(Squamata: Gekkonidae) from Panay Island ... - Semantic Scholar

Copeia, 2007(2), pp. 413–425

A New Species of Luperosaurus (Squamata: Gekkonidae) from Panay Island, Philippines, with Comments on the Taxonomic Status of Luperosaurus cumingii (Gray, 1845) MAREN GAULKE, HERBERT ROESLER,

AND

RAFE M. BROWN

We describe a new species of Luperosaurus on the basis of three recently collected specimens from Panay Island, Philippines. The new species is phenotypically similar to L. cumingii, but differs from it by the absence of enlarged ventrolateral tail tubercles and spinose scales on the edges of the anterior cutaneous expansions of the limbs, by having fewer middorsal scales, fewer ornamented tubercles on the head, neck, and body, differences in ventral color pattern, and a larger maximum body size. Examination of all available specimens of L. cumingii demonstrates that this species exhibits a high degree of morphological variation correlated with patterns of lineage isolation on several Pleistocene aggregate island complexes (biogeographical subprovinces) of the Philippines. Although it is suspected that L. cumingii comprises a complex of multiple species, current sampling limitations prevent taxonomic subdivision beyond the recognition of the new species described here. The name L. cumingii probably should only be applied to populations from Luzon Island. Luperosaurus cf. cumingii from Camiguin Sur and Lubang Island may represent additional unnamed species.

HE enigmatic Southeast Asian gekkonine genus Luperosaurus is characterized by a unique combination of morphological features, including moderately to strongly developed cutaneous folds on the posterior and, in some species, anterior border of the limbs, broadly dilated digits, a lack of enlarged postmental scales, small scales encircling the tail (differentiated subcaudals absent), and well-developed interdigital webbing (Brown and Alcala, 1978; Russell, 1979; Brown et al., 2000a). Until 1979, Luperosaurus was considered a Philippine endemic, but subsequent species discoveries on landmasses of the Sunda Shelf (Hall, 1996; Voris, 2000) increased species diversity. Since Gray’s description of L. cumingii (Gray, 1845), nine additional species have been described, seven of which are still recognized: L. joloensis, L. macgregori, and L. palawanensis from the Philippines (Taylor, 1918, 1922; Brown and Alcala, 1978; Brown and Diesmos, 2000), L. browni from the Malay Peninsula and North Borneo, L. brooksii (formerly Gekko brooksii) from Sumatra (Russell, 1979), L. yasumai from Borneo (Ota et al., 1996), and L. iskandari from Sulawesi (Brown et al., 2000a). The remaining two species include Taylor’s (1962) L. amissus, which was placed in the synonymy of Gekko hokouensis (Brown and Alcala, 1978; Ota et al., 1989) and L. serraticaudus of Ota et al. (1989), which was placed in the synonymy of L. browni (Brown et al., 2000a). Luperosaurus is one of the least known gekkonid genera in Asia (Manthey and Grossmann, 1997; Brown and Diesmos, 2000; Henkel and

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Schmidt, 2003). Except for L. cumingii, each of the eight species is represented by only one to three specimens, and almost nothing is known of their intraspecific variability and natural history (Brown and Diesmos, 2000). It is presumed that all live in the high forest canopy and that this microhabitat preference contributes to the scarcity of specimens in collections (Ota et al., 1996; Brown and Diesmos, 2000; Brown et al., 2000a). Ota et al. (1996) noted the morphological variation within Luperosaurus and distinguished stout-bodied species from more slender-bodied species. The phylogenetic analysis of Brown et al. (2000a) supported this morphological distinction in body shape and placed a newly discovered species, L. iskandari, in the ‘‘slender-bodied’’ clade (together with L. browni and L. brooksi). The remaining species belonging to the ‘‘stoutbodied’’ clade included four Philippine species (L. cumingii, L. joloensis, L. palawanensis, and L. mcgregori) and L. yasumai from Borneo. The relationship of the genus to other gekkonine genera has remained uncertain (Russell, 1979; Ota et al., 1996; Brown et al., 2000a) although Russell and Bauer (2002) discuss the presumed monophyly of Underwood’s Gekko Group (Gekko, Gehyra, Hemiphyllodactylus, Lepidodactylus, Luperosaurus, Perpchirus, Pseudogekko, and Ptychozoon). Recently, a species of Luperosaurus was discovered in the northwest corner of Panay Island in the central Philippines. The first specimens collected were juveniles and were provisionally identified as L. cumingii (Gaulke, 2003; Gaulke et al., 2003). However, when adults became avail-

2007 by the American Society of Ichthyologists and Herpetologists

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able, it became clear that the Panay population possessed distinctive characteristics that set them apart from other Philippine populations. In this paper, we describe the new Luperosaurus species from Panay Island, discuss the taxonomic status of L. cumingii, and comment on variation that we expect will eventually lead to the recognition of additional species. MATERIALS AND METHODS Measurements (to the nearest 0.1 mm) of preserved museum specimens (institutional acronyms follow Leviton et al., 1985) were taken with dial calipers. Standard scale counts (Brown and Alcala, 1978; Brown et al., 2000a) are presented as right/ left for bilaterally symmetrical meristic characters. All authors examined specimens, but in an effort to reduce inter-observer bias (Lee, 1990; Hayek et al., 2001), only data scored by MG are presented. Characters (following Brown and Alcala, 1978; Brown et al., 2000a) and their abbreviations include: eye–tympanum distance from the posterior border of eye to the anterior edge of the tympanun (ETD), eye diameter (ED), snout length from the tip of the snout to the anterior border of the eye (SNL), maximal head depth (HD; taken at the midpoint behind the orbits), head length from the tip of the snout to the posterior border of auricular opening (HL), maximal width of the head (HW; at the widest point of the temporal region), diameter of the auricular opening (AOD), snout–vent length (SVL), and tail length (TL). Sex and maturity were determined by gonadal inspection when possible and/or by the scoring of prominent secondary sexual characteristics (Brown and Alcala, 1978; Brown et al., 2000a) when dissection was not possible. For comparison of described taxa with the new species, we attempted to verify the taxonomic status of the various populations referred to L. cumingii, adopting for this purpose the General Lineage Species Concept (GLC) of de Queiroz (1998, 1999) as the natural extension of the Evolutionary Species Concept (Wiley, 1978). Application of lineage-based species concepts to island endemics is straightforward because of the known history of isolation of island populations (Brown et al., 2000b; Brown and Diesmos, 2002; Brown and Guttman, 2002). We comment below (see also Discussion) on populations that seemed problematic during the process of recognizing diagnosable species that are distinct evolutionary lineages. RESULTS One clutch containing two eggs was collected on the northwest Peninsula of Panay Island (Fig. 1) on 30 May 2002, and the eggs hatched

Fig. 1. Map of the Philippines (darkly shaded islands) with outlines of Pleistocene Aggregate Island Complexes (PAICs) indicated (light gray) by tracing submarine 120 m bathymetric contours. Labeled islands are discussed in text; star indicates type locality of Luperosaurus corfieldi. Known localities for Luperosaurus cumingii are presented as circles; squares denote taxonomically uncertain specimens provisionally assigned to ‘‘Luperosaurus cf. cumingii.’’

on 9 July 2002. The first hatchling (male, paratype PNM 7920) died after 3 mo, and the second specimen (male, holotype PNM 7919) was preserved after 7 mo in captivity. On 5 May 2003, a second egg clutch of the new species was discovered at the same area, and the eggs hatched on 9 June 2003. One specimen died on 26 January 2004 and was preserved (female, paratype PNM 8489); the other one was released as an adult in the type locality following the collection of morphometric data. An additional adult specimen was captured, measured, and released in the type locality. In total, we examined one juvenile and four adult specimens (three of which are now deposited in PNM) of the new species. The foremost taxonomic issue with respect to the recognition of the new species described below was to distinguish it from L. cumingii. Because specimens from four of the Pleistocene Aggregate Islands of the Philippines (PAICs; Fig. 1) are currently referred to L. cumingii, we

GAULKE ET AL.—NEW PHILIPPINE LUPEROSAURUS suspect that these will eventually be recognized as four separate species, and so identifying the true population to which the name ‘‘L. cumingii’’ applies was paramount. The type locality for the species is uncertain: the type specimens (BMNH 1946.8.22.41–42) are only accompanied by the locality data ‘‘The Philippines,’’ and thus, the origin of the specimens cannot be used to identify which of the four putative lineages should be regarded as true L. cumingii. Brown and Alcala (1978) based their re-description of L. cumingii on five specimens (BM 1946.8.22.41–42, CAS-SU 24394–95, SM 4181) and, on the basis of two hatchling specimens (CAS-SU 24394–95) collected on Negros, recommended restricting the type locality for the species to this island. We question the advisability of following this recommendation for two reasons. First, the hatchling specimens do not exhibit the prominent diagnostic adult characters that we now know distinguish the species (ornamental scalation of the body, cutaneous expansions, and ventrolateral tail edge; adult color pattern; adult maximum body size) and thus cannot be used to distinguish between the four possible lineages within L. cumingii or convincingly match the Negros population to the type material. Second, the types of L. cumingii are still available (Fig. 2) and can be used to match morphological characteristics observed in the type series to new adult specimens from a known locality. Thus, among all available specimens of ‘‘L. cumingii,’’ we note that only the L. cumingii types (BM 1946.8.22.41–42) and recently collected adult specimens from Luzon (e.g., UF 77829, TNHC 61910, SMF 9044 [5‘‘SM 4181’’ as reported by Brown and Alcala, 1978]) uniquely possess the following combination of character states: (1) moderate maximum body size in adults, (2) strongly spinose tubercles along the ventrolateral edge of the tail, (3) well developed enlarged spinose scales along the edges of the cutaneous expansions bordering the limbs, (4) heterogeneous ornamental tubercles present on the neck and body, and (5) a strongly patterned ventral coloration of bright yellow with distinct, thin, transverse black bands (Fig. 2) that remains prominent in preservative. In summary, based on our comparison of the actual L. cumingii type specimens to newly collected material from Luzon, we have no doubts that Cuming’s specimens originated on Luzon and not Negros. Remaining specimens include four from Negros (CAS 182570, CAS-SU 24394–95, and SUR 2211), specimens of undetermined affinities from Lubang Island (PNM 7242), and Camiguin Sur Island (NMW 17985:1), and a single specimen lacking locality data other

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than ‘‘The Philippines’’ (ZMB 5578). These we refer to Luperosaurus cf. cumingii with the caveat that we expect Lubang and Camiguin Sur populations to eventually be recognized as two additional species and the Negros sample to eventually be referable to the new species described below (see Discussion). Luperosaurus corfieldi, new species Figures 3A, 4B, 5A, 6A–B Holotype.—PNM 7919 (Fig. 3A), adult male, Philippines, Panay Island, Aklan Province, Municipality of Buruanga, Barangay Tagosip, 11u49.39N, 121u58.29E, 450 m above sea level, collected as egg 30 May 2002 (preserved as adult 30 Dec. 2003), M. Gaulke. Paratypes.—PNM 7920, juvenile male, same locality and collecting date as holotype; PNM 8489, adult female (Fig. 4B), same locality as holotype, 26 Jan. 2004, S. Eickermann. Diagnosis.—A large species of Luperosaurus with a generally robust and stout body shape (Fig. 4B); SVL of adults 70.0–95.0 mm. The new species is characterized by wide cutaneous expansions on anterior and posterior edges of forelimbs (Fig. 4B) and posterior edges of hindlimbs, narrower folds on anterior edges of hindlimbs; interdigital webbing well developed; tail non-depressed, lacking spinose ventrolateral tail tubercles or denticulate tail lobes (Fig. 6A); body scales small, granular, interspaced with few, slightly enlarged dorsolateral tubercles (Figs. 3A, 5A); midbody scale rows 165; subdigital scansors 13–15 beneath first toe and 16–19 beneath fourth toe; 16 preanal pores in males, 11 in females; venter immaculate to dark yellow with faded, diffuse, and broad brown bands that are interrupted by a row of brownish spots along midline (Fig. 6A, 6C). Luperosaurus corfieldi differs from all other species of Luperosaurus by the absence of lateral tail tubercles or spines and/or denticulate tail lobes (Fig. 6A, 6C). It differs from L. brooksii, L. browni, and L. iskandari by the robust body shape (vs. habitus slender) and further from L. brooksii and L. browni by a subcylindrical (vs. depressed) tail. From L. yasumai, L. macgregori, L. palawanensis, and L. joloensis, L. corfieldi differs by its much larger size. Luperosaurus corfieldi has fewer preanal (or preanofemoral) pores (16) than L. brooksii (40), L. browni (28–32), L. palawanensis (28–32), or L. joloensis (30 or 31). The new species has a greater number of scansors beneath toes I (13– 15) and IV (16–19) than L. yasumai (8 scansors

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Fig. 2. Dorsal (A) and ventral (B) aspect of one of the syntypes (BMNH 1946.8.22.41; SVL 85.0 mm) of Luperosaurus cumingii and ventrolateral aspect of a live adult female of L. cumingii (TNHC 61910; SVL 75.2 mm) from Mt. Malinao, Albay Prov., Bicol Peninsula, Luzon Isl. Note similar heterogeneous scalation between the type (A) and the live specimen (C), the presence of spinose scales on the margins of cutaneous expansions of the limbs and ventrolateral tail edge, and the identical ventral color pattern (B: faded in the type; bright yellow with distinct black transverse bands in life).

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Fig. 3. (A) Luperosaurus corfieldi holotype as a subadult (PNM 7919; SVL 62.0 mm; preserved six months later as adult); (B) Adult female L. cumingii (TNHC 61910; SVL 75.2 mm) from Mt. Malinao, Albay Prov., Bicol Peninsula, Luzon Isl. Note prominent spinose ventrolateral tail tubercles and heterogeneous nuchal and body tubercles in L. cumingii (B).

beneath toe I, 7 beneath toe IV), L. macgregori (10 or 11 scansors beneath toe I, 12–14 beneath toe IV), L. palawanensis (9–11 scansors beneath toe I, 12–13 beneath toe IV), and L. joloensis (8 or 9 scansors beneath toe I, 9–13 beneath toe IV). Luperosaurus corfieldi differs from L. cumingii by the absence (vs. presence) of enlarged lateral tail tubercles, the absence (vs. presence) of spinose

scales on the edge of the anterior cutaneous expansions of the limbs, a lower number of middorsal scales (165 vs. 182–194 in Luzon populations of L. cumingii), a generally smooth, non-tuberculate appearance (vs. dorsal and lateral body scalation heterogeneous with interspersed enlarged convex to spinose tubercles; Figs. 3–5) and a larger maximum body size (Table 1).

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Fig. 4. Adult specimens of (A) Luperosaurus cumingii (TNHC 61910; SVL 75.2 mm) and (B) L. corfieldi (PNM 8489; SVL 86.0 mm) from Luzon and Panay islands, respectively. Note heterogeneous, tuberculate appearance of L. cumingii (A; especially nuchal region, edge of anterior forearm cutaneous expansion, and ventrolateral tail margin) and smooth, non-tuberculate appearance of L. corfieldi (B; ornamental spinose tubercles absent).

Fig. 5. Details of head scalation in (A) L. corfieldi (PNM 8489; HL 20.4 mm) and (B) L. cumingii (TNHC 61910; HL 19.9 mm). Note smooth, non-tuberculate appearance of posterolateral region of head and nuchal region of L. corfieldi and the presence of enlarged, spinose nuchal tubercles in L. cumingii.

Fig. 6. Ventral aspect of (A) adult female paratype of L. corfieldi (PNM 8489). Note complete absence of ventral transverse banding and smooth, nontuberculate appearance of ventrolateral body margins. Characteristics of preanofemoral regions and basal portions of tail in (B) L. corfieldi male holotype (PNM 7919) and (C) L. cumingii (TNHC 61910). Note differences in ventral tail color pattern and complete absence of ventrolateral tail tubercles in L. corfieldi (B; dark arrows) vs. tubercles present in L. cumingii (C; white arrows).

Description of holotype.—(Fig. 3A) Adult male with robust habitus; SVL 70.0 mm (71.5 before preservation); TL 61.0, HL 17.4, HW 14.0, HD 9.0, ED 4.0, AOD 1.0, SNL 9.0, ETD 6.8, head and body almost same width; head only scarcely distinct from neck due to the presence of well developed endolymphatic sacs; trunk stout, venter flat, tail not depressed (subcylindrical), not basally thickened and without basal constriction; SVL/TL 1.2, SVL/HL 4.0, HL/HW 1.2, HL/HD 1.6, SNL/ETD 1.2; rostral twice as broad as high, three times as wide as mental, without median cleft; supralabials 16/16, infralabials 14/ 13; narrow skin fold present along ventrolateral edge of both sides of lower jaw; scales surrounding nostril concave, protuberant, including rostral, first supralabial, supranasal, and postnasal; right supranasal narrow and oblong, left supranasal equivalent to adjacent postnasal; internasal single, wider than long, bordered posteriorly by four slightly enlarged scales; mental pentagonal,


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TABLE 1. MEASUREMENTS AND SCALE COUNTS OF Lupersosaurus corfieldi (PNM 7919, 7920, 8489), L. cumingii (SMF 9044 ZMB 5578, BMNH 1946.8.22.41), AND L. cf. cumingii (NMW 17985.1, PNM 7242). Specimen Island

Sex SVL TL HL HW HD SNL ETD SVL/TL SVL/HL HL/HW HL/HD SNL/ETD Supralabials Infralabials Nasals Internasals Postmentals Scales behind postmentals Midbody scales Scansors finger I Finger II Finger III Finger IV Finger V Scansors toe I Toe II Toe III Toe IV Toe V Preanal pores Postcloacal spurs Dorsal tail scale rows on whorl 3 Ventral tail scale rows on whorl 3

PNM 7919 Panay

PNM 7920 Panay

PNM 8489 Panay

SMF 9044 Luzon

ZMB 5578 ‘‘Philippines’’

BMNH 1946.8.22.41

NMW 17985.1 Camiguin Sur

PNM 7242 Lubang

Male 70.0 61.0 17.4 14.0 9.0 8.0 6.8 1.2 4.0 1.2 1.9 1.2 16/16 14/13 4/3 1 2 5

Male 44.0 30.0 12.3 9.6 6.1 6.1 4.6 1.47 3.58 1.28 2.02 1.33 — — — — 3 5

Female 86.0 66.8 20.4 18.2 10.0 9.2 7.2 1.3 4.2 1.1 2.0 1.3 14/15 12/13 3/3 1 2 4

Male 67.8 40.0* 17.4 13.6 7.7 8.0 7.3 — 3.9 1.3 2.3 1.1 16/— 15/14 3/3 2 2 5

Female 63.7 41.0 16.6 12.2 8.2 7.6 6.4 1.6 3.8 1.4 2.0 1.2 14/14 15/15 3/3 1 2 5

Female 85.0 44.0** 21.6 17.0 — — — — 3.9 1.3 — — 17/18 15/16 3/3 2 3 —

Female 61.0 38.0 16.0 10.6 7.6 7.2 5.8 1.6 3.8 1.5 2.1 1.2 14/15 15/16 3/3 2 2 4

Male 65.5 47.5 17.0 12.4 8.0 8.1 5.9 1.4 3.9 1.4 2.1 1.4 15/14 13/14 3/4 3 3 5

165 13/13 13/16 17/17 16/20 13/14 15/14 17/15 19/20 16/17 14/13 16 3/2 12

— 14/14 13/14 19/15 21/19 14/14 —/13 —/16 —/23 —/19 14/15 16 2/2 —

165 12/11 13/13 16/16 15/19 13/14 14/14 14/15 17/17 16/16 15/14 11*** 1/1 10

182 11/11 12/12 15/14 15/15 14/12 13/12 13/13 16/16 13/14 11/12 19 1/1 11

182 10/10 12/12 15/16 17/17 12/11 11/10 13/12 17/17 14/13 11/11 0 2/2 11

— 11/9 12/12 16/15 16/14 12/12 12/12 13/13 18/16 13/14 11/12 13*** 1/1 —

194 10/11 12/11 14/13 14/12 11/11 12/11 13/13 15/14 12/13 10/11 0 2/2 11

164 10/10 11/11 13/14 14/16 12/12 10/10 12/12 16/15 13/14 12/12 15 1/1 9

8

—

8

8

7

6

7

8

* Distal 13.9 mm regenerated. ** Distal 14.6 mm regenerated. *** Pores not forming a confluent line; preanal series interrupted by undifferentiated scales.

wider than long, smaller than adjacent first infralabials; postmentals hexagonal, wider than long; five small scales bordering postmentals; snout scales granular, 1.5–2.0 times larger than head scales; scales bordering supralabials flat, 2– 3 times as large as median snout scales; head and nuchal scales granular; lateral nuchal tubercles convex to conical, twice as large as adjacent median nuchal scales; auricular opening round, dorsal border partially covered by a small skin fold. Anterior chin scales flat, smooth, cycloid, nonimbricate, becoming granular posteriorly; gular scales as large or slightly larger than head

scales; ventral head, gular, and pectoral region scales flat, smooth, imbricate, 2–3 times larger than gulars; middorsals in 165 scale rows; dorsals granular, as large as median snout scales, occasionally separated by small interstitial granules; no median dorsal tubercles; a few scattered dorsolateral tubercles convex to conical, 1.5 times larger than dorsals; lateral body scales subimbricate, 1.5 times larger than dorsals; ventrals flat, smooth, anteriorly subimbricate, posteriorly imbricate; largest ventrals three times as large as dorsals. Limbs with wide cutaneous expansions; cutaneous folds posterior to hindlimbs larger than

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those posterior to forelimbs; folds anterior to forelimbs larger than those anterior to hindlimbs; cutaneous expansions with subimbricate scales on dorsal and ventral sides, and without enlarged spinose scales along the edges. Dilated digits of feet and hands two-thirds webbed; interdigital webbing notched medially; subdigital scansors wide, undivided; distal scansors straight, penultimate 2–3 scansors strongly arched, proximal scansors slightly bowed; subdigital scansor numbers per digit of manus: (digit presented in parentheses) 13/13 (I), 13/16 (II), 17/17 (III), 16/20 (IV), 13/14 (V); subdigital scansors of pes: 15/14 (I), 17/15 (II), 19/20 (III), 16/17 (IV), 14/13 (V); subdigital scansors followed by a few rows of enlarged, slightly widened scales. Digits II–V clawed; terminal, claw-bearing phalanges depressed, rising free at distal ends of dilated portions; Digit I (inner digit) with strongly enlarged cycloid scale on dorsal tip of distal phalanx. Preanal pores 16, arranged in a slightly bowed row on an integumentary bulge (Fig. 6B); postcloacal spurs 3/2, well developed, conical. Tail subcylindrical, without basal swelling or constriction, only very slightly ventrally flattened at base, more depressed towards tip; tail segmented in whorls (annuli) with 12 dorsal and 8 ventral scale rows at Whorl III; dorsal tail scales convex, arranged in regular transverse rows, twice as large as dorsal body scales; scales of posterior edge of each caudal annulus enlarged, but without tubercles or spinose scales, or skin flanges; subcaudals flat, smooth, subimbricate, twice as large as dorsal tail scales; dorsolateral longitudinal edge of tail rounded, lacking differentiated scales, enlarged tubercles, or spinose projections. Coloration in life.—(Figs. 3A, 4B) Dorsum brown, with three darker transverse chevrons between fore- and hindlimbs. Depending on the substrate on which the specimen rests, the dorsum is crossed by either wide dark brown transverse chevrons, giving the lizard a very dark appearance, or by thin black wavy lines with pale brown interspaces more prominent, giving the lizard a much lighter overall appearance. On a very light background, specimens become light greybrown, with darker bands hardly noticeable; on a dark background specimens become dark reddish-brown. Tail either dark brown with six pale brown rings, or pale brown with six dark brown rings. Dorsal surfaces of head and extremities mottled dark and light brown. Circumorbitals bright orange. Venter bright yellow with irregular, medially interrupted brown ventrolateral bands. Ventral surfaces of head and

limbs yellow, with irregular brown mottling. Iris light tan to silver, with a fine dark brown reticulate network (Fig. 5A). Color photographs in life are presented by Gaulke (2003:figs. 13a, 13b) and Gaulke et al. (2003:figs. 5, 6). Coloration in preservative.—Upper side of head dark purplish grey, snout with fine dark flecks and reticulations; a V-shaped marking between eyes; two dark-grey flecks posterior to eyes; posterior part of head with an irregular transverse band; lateral sides of lower jaw with faint reticulation. Dorsum and dorsal side of tail dark grey; three grayish brown irregular transverse bands across dorsum: one at midbody and another above the insertions of fore- and hindlimbs; dorsal tail coloration banding faintly banded with alternating light and dark brown; lateral sides of tail with narrow, interrupted, blackish brown stripes; upper extremities bluish grey, with dark grey reticulation, fine stripes, and flecks; throat pale yellowish white, flecked with brighter yellow; venter yellowish white with three sepia bands posteriorly; ventral surfaces of limbs pale brown; ventral side of tail grayish brown with 12 narrow, interrupted irregular bands. Variation.—Paratypes (PNM 7920, PNM 8489) agree well with the holotype (Table 1), but differ by the slight presence of additional enlarged lateral tubercles on the neck and body. PNM 7920 has three enlarged imbricate scales on the ventral side of the left thigh, arranged in a linear line. The head of this juvenile male is relatively larger than the heads of the adults (SVL/HL 5 3.58). The color pattern of PNM 7920 is darker than that of the holotype. The venter of this specimen has ten diffuse bands, and the ventral side of the tail is also prominently banded. Its venter as a juvenile was orangish brown in life. The ventral banding of the female was also much less distinct in life and faded completely following preservation. In life, the relatively bright coloration of the circumorbital ring was much less prominent in the female than in the males. In life, ventral coloration (yellow or orange) becomes more prominent in more active animals. In the female the preanal scales have a small pore in their middle, and they are not situated on an integumentary bulge. Two female specimens that were released at the type locality had SVLs of 95 mm. They are the largest individuals recorded to date. Distribution.—Luperosaurus corfieldi is only known from the type locality, a forested area in the center of the NW Panay Peninsula (Fig. 1). Based


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on geology and channel depth (Heaney, 1985; Voris, 2000) we expect that this species could be (or may have once been) distributed in forested areas throughout the Visayan PAIC islands of Panay, Negros, Cebu, Guimaras, and Masbate.

diversity research and nature conservation in the Philippines by taking patronage of this endemic Philippine species.

Reproduction, ecology, and behavior in captivity.— Both egg clutches were found in the same area beside a small river on the edge of a slightly disturbed, primary lowland forest at an elevation of 450 m above sea level. They were attached to the inner, sunken part of dead Pandanus leaves laying on the ground, and it is presumed that they were deposited on these leaves when they were still attached to the tree at an undetermined height in the canopy. The hard-shelled, round, white eggs were adhered together and had a diameter of 12–13 mm each. The first clutch (collected on 30 May 2002) hatched after six weeks, the second clutch (collected on 5 May 2003) after five weeks. The hatchlings were 34– 35 mm SVL, and 26–31 mm TaL. Young hatchlings had highly developed camouflage ability, ranging from almost patternless light grey to a dark reddish brown depending on perch substrate. In captivity specimens were placid, almost catatonic. Usually individuals moved very slowly when climbing, with the venter pressed close to the surface of the branch or other perch. Only on the ground did individuals move on erect limbs with the venter held away from the ground. The tail is prehensile and anchors the body while climbing. Captive specimens preferred crevices in bark when inactive during the day and became active during late afternoon. Activity was observed throughout the night, but specimens generally remained quite still, even during (subjectively characterized) ‘‘peak activity’’ hours. When captive specimens first observed insect prey, they approached very slowly and eventually caught food items with fast movement of the head. Both males and the female vocalized in captivity. Vocalizations resembled those produced by Hemidactylus frenatus and sounded like ‘‘tschek-tschek.’’ Two females kept in the same cage demonstrated few interactions, although they approached each other occasionally and licked each other’s heads. Aggressive behavior was not observed. Possibly sympatric (but not necessarily syntopic) gekkonid species include Gekko gecko, G. mindorensis, Cosymbotus platyurus, Cyrtodactylus philippinicus, C. anulatus, Gehyra mutilata, Hemidactylus frenatus, H. stejnegeri, Lepidodactylus planicaudus, Hemiphyllodactylus insularis (Ferner et al., 2001), and Pseudogekko cf. brevipes (Gaulke et al., 2003).

The description of the new species of Luperosaurus brings the total number of species to ten (see also Brown et al., 2007). However, we suspect that additional species will be described in the near future, with greater attention to sampling and evaluation of appropriate characters for distinguishing evolutionary lineages in these rare geckos. There are substantive taxonomic problems associated with the current recognition of Luperosaurus cumingii as a ‘‘widespread’’ endemic Philippine species. Specimens previously referred to L. cumingii have been collected on the islands of Luzon, Negros, Panay, Camiguin Sur, and Lubang (Fig. 1). Given the highly partitioned nature of biodiversity in the Philippines (Heaney, 1985; Heaney and Regalado, 1998; Brown et al., 2002), it is improbable that a single species (L. cumingii sensu lato) is widely distributed in isolated populations that are separated from one another on four of the seven Pleistocene Aggregate Island Complexes (Brown and Diesmos, 2002; Diesmos et al., 2002) of the Philippines. Almost all previous comprehensive phylogenetic studies of closely related Philippine species have found highly partitioned groups of endemic species, the distributional boundaries of which coincide reasonably well with former Pleistocene sea shores (McGuire and Alcala, 2000; Brown and Guttman, 2002; Evans et al., 2003). Given a known history of isolation (Heaney, 1985; Brown and Diesmos, 2002), a more sensible taxonomic arrangement might be to recognize each isolated population as a distinct species. Under this more liberal application of a lineage-based species concept, we recognize the population from S. Luzon and its satellite island Polillo as L. cumingii (see Results for justification), with the following populations additionally recognized as separate and distinct lineages: the Visayan PAIC (Panay + Negros islands) population, the Lubang PAIC (Lubang Isl.) population, and the Mindanao PAIC (associated with Camiguin Sur) population. Such an arrangement is logically consistent with known evolutionary history of these isolated populations and reflects the status of each as a clearly isolated, allopatric lineage that possesses predictable evolutionary fates (Brown and Diesmos, 2002; Brown and Guttman, 2002). However, we are not inclined to indiscriminately recognize three additional species in the L. cumingii

Etymology.—We are pleased to name the new species for Charles Corfield for supporting bio-

DISCUSSION

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complex at the present time for three reasons. First, we have too few adults from the Camiguin Sur and Lubang populations to distinguish interfrom intraspecific variation and reliably diagnose these lineages from conspecifics. Second, within one of the lineages (the Visayan PAIC [Fig. 1]: Panay, Negros, Cebu, Guimaras, and Masbate), several specimens are incomplete and lack diagnostic features. The available Negros Island specimens lack complete original tails and thus cannot be reliably assigned to species because the disposition of ventrolateral tail tubercles cannot be assessed. This is particularly unfortunate because we would expect, on the basis of channel depth between the islands and their known history of Pleistocene connectivity (Heaney, 1985; Hall, 1998; Voris, 2000; Brown and Diesmos, 2002) that Negros and Panay island populations would be conspecific. We expect that the Negros population will eventually be assigned to the new species described here, but, at present, we are unable to confidently or objectively arrive at this conclusion. Finally, we acknowledge the possibility that recent dispersal might have led to widespread species distributions that do not correspond well with the formation of Philippine PAICs. Given the currently available material and low probability of obtaining additional specimens in the near future, we were forced to compromise by restricting our concept of L. cumingii to populations of S. Luzon and satellite islands and describing the Panay Island sample as a new species. At present, we hold in abeyance taxonomic decisions regarding the Negros, Lubang, and Camiguin Sur populations until further specimens become available that would allow us to confidently assign these populations to a current taxon or diagnose them as additional species. When considering the entire range of morphological variation in specimens currently assigned to L. cumingii, L. cf. cumingii, and L. corfieldi, we note that some characters distinguish what we consider to be evolutionary lineages while other characters that have been emphasized by previous authors do not. For example, traditionally emphasized characters such as the number and arrangement of preanal pores, the arrangement of snout scales, and number of subdigital scansors do not significantly vary between putative evolutionary lineages of ‘‘L. cumingii.’’ In contrast, specimens from different islands exhibit variation in the number and form of dorsal, lateral, and nuchal tubercles, the development of interstitial granules, the width of cutaneous expansions bordering the limbs, the presence or absence of spinose scales along

these cutaneous expansion, and color pattern. The rather short type description and some consecutive descriptions of L. cumingii include no reference to dorsal or lateral tubercles (Gray, 1845; Boulenger, 1885; Taylor, 1922; Alcala, 1986); however, the illustration of one syntype (in Boulenger, 1885; reproduced in Taylor, 1922 and Alcala, 1986) shows prominent, spinose lateral tail tubercles. Ornamental body tubercles have been described elsewhere by Brown and Alcala (1978), Russell (1979), Ota et al. (1996), and Brown et al. (2000a) and vary from flat, to convex, to conical, to highly spinose and occasionally recurved. In the specimen from Camiguin Sur (in NMW 17985:1), convex dorsal body tubercles are positioned on small epidermal swellings. In Luzon L. cumingii (e.g., TNHC 61910, SMF 9044), tubercles of the nuchal region, body, ventrolateral tail, and edge of cutaneous expansions bordering the limbs are conical or extremely spinose. In the Lubang Island specimen (PNM 7242), no nuchal or dorsolateral body tubercles are present, but the specimen does possess a ventrolaterally spinose tail. Small interstitial granules surrounding the dorsal scales are well developed in BMNH 1946.8.22.41 (syntype, presumably from Luzon), SMF 9044 (Central Luzon), NMW 17985:1 (Camiguin Sur), and PNM 7242 (Lubang), although they are hardly discernible in ZMB 5578 (no locality data). Only one individual (PNM 7242 from Lubang) varies significantly in extent of cutaneous expansion bordering the limbs. In this specimen, cutaneous expansions bordering the limbs are much narrower than in the other known specimens. This observation (together with the absence of nuchal and dorsolateral body tubercles) would tend to support the recognition of the Lubang Isl. population as a distinct species, but we take no such action at this time due to the presence of only a single small male specimen available in collections. Due to the paucity of material and the unknown origin of some of the specimens, we are unable to determine at present whether some of the above-mentioned differences are of taxonomic importance. At present, we have no doubt regarding the mutually exclusive diagnoses of L. cumingii (in the strict sense: from Luzon and its satellite islands) and L. corfieldi from Panay. Remaining populations to consider include those from Lubang Island, the Visayan PAIC (Negros), and the Mindanao PAIC (Camiguin Sur; Heaney and Regalado, 1998; Kennedy et al., 2000; Brown and Diesmos, 2002). Given this wide distribution, spanning four Philippine

GAULKE ET AL.—NEW PHILIPPINE LUPEROSAURUS Pleistocene Aggregate Island Complexes, it seems likely that additional lineages will soon be recognized as endemic to some of these isolated island landmasses that are well known for their non-overlapping suites of endemic taxa (Brown and Diesmos, 2002; Brown and Guttman, 2002; Evans et al., 2003). The discovery of Luperosaurus corfieldi on NW Panay further emphasizes the high degree of faunal diversity and endemism of the Visayan PAIC. In recent years many new amphibian and reptile species have been discovered on Panay (Brown et al., 1997, 1999, 2001; Ferner et al., 1997, 2001; Gaulke and Curio, 2001; Gaulke, 2002; Gaulke et al., 2003, 2004), and our experience of the last decade suggests that many additional unrecognized species await discovery if naturalists can overcome logistical obstacles to conducting biodiversity research in this part of the Philippines. Great strides can be made towards sustainable development and conservation of Panay’s natural resources if policy makers, indigenous peoples’ organizations, and conservation biologists can work together to make use of the last decade’s accumulated biodiversity research and put an end to destructive exploitation of the remaining forests on Panay. MATERIAL EXAMINED Luperosaurus cumingii: Philippines, Luzon Isl., Camarines Sur Province, Municipality of Caramoan, Anuling Mountain: UF 77829; Albay Province, Municipality of Tiwi, Barangay Banhaw, Sitio Purok 7, Mt. Malinao, 550 m above sea leve: TNHC 61910; ‘‘Central Luzon, Philippines:’’ SMF 9044; ‘‘Philippines:’’ BMNH 1946.8.22.41 (type series). Luperosaurus sp. (probably cumingii): ‘‘Philippines’’ (specimen probably from Luzon): ZMB 5578; Philippines, Polillo Isl., vicinity of Polillo Town: CAS 62453–54. Luperosaurus cf. cumingii: Philippines, Camiguin Sur Island: NMW 17985:1. Luperosaurus cf. cumingii: Philippines, Mindoro Occidental Province, Lubang Island: PNM 7242. Luperosaurus cf. cumingii: Philippines, Negros Isl., Negros Oriental Prov., Municipality of Valencia, Lake Balinsasayao: SUR 2211; Saksak Cr., Mt. Cuernos de Negros, ‘‘Camp Lookout:’’ CAS-SU 24394–95; Negros Isl., Lake Balinsasayao: CAS 182570. L. joloensis: Philippines, Mindanao Isl., ‘‘Cotobato Coast:’’ MCZ 26118; Jolo Isl., Siet Lake: CAS 60675 (paratype). L. macgregori: Philippines, Calayan Isl.: CAS-SU 6263 (paratype); Babuyan Island group, Barit Isl. (near Fuga Isl.): USNM 508306–508308. L. palawanensis: Philippines, Palawan Isl., Thumb Peak, about 7 km NW of Iwahig: CAS 136740 (paratype); Palawan Isl., Malatgaw River, SE of

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Thumb Peak, about 3.5 km WNW of Iwahig: CAS 134207 (holotype). L. yasumai: Indonesia, Kalimantan (Borneo Isl.), Bukit Soeharto Experimental Forest, 45 km SSW of Samarinda: KUZ 30408 (holotype). L. brooksii: Indonesia, Sumatra Isl., Benkuelen, Lebong Tandai: BMNH 1920.1.16.2 (holotype). L. browni: Malaysia, peninsular Malaysia, Selangor, Ulu Gombak forest reserve, 35 km N of Kuala Lumpur: FMNH 185106 (holotype); Malaysia, Sarawak (Borneo Isl.), Lambir National Park: KUZ 12835 (L. serraticaudus holotype). Gekko hokouensis: ‘‘Philippines:’’ FMNH 17812 (L. amissus holotype). P. rhacophorus: Malaysia, Sabah, Mt. Kinabalu: KUZ 35118. ACKNOWLEDGMENTS The work of the Philippine Endemic Species Conservation Project of the Frankfurt Zoological Society is formalized under the aegis of a Memorandum of Agreement with the Department of Environment and Natural Resources (Quezon City, Philippines) wherein we are especially grateful to C. Custodio for years of logistical advice and support. The help of the Protected Areas and Wildlife Bureau (R. Bayabos and W. Pollisco) and J. Amador (DENR Region VI) is gratefully acknowledged. We are grateful to A. Alcala and C. Dolino of Silliman University, Dumaguete; G. Koehler and M. Laudahn of the Forschungsinstitut Senckenberg, Frankfurt; C. McCarthy of the British Museum (Natural History), London; R. Sison and A. Diesmos of the Philippine National Museum, Manila; F. Tiedemann of the Naturhistorisches Museum Wien; R. Guenther of the Museum fuer Naturkunde der Humboldt–Universitaet zu Berlin; and D. Cannatella and T. LaDuc of the Texas Memorial Museum for the loan of material or the permission to examine specimens under their care. Support for RMB’s fieldwork was provided by the Society of Systematic Biologists, the Society for the Study of Amphibians and Reptiles, the American Society of Ichthyologists and Herpetologists, the University of Texas, the University of Kansas, and the National Science Foundation. S. Eickermann helped during one field survey on Panay and collected Luperosaurus eggs. We thank C. Pioquinto, T. Ziegler, E. Lehr, C. Siler, L. Trueb, and J. Weghorst for comments on earlier drafts of this paper. This is publication No. 52 of the Philippine Endemic Species Conservation Project. LITERATURE CITED ALCALA, A. C. 1986. Guide to Philippine Flora and Fauna. Vol. X. Amphibians and Reptiles. Natural Resources Management Center, Ministry of Natu-

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(MG) GEOBIO-CENTERLMU, RICHARD-WAGNERSTRASSE 10, 80333 MUNICH, GERMANY ; (HR) STAATLICHE NATURHISTORISCHE SAMMLUNGEN DRESDEN, MUSEUM FUER TIERKUNDE, FORSCHUNGSSTELLE A. B. MEYER BAU, KOENIGSBRUEDER LANDSTRASSE 159, 01109 DRESDEN, GERMANY ; AND (RB) NATURAL HISTORY MUSEUM, BIODIVERSITY RESEARCH CENTER, AND DEPARTMENT OF ECOLOGY AND EVOLUTIONARY BIOLOGY, UNIVERSITY OF KANSAS, DYCHE HALL, 1345 JAYHAWK BOULEVARD, LAWRENCE, KANSAS 66045. E-mail: (MG) [email protected]; (HR) herbertroesler@aol. com; and (RMB) [email protected]. Send reprint requests to MG. Submitted: 22 June 2005. Accepted: 18 Dec. 2006. Section editor: T. W. Reeder.