Goniurosaurus catbaensis, Ziegler, Thomas, Truong, Nguyen Quang, Schmitz, Andreas, Stenke, Roswitha & Rösler, Herbert, 2008

Ziegler, Thomas, Truong, Nguyen Quang, Schmitz, Andreas, Stenke, Roswitha & Rösler, Herbert, 2008, A new species of Goniurosaurus from Cat Ba Island, Hai Phong, northern Vietnam (Squamata: Eublepharidae), Zootaxa 1771, pp. 16-30 : 18-26

Goniurosaurus catbaensis sp. n.

Holotype. IEBR A.0717, an adult male ( Figs. 1–2 ), from Cat Ba Island, Cat Hai District, Hai Phong City, northern Vietnam, collected by Roswitha Stenke, Nguyen Quang Truong and Thomas Ziegler on 23 May 2007 near Tra Bau Ranger Station in Ong Bi valley at an altitude of ca. 10–20 m above sea level.

Paratypes. All from Cat Ba Island, Cat Hai District, Hai Phong City, northern Vietnam, collected by Roswitha Stenke, Nguyen Quang Truong and Thomas Ziegler at an altitude of ca. 10–30 m above sea level: MHNG 2699.49, a subadult to adult male ( Fig. 3 ), coll. 20 May 2007 in the surroundings of Trung Trang Cave; VNUH 210507, an adult female ( Fig. 4 ), coll. 21 May 2007 near Ang Dai valley; and ZFMK 87056, an adult male, coll. 23 May 2007 near Tra Bau Ranger Station in Ong Bi valley.

Diagnosis. A species of the genus Goniurosaurus , characterized by a combination of the following characters: 1) gracile body and limbs; 2) thin, posteriorly protracted nuchal loop; 3) three (or four) thin immaculate dorsal body bands between limb insertions, without dark spotting and with dark, narrow border surrounding the body bands; 4) mottled dorsal surface and maculate limbs; 5) head pattern consisting of a dark marbling; 6) iris orange-brown; 7) lack of postrostral (internasal) scales (supranasals meeting in midline behind the rostral suture); 8) 5–6 nasal scales surrounding the nare; 9) greatly enlarged row of supraorbital tubercles; 10) outer surface of the upper eyelid composed of granular scales of about the same size of those on top of head and with a longish row of 6–9 enlarged tubercles; 11) 52–55 eyelid fringe scales; 12) 8–9 supralabials, and 6–8 sublabials; 13) granular body scales, with 8–11 granular scales surrounding the dorsal tubercles; 14) deep axillary pockets; 15) long, thin digits with wide subdigital lamellae and claws being sheathed by four scales; 16) 16–21 precloacal pores.

Description of holotype. Adult male; SVL 111.5 mm; TaL 101.5 mm; SE 12.6 mm; EE 11.5 mm; HW 20.7 mm; HH 12.6 mm; HL 30.6 mm; SVL: TaL 1.10; SVL: HL 3.64; HL: HW 1.48; HL: HH 2.43; SE: EE 1.10; head triangular, wider than neck, covered with uniform granular scales interspersed with tubercles in the temporal and occipital regions; scales of rostrum slightly larger and flatter; conspicuous row of enlarged supraorbital tubercles; middorsal portion of rostral partially sutured dorsomedially, bordered laterally by first supralabial, dorsolaterally by prenasal, and dorsally by two supranasals; postrostral (internasal) scales are lacking; external nares bordered by five nasals each: anteriorly by prenasal and supranasal, dorsally by supranasal, posteriorly by two slightly enlarged postnasals and one smaller granular scale, and ventrally by the prenasal; prenasals with long recurved ventral portion; supranasals triangular, meeting in midline behind rostral suture; supralabials nine / eight, grading into granular scales posteriorly; eyes relatively large, pupils vertical; eyelid fringe scales (ciliaria) 54 / 52, those of upper eyelid slightly enlarged; outer surface of upper eyelid composed of granular scales of about the same size of those on top of head, including a longish row of ca. six enlarged tubercles; a fold of skin originating in the suborbital region extends posteroventrally across the angle of the jaw; external auditory meatus elliptical with long axis directed dorsoventrally; tympanum deeply recessed; mental triangular, bordered laterally by first infralabial and posteriorly by three postmentals; postmentals bordered by six gular scales; infralabials eight / eight, grading posteriorly into smaller scales; gular scales juxtaposed and granular, abruptly grading posteriorly into flat hexagonal pectoral scales and even larger ventral scales. Neck narrower than body, covered with uniform granular scales interspersed with several sharply pointed conical tubercles on nape; tubercles on flanks conical, those of vertebral region somewhat lower in profile; body tubercles not increasing in size posteriorly, grading into somewhat more pointed caudal tubercles; dorsal body tubercles surrounded by 10 granular scales; 23 dorsal tubercle rows at midbody; 33 paravertebral tubercles between limb insertions, distinct vertebral row of scales absent. Limbs relatively long and thin, covered dorsally with granular scales interspersed with several tubercles and ventrally with flat juxtaposed to subimbricate scales; dorsal granular scales grade into slightly flattened subimbricate scales on top of pes and manus; hind limbs slightly larger than forelimbs; larger granular scales on ventral surfaces of pes and manus; deep axillary pockets present; subdigital lamellae wide, 12 / 12 on first finger, 16 / 15 on second finger, 18 / 17 on third finger, 18 / 19 on fourth finger, 19 / 18 on fifth finger, as well as 11 / 12 on first toe, 17 / 15 on second toe, 19 / 20 on third toe, 23 / 24 on fourth toe, and 21 / 22 on fifth toe; digits laterally compressed, increasing in length from first to fourth, fifth shorter than fourth. Body relatively long and thin, covered with granular scales grading ventrally into larger hexagonal flattened subimbricate ventral scales; 127 scales around midbody; larger ventral scales grade abruptly into smaller granular scales immediately anterior to the vent at the level of the preanal pores; 16 preanal pores in a transverse continuous series beginning and ending somewhat beyond the hind limb insertions; region posterior to vent covered with flat juxtaposed scales and greatly swollen, containing three enlarged tubercles (postcloacal tubercles) laterally on each side at the level of the vent. Tail long and thin, thickest at base, anteriorly with whorls; dorsal tail scales flat, smooth, up to 1.5 times the size of dorsal scales, arranged in more or less regular transverse rows; the first tail whorl with five, the fourth and eighth with six scale rows; the first whorl with six, the fourth whorl with five, and the eighth whorl with one tubercles in a transversal series; caudal tubercles occur in hemiwhorls, being present only dorsally and fading laterally and posteriorly; ventral caudals larger than dorsal caudals.

Coloration in life: Dorsal ground colour of head, body and limbs grey-brown to pale brown and mottled with dark brown blotches; iris orange-brown; thin posteriorly tapered dull yellow nuchal loop present, anterior ends inserting on lower eyelids, edged dorsally and ventrally in dark brown; three thin immaculate yellowish bands on body between limb insertions, and another one on tail base, all edged anteriorly and posteriorly by thin dark brown bands; ground colour of tail grey brown to dark brown, tail base mottled; five immaculate white caudal bands edged anteriorly and posteriorly in solid black; ventral surfaces of head, body and limbs dull white and immaculate except for few dark spots on limbs; weak brown lateral spotting in gular region and on venter and limbs; subcaudal region dark brown with distinct lighter interspaces between caudal bands; caudal bands complete ventrally except for first.

Variation of paratypes. The paratype series largely corresponds to the description of the holotype. For measurements, proportions and scalation data of the paratypes, which all have regenerated tails, see Table 1 . However, the external nares of the specimens VNUH 210507 and MHNG 2699.49 are bordered by six instead five nasals each: anteriorly by prenasal and supranasal, dorsally by supranasal, posteriorly by two slightly enlarged postnasals and two smaller granular scales, and ventrally by the prenasal. In addition, the surface of the upper eyelid of the paratype series contains a longish row of up to nine enlarged tubercles, and the postmentals are bordered by up to eight gular scales. With respect to the immaculate yellowish body bands between limb insertions, specimens VNUH 210507 and MHNG 2699.49 show a somewhat differing banded pattern: MHNG 2699.49 has an additional yellowish-orange band in front of the last (and in addition incomplete) band, which consists of a series of three large light blotches, and VNUH 210507 has an additional, incomplete band that is medially fused with the last body band. VNUH 210507 and MHNG 2699.49 also have only four immaculate white caudal bands. The light caudal bands are complete ventrally in all paratypes.

Further records. Five further specimens could be observed on Cat Ba Island. Three of them were collected, and two of them released after photography ( Fig. 5 ). The remaining specimen is currently kept alive for behavioural studies in Vietnam and will subsequently be donated to the scientific collection of the Cat Ba National Park. All three collected specimens corresponded in external morphology and colour pattern with the holotype description. The latter three specimens had three thin immaculate yellowish bands on body between limb insertions and lacked one or more postrostral (internasal) scales, thus having the supranasals in broad contact behind the rostral suture.

Etymology. The specific name refers to the type locality Cat Ba Island. The scientific name emphasizes the importance and uniqueness of the Biosphere Reserve Cat Ba Archipelago, Hai Phong, which also houses an endemic primate, the Cat Ba langur Trachypithecus p. poliocephalus ( Roos et al. 2001). We propose the standard name “Cat Ba Leopard Gecko”.

Comparisons. The new species most closely resembles G. l u i i, from which it consistently differs by lacking postrostral (internasal) scales and by having the supranasals meeting in midline behind the rostral suture ( Fig. 6 ): Besides the G. l u i i listed in Table 2 we studied thirteen additional G. luii from continental China and Vietnam (i.e., another record from Cao Bang Province, Vietnam, and three specimens from China kept in the Cologne Zoo, as well as nine Chinese specimens from the private holdings of one of the authors [HR]), which all had the supranasals separated by at least one postrostral scale (see holotype description in Grismer et al. 1999: 383), whereas the type series and further records of G. catbaensis sp. n. (n = 7) consistently lacked a postrostral and thus had the supranasals in broad contact. Also the Chinese specimen depicted on page 159 in Seufer et al. (2005) shows the supranasals being separated by at least one postrostral scale. G. catbaensis sp. n. further differs from G. l u i i in having the outer surface of the upper eyelid composed of granular scales of about the same size of those on top of head (versus small granular scales one-half the size of those on top of head in G. luii , see Grismer et al. 1999: 383), including a longish row of ca. six to nine enlarged tubercles (which is lacking in G. luii ). In addition, the type series of G. catbaensis sp. n. had fewer dorsal tubercles in the eighth caudal whorl compared with G. luii (one versus 3–6), fewer precloacal pores (5–21 versus 21–29 in G. luii ), a lower number of nasal scales surrounding the nare (5.1±0.4 versus 6.8± 0.8 in G. l u i i), fewer supra- and sublabial counts (8–9 and 6–8 versus 9–12 and 8–11 in G. l u i i), and a lower count of granular scales surrounding the dorsal tubercles (9.8±1.3) versus 12.2± 1.3 in the type series of G. l u i i and versus 12.3± 1.3 in the four continental G. l u i i listed in Table 2 . Concerning the colour pattern, it is most obvious that the nuchal loop is thinner and much more tapered backwards (v-shaped) in G. catbaensis sp. n. than it is the case in G. luii ; furthermore, the dark head pattern rather consists of a dark marbling in G. catbaensis sp. n. instead of dark brown blotches in G. l u i i.

G. catbaensis sp. n. differs from all other species of Goniurosaurus in having a greatly enlarged row of supraorbital tubercles. With respect to the remaining Vietnamese species, G. catbaensis sp. n. additionally differs from G. a r a n e u s in lacking wide body bands and nuchal loop, in having a mottled dorsal surface, which is nearly immaculate and lacking mottling in G. araneus , in having maculate versus nearly immaculate limbs, having fewer precloacal pores (16–21 versus 23–29), and having granular body scales as opposed to elongate dorsal body scales. G. catbaensis sp. n. differs from G. lichtenfelderi in having a gracile body and limbs as opposed to being more compact and robust, having 3–4 versus two dorsal body bands, dark, narrow border surrounding the body bands as opposed to a wide border, a posteriorly protracted nuchal loop as opposed to being rounded posteriorly, and deep axillary pockets.

C. catbaensis sp. n. differs from C. hainanensis in having a gracile body and limbs versus more compact and robust, having 3–4 versus two dorsal body bands, lacking dark spotting in body bands, having a posteriorly protracted nuchal loop as opposed to being rounded posteriorly, deep axillary pockets, and 16–21 precloacal pores as opposed to 26–32.

C. catbaensis sp. n. differs from C. bawanglingensis by lacking a more robust body stature, by having long, thin digits as opposed to short, robust digits, having immaculate body bands as opposed to a boldly mottled dorsal ground colour with dark spotting in the body bands, by having the outer surface of the upper eyelid composed of granular scales of about the same size of those on top of head (versus granular scales being smaller than the granular scales on the top of the head), by having 16–21 precloacal pores as opposed to 37– 46, and by having a lower number of eyelid fringe scales (52–55 versus 56–67).

G. catbaensis sp. n. further differs from the G. k u ro i w a e group in having a gracile body and limbs as opposed to being more compact and robust, dark, narrow border surrounding the body bands as opposed to a wide border, a posteriorly protracted nuchal loop as opposed to being rounded posteriorly, having precloacal pores and deep axillary pockets, having wide versus narrow subdigital lamellae, and claws being sheathed by four scales, as opposed to being unsheathed and surrounded by 6–8 scales ( Grismer et al. 1999, 2002).

Distribution. The new species is known only from Cat Ba Island ( Figs. 7–8 ).

Natural history. G. catbaensis sp. n. was found active at night in limestone forest ( Fig. 5 ). The geckos were mainly found on the forest ground but some of them also on limestone cliffs in up to 2–3 m height. Specimens were found in the surroundings of large limestone caves ( Fig. 9 ) with parts of primary forest vegetation and in the vicinity of primary shrub vegetation on limestone. Once disturbed, the geckos tried to escape into rock holes and crevices.

Figures

FIGURE 1. General aspect of the adult male holotype of Goniurosaurus catbaensis sp. n. (IEBR A. 0717) from Cat Ba Island, Hai Phong, Vietnam. Photograph by Thomas Ziegler

FIGURE 2. Portrait of the adult male holotype of Goniurosaurus catbaensis sp. n. (IEBR A. 0717) from Cat Ba Island, Hai Phong, Vietnam. Photograph by Thomas Ziegler.

FIGURE 3. Subadult to adult male paratype of Goniurosaurus catbaensis sp. n. (MHNG 2699.49) from Cat Ba Island, Hai Phong, Vietnam. Photograph by Thomas Ziegler.

FIGURE 4. Adult female paratype of Goniurosaurus catbaensis sp. n. (VNUH 210507) from Cat Ba Island, Hai Phong, Vietnam. Photograph by Thomas Ziegler.

FIGURE 5. Goniurosaurus catbaensis sp. n. active at night in its habitat in the surroundings of Trung Trang Cave. Photograph by Thomas Ziegler.

FIGURE 6. Snout tip of the holotype of Goniurosaurus catbaensis sp. n. (IEBR A. 0717) from Cat Ba Island, Hai Phong (left), and of G. luii (ZFMK 87057) from Cao Bang Province, continental Vietnam (right); contrary to the situation in G. luii, postrostral (internasal) scales are lacking in G. catbaensis sp. n. and the supranasals are in broad contact behind the rostral suture. Photograph by Thomas Ziegler.

FIGURE 7. Map showing the type locality of Goniurosaurus catbaensis sp. n. (IEBR A. 0717) in the Gulf of Tonkin in northeastern Vietnam.

FIGURE 8. Shoreline of Cat Ba Island with its characteristic limestone formations. Photograph by Thomas Ziegler

FIGURE 9. Entrance of Trung Trang Cave within the habitat of Goniurosaurus catbaensis sp. n. on Cat Ba Island. Photograph by Thomas Ziegler

Tables

TABLE 1. Measurements (in mm), proportions and pholidosis (minimum-maximum [mean ± standard deviation]) of the type series of Goniurosaurus catbaensis sp. n. (n = 4). Abbreviations are as follows: n = number; for remaining abbreviations see material and methods.

	G. c a tb ae n si s sp. n. males (n = 3)	G. c a tb ae n si s sp. n. female (n = 1)
SVL	84.7–111.5 (101.7±14.8)	104.5
TaL	52.5–101.5(71.0±26.6)	59.5
SE	9.8–12.6 (11.7±1.6)	10.8
EE	8.5–12.3 (10.8±2.0)	9.8
HW	16.2–21.6(19.5±2.9)	19.6
HH	10.1–14.3 (12.3±2.1)	12.0
HL	23.1–30.6(28.0±4.2)	26.6
SVL:TaL	1.10 in the holotype	(tail regenerated)
SVL:HL	3.61–3.67 (3.64±0.03)	3.93
HL:HW	1.43–2.11 (1.67±0.38)	1.36
HL:HH	2.29–2.43 (2.37±0.07)	2.22
SE:EE	1.02-1.15 (1.09±0.07)	1.10
SPL	8–9(8.8±0.4)	8/9
SBL	6–8 (7.7±0.8)	8
N	5–6(5.2±0.4)	5
IN	0	0
CIL	52–55(53.7±1.0)	55
PM	2–3 (2.7±0.6)	3
GP	6–7 (6.7±0.6)	8
DTR	23–25 (23.7±1.2)	25
GST	8–11 (9.7±1.5)	10
TL	33–34 (33.3±0.6)	35
MB	112–127 (119.3±7.5)	119
LD1	12	11
LD4	18–19 (18.5±0.6)	17/18
LT1	11–12 (11.5±0.6)	11
LT4	22–24 (23.3±0.8)	23/24
PP	16–21 (18.7±2.5)	5
PAT	2–3 (2.7±0.5)	3
DT1	6–8 (7.3±1.2)	7
DT4	5 in the holotype	(tail regenerated)
DT8	1 in the holotype	(tail regenerated)

TABLE 2. Measurements (in mm), proportions and pholidosis (minimum-maximum [mean ± standard deviation]) of continental Goniurosaurus luii from Cao Bang Province, northeastern Vietnam (n = 2: VNUH 0 10205, ZFMK 87057), from the animal trade (n = 2: CPHR 1924, ZFMK 87058), and from Pingxiang and Longzhou in China (n = 6: after Grismer et al. 1999; the locality Hainan island proved to be wrong according to Seufer et al. 2005: 157). Abbreviations are as follows: n = number; for remaining abbreviations see material and methods.

	VNUH 0 10205 after Vu et al. (2006)	ZFMK 87057	CPHR 1924	ZFMK 87058 after Grismer et al. (1999)
Sex SVL	Female 119.0	Female 105.5	Male 115.5	Male both sexes 80.5
TaL	67.0	99.5	96.5	58.4
SE EE	12.8 11.7	11.8 10.3	12.5 11.5	7.7 7.5
HW	20.0	19.6	22.3	14.0
HH HL	13.4 31.3	12.3 28.0	14.4 28.5	8.3 20.3
SVL:TaL	1.78	1.06	1.20	1.38
SVL:HL HL:HW	3.80 1.57	3.77 1.43	4.05 1.28	3.97 1.45
HL:HH	2.34	2.28	1.98	2.45
SE:EE SPL	1.09 9	1.15 9	1.09 9/10	1.03 10/9 9–12 (9.5±0.6)
SBL	8/9	8/9	8/9	9/8 9–11 (10.0±0.6)
N IN	2	6/7 2	8/7 1	6/7 1 1
CIL		53/52	63	61/62 57–61(59.5±1.9)
PM GP	4	5 11	6 8	2 2–4 (3±0.9) 7
DTR		24	22	22
GST TL	11–12 33	12 32	12 33	14 9–14 (12.2±1.3) 29 33–34 (33.8±0.8)
MB	139	127	120	131 119–144 (134.5±12)
LD1 LD4	11 19/18	10/11 17/20	11/12 18/17	9/10 18/19
LT1	12	11	11/13	9/11
LT4 PP	22/23	24/23 0	23/22 21	20/21 21–24 (23.5±1.4) 27 23–29 (26.0±2.6)
PAT		2	2 /3	2
DT1 DT4		8 4	10 5	6 4
DT8		3	6	3