Lepidodactylus buleli, Ineich, Ivan, 2008

Ineich, Ivan, 2008, A new arboreal Lepidodactylus (Reptilia: Gekkonidae) from Espiritu Santo Island, Vanuatu: from egg to holotype, Zootaxa 1918, pp. 26-38 : 29-35

Lepidodactylus buleli , new species

Figures 1–5

Holotype. MNHN 2008.0004, an adult male from the forest about two walking hours east of Penaoru Village(14.95894° S, 166.63290° E), collected as an egg on 14 November 2006 by Ivan Ineich. It was euthanized with sodium pentobarbitol injection in March 2008 after numerous photographs were taken. A sample of tissue was preserved from the ventral side of its right thigh.

Paratypes.- None.

Etymology. The specific epithet buleli is given by the author as a reference to a personal and private story and has no particular signification related to the species, its characteristics, geographical origin or biology. It is a noun in apposition and not formed from the masculine personal name ‘Bulel’ in the genitive singular case.

Diagnosis. An intermediated-sized Lepidodactylus , body slightly depressed dorso-ventrally, limbs overlapping when adpressed. Two or three deep notched or divided subterminal scansors of second through fifth digits of the fore- and hindfeet (thus belonging to Group II of Brown and Parker 1977). It differs from other Group II members by the following combination of characters: moderately dilated digital pads of pes and manus, lamellae covering most of elongate toe length, presence of one enlarged cloacal spur followed by 2–3 smaller spurs on each side (fig. 5), a short and wide snout (fig. 1), an elevated number of midbody scale rows, uniform lemon-yellow upper- and lower labials and a yellowish head area mixed with purplish dark brown particularly between eye and in the tympanic region (fig. 4), whitish chin and throat (fig. 3), and a distinctive anchor-like yellowish to cream dorsal pattern at the base of tail (fig. 5).

Description of holotype (measurements in mm, after preservation). Snout-vent length 37.5; head length 9.06; head width 6.28; head height 4.68; jaws length from posterior border to snout end left/right 6.44/6.75; snout-eye length 4.33; naris-eye length 3.07; naris circular about 0.48 × 0.40; orbit diameter 2.26; eye-ear length 3.43; snout width (internasal distance) 1.82; ear opening length × width 1.22 × 0.80; interorbital width 3.80; snout-forelimb length 14.44; trunk length 17.37; length of hind limb 16 (62.8 % of axilla-groin distance); length of fore limb 11; crus length 4.69; tail length 40 (entire); tail width 3.59; tail depth 2.93.

Snout rounded (fig. 4), rather short and wide; snout scales (scales touching rostral between left and right nares) 5; supranasals separated by three scales in contact with rostral; rostral entering nares, broader than high, 2.03 × 0.92 (width about 2.2 times height); no rostral cleft; nares bordered by five scales: one postnasal, and two supranasals (in total three nasals), one rostral, and first supralabial; five scales touching rostral between left and right nares; 39–40 interorbital scales (several counts); 11 left and 10 right supralabials, 8th below centre of eye; 10 left and 10 right infralabials; mental scale distinct, triangular, its anterior width nearly equals to midline length (0.91 × 1.08); 17 chin scales and 10 postmentals. Mental followed posteriorly by a pair of enlarged plates also in contact with infralabial I on each side, themselves followed laterally on each side by another enlarged plate, slightly shorter, in contact with infralabial I or its suture with infralabial II, and with infralabial II.

Body slightly depressed; 147 rows of scales (3 counts: 144, 146, 152) around midbody, grading into granular scales on lower lateral surfaces; dorsal and lateral scales granular, without enlarged tubercles, and in juxtaposition; ventral scales almost flat, cycloid, 2–3 times larger than dorsal scales; limbs well developed (figs. 1–3); digital lamellae ventrally covering nearly all of forefoot digits and digits I–II of hindfoot, about ¾ of digits III–V of hindfoot; all digits of fore- and hindfoot clawed except the first; ultimate claw-bearing and penultimate phalanges of fore and hind digits raised above pad and extending over distal edge of pad; fore- and hindfoot webbing slight (less than basal 20% of 2nd digit’s length); breadth of fourth toe about 28–31 % of its length; breadth of fourth finger 31–32 % of its length; left/right scansors covering underside of toes to base, 8/8 under first toe, 10/10 under second toe, 13/12 under third toe, 14/13 under fourth toe, 10/10 under fifth toe; left/right scansors covering underside of fingers to base, 8/8 under first finger, 10/11 under second finger, 10/11 under third finger, 14/14 under fourth finger, 6/5 under fifth finger; fourth toe lamellae: one entire, two divided, ten entire and 1–2 enlarged but not touching the toe border laterally; percentage of digit and toe covered by entire lamellae (right side only) respectively 80–90 % and 75–80 %; precloacal and femoral pore rows absent, as are enlarged adjacent scales.

Tail entire, subcylindrical throughout length, gradually tapering to a blunt tip; lateral margins without spines or skin flanges; scales on tail annulate, squarish or rectangular, larger ventrally than dorsally, subcaudal scales about 1.5 times belly scales; base of tail distinctly swollen by hemipenes; one large and about two smaller blunt cloacal spurs on each side.

Color in life (based on laboratory observation and numerous color photographs of holotype taken at different times). Coloration is very variable depending on the physiological and psychological situation of the animal or its following response to light intensity. Dorsal ground color is variable from dark, purple brownish to pinkish, with distinct more or less circular vivid black spots in seven medio-dorsal paired rows from neck to tail base (fig. 1). The first two pairs of middorsal black spots are more widely separated between them than the five following pairs. The first pair, anteriorly, medium in relative size, is located just in front of anterior members. The second anterior pair is the largest, nearly circular, and located just behind anterior members. The third pair, like the four following pairs, is more centred against the mid-back, thus narrower. Pairs 4 and 5 are identical to pair 3. Pair 6 is located at the level of posterior limbs. The last (7th) pair is located just behind posterior members and has also a similar size than former pair, but is followed anteriorly by oblique darker lines giving the whole an open V-shape. That black pair is followed by a darker brownish square patch. Those seven pairs of dark black spots have also sometimes irregular lateral black smaller spots. If interpreted as the W-chevron considered by Ineich (1988) for the Lepidodactylus lugubris unisexual-bisexual species complex’s dorsal pattern, black spots pairs 1 and 2 are on lateral tops of the W, whereas pairs 3 to 7 are located on both basal points of the W. The oblique dark lines following black spots of pair 7 correspond to the W lateral lines running from bottom to top.

Anterior and posterior of pair 7 is a lighter shape representing a shape like an anchor (fig. 5). That shape was almost yellowish or cream white in live holotype, depending on the gecko’s state. The anchor began posterior to black spots pair 6, at the level of the first light annuli on tail base and the round hook part of the anchor is located at the second light tail dorsal ring. That shape is clearly visible and may constitute a good diagnostic character for the species, in combination with other characters. Dorsum with 5 lighter more or less square-shaped purplish zones medially between pairs 2 and 7 of black spots. Flanks are always darker than dorsum, covered with darker marks and some whiter spots like ocelli. Each ocellus covers about 20 granules. In total, on the right side five to six ocelli are visible. Each dorsal black spots pair from 2 to 5 is laterally followed on flanks by a brownish rectangular dark brown mark. Venter is cream white with small light brown spots more developed on sides. Supra- and infralabials are brightly lemon-yellow and without any darker mark. Eye is not clearly encircled by a lighter coloration, but the whole peri-ocular area including labials is lemon-yellowish, as is the tympanum area, but that yellowish coloration is mostly developed in the posterior head part (fig. 4).

Eye is surrounded by 5 dark reddish brown radiating bands (fig. 4). Eyes are brownish orange with reticulations. Upper head ground colour is sometimes dark brown, purplish like the back (when animal is not too darkened), or even more or less yellowish. A first median V shaped dark mark, pointing posterior, can be seen medially on the snout in front of eyes. Tail is annulated with cream and dark rings. Note however that annuli can be more or less evident according to the gecko owing to physiological and psychological state. There are 11 dark dorsal tail annulations and 12 lighter cream annulations posterior to tail base. Light rings are about 2/ 3 to 3/4 as wide as dark ones. Ventrally tail is lightly orange coloured. Throat is whitish to yellowish. Mouth interior colour was not checked.

Colour of holotype in preservative. Colour of the preserved holotype is very reduced and retains only a small fraction of coloration in life (fig. 2). On upper head the snout anterior opened V mark is still discernable. The interocular dark band also is reduced but still visible. Between eyes there is just a small black spot conserved and another one in the neck area. The dark band in front of the eyes is visible. Middorsal black spot pairs are variably visible. Pair 1 is weak, pair 2 very evident and still heavy black, pair 3 is visible with lateral dark expansions, the same for pair 4 with even larger lateral expansions, another large lateral dark expansion is visible between dorsal black pairs 4 and 5 and a last one dorso-ventrally at pair 5 level. Between black spot pairs 3 to 5 one supplementary black mark can be seen. Pair 6 has reduced lateral dark expansions and pair 7 has clearly line prolongations forming together an opened V shaped black mark pointing posteriorly at tail base. Dorsal coloration is whitish with some brownish patches. Tail is dorsally brownish and upper thighs white. Ventrally, tail is darker than venter but still not dark except its end (fig. 3). Anchor-shape mark is still discernable but no more as evident. Venter is white with fine speckles mostly on sides and more evident in the area between anterior and posterior members (fig. 3); each speckle covering 1 to 2 ventral scales. Throat is weakly speckled. MNHN 2006.0544 (female) also present typical disseminated minute speckles on venter and throat.

Distribution. The species is known presently only by two specimens from its type locality, primary forest at about 630 m elevation, east of the Village of Penaoru, on the west dry coast of Espiritu Santo, Vanuatu.

Natural history. The egg giving birth to holotype was taken in a myrmecophilous plant ( Hydnophytum sp., Rubiaceae ) located at about 22 meters high on a gymnosperm tree trunk ( Decussocarpus sp.) in primary dry forest at 630 m elevation. The epiphytic plant had an ellipsoidal shape and a size of about 30 × 25 cm and had numerous holes inside. It was collected by Bruno Corbara and his team using a professional climber. The egg was later hatched in captivity in Paris and grown until adult size by the gecko breeder Francis Girard. A gravid female ready to lay its eggs was also collected on the same epiphytic plant. The eggs of Gehyra vorax Girard (non adhesive) were found in the same plant, just beneath L. buleli adhesive eggs (fig. 6). Interactions between ants living in myrmecophilous plants and L. buleli are unknown. In any case, large areas of such plants are not occupied by ants, thus leaving enough space for geckos. Co-existence of geckos inside myrmecophilous plants on Santo will be reviewed in another paper (Corbara & Ineich in prep.). Lepidodactylus buleli is clearly an arboreal species, according to its collection site, above 20 m high on a tree, but also to its resting position with curled tail and its habits, to easily jump from hand or from support to the soil when handled. The holotype often vocalized when kept captive and disturbed.

Reproduction. The female was gravid on 14 November 2006, and its both eggs were ready to be laid according to their size and shell state. We measured 16 eggs referable to L. buleli collected inside an epiphytic myrmecophilous plant from Penaoru. Comparison with 15 eggs of L. vanuatuensis shows them to be significant larger ( Table 1 ).

Comparison. Lepidodactylus buleli belongs to Group II as defined by Brown and Parker (1977). It can be easily distinguished from the sympatric L. vanuatuensis by its narrower toes and fingers, lamellae extending more to the bases of fingers and toes, a more elongate and ovoid tympanum (more circular by L. vanuatuensis ), less elongate and more massive head, presence of two enlarged median postmentals in contact with mental plate, grey brownish speckled venter, smaller head granules, and particular colouration (see above). When regarding number of scale rows at midbody, L. buleli , with the highest number of such scales (147), can be easily distinguished from most Group II species: L. gardineri Boulenger, 1897 (103–118), L. guppyi Boulenger, 1884 (110–138), L. intermedius (121), L. lombocensis (110–112), L. novaeguineae Brown and Parker, 1977 (108–125), L. shebae (91), and L. tepukapili (105–118) (see Table 2 ).

By its scalation, L. buleli is most similar to L. pulcher Boulenger, 1885 , from Wild Island off the northwest coast of Manua Island (type locality), and Plot and Lengendrowa Islands on the southeast coast, Admiralty Islands, Papua New Guinea. Records from the Huon Peninsula need confirmation ( Bauer & Henle 1994: 162). With that species it shares an elevated number of midbody scale rows and IOS ( Table 2 ). It can however be distinguished from L. pulcher by its lower count of fourth-toe lamellae, its slight toe webbing, the lack of typical heavy spotted head pattern ( Boulenger 1885) and presence of dark medio-dorsal black spots even present in preserved specimens (see Plate XIII, fig. 5 of Boulenger 1887a illustrating L. pulcher ), shorter and narrower head and its colouration; also by the presence of several cloacal spurs vs. only one in L. pulcher (see Ota et al. 1995). It also differs from L. guppyi by several scalation characters (tabl. 2) and its more flattened and less elongated body shape, colour pattern and interorbital dark band (see pl. 19–20 in McCoy 2006).

Figures

FIGURE 1. Lepidodactylus buleli sp. nov., general view of the alive holotype (MNHN 2008.0004).

FIGURE 2. Lepidodactylus buleli sp. nov., dorsal view of the preserved holotype (MNHN 2008.0004).

FIGURE 3. Lepidodactylus buleli sp. nov., ventral view of the preserved holotype (MNHN 2008.0004).

FIGURE 4. Lepidodactylus buleli sp. nov., head of the alive holotype (MNHN 2008.0004) showing the star-shaped dark markings around eye.

FIGURE 5. Lepidodactylus buleli sp. nov., holotype (MNHN 2008.0004). The anchor-shaped typical marking at the base of tail is cream yellowish coloured. Cloacal spurs are clearly evident.

Tables

TABLE 1. Egg size comparison (mm) between L. vanuatuensis (Tasmate Village, west coast of Espiritu Santo) and L. buleli (Penaoru Forest, west coast of Espiritu Santo).

	L. vanuatuensis	L. buleli
Number of eggs	15	16
Mean width (range)	5.95 ± 0.17 5.69–6.20	7.46 ± 0.30 6.88–7.95
Mean height (range)	8.46 ± 0.41 7.61–9.29	10.36 ± 0.64 9.04–11.47

TABLE 2. Summary of selected scalation and morphometric data of adult Group II Lepidodactylus. Abbreviations are defined in Material and Methods. Ranges of the data are presented when possible; all values are for both sexes mixed, except males only for cloacal spurs. Data after Brown & Parker (1977), Ota & Hikida (1988), Ota et al. (1995, 1998), Zug et al. (2003), Buden (2007).

Lepidodactylus	MSR	TIVS	IOS	UL	LL	WEB	CS
L. buleli	147	13–14	39–40	10 to 11	10 to 11	less 20 %	+
L. gardineri	103 to 118	11 to 17	29 to 39 / 31 to 35	11 / 8 to 10	?	less 20 %	+
L.guppyi	117 to 138 / 110 to 133	11 to 13 / 9 to 13	29 to 39 / 33 to 37	11–13 / 8–11	10 to 13	25 to 33 %	+
L. intermedius	121	8	?	10 to 11	?	20%	+
L. lombocensis	110 to 112	8 to 9	?	9 to 10	?	less 20 %	+
L. novaeguineae	124 to 132 / 108 to 125	11 to 16 / 10 to 16	29 to 39 / 32 to 39	9 to 12 / 8 to 10	11	25 to 33 %	+
L.oligoporus	130 to 134	15 to 19	34 to 35	10 to 12	10 to 12	25 to 33 %	no
L. paurolepis	98 to 110	12 to 14	29 to 39 / 31 to 32	11 to 12	9 to 11	less 20 %	+
L. pulcher	143 to 167	17 to 20	39 to 46	10 to 12	12	25 to 33 %	+
L. shebae	91	11	27	10	9	20 to 33 %	?
L. tepukapili	100 to 118	12 to 15	29 to 35	7 to 9	8 to 9	less 20 %	+
L. vanuatuensis	99 to 114 / 91 to 118	10 to 12	35 to 38 / 31 to 35	10 to 12 / 9 to 10	10 to 11	less 20 %	+