Two new cryptic species of the Cyrtodactylus irregularis complex (Squamata: Gekkonidae) from southern Vietnam Author Nazarov, Roman Author Poyarkov, Nikolay A. Author Orlov, Nikolai L. Author Phung, Trung My Author Nguyen, Tao Thien Author Hoang, Duc Minh Author Ziegler, Thomas text Zootaxa 2012 3302 1 24 journal article 10.5281/zenodo.209998 efadc615-71e8-4474-afed-7f073917e179 1175-5326 209998 Cyrtodactylus bugiamapensis sp. nov. Holotype . Adult male ZMMU R- 13366 collected by Nikolay Poyarkov in June 2011 within Bu Gia Map National Park, Dak Ka stream valley ( 12°12’N ; 107°12’ E ), Bu Gia Map commune, Bu Gia Map District, Binh Phuoc Province, southern Vietnam , at an altitude of ca. 354 m a.s.l. ( Fig. 3 ). Paratypes . One adult male ZMMU R- 13367-1, three adult females ZISP 26323; ZMMU R- 13367-2; ZMMU R- 13367-3, and one juvenile ZMMU R- 13367-4 collected by Nikolay Poyarkov during 2009-2011 , the same collection data as for the holotype ; two adult females ( IEBR A.2011.3 [field number Tao 993], VNMN 994, and a subadult male ( ZFMK 92323 [field number Tao 995]), collected by Trung My Phung in May 2010 in Bu Gia Map commune, Bu Gia Map District, Binh Phuoc Province, southern Vietnam . Diagnosis. A medium sized Cyrtodactylus , SVL 58.6–76.8 mm , TailL 65.3–83.0 mm; body slender, limbs and digits moderately long, original tail relatively thin, longer than the body; two pairs of postmental scales, first pair in broad contact with one another, size of the second pair about half of first pair; nostrils surrounded by supranasal, rostral, first supralabial, and three (rarely four) small postnasal scales; dorsal tubercles enlarged, conical, and with keels, in 20–24 irregular longitudinal rows; lateral folds weakly developed; 36–46 ventral scales between lateral folds; 164 – 205 ventral scales between postmental scales and cloaca; males with 7 – 11 precloacal pores in single ^- shaped series; precloacal groove and femoral pores absent; 6 – 8 implicit enlarged femoral scales on each side; midventral subcaudal scales uniform, larger than the scales on the caudal flanks, without transversely enlarged plates; only tail base with tail segments, tail-base whorls without prominent spurs; average number of subdigital lamellae on the fourth finger is 15.9 and on the fourth toe 18.3; ground coloration of back light yellow to light brown (light beige to brown in preservative); dorsal pattern consisting of unclear transversal bands formed by irregular roundish to oblong, dark brown spots; dark neck band, which can be medially divided, narrow, U-shaped, extending towards ear and orbit; dorsal head surface with few irregular dark spots; all dark dorsal pattern with distinct white bordering; tail with 8 – 11 dark transversal bands, which fade on the ventral side. Description of Holotype . Medium sized (SVL 64.8 mm , TailL 66.0 mm), HeadL 17.7 mm , HeadW 12.0 mm, HeadH 7.3 mm , SnEye 7.0 mm, OrbD 3.8 mm , EarL 1.1 mm , EyeEar 5.2 mm ; proportions are as follows: SVL/ HeadL 3.66, HeadL/HeadW 1.47, HeadL/HeadH 2.42, SnEye/EyeEar 1.34. Rostral is large, somewhat wider than high (RW 2.8 mm , RH 1.9 mm , RW/RH 1.4) with an inverse Y-shaped median suture ( Fig. 4 c); supralabials 11/10; scales between orbit and the seventh supralabial 3/4, small; infralabials 10/9; nares surrounded by rostral anteriorly, first supralabial laterally, supranasal and 4 nasals posteriorly; rostral about 5 – 6 times larger than supranasal; supranasals separated from each other by intersupranasal (two times smaller than supranasals); snout scales medially granular, those in contact with the supralabials are flattened and about 2 – 3 times larger than the medial ones; upper anterior ciliaries two times larger than posterior ones; head scales granular, some smaller than the median snout scales; dorsum of head and temporal region with rounded, keeled tubercles, that are three times larger than the surrounding scales; mental pentagonal, as wide as rostral ( Fig. 4 b); three pairs of enlarged postmentals, longer than wide, first pair in broad contact; dorsal scales granular, 3 – 4 times smaller than the ventral scales; dorsal tubercles round, conical, keeled, surrounded by 9 – 12 granular scales, tubercles forming about 20 irregular longitudinal rows at midbody; ventral scales smooth, 39 longitudinal rows at midbody; lateral folds weakly developed, marked by small light spots; dorsal surface of fore and hind limbs with granular scales and strongly keeled conical tubercles; fingers and toes without web, basal lamellae more rounded than on distal surface of digits, numbering 9 under first finger, 16/15 under fourth finger, 10 under first toe, and 18 (8 basal and 10 distal lamellae) under fourth toe; V – shaped row of precloacal scales with 8 pores, and three additional precloacal pores on enlarged scales below ( Fig. 4 d); enlarged femoral scales (without pores) are present; three pairs of enlarged postcloacal tubercles; first tail third with whorls, dorsally covered by convex keeled tubercles; posterior part of tail covered by flattened and rounded scales; subcaudals without enlarged plate row, flat, smooth, imbricate, about two times larger than dorsal scales on tail. TABLE 3. Measurements and selected morphological characters of the type series of Cyrtodactylus bugiamapensis sp. nov. ; f = female, j = juvenile, m= male. HOLOTYPE PARATYPES

ZMMU R-13366	ZMMU ZISP 26323 R-13367-1	ZFMK 92323	VNMN 944	IEBR A.2011.3	ZMMU R-13367-2	ZMMU R- 13367-3	ZMMU R-13367-4
Sex SVL TailL	M 64.8 66	M F* 64.9 73.1 65* 49*	M 58.6 65.3	F 72.3 73	F 76.5 55*	F 76.8 83	F 61.2 71.5	JUV 39.5 45
TrunkL HeadL	27.6 17.7	28 33.3 17.6 20	24.3 16.7	30.7 20.8	32 21.4	34.7 21	26.7 18.3	16.2 12.5
HeadW HeadH SnEye OrbD	12 7.3 7 3.8	12.6 13.1 7.7 8 7.2 8 4.2 4.5	10.7 6.7 6.5 3.2	12.6 8 7.8 3.8	13.7 8.9 8.1 3.8	13.9 8.6 8 4.8	11.4 7 6.8 3.8	7.3 4.5 4.3 2.4
EarL EyeEar	1.1 5.2	0.9 1.2 5.5 6.4	1 4.3	1.1 6	1.4 6.5	1.4 6.8	0.8 5.4	0.7 3.6
LS ForeaL FemurL Crus L LD4A LD4P	9.7 9.1 11.8 9.3 5.6 7.1	10 10.8 9 10 12.2 13.3 9.2 11 5.9 5.3 7.2 7.2	9.7 8.2 11.3 9.2 5.3 6.2	11.7 10 13.6 11.3 6.5 8.6	11.7 10.8 13 11 6.2 7.6	11.6 10.6 13.4 11.8 5.6 7.9	8.3 8.2 11.3 9.1 5.8 7.0	6.4 5.5 7.3 6.7 3.3 4.5
V SLB TubL/TubW SLH/SAH	39 178 20/46 108/86	42 38 188 164 23/46 22/42 99/89 96/76	46 201 20/43 106/88	36 189 20/45 102/82	42 205 22/44 95/82	40 189 24/46 96/80	22/46 101/78	43 181 20/42 110/95
LF4	15	16 16	16	15	16	16	17	16
LT4	18	17 17	18	19	19	20	20	17
PP	8(+3)***	7 7*	8	0	0	7*	**	0
SL	11/10	10/10 10/9	10	10	9	9/10	10/10	10/9
IL	10/9	9/8 8/8	9	9	8	8/8	8/9	9/9

* - perforated enlarged precloacal scales **- damaged precloacal region *** - see description of holotype Coloration: Dorsal head surface brown with three roundish dark brown spots with contrasted white edge in the occipital region and two irregular patches on the tympanum region; nuchal band not broadened posteriorly, dark brown with a distinct white margin, extending from the neck to the posterior margins of eyes; labials brownish grey with white dots. Dorsum brownish grey with seven irregular, dark transverse bands, with light margins; flanks greyish white. Venter white, the lower side of toes and fingers grey; dorsal surface of limbs and digits brownish with irregular bands. Tail dorsum with 11 dark brown bands which are wider than the light grey bands in-between. Ventral side of tail dark grey with light spots that are becoming more distinct posteriorly. For coloration in life see Fig. 3 . During daytime, the colour is darker, making the colour pattern more indistinct. Variation of paratypes . For the variation of the type series see Table 4 and Fig. 4 a. The dorsal pattern is somewhat variable and two paratypes did not show additional pores below the V-shaped row of precloacal pores. Sexual dimorphism is well developed, males are smaller then females, precloacal pores are present in males only and males have more developed postcloacal spurs. FIGURE 3. Holotype of Cyrtodactylus bugiamapensis sp. nov. from Bu Gia Map National Park, southern Vietnam. Comparisons with Vietnamese species. Cyrtodactylus bugiamapensis sp. nov. differs from C. badenensis Nguyen, Orlov & Darevsky , C. bichnganae Ngo , C. caovansungi Orlov, Nguyen, Nazarov, Ananjeva & Nguyen , C. chauquangensis Hoang, Orlov, Ananjeva, Johns, Hoang & Dau , C. condorensis (Smith) , C. cucphuongensis Ngo & Chan , C. eisenmanae Ngo , C. grismeri Ngo , C. hontreensis Ngo, Grismer & Grismer , C. huongsonensis Luu, Nguyen, Do & Ziegler , C. intermedius (Smith) , C. martini Ngo, C. nigriocularis Nguyen , C. paradoxus (Darevsky & Szczerbak) , C. phongnhakebangensis Ziegler, Rösler, Herrmann & Vu , C. phuquocensis Ngo, Grismer & Grismer, C. takouensis Ngo & Bauer , C. roesleri Ziegler, Nazarov, Orlov, Nguyen, Vu, Dang, Dinh & Schmitz , and C. yangbayensis Ngo & Chan , by the absence of transversally enlarged subcaudals. From the Vietnamese Cyrtodactylus species, which equally lack transversally enlarged subcaudals ( C. cattienensis Geissler, Nazarov, Orlov, Böhme, Phung, Nguyen & Ziegler , C. cryptus Heidrich, Rösler, Vu, Böhme & Ziegler , C. huynhi Ngo & Bauer , C. irregularis Smith , C. pseudoquadrivirgatus Rösler, Nguyen, Vu, Ngo & Ziegler , and C. ziegleri Nazarov, Orlov, Nguyen & Ho ), the new species differs as follows. From C. huynhi the new species differs by lacking femoral pores (3 – 8 pores present in C. huynhi ). Cyrtodactylus bugiamapensis sp. nov. differs from C. cryptus and C. pseudoquadrivirgatus by enlarged femoral scales (absent in both species versus 6 – 8 in Cyrtodactylus bugiamapensis sp. nov. ) and the dorsal pattern (blotched in the new species versus banded in C. cryptus ). Cyrtodactylus bugiamapensis sp. nov. is most similar to the members of the C. irregularis complex. Cyrtodactylus bugiamapensis sp. nov. differs from C. pseudoquadrivirgatus by the presence of enlarged femoral scales and by a continuous dark neck-band with distinct white borders, which is absent in C. pseudoquadrivirgatus . Cyrtodactylus bugiamapensis sp. nov. differs from C. irregularis s. str. by the absence of large spurs on the tail-base whorls (which are present in C. irregularis ), by a thin tail which is longer than the body (index SVL/TailL = 0.91, n=6) versus a comparatively thick tail in C. irregularis which is shorter than the body (index SVL/Lcd = 1.1, n=3) and by its smaller maximum size (SVL 76.8 mm versus 86 mm in C. irregularis ). From the likewise phenetically similar C. cattienensis , Cyrtodactylus bugiamapensis sp. nov. differs by the dorsal pattern (which consists of separate roundish dark-brown spots with yellowish-white borders in the new species versus dark-brown pattern without light borders in C. cattienensis ), and by its larger size (SVL 58.6 – 76.8 mm versus 49.9 – 70.0 mm in C. cattienensis ). From C. ziegleri , the new species can be distinguished by its different dorsal pattern (blotched in the new species versus predominantly banded in C. ziegleri ) and by its smaller size (SVL 74 – 76.8 mm versus 84.6 – 93 mm in C. ziegleri ). In addition, the average number of subdigital lamellae on the fourth finger and toe in Cyrtodactylus bugiamapensis sp. nov. is 15.9 and 18.3, versus 17.1 and 22.1 in C. ziegleri . Moreover, Cyrtodactylus bugiamapensis sp. nov. has shorter limbs (index SVL / fore limbs 2.65 and SVL/ hind limbs 2.26, n=9) versus relatively elongated limbs in C. ziegleri (index SVL / fore limbs 2.51 and SVL/ hind limbs 2.17, n=8). For more detailed comparisons see Table 6. FIGURE 4. Part of the type series of Cyrtodactylus bugiamapensis sp. nov. from Bu Gia Map National Park, southern Vietnam, showing variation in dorsal colour pattern (A). Holotype of Cyrtodactylus bugiamapensis sp. nov. : mental scalation (B), rostral scalation (C) and precloacal region (D). Comparisons with congeners from Southeast Asia. Cyrtodactylus bugiamapensis sp. nov. differs from other mainland congeners by the following characters. By the absence of transversely enlarged subcaudal plates the new species may be distinguished from: C . aequalis Bauer , C. annandalei Bauer , C. aurensis Grismer , C. baluensis (Mocquard) , C. chanhomeae Bauer, Sumontha & Pauwels , C. consobrinus (Peters) , C. feae (Boulenger) , C. ingeri Hikida , C. jarujini Ulber , C. lomyenensis Ngo & Pauwels , C. malayanus (de Rooij), C. oldhami (Theobald) , C. pageli Schneider, Nguyen, Schmitz, Kingsada, Auer & Ziegler , C. peguensis (Boulenger) , C. russelli Bauer , C. sumonthai Bauer, Pauwels & Chanhome , and C. teyniei David, Nguyen, Schneider & Ziegler. By the absence of a precloacal groove, Cyrtodactylus bugiamapensis sp. nov. differs from the following Southeast Asian mainland species: C. marmoratus (Gray) , C. pulchellus Gray , and C. rubidus (Blyth) . The new species can be distinguished from the following species by the presence of enlarged femoral scales: C. brevidactylus Bauer , C. buchardi David, Teynie & Ohler , C. chrysopylos Bauer , C. gansi Bauer , and C. elok Dring. FIGURE 5. Type locality of Cyrtodactylus bugiamapensis sp. nov. : Dak Ka stream valley, environs of the second forest station in Bu Gia Map National Park, Bu Gia Map community, Bu Gia Map District, Binh Phuoc Province, southern Vietnam. Etymology. The specific epithet is a Latinized toponymic adjective and commemorates the type locality of the new species, the tropical monsoon forest of Bu Gia Map National Park. As common names we propose Bu Gia Map Bent–toed Gecko (in English), and Thach sung ngón bu gia map (in Vietnamese). Habitat description . Specimens were collected in lowland tropical forest in the valley of Dac Ka stream ( Fig. 5 ). The forest condition varies from medium disturbed parts along the road and around the ranger station surrounded with non-disturbed parts of primary tropical forest. This area is quite hilly, with Dac Ka river tributaries forming numerous gorges. The forest floor is covered with numerous rocks of volcanic tuff; the canopy is closed for large parts. The forest composition is dominated by Dipterocarpaceae , Fabaceae , Poaceae and Magnoliaceae trees. Disturbed forest areas house bamboo and rattan palm bushes. The geckos were found at day time under tree logs or pieces of volcanic tuff, and at night on the tree trunks on approximately 1 – 2 m above the ground. One specimen was caught in the ruins of a brick building under ceramic tiles. Distribution. So far, Cyrtodactylus bugiamapensis sp. nov. is known only from Bu Gia Map National Park (Bu Gia Map Commune, Bu Gia Map District, Binh Phuoc Province) and from the neighboring areas of Dak Nong Province (Quang Truc Commune, Tuy Duc District) (see Fig.1), southern Vietnam . Considering the geographic position of this area, records of Cyrtodactylus bugiamapensis sp. nov. from the adjacent Cambodia (Mondolkiri Province) are anticipated. Phylogenetic position. According to mtDNA data, Cyrtodactylus bugiamapensis sp. nov. belongs to a mountain species group of the Cyrtodactylus irregularis complex and is sister species to C. ziegleri ( p -distance between the two species is 7.39% of substitutions based on COI partial sequences).