Cyrtodactylus nuaulu, Oliver, Paul, Edgar, Paul, Iskandar, Djoko T. & Lilley, Ron, 2009

Oliver, Paul, Edgar, Paul, Iskandar, Djoko T. & Lilley, Ron, 2009, A new species of bent-toed gecko (Cyrtodactylus: Gekkonidae) from Seram Island, Indonesia, Zootaxa 2115, pp. 47-55 : 48-54

Cyrtodactylus nuaulu sp. nov.

Figures 1–2

Holotype: MZB lace 2326 (F-num S80) adult male, 26/08/87, ~ 50m asl, Solea (2°51' S, 129° 39' E), Manusela National Park, central Seram Island, Maluku Province, Indonesia, collected by P. Edgar and R. Lilley.

Paratypes: MZB lace 2325 (F-num S52) adult male, 20/10/87, MZB lace 2327 (F-num S152) adult female, 13/09/87 with same locality and collector details as holotype; MZB lace 2328 (F-num S203) adult female 20/10/87 and MZB lace 2329 (F-num S227) juvenile 29/10/87 from Waikawa, Manusela National Park, 3 km from Saunulu Village, inland from Japutih (3°17' S 129°31' E) southern Seram Island, Maluku Province, Indonesia.

Diagnosis. Cyrtodactylus nuaulu sp. nov. can be distinguished from all other Melanesian and Wallacean Cyrtodactylus by the following unique combination of character states: moderate size (SVL up to 88.5mm); relatively slender body with robust head (HW/SVL 0.187–0.195); deep precloacal groove with low number of pores (6); femoral pores absent; subcaudal scales granular, not transversely enlarged; small dentate tubercles over the dorsum and along the lateral fold; whorls of prominent dentate tubercles extending to the tip of the tail; and dorsal colouration consisting of relatively few large indistinct transverse and/or longitudinal dark brown blotches.

Description of holotype. A moderately large (81.7 SVL mm) and slender gecko. Head long (HL/SVL 0.253), moderately wide (HW/HL 0.744) and distinct from neck. Snout tapering to relatively blunt tip in dorsal profile, relatively long (longer than eye diameter), loreal region weakly inflated, interorbital region and top of snout strongly concave, canthus rostralis smoothly rounded. Eyes very large with vertical pupil. Supracillaries extending from anterior-ventral to posterior-dorsal edge of eye, longest at the anterior-dorsal corner. Ear opening small, dorso-ventrally flattened, and oriented at 45 degrees to apex of rictus with dorsal edge posteriormost, bordered by small indistinct ventral skinfold.

Rostral approximately twice as wide as high, with short medial suture, widest at the ventral edge of the nares, bordered dorsally by right supranasal, larger rounded internasal, and damaged left supranasal. Nares bordered by first supralabial, rostral, first supranasal (point contact only) and series of three elongate postnasals. Supralabials: 10 on right lip and 11 on left, 7 or 8 to midpoint of eye, supralabials anterior to eye much broader than high and bordered dorsally by a single series of variably sized enlarged scales. Head scales small and granular, temporal and nuchal scales smaller than those on snout, scattered small conical tubercles on temporal and nuchal regions. Infralabials to rictus: 9 on right, 10 on left, all much broader than high, bordered by several rows of enlarged scales grading into small and granular gular scales. Mental triangular, approximately as wide as long, bordered by first infralabials and two diamond-shaped postmentals in contact for approximately 50% of their length.

Body elongate (TrL/SVL 0.494) with moderately distinct ventrolateral folds. Moderately tuberculate, lateral fold with distinct dentate tubercles separated from each other by 2–6 granules, posterior tubercles closer together. Dorsum with approximately fourteen rows (including lateral fold) of dentate tubercles. Dorsal scales small and granular. Ventral scales much larger than dorsal scales, increasing in size medially, arranged in approximately 51 rows at midpoint of body. Prominent raised straight precloacal groove surrounded by several rows of enlarged ventral scales, largest scales at base of groove containing 3 precloacal pores on each side. Distinctly enlarged rows of femoral scales absent.

Forelimbs relatively elongate (FA/SVL 0.158), hindlimbs much more robust than forelimbs (CS/SVL 0.191). Lateral and dorsal surfaces of limbs with rows of dentate tubercles. Digits long and well developed, inflected at basal interphalageal joints; subdigital lamellae smooth, rounded, undivided and expanded proximal to joint inflection; large recurved claws sheathed by a dorsal and ventral scale. Slight basal webbing between digits II–IV on both manus and pes; lamellae on digits I /IV of manus 8/15R 10/15L and pes 12/17R 9/ 15L.

Tail original, long and slender, tapering to point with distinct lateral groove extending approximately two thirds its length. Caudal scales granular, increasing in size ventrally, numerous rows of prominent dentate tubercles extending along all surfaces of tail for its full length, including 2 ventral rows. Hemipenal bulge swollen and prominent, left hemipenis everted, two enlarged postcloacal tubercles present at base of tail.

Colouration. Dorsal ground colour light brownish grey with extensive fine brown flecking; a pair of almost continuous (broken just anterior to the hindlimbs) dark brown dorsolateral streaks extend from behind the eye to base of the tail; 2 darkish brown transverse blotches between the fore and hindlimbs. Nuchal region light brown with dark brown V-shaped blotch. Lateral regions light grey with small amounts of dark brown pigmentation forming indistinct scattered blotches and a faint discontinuous brown ventro-lateral stripe between fore and hindlimbs, strongest anteriorly and becoming very indistinct posteriorly. Venter grey with scattered light brown speckling, densest between the forelimbs.

Head light grey dorsally, bordering relatively sharply against brown nuchal region, three small dark brown blotches forming Y-shape just posterior to orbital, additional dark brown longitudinal blotches medial to both eyes, above the rostrum and along the dorsal edge of the off-white supralabials. Supracillaries off-white and dark brown. Broad indistinct lateral band extends from orbital, above ear, and joins dorsolateral bands on the body. Infralabials off-white with scattered small dark brown blotches, ventral surface of lower jaw yellowishgrey with scattered indistinct greyish-brown flecking.

Limbs dorsally light greyish brown with wide indistinct dark brown bands; bands on hindlimbs much lighter than forelimbs; digits mottled light and dark brown, with off-white bands proximal to claws. Ventrally limbs and digits greyish yellow with extensive brown speckling and blotching, especially on the crus. Tail with six wide dark brown bands and six (including tail tip) narrower light greyish to off-white bands; posteriormost dark bands slightly broken up by small amounts of light grey mottling.

Variation. Comparative mensural and meristic data for the holotype and paratypes is given in Table 1 . While broadly consistent, the colouration of the adult types shows significant differences ( Fig. 2 ). All adult paratypes possess three dark to very dark brown transverse bands or blotches between the fore and hindlimbs (versus two in the holotype). The shape and definition of these marking also varies considerably; the holotype is the only specimen with dark dorsolateral stripes that extend to the legs, on other specimens these stripes tend to be lighter and at most extend to the midpoint of the body. The extent and darkness of other dorsal marking also varies, although all specimens possess a dark nuchal V- or Y-shape and at least some dark mottling or blotches between the eyes. The base dorsal colour of MZB lace 2328 is more brownish than grey. All types have at least some brown mottling on the ventral and lateral surfaces, however the density and extent of pigmentation again varies (the venter of MZB lace 2335 is particularly densely flecked).

On the two adult individuals with original tails the number of bands varies from five to six (dark bands) to four to five (light bands); the tip of the original tail is dark in one of the paratypes (MZB lace 2325) as opposed to light in the holotype (MZB lace 2326). MZB lace 2327 has an almost complete original tail (1.5cm regrown at the tip) which was broken at collection, with five light and five dark bands. Approximately 60% of the tail on paratype MZB lace 2328 is original with three off white bands and two greyish brown bands; the regrown section is yellowish off-white with extensive brown pigmentation, no tubercles and uniform scalation.

MZB lace 2329 is a recently hatched juvenile, with original tail that was broken at collection and a circular patch of skin missing from around the left ear. The dorsal pattern on this specimen is simplified relative to the adult types and consists of two brown longitudinal dorsolateral streaks, a dark brown nuchal blotch and small brown blotch at the midpoint of the hindlegs. The lateral and ventral regions are also relatively plain with less brown pigmentation than the other types.

Comparisons. The whorls of dentate tubercles extending to the tip of the original tail readily distinguish C. nuaulu sp. nov. from its two recognised Moluccan congeners; C. deveti and C. halmahericus . Cyrtodactylus deveti can be further distinguished by its more robust build, absence of a precloacal groove, possession of several rows of enlarged femoral scales, and much larger ventral caudal scales. Cyrtodactylus halmahericus , collected both sympatrically and on other Moluccan islands, are smaller (<74mm), have a long continuous series of precloacal and femoral pores (21–26), have at most a few small spines and tubercles on the anterior portion of tail only, lack enlarged ventrolateral tubercles at any point on the tail and have a dorsal body pattern consisting of more than three transverse dark brown bands or series of blotches.

MZB2326 MZB2325 MZB2327 MZB2328 MZB2329

holotype paratype paratype paratype paratype Populations of small Cyrtodactylus ascribed to C. papuensis (Brongersma) from New Guinea and surrounding islands, lack enlarged tubercles on the tail and dorsum, are smaller (<70mm), and often have a distinct dorsal pattern including dark cross-bands. The new species can be distinguished from all other Melanesian Cyrtodactylus by the presence of a precloacal groove. It can be further distinguished from the only other Melanesian species with enlarged tubercles along the tail, Cyrtodactylus serratus Kraus , (and related members of the C. loriae and C. louisiadensis species groups sensu Rösler et al. 2007) by its relatively small size and very small ventral caudal scales.

Cyrtodactylus darmandvillei (Weber) from the Lesser Sundas lacks a precloacal groove, is much more robust and has much larger trihedral tubercles across the dorsum (see figure in De Rooji 1915). Cyrtodactylus wetariensis (Dunn) , also from the lesser Sundas, is smaller <70mm, possesses large trihedral dorsal tubercles and has 12–13 femoral pores on each hindlimb ( Dunn 1927). Cyrtodactylus spinosus Linkem et al. from Sulawesi has long thin spines along the lateral fold, on the tail and on the postantefemoral region of the hindlimbs ( Linkem et al. 2008). Other Indonesian Cyrtodactylus from islands to the west and north of Seram lack rings of enlarged tubercles extending the full length of the tail. Of the most geographically proximate species, C. marmoratus (Gray) (Java) and C. fumosus (Müller) (Java and Sulawesi) can be distinguished by possessing a femoral pore series, while C. jellesmae (Boulenger) and C. wallacei Hayden et al. from Sulawesi lack a precloacal groove; all these species also have far more extensive and darker transverse banding or blotching across the dorsum ( Hayden et al. 2008).

Distribution and Natural History. Cyrtodactylus nuaulu sp. nov. is only known from two localities in lowland rainforest from the island of Seram. The forest around the type locality was dominated by Eucalyptus deglupta and Shorea sp. All adult types were collected by hand at night between 50–200cm above the ground, head down on trees and vines. The juvenile paratype was collected in a pit-trap at night. Other geckos collected in sympatry or near sympatry were Cyrtodactylus halmahericus , a probably undescribed small (<41mm) Nactus sp. (listed as Cyrtodactylus pelagicus in Edgar & Lilley 1993) and Gehyra mutilata . Both collection localities are in Manusela National Park, which at the time of collection, was being threatened by illegal logging and conversion of forest to gardens.

Eytmology. Named in honour of the Nuaulu people of south Seram.

Cyrtodactylus nuaulu sp. nov. shows a combination of morphological characters that is very distinctive and does not strongly suggest a close relationship with any described Cyrtodactylus . While biogeographic studies and field surveys have found that the mammal, bird and frog fauna of Seram have a strong Melanesian influence ( Edgar & Lilley 1993; Helgen 2003), most Melanesian Cyrtodactylus lack a precloacal groove, possess obvious femoral pores and have a relatively robust build. We tentatively suggest that Cyrtodactylus nuaulu sp. nov. is most likely to be allied to a number of smaller bodied, relatively gracile taxa with precloacal grooves distributed both to the west (C. cf marmoratus ) and east ( C. halmahericus , C. papuensis ) of Wallace's Line. Nonetheless, we emphasise that the distinctive morphology of this species makes this association tenuous at best.

Seram and surrounding islands (especially Buru) are recognised areas of endemism for birds ( Stattersfield et al. 1998) and mammals (Flannery 1995; Helgen 2003). With the description of Cyrtodactylus nuaulu sp. nov. (known only from Seram) four reptile species are known to be endemic to this region of the Maluku Islands. The other endemic and near endemic species (also found on the neighbouring small island of Ambon) are the recently described monitor Varanus ceramboinensis ( Philipp et al. 1999) , Carlia leucotaenia and Calamaria ceramensis . Not surprisingly given the biogeographic position of Seram, these endemics include representatives of both Australasian ( Carlia ) and Asian ( Calamaria ) groups. As with mammals and birds, it seems that the herpetofauna of Seram and surrounding islands includes a small but significant and biogeographically interesting endemic component. It would also seem likely that further taxonomic and survey work in Seram and surrounding islands will add to this total. In particular the neighbouring large island of Buru remains very poorly explored and should be a priority for further surveys.

Figures

FIGURE 1. Details of holotype (MZB lace 2326) of Cyrtodactylus nuaulu sp. nov. A) dorsal view of whole animal B) lateral view of head and neck C) ventral view of head and neck D) ventral view of cloacal region and E) anterior section of tail showing whorls of enlarged dentate tubercles and lateral groove. Scale = 10 mm.

FIGURE 2. Dorsal view of type series of Cyrtodactylus nuaulu sp. nov. From left to right MZB lace 2325, MZB lace 2326 (holotype), MZB lace 2327, MZB lace 2328, MZB lace 2329. Scale = 20 mm.

Tables

TABLE 1. Measurements (in mm) and scale counts for the type series of Cyrtodactylus nuaulu sp. nov.

Sex	m	m	f	f	juv
SVL	81.7	86.3	88.4	76.9	35.7
TL	106.0	107.0	101.0	78.5	41.0
Original tail	100%	100%	85%	60%	100%
Regrown tail	na	na	15%	40%	na
TrL	38.1	42.6	43.9	37.4	16.5
HL	21.6	21.9	23.3	20.3	10.1
HW	15.8	16.2	17.0	14.6	7.0
HH	9.9	9.9	11.0	8.4	4.2
EYE	4.8	5.5	5.4	4.9	2.8
FA	13.3	13.6	14.1	13.4	7.2
EN	7.4	7.9	8.5	7.1	3.7
IN	2.6	2.7	3.0	2.4	1.4
EAR	1.3	1.9	2.1	1.3	na
CS	15.6	16.5	16.4	15.4	7.0
SUPR(R)	10	12	12	11	12
SUPR(L)	11	11	11	11	12
INFR(R)	9	9	10	10	10
INFR(L)	10	9	10	10	9
POMEN	2	2	2	2	2
DTR	13–14	14	14	13–14	15–16
LTTUBR	25	26	28	23	20
LTTUBL	25	26	26	22	21
VENT	51	48	55	49	na
LAM1R	9	11	10	11	11
LAM1L	12	11	11	11	12
LAM4R	15	19	18	20	18
LAM4L	17	20	19	19	19
PORES	6	6	0	0	na
PATUB	2	2	2	2	1