A) Closely arranged, trihedral, raised, and strongly keeled dorsal tubercles at midbody; B) eidonomy of precloacofemoral scales (three series), precloacal groove, and posterior precloacal scales; C) gular region, showing the presence of enlarged 2nd postmentals (photos and drawings of SMF 22476, holotype ). Cyrtodactylus fumosus : D) Widely scattered, roundish, flat, and smooth dorsal tubercles at midbody; E) eidonomy of precloacofemoral scales (only one series distinctly enlarged), precloacal groove, and posterior precloacal scales; F) gular region, indicating the absence of enlarged 2nd postmentals (D, F = NMB-REPT 2662, holotype ; E = BMNH 1896.12.9.3). Cyrtodactylus marmoratus : G) Closely arranged, slightly raised and keeled dorsal tubercles at midbody; H) eidonomy of precloacofemoral scales (three series) and precloacal groove, with posterior precloacal scales absent; I) gular region, indicating the absence of enlarged 2nd postmentals and the presence of a single pair of enlarged gular scales bordering the single pair of enlarged postmentals posteriorly (G = MTKD 8903; H, I = RMNH.RENA 2710a.1, lectotype ). Photos by Sven Mecke; line drawings by Felix Mader (based on photos by Sven Mecke). Description of the holotype . General habitus, metrics (in mm) and ratios. Adult male; SVL = 68.1; AxialL = 31.9; TailL = 61.7 (tail regenerated); ArmL = 25.9; LegL = 36.3; HeadL = 18.1; HeadW = 10.6; HeadH = 7.0; SnoutL = 7.6; OrbEarL = 5.9; OrbD = 3.7; EarL = 1.3; head length moderate (HeadL/SVL = 0.27); head rather wide (HeadW/HeadL = 0.59), clearly depressed between eyes, distinct from neck; snout rather elongate (SnoutL/ HeadL = 0.42), much longer than OrbD (SnoutL/OrbD = 2.10), canthus rostralis distinct; fore- and hindlimbs of moderate size (ArmL/SVL = 0.38; LegL/SVL = 0.53), without webbing between digits; relative length of fingers = IV> III> V> II> I, relative length of toes = IV> III> V> II> I. Scalation. Dorsal scales rounded and granulate, interspersed with distinctly enlarged, trihedral, raised, strongly keeled, and irregularly arranged dorsal tubercles (Figure 3A); 19 DTR; 26 PVT; latero-dorsal tubercles most strongly keeled, tubercles on lateral portion of the trunk and PVT smaller and feebly keeled; tubercles on base of the tail largest, strongly keeled, pointed and elongate, in eight rows (the longer part of the tail is regenerated, without tubercles present); tubercles on forelimbs small, most prominent on the antebrachium; tubercles on hindlimbs similar in size and shape to latero-dorsal tubercles. Ventral scales distinctly larger than dorsals, juxtaposed; 38 VS; three series of enlarged PFS, lowest series possessing 40 scales, bearing 37 pores; pore series interrupted by a single, enlarged, pore-less PFS on the left femur (PFS at level of this pore-less scale irregularly arranged and/or smaller; due to an aberration likely caused by an injury); scales immediately posterior to the precloacal groove (posterior precloacal scales) enlarged, arranged in a chevron-like shape consisting of three series (from anterior to posterior: nine scales/ seven scales/ three scales) ( Figures 1 B & 3B); two domed PCT; number of lamellae under fingers: I 11, II 12 , III 14, IV 15 , V 15; number of lamellae under toes: I 14, II 16 , III 18, IV 20 , V 19. Rostral shield rectangular, about 0.6 time high as wide, partly divided by a suture dorsally, in contact with 1st SupraLab, two rostronasals and a single InterNas; nostril surrounded by rostral, 1st SupraLab, three post-nasals, and a single rostro-nasal; R10 L10 SupraLab1, R6 L5 SupraLab2, separated from the orbit by 2–3 rows of small granular scales; R9 L9 InfraLab, bordered by two rows of scales larger than granular scales on the throat; cephalic scales small, rounded, granulate and juxtaposed; tubercles on occiput and neck raised, bearing an apex; 43 IOS; 26 SC; mental triangular, wider than long (MentalL/MentalW = 1.4); one pair of enlarged 1st postmentals, followed by a pair of enlarged 2nd postmentals (Figure 3C); pair of 1st postmentals bordered by mental, 1st InfraLab, enlarged 2nd postmentals, and four GulS (Figure 3C); scales on throat minute, rounded. Coloration. In preservative, ground color of dorsal surface of head and body Drab (19); head with indistinct Walnut Brown (27) colored reticulum; a Burnt Sienna (38) stripe running from the posterior border of the orbit to a point above the ear opening; light Pale Buff (1) labial scales and postmentals strongly stippled with darker color; dorsum with irregular, faint Warm Sepia (40) blotches, most visible on neck and at level of posterior margin of the forelimbs, between hindlimbs and base of tail; ground color of dorsal surface of limbs like body; venter, throat and lower surface of limbs uniformly Pale Buff (1), heavily dotted and stippled with dark markings of different size; color of regenerated tail Pale Pinkish Buff (3). Variation. Paratypes similar to holotype except as follows: Paratype SMF 22477 (adult male; Figure 2 A) with 17 DTR; 29 PVT; dorsal scales on original part of tail arranged in whorls, each ending in a row of four tubercles; 36 VS; a series of 38 PFS bearing 38 PFP in a continuous series; posterior precloacal scales arranged in a chevronlike shape consisting of four series of scales (from anterior to posterior: nine scales/ nine scales/ six scales/ three scales); three domed PCT; number of lamellae under fingers: I 14, II 14 , III 14, IV 15 , V 15 (counted on left side of the body; some fingers on right side damaged); number of lamellae under toes: I 12, II 15 , III 16, IV 17 , V 15; R11 L10 SupraLab1, R5 L5 SupraLab2; R9 L10 InfraLab; 27 SC; a single InterNas; a second dark stripe running from nostril to occiput, interrupted only by the orbit; a dark V-shaped collar present, followed by nine pairs of dark paravertebral blotches, larger than lateral ones, which may be fused to form bars anteriorly. Paratype SMF 22478 (adult female; Figure 2 B) with 17 DTR; 33 PVT; dorsal scales on original part of tail arranged in whorls, each ending in a row of four to six tubercles; 36 VS; 38 PFS, lacking pores; no sign of a precloacal groove; posterior precloacal scales arranged in a chevron-like shape consisting of three series of scales (from anterior to posterior: nine scales/ seven scales/ five scales); two domed PCT; number of lamellae under fingers: I 13, II 14 , III 18, IV 16 , V 13; number of lamellae under toes I 13, II 15 , III 19, IV 17 , V 19; R10 L11 SupraLab1, R5 L6 SupraLab2; R8 L10 InfraLab; 39 IOS; 33 SC; three InterNas; eight pairs of dark paravertebral blotches, which may be fused to form bars; original part of tail with six Burnt Sienna (38) colored bands. Paratype SMF 22479 (unsexed juvenile; Figure 2 C) with 15 DTR; dorsal surface on entirely original tail with scales arranged in whorls, ending in a row of tubercles, tubercles present on proximal quarter of tail only; 35 VS; 38 slightly enlarged PFS, lacking pores; no sign of a precloacal groove; posterior precloacal scales arranged in a chevron-like shape consisting of three series of scales (from anterior to posterior: eight scales/ eight scales/ five scales); two domed PCT; number of lamellae under fingers: I 14, II 15 , III 17, IV 18 , V 15; number of lamellae under toes: I 12, II 14 , III 16, IV 17 , V 15; R11 L10; SupraLab1, R5 L6 SupraLab2; R9 L8 InfraLab; 36 IOS; 28 SC; a single InterNas; a second dark stripe running from nostril to occiput, interrupted only by the orbit; a dark Verona Brown (37) V-shaped collar present; tail with 15 Verona Brown (37) colored bands, becoming Burnt Sienna (38) distally. Etymology. The specific epithet is a Latinized, toponymic adjective referring to the type locality Klakah (Lumajang Regency, Jawa Timur Province, Java, Indonesia ) of the new species. Distribution and Natural History. Cyrtodactylus klakahensis is currently only known from its type locality, Klakah, Lumajang Regency, Jawa Timur Province, Java, Indonesia ( Figure 4 ). Klakah is located in the lowlands (elevation ca. 200 m ) between the Tengger and Iyang-Argapura mountain massifs. Although the species range is probably not restricted to Klakah, it may exhibit a relatively limited distribution in central Jawa Timur Province and therefore should be regarded as endemic to the region, until evidence to the contrary becomes available. FIGURE 4. Map of Java illustrating the type-, potential type-, and paratype localities of Cyrtodactylus spp. The black triangle marks the type locality of C . klakahensis . The black diamond marks the type locality of C . semiadii (paratype locality identified by a white diamond). The exact type locality of C . marmoratus in Java is unknown, but the type specimens were, in all probability, collected from western Java. The potential type locality (indicated by a black star) may lie within the mountainous area around Bogor, but a revision of the C . marmoratus species-complex is needed to narrowly restrict the type locality of C . marmoratus sensu stricto. Base map modified from Wikipedia by Sven Mecke. Remarks on the taxonomy of Cyrtodactylus fumosus . The bent-toed gecko species Cyrtodactylus fumosus (proposed vernacular name: Sulawesi bent-toed gecko) was described by Friedrich (‘Fritz’) Müller ( 1834–1895 ) based on a single adult female (NMB-REPT 2662; Figure 5 ) collected by Karl Friedrich (‘Fritz’) Sarasin ( 1859– 1941 ) and Paul Benedict Sarasin ( 1856–1929 ) in the “Bulawa Mountains” (North Sulawesi, elevation ca. 1200 m ) (Müller 1895; the 1894 volume was issued in 1895). In his second article published in the Verhandlungen der naturforschenden Gesellschaft Basel (Reptilien und Amphibien aus Celebes, part II), Müller (1895b) mentioned a second specimen of the species (NMB-REPT 2663) from “Masarang” but referred to NMB-REPT 2662 as “ Original-Exemplar ” (i.e., the holotype ). We therefore recognize NMB-REPT 2662 as the only type specimen (see Brongersma 1934 ; Kramer 1979 ; Koch 2012 ) in disagreement with de Rooij (1915:17) who referred to an unspecified number of type specimens in a footnote. Boulenger (1897) provided a more detailed description of C . fumosus based on four specimens, including NMB-REPT 2662–63 and two additional North Sulawesi specimens donated by Paul Sarasin, BMNH 1895.2.27.7 and BMNH 1896.12.9.3, and corrected the type locality from Boelawa Mountains (= Huidu Matabulawa) to Bone Mountains (= Pegunungan Bone). De Rooij (1915) recorded C . fumosus for Java based on a single specimen but Brongersma (1934) subsequently referred to this apparently misidentified voucher as C . marmoratus and restricted C . fumosus to Sulawesi. Mertens (1934) once again listed C . fumosus as occuring on Java (Klakah, Lumajang) and clearly indicated that this assignment would be provisional with the genus being in need of a revision. Several subsequent authors have listed C . fumosus as part of the Javanese herpetofauna (e.g., Manthey & Grossmann 1997 ; Hayden et al . 2008 ; Oliver et al . 2009 ; Das 2010 ; Riyanto et al . 2014 ) but these records seem to be either based on erroneous data provided in the literature (e.g., de Rooij 1915 ; Das 2010 ) and/or misidentified specimens. Linkem et al . (2008) did not consider C . fumosus a valid species, stating that it had been synonymized with C . marmoratus . However, Brongersma (1934) did not synonymize C . fumosus with C . marmoratus but indicated both taxa were distinct (see also Brongersma 1953 ; Koch 2012 ). FIGURE 5. Holotype of Cyrtodactylus fumosus (NMB-REPT 2662; adult female) in (A) dorsal, (B) lateral, and (C) ventral view. Photos by Sven Mecke. There is considerable confusion in the literature as to whether Cyrtodactylus fumosus possesses a precloacal groove in males. Cyrtodactylus fumosus was referred to as a species lacking a precloacal groove by Rösler et al . (2007) , Grismer & Norhayati (2008) , Welton et al . (2009) , Chan & Norhayati (2010) , and Grismer et al . (2012) . Boulenger (1897) , who examined C . fumosus specimens from North Sulawesi (Bone Mountains, Masarang, Rurukan; all collected by Fritz and Paul Sarasin), reported that male individuals of this species did possess a precloacal groove. We examined the adult male specimen (BMNH 1896.12.9.3) used by Boulenger for his species account of C . fumosus and confirm the presence of a precloacal groove in this species. A precloacal groove, however, may be only weakly defined in subadult male specimens, as seen in NMB-REPT 2663. It appears that several species may be masquerading under the name Cyrtodactylus fumosus , both in the Greater and Lesser Sunda Islands ( Riyanto et al . 2014 ; pers. obs.). Above, we were able to demonstrate unequivocally that the specimens from Klakah, Lumajang Regency, Java, originally referred to as C . fumosus by Mertens (1934) , represent a new species. The populations on Bali referred to as C . fumosus by Mc Kay (2006) and additional forms from Java are likely also new species (Mecke et al ., in prep., Riyanto et al ., in prep.). Based on a photograph of a strongly tuberculated individual of C . cf. fumosus in Koch (2012: 151) from North Sulawesi, we assume that Sulawesi C . fumosus populations may represent a species complex as well. The taxon is in dire need of a revision, with a thorough re-description of C . fumosus sensu stricto currently underway (Hartmann et al ., in prep.).