A new species of the Cyrtodactylus irregularis complex (Squamata: Gekkonidae) from southern Vietnam Author Geissler, Peter Author Nazarov, Roman Author Orlov, Nikolai L. Author Böhme, Wolfgang Author Phung, Trung My Author Nguyen, Truong Quang Author Ziegler, Thomas text Zootaxa 2009 2161 20 32 journal article 10.5281/zenodo.188976 14a08276-99c2-40ba-bece-093572f2b295 1175-5326 188976 Cyrtodactylus cattienensis sp. nov. Holotype . Adult male ( IEBR A.0856) collected on 1 August 2008 by Peter Geissler within Cat Tien National Park ( 11°27’N ; 107°20’E ), Dong Nai Province, southern Vietnam , at an altitude of ca. 120 m a.s.l. Paratypes . Two adult males ( IEBR A.0845, VNUH 2008.0520) and two adult females ( IEBR A.0843- A.0844) from Nui Dinh, Ba Ria – Vung Tau Province, southern Vietnam , coll. Phung My Trung on 20 May 2008 ; one male ( ZFMK 88095), three females ( ZFMK 88090-88091, IEBR A.0855), and three juveniles ( ZFMK 88092-88094) from Cat Tien National Park, Dong Nai Province, coll. Peter Geissler, 23 July to 3 August 2008 ; one male ( IEBR 656) from Cat Tien National Park, Dong Nai Province, coll. Paul Moler on 20 May 2001 ; one male ( IEBR A.0854) and one female ( VNUH 2008.09.07) from Vinh Cuu Nature Reserve, Dong Nai Province, coll. Phung My Trung and Nguyen Thanh Binh on 9 July 2008 ; two males ( ZMMU R- 11444.1, R- 11444.2), coll. Alex Borisenko and Nataly Ivanova, 12–21 December 2002 , one male ( ZMMU R- 11926), coll. Vladimir Bobrov on 17 May 2004 , and one female ( ZMMU R- 11189), coll. Alex Borisenko on 9 December 2004 , all from Cat Tien National Park, Dong Nai Province; one male ( ZISP 25216) and one female ( ZISP 25217) from Cat Tien National Park, Dong Nai Province, coll. Nikolai Orlov in 2006. Diagnosis. A small slender Cyrtodactylus with a maximum SVL of 69 mm , distinguished from all congeners by the combination of the following characters: 1) rounded body in cross-section, not depressed; 2) neck band extending to the posterior margin of the eye; 3) trunk, tail, and limbs bearing irregularly shaped bands (4–6 on trunk, and 4–12 across the tail); 4) tubercles present on the head, body, limbs and tail; 5) dorsal tubercles in 16–22 irregular longitudinal rows; 6) ventrals in 28–42 longitudinal rows at midbody; 7) lateral folds weakly developed or absent and lacking enlarged tubercles; 8) tail base bearing scales arranged in whorls, not depressed nor enlarged; 9) two or three postcloacal spurs in both males and females; 10) precloacal pores 6–8, in an angular row in males and a patch of enlarged precloacal scales in both sexes; 11) slightly enlarged femoral scales present in both males and females, numbering 3–8 on each hindlimb; 12) femoral pores absent; 13) precloacal groove absent; 14) subcaudal scales not enlarged; 15) fourth toe with 14–19 subdigital lamellae. Description of holotype . Size small (SVL 49.9 mm , TL 59.3 mm ), HL 13.4, HW 10.0 mm, HH 6.0 mm, SE 5.9 mm , EE 4.0 mm, ED 3.1 mm , EAD 0.9 mm ; proportions are as follows: SVL/HL 3.72, HL/HW 1.35, HL/HH 2.23, SE/EE 1.49, ED/EAD 3.34. Rostral wider than high (RW 2.3 mm , RH 1.0 mm, RW/RH 2.3) with an inverse Y-shaped median suture; supralabials 10/9; scales between orbit and the seventh supralabial 3/3, small; infralabials 8/9; nares bordered by rostral anteriorly, first supralabial laterally and four nasals posteriorly; rostral approximately four times larger than supranasal and 2.5 times larger than intersupranasal; supranasals separated from each other by an intersupranasal; medial snout scales granular, those in contact with the supralabials flattened and approximately 2–3 times larger than the medial scales; upper anterior ciliaries two times larger than posterior cilaries; head scales granular, one-half as large as median snout scales; centre of dorsal head and temporal region with rounded, smooth tubercles, three times larger than the surrounding scales; mental triangular, as wide as rostral; one pair of enlarged postmentals, trapezoidal, longer than wide, bordering with mental anteriorly and followed by seven pairs of enlarged gular scales, the anteriormost pair largest and surrounded by nine granular scales; gular pairs from second to seventh separated from infralabials by a row of scales; dorsal scales granular, as large as medial snout scales; dorsal tubercles round, conical, keeled anteriorly, surrounded by 9–10 granular scales, tubercles forming 17 longitudinal rows at midbody; ventral scales smooth, medial scales three times larger than the dorsal granules, 38 longitudinal rows at midbody; lateral folds absent; upper and lower arm lacking enlarged tubercles, covered with slightly enlarged granules; dorsal hind limb covered with granular scales and scattered, smooth, conical tubercles; four enlarged femoral scales on each hind limb; fingers and toes lacking webbing; basal lamellae basally more rounded than distally, numbering eight under first finger, 16 under fourth finger, nine under first toe, and 17 (6 basal and 11 distal lamellae) under fourth toe; claws surrounded by a small scale on upper and a large scale on lower sides; six contiguous precloacal pores in a right angle; precloacal region covered by a patch of 18 enlarged scales; two pairs of slightly enlarged postcloacal tubercles on lateral surface of hemipenial swelling; dorsal tail base whorls covered by convex scales, posterior part of tail covered by flattened, rounded scales, twice as large as medial snout scales; first four whorls with tubercles (numbering 10/4/4 /4), tubercles keeled anteriorly; subcaudals not enlarged, flat, smooth, imbricate, about two times larger than scales on dorsum of tail. FIGURE 1. a) Cyrtodactylus cattienensis sp. nov. in life (holotype: IEBR A.0856) showing light color pattern at night and b) Ventral view of the cloacal region with everted hemipenes (paratype: ZFMK 88095). TABLE 1. Measurements and selected morphological characters of the type series of Cyrtodactylus cattienensis sp. nov. (f: female, j: juvenile, m: male, measurements in mm, -: regenerated or broken tail) a) holotype and male paratypes b) female and juvenile paratypes

Sex	0856. A IEBR m	0845. A IEBR m	0854. A IEBR m	656 IEBR m	11444.1 R- ZMMU m	11444.2 R- ZMMU m	11926 R- ZMMU m	25216 ZISP m	2008.0520 VNUH m	88093 ZFMK m	88095 ZFMK m	± deviation value Mean Standard n = 11
SVL	49.9	54.6	61.7	52.6	58	56	59	57	58.7	43.5	58.2	55.4 ± 5.11
TL	59.3	58.9	-	54.7	62	55	61	57.5	-	51.2	-	57.8 ± 3.75 (n = 8)
EAD	0.9	0.9	1.2	1.5	1	1	0.8	0.8	1.0	0.9	1.1	1.0 ± 0.20
ED EE	3.1 4	3.1 4.7	4.2 5.3	3.4 4.5	3.5 4.4	3.4 4.4	3.4 4.4	3.6 4.3	3.7 4.9	3 4.2	4 5.4	3.5 ± 0.37 4.6 ± 0.44
HH	6	6.2	8.1	5.9	7	7	6.8	6.4	7.4	5.1	6.2	6.6 ± 0.82
HL HW	13.4 9.6	15.3 10.2	17.8 13.1	14.1 10.2	17.5 11.3	17.4 11.2	17.2 11.3	16.5 10	17.2 11.2	13.4 8.5	17.1 10.8	16.1 ± 1.71 10.7 ± 1.18
IND	1.8	1.9	2.1	2.2	2.5	2.4	2.3	2.4	2.2	2.3	2.3	2.2 ± 0.21
SE AG	5.9 18.7	6.3 22.6	7.9 23.1	6.3 24.1	6.5 25.3	6.3 22.3	6.7 25.2	6.2 23.4	7.3 23.1	5.5 17.6	6.1 23.6	6.5 ± 0.66 22.6 ± 2.42
FEL	9.2	10.3	11.8	8.6	11.2	10.7	11.2	10.6	10.2	7.2	10.9	10.2 ± 1,34
FL GBLL	7 5	9 4	10 6	8 7	9.2 7	8.7 9	9 7	8 5	9 5	6 6	9 6	8.3 ± 1.05 6.1 ± 1.38
GBUL	8	6	8	10	9	10	9	7	8	8	7	8.2 ± 1.25
SBL IN	8 1	8 1	8 1	9 3	9 2	9 1	7 1	8 2	10 3	9 3	10 1	8.6 ± 0.92 1.7 ± 0.91
IO	18	19	17	17	40	47	41	41	17	19	18	26.7 ± 12.45
S5S SL	51 10	49 8	44 10	50 11	49 10	50 10	50 10	44 9	44 9	46 11	50 11	47.9 ± 2.81 9.9 ± 0.94
GST	9	10	9	9	10	10	10	10	10	9	9	9.5 ± 0.52
V DTR	38 17	38 17	35 19	34 21	37 20	42 20	36 18	34 18	33 16	35 19	28 19	35.5 ± 3.53 18.5 ± 1.51
EFS	4	4	5	5	6	6	7	7	6	4	8	5.6 ± 1.36
LF4 LT4	16 17	14 17	16 17	15 14	14 18	15 16	13 15	14 16	12 15	16 17	16 18	14.6 ± 1.36 16.4 ± 1.29
EPS	18	19	17	20	10	9	10	8	15	16	21	14.8 ± 4.75
PP	6	7	6	8	6	6	7	6	7	6	7	6.5 ± 0.69

7 . 0 Coloration in alcohol. Head dorsally brownish grey with seven irregularly shaped dark brown patches; nuchal band broadened, dark brown and without a white margin, extending from the neck to the posterior margins of eyes; labials lighter with black dots. Dorsum brownish grey with six irregular, dark brown transverse bands, bands without white margins; first band on the shoulder X-shaped; the last dorsal band located in the sacral region; dark bands interrupted by light grey patches on lateral sides; flanks greyish white. Venter cream, the lower side of toes and fingers grey; dorsal limbs and digits grey with white spots. Dorsal side of tail with 12 dark brown bands which are wider than the light grey bands in-between. Ventral side of tail dark grey with white spots that are becoming darker posteriorly.

Sex	0844. A IEBR f	0843. A IEBR f	0855. A IEBR f	11189 R- ZMMU f	25217 ZISP f	2008.09. VNUH f	88090 ZFMK f	88091 ZFMK f	± value deviation Mean Standard n = 8	88092 ZFMK j	88094 ZFMK j
SVL TL	47.7 51	67.2 64.7	63.4 -	69 62	58.4 59	64.0 -	64.8 -	49.2 54.5	60.5 ± 8 58.2 ± 5.5 (n = 5)	42 49.2	36.2 34.4
EAD ED	1.4 3.5	1.2 4.1	1.3 4.1	1.2 4	1 3.4	1.1 3.8	1.5 3.1	1.1 3.2	1.2 ± 0.2 3.7 ± 0.4	1.3 2.8	0.8 2.6
EE	4.3	6.7	5.3	5.3	4.5	5.8	5.4	3.8	5.1 ± 0.9	4.1	3.7
HH HL	5.2 14.2	7.3 20	7.1 18.4	7.8 19.5	6.8 16.8	7.8 18	7.3 18.7	5.8 14.8	6.9± 0.9 17.6± 2,1	4.8 11.7	4.5 11.3
HW	8.8	12.5	12.3	13.3	11	12.7	11.9	9.6	11.5± 1.6	8.6	7.3
IND SE	1.8 6	2.4 7.6	2.5 6.9	2.4 7.3	2 6.2	2.5 8.2	2 7	2.3 6.2	2.2 ± 0.3 6.9 ± 0.8	2.4 5.3	2.0 4.8
AG	19.2	25.9	27.8	30	24.3	27.1	28.1	20.4	25.4± 3.8	17.1	14.5
FEL FL	8.5 7	11 10	10.1 8	11.5 9.8	11 9.2	11.8 10	11.8 10	10.1 7	10.7 ± 1.1 8.9 ± 1.2	8.9 6	6.8 6
GBLL	6	5	6	7	6	6	6	5	5.9 ± 0.64	7	5
GBUL SBL	8 10	7 9	7 9	9 8	9 9	9 9	9 9	7 8	8.1 ± 0.99 8.9 ± 0.64	7 9	7 8
IN	4	1	2	1	1	3	1	1	1.75 ± 1.65	1	1
IO S5S	19 46	19 50	17 45	38 47	48 48	16 47	18 45	17 44	24 ± 12.07 46.5 ± 1.93	16 47	17 50
SL	10	9	11	10	10	10	11	10	10.1 ± 0.64	10	10
GST V	9 41	10 34	9 31	10 39	10 36	10 42	9 35	9 41	9.5 ± 0.53 37.4 ± 3.96	9 36	10 33
DTR	16	19	19	22	18	18	20	18	18.8 ± 1.75	19	18
EFS LF4	3 14	5 14	4 14	7 16	6 14	5 15	4 14	4 15	4.8 ± 1.28 14.5 ± 0.76	5 15	5 15
LT4	15	15	18	19	17	17	18	17	17 ± 1.41	17	16
EPS PP	16 0	14 0	19 0	10 0	10 0	20 0	20 0	20 0	16.1 ± 4.35 0	15 0	14 0

For coloration in life see Fig. 1 . During daytime, the colour is darker, making the colour pattern more indistinct. FIGURE 2. a) Habitat of Cyrtodactylus cattienensis sp. nov. showing secondary lowland rainforest, dominated by Lagerstroemia sp., b) Two call frequency structure types of male acoustic signals of Cyrtodactylus cattienensis sp. nov. , and c) Sonogram showing the male acoustic signal of Cyrtodactylus cattienensis sp. nov. consisting of a long series of call groups. FIGURE 3. Map showing the currently known localities of Cyrtodactylus cattienensis sp. nov. in Vietnam Variation of paratypes . For the variation of paratypes see Table 1 . Males are somewhat smaller than the females (SVL males 55.4 ± 5.11 mm , n = 11; females 60.5 ± 8.0 mm, n = 8). Paratypes ZFMK 88090 and ZFMK 88094 have dark dorsal bands longitudinally connected to each other, thus enclosing brownish grey patches (three in ZFMK 88090 and five in ZFMK 88094). Paratype IEBR A.0854 shows a considerably lighter (cream) ground colour between the dorsal bands. The paratypes IEBR A.0854-A.0855, VNUH 2008.09.0 7, and ZFMK 88090 have regenerated tails without dark brown bands. Paratype IEBR 656 was fixed in formalin and thus shows a yellowish-brown ground colour instead of a greyish one. Paratype ZFMK 88091 shows a bifurcation at the posterior 10 mm of its tail. The hemipenes of paratype ZFMK 88095 are partially everted. Their apex is thickened and deeply bifurcated. Sulcus spermaticus thin, deep, bordered by bulging lips; the sulcus bifurcates in the upper truncus region. Apical lobes with serrated skin rims. Comparisons. Comparisons are based on the original descriptions or descriptions provided in broader faunal and taxonomic publications (e. g., Smith 1920 , 1921 , 1935 ; Taylor 1963 ; Ulber 1993 ; Darevsky & Szczerbak 1997 ; Das 1997 ; Manthey & Grossmann 1997 ; Chan-ard et al. 1999 ; Das & Lim 2000 ; Ziegler et al. 2002 ; Bauer 2002, 2003; Bauer et al. 2002, 2003; David et al. 2004; Pauwels et al. 2004 ; Batuwita & Bahir 2005 ; Grismer 2005; Grismer & Leong 2005 ; Kraus & Allison 2006 ; Nguyen et al. 2006 ; Youmans & Grismer 2006 ; Grismer et al. 2007 , 2008; Heidrich et al. 2007 ; Hoang et al. 2007 ; Kraus 2007 ; Orlov et al. 2007; Grismer & Norhayati 2008 ; Nazarov et al. 2008 ; Ngo 2008; Ngo & Bauer 2008 ; Ngo et al. 2008; Rösler & Glaw 2008 ; Rösler et al. 2008). In comparison with Vietnamese congeners, Cyrtodactylus cattienensis sp. nov. is distinguished from C. badenensis , C. caovansungi , C. condorensis , C. eisenmani , C. grismeri , C. hontreensis , C. intermedius , C. nigrocularis , C. paradoxus , C. phongnhakebangensis , and C. takouensis by the absence of transversally enlarged subcaudals. Cyrtodactylus cattienensis sp. nov. differs from C. chauquangensis and C. cryptus by having enlarged precloacal scales (absent in latter species). The new species further differs from C. chauquangensis by having a smaller SVL ( 43.5–69 mm versus 90.9–99.3 mm ) and lacking precloacal pores in females (present in females of C. chauquangensis ). Cyrtodactylus cattienensis sp. nov. can also be distinguished from C. cryptus by having fewer ventrals in longitudinal rows (28–42 versus 47–50), a lower number of precloacal pores in males (6–8 versus 9–11), and fewer subdigital lamellae on fourth toe (14–19 versus 20–23). Cyrtodactylus cattienensis sp. nov. differs from C. huynhi by having fewer ventrals in longitudinal rows (28–42 versus 43–46) and lacking femoral pores in males (1–5 pores on each thigh in males of C. huynhi ). Cyrtodactylus cattienensis sp. nov. is most similar to the members of the C. irregularis complex. However, the new species differs from C. pseudoquadrivirgatus by having a continuous nuchal band and enlarged femoral scales (nuchal band interrupted and enlarged femoral scales absent in C. pseudoquadrivirgatus ). The new species is distinguished from C. irregularis and C. ziegleri by having a smaller SVL ( 43.5–69 mm versus 72–86 mm in C. irregularis and 84.6–93 mm in C. ziegleri ). Cyrtodactylus cattienensis sp. nov. further differs from C. irregularis by having smooth enlarged femoral scales (enlarged femoral scales containing pits in C. irregularis ), by lacking light margins around the dark patches on the back and enlarged spurs on the tail-base whorls (present in C. irregularis ). The new species also differs from C. ziegleri by lacking femoral pores in males (present in males of C. ziegleri ). For further comparisons see Table 2. We also compare Cyrtodactylus cattienensis sp. nov. with its remaining congeners from the neighbouring countries in mainland Southeast Asia whose species lack enlarged subcaudal scales or femoral pores in both sexes. Cyrtodactylus cattienensis sp. nov. differs from C. angularis and C. feae by having discontinuously enlarged precloacal-femoral scales; the new species differs from C. aurensis , C. gansi , C. pulchellus , C . semenanjungensis , C. stresemanni , and C. tiomanensis by lacking a precloacal groove; from C. ayeyarwadyensis by having a lower count of precloacal pores in males (6–8 versus 10–28 in C. ayeyarwadyensis ); from C. brevidactylus in dorsal pattern (5–7 irregular dark bands versus 3–4 black dorsal blotches in C. brevidactylus ) and by having enlarged femoral scales; from C. buchardi by having precloacal pores in males (absent in males of C. buchardi ); from C. chrysopylos by the absence of a single, much larger pore bearing scale posterior to the precloacal pore series (present in C. chrysopylos ); from C. consobrinus by having a lower number of ventrals in longitudinal rows at midbody (28–42 versus 65–70 in C. consobrinus ); from C. elok by having a higher count of dorsal tubercles (16–22 versus 6–10 in C. elok ) and a lower number of ventrals in longtudinal rows at midbody (28–42 versus 44 in C. elok ); from C. interdigitalis and C. sumonthai by lacking well developed lateral folds with large tubercles (present in latter species); from C. jarakensis in the presence of enlarged precloacal and femoral scales (absent in C. jarakensis ); from C. lateralis by having a lower number of ventrals in longitudinal rows at midbody (28–42 versus 60–64 in C. lateralis ); from C. quadrivirgatus by having a higher count of precloacal pores in males (6–8 versus 4 in C. quadrivirgatus ); from C. sworderi by having enlarged femoral scales (absent in C. sworderi ); and from C. wakeorum by having a lower count of longitudinal dorsal tubercle rows (16–22 versus 24 in C. wakeorum ). Distribution. Cyrtodactylus cattienensis sp. nov. is currently known from Dong Nai Province (Cat Tien National Park and Vinh Cuu Nature Reserve) and Ba Ria – Vung Tau Province (Nui Dinh) ( Fig. 3 ). Etymology. We name this species after Cat Tien National Park where the holotype and a part of the type series were collected and which represents the first locality where the new Cyrtodactylus species was discovered. The National Park was established in 1978 and it represents one of the largest remaining and most important lowland forest areas in Vietnam with a total land area of 71.920 hectares and the protection of its in part unique biodiversity deserves special attention. Ecological notes. Most of the observations took place at night. The specimens were found on large leaves of scrub vegetation and young trees in lowland forest, at a height of about 1 m above ground (even during rain). A few specimens were found during the day, beneath volcanic rocks or below the bark of rotten trees. Bioacoustic notes. The vocal repertoire of the new species consists of two signal types : 1) Squeak: an acoustic signal of discomfort, produced by males, females and juvenile specimens when they are stressed (e. g., during capture); 2) Male signal ( Fig. 2 b). This acoustic signal can only be produced by males. It consists of a long series (6.55– 22.36 s) of call groups ( Fig. 2 c). The single call groups consist of 1–4 calls each. The intervals between call groups within a series vary from 0.85– 1.25 s. Intervals usually increase by the end of the series. At the beginning of a series the call loudness is -45dB re 0dB and distinctly increases towards the end of the series (-23.5 dB re 0dB). We observed two call frequency structure types ( Fig. 2 b): 1) single call and first call in a call group, respectively ( Fig. 2 c): these calls are frequency modulate and always consist of three harmonics (405, 1395, and 1825 kHz), the dominant average frequency is 900 kHz and 2) subsequent calls in a call group: these calls are also frequency modulate, but include two or three harmonics (1285, 2052, and 3180 kHz), the dominant average frequency is 568 kHz.