Herpetologica, 64(3), 2008, 305–320 E 2008 by The Herpetologists’ League, Inc. A NEW GEKKO FROM THE BABUYAN ISLANDS, NORTHERN PHILIPPINES RAFE M. BROWN1,4, CARL H. OLIVEROS1,2, CAMERON D. SILER1, AND ARVIN C. DIESMOS3 1 Natural History Museum and Biodiversity Research Center, Department of Ecology and Evolutionary Biology, The University of Kansas, Lawrence, KS 66045-7561, USA 2 ISLA Biodiversity Conservation; 9 Bougainvillea St, Manuela Subdivision, Las Pinas City 1741 Philippines 3 National Museum of the Philippines, Rizal Park, Padre Burgos Ave. Ermita 1000, Manila, Philippines ABSTRACT: We describe a new species of gekkonid lizard on the basis of 21 recently acquired specimens from Babuyan Claro Island, Babuyan Islands group, northern Philippines. The new species differs from other Philippine Gekko by characteristics of external morphology, color pattern, and body size. The new species has been found low on trunks and buttresses of mature closed-canopy climax forest trees at low elevation near the island’s coast. It is known from only Babuyan Claro Island and is likely endemic to this single small, isolated landmass. The remaining gekkonid fauna of the Babuyans and Batanes island groups is understudied and in need of comprehensive review. Key words: Babuyan Claro; Babuyan Islands; Gekkonidae; New species; Philippines PHILIPPINE lizards of the family Gekkonidae Philippine members of the genus Gekko are represented by 10 genera and 37 species: include seven endemic species (G. athymus, Cyrtodactylus (4), Gekko (9), Gehyra (1), G. ernstkelleri, G. gigante, G. mindorensis, G. Hemidactylus (5, including H. platyurus, a palawanensis, G. porosus, and G. romblon; species formerly assigned to Cosymbotus), Brown and Alcala, 1978; Brown et al., 2007; Hemiphylodactylus (1), Lepidodactylus (6), Roesler et al., 2006; Taylor, 1922a), two Luperosaurus (6), Pseudogekko (4), and Pty- species shared with neighboring countries chozoon (1) (Brown and Alcala, 1978; Brown (G. gecko and G. monarchus; Manthey and et al., 2007; Gaulke et al., 2007; Taylor, Grossman, 1997; Wermuth, 1965), and one 1922a,b). Although the lack of definitive species (G. hokouensis) that is probably characters distinguishing the genera Lupero- included in the country’s gekkonid fauna in saurus and Gekko have clouded generic error (Brown and Alcala, 1978; Ota et al., boundaries (Brown and Alcala, 1978; Brown 1989). et al., 2000a; Brown et al., 2007; Russell, Several additional Philippine gekkonid taxa 1979), most authors have applied the name presently await description, including multiple Gekko to Philippine species that possess (1) unnamed species from the northern Philip- moderate to large body size and longer, more pines. During recent biological inventory slender limbs; (2) near or complete absence of work in the Babuyan archipelago, we discov- interdigital webbing; (3) a lack of cutaneous ered a distinctive species of Gekko on expansions, save for ventrolateral adipose Babuyan Claro Island, now represented in folds and very slight expansions bordering U.S. and Philippine collections by 21 speci- the posterior margins of the hind limbs; (4) mens. In this paper we describe the new convex to conical enlarged dorsal tubercles species and discuss morphological diversity, arranged in longitudinal rows; (5) enlarged, patterns of gekkonid endemism, and probable plate-like subcaudals; (6) differentiated, slen- processes of evolutionary diversification in the der, and elongate postmentals; and (7) minute Philippine island radiation of Gekko. and non-imbricate rows of dorsal scales between tubercle rows (minute dorsals differ- MATERIALS AND METHODS entiated from the enlarged, imbricate scales of We (RMB and CDS) collected data from the venter (Brown and Alcala, 1978; Brown et fluid-preserved specimens deposited in U.S. al., 2000a; Brown et al., 2007). and Philippine collections (see Acknowledg- ments). Sex was determined by inspection of 4 CORRESPONDENCE: e-mail, email@example.com gonads or by scoring of prominent secondary 305 306 HERPETOLOGICA [Vol. 64, No. 3 sexual characteristics (Brown, 1999; Brown et tions where the hypothesis of a distinct al., 1997, 2000a;) when dissection was not evolutionary lineage cannot be rejected. possible. Measurements (to the nearest 0.1 mm) were taken with Fowler digital SYSTEMATICS calipers following character definitions by Gekko crombota sp. nov. Ota and Crombie (1989), Brown et al. Figs. 2–5 (1997), Brown (1999), and Brown et al. (2007). Character abbreviations are: snout– Holotype.—PNM 9280 (Field no. RMB vent length (SVL); tail length (TL); head 5954; formerly KU 304833), an adult male length, (HL); head width (HW); head depth collected at night (2125 h on 13 March 2005) (HD); snout length (SNL); eye diameter, on the buttress of a large dipterocarp tree (ED); eye–narial distance (END); auricular by RMB at an area known locally as opening diameter (AO); inter-narial distance ‘‘Asked,’’ Barangay Babuyan Claro, Munici- (IND); inter-orbital distance (IOD); axilla– pality of Calayan, Cagayan Province, W groin distance (AGD); femur length (FL); coast of Babuyan Claro Island, Philippines tibia length, (TBL); Toe I length (TiL); Toe IV (19.5041u N, 121.9120u E; 20 m above sea length (TivL); tail width (TW); and tail depth level). (TD); number of supralabials (SUL) and Paratopotypes.—KU 304825 adult male; infralabials (IFL) to the center of the eye KU 304808, 304814, 304821, 304848, PNM and posteriorly to the point at which point 9281–82 (formerly KU 304829, 304832), two labials are no longer differentiated; enlarged male juveniles and four juveniles of undeter- circumorbitals dorsoanterior to orbit (CO); mined sex; KU 304807, subadult female; KU modified spiny circumorbitals (cilaria;) dorso- 304809, 304826, 3043830, 304836, 304845, posterior to orbit (SC); differentiated preanal PNM 9283–84 (formerly KU 304847 and pore-bearing scales (PS); femoral pore-bear- 304849), adult females, collected by RMB, ing scales (FS); differentiated subdigital scan- CO, Jason Fernandez and Boying Fernandez; sors on fingers 1–5 (FS I–V); subdigital same date, locality, and circumstances of scansors on toes 1–5 (TS I–V); midbody capture (19:45–22:45 h) as holotype. ventral transverse scale rows counted between Other paratypes.—PNM 9095–96 (Field lateral body folds (MBVS); midbody dorsal Nos. MGDP 110 and 113), adult females; transverse scale rows counted across dorsum and PNM 9097 (Field No. MGDP 111), adult between lateral body folds (MBDS); midbody male, collected by CO and Marisol Pedigrosa, transverse tubercle rows counted across dor- 23 April 2004 at Ayumit, Municipality of sum between dorsolateral body folds (MBTR); Calayan, Cagayan Province, Babuyan Claro paravertebrals between midpoints of limb Island (19.545u N, 121.958u E, 360 m); PNM insertions (PVS); ventrals within AGD (count- 9098, subadult male, collected by CO and ed midventrally between limb insertion; VS) Marisol Pedigrosa, 27 April 2004 at Corog, paravertebral tubercles within AGD (counted Municipality of Calayan, Cagayan Province, along right side of vertebral axis between limb Babuyan Claro Island 19.489u N, 121.948u E, insertions; PVT); tail annuli (in complete tails; between sea level and 100 m; PNM 9090 TA); and subcaudals (beneath complete tails; (Field No. MGDP 119), adult male, collected SC). For the recognition of the new species, by CO and Marisol Pedigrosa, 27 April 2004 at we adopted the General Lineage Species Rakwaranom, Municipality of Calayan, Caga- Concept (GLC) of de Queiroz (1998, 1999) yan Province, Babuyan Claro Island 19.561u as the natural extension of the Evolutionary N, 121.953u E, between sea level and 100 m. Species Concept (Wiley, 1978). Application of Diagnosis.—Gekko crombota differs from lineage-based species concepts to island all other species of Philippine Gekko (i.e., G. endemics is straightforward because of the athymus, G. ernstkelleri, G. gecko, G. gigante, known history of isolation of island popula- G. mindorensis, G. monarchus, G. palawa- tions (Brown and Diesmos, 2002; Brown and nensis, G. porosus, and G. romblon) in the Guttman, 2002). We consider as new species following combination of features (1) large morphologically diagnosable, isolated popula- body size (SVL 85.5–117.9 for adult males; September 2008] HERPETOLOGICA 307 FIG. 1.—Map of the northern Philippines (inset) showing the type locality (1: Asked) of Gekko crombota on the SE coast of Babuyan Claro Island (A). Additional collection localities include (2) Corog, (3) Rakwaranom, and (4) Ayumit. 85.1–106.9 mm for females); (2) dorsum similar Philippine congener, Gekko porosus medium brown to gray, marked with cream (Batan and Itbayat islands, of the Batanes colored, transverse, tri-lobed bars with a black Islands group, north of the Babuyans), by its anterior border; (3) high numbers of dorsal attainment of a larger maximum male body body scales (107–132 transverse midbody size (males: SVL 85.5–117.9 vs. 91.0–96.7 in scales; 192–226 paravertebrals); (4) high G. porosus; Fig. 5); swollen (presumptively numbers of sharply conical dorsal body hypertrophied) adductor and temporal mus- tubercle rows (17–21 midbody; 27–33 para- culature of the head, leading to a wider, vertebrally); (5) preanofemorals arranged in a vaguely triangular head shape (Fig. 3B,C; non-continuous series (preanals and femoral versus adductor and temporal musculature pore-bearing scale series separated by one or distinctly less enlarged in G. porosus, head two non-pored scales in 75% of specimens) of more slender); brown to gray dorsal colora- 58–74 differentiated, slightly enlarged scales. tion, interrupted by distinct, tri-lobed, cream- Comparison with similar species.—Gekko colored bars (Fig. 2; versus indistinct dark crombota differs from its phenotypically most transverse bands and/or circular vertebral 308 HERPETOLOGICA [Vol. 64, No. 3 FIG. 2.—Photographs of northern Philippine Gekko in life. (A, B) Adult female Gekko crombota (KU 304849; SVL 5 111.8 mm); (C, D) juvenile G. crombota of undetermined sex (KU 304829; SVL 5 58.5 mm); (E) Adult presumably male Gekko porosus and (F) juvenile G. mindorensis of undetermined sex; both photographed (specimens not collected) in Crystal Cave, Batan Island, 2006, by RMB. blotches in G. porosus); a greater number 17–24); separation (1 or 2 undifferentiated (107–132) of transverse midbody dorsal scales scales) between the preanal and femoral series (vs. 88–103 in G. porosus); a greater number of pore-bearing scales (in 75% of specimens; (192–226) of paravertebral scales (vs. 173–191 vs. absent in G. porosus); the presence of in G. porosus); the presence of sharply fewer (58–74) preanofemoral pore-bearing protuberant (vs. merely convex or slightly scales (vs. 74–82 in G. porosus); and the raised) dorsal tubercles; a greater number absence of a modified distal femoral pore- (17–21) of midbody dorsal tubercle rows (vs. bearing patch (vs. present in G. porosus, 15–17 in G. porosus); and a greater number composed of a short series of 2 or 3 rows of (27–33) of paravertebral tubercle rows (vs. pore-bearing scales (Fig. 4C). September 2008] HERPETOLOGICA 309 FIG. 3.—Lateral (A), dorsal (B), and ventral (C) scalation of head of holotype of Gekko crombota (male PNM 9280). Scale bar 5 5 mm. 310 HERPETOLOGICA [Vol. 64, No. 3 FIG. 4.—(A) Ventral view of left hand of holotype of Gekko crombota (male PNM 9280; Scale bar 5 5 mm); (B) left side of preano-femoral pore-bearing scale series of holotype of Gekko crombota (male PNM 9280) and (C) pore-bearing series of an adult male of Gekko porosus (male USNM 266517; Scale bar 5 5 mm). Note single undifferentiated, non- pore-bearing scale interrupting pre-anal and femoral series in G. crombota (arrow). September 2008] HERPETOLOGICA 311 FIG. 5.—Photographs of left side of heads of preserved specimens of (upper) Gekko porosus (juvenile female holotype CAS 60526), (middle) Gekko porosus adult male (USNM 266517), and (lower) Gekko crombota (adult male holotype, PNM 9280); scale bar 5 5 mm. The new species differs from Gekko mon- crombota from G. monarchus, G. gigante, G. archus, G. mindorensis, G. romblon, G. ernstkelleri, G. athymus, and G. gecko (Ta- gigante, G. ernstkelleri and G. palawanensis ble 1). The presence of separated preanal and in having a larger maximum male body size femoral pore-bearing scales is shared at (85.5–117.9 mm SVL; Table 1); the only variable frequencies (Table 1) by at least G. Philippine species approaching the body size crombota, G. romblon, G. gigante, and G. of G. crombota are G. gecko (SVL 120.0– palawanensis and distinguishes these species 153.8 mm) and G. athymus (99.2–119.9). The from G. porosus, G. monarchus, G. min- number of preanofemorals distinguishes G. dorensis, G. ernstkelleri, G. athymus, and G. 312 HERPETOLOGICA [Vol. 64, No. 3 FIG. 6.—Photographs of the coastal forest habitat of G. crombota on Babuyan Claro Island. View of uninhabited east coast of the island (A) and view of the type locality (Asked) from the southeast side of the island (B). In both images, thick volcanic igneous layers are in the foreground, intact coastal forest can be seen above the uninhabited beach, and the largely forested Mt. Pangasun is visible in the background. Photos courtesy of G. Broad. TABLE 1.—Distribution of selected diagnostic characters in Gekko crombota and other Philippine species of Gekko. For simplicity, bilaterally symmetrical characters are presented for the left side only. Measurements are presented in mm and all specimens (with the exception of the G. porosus holotype) are considered adults. Data from seven juvenile paratypes of G. crombota were exclude. September 2008] crombota porosus holotype porosus adults monarchus mindorensis romblon gigante ernstkelleri palawanensis athymus gecko n5 4m; 9f; 1f (juv) 3m 4m; 12f 22m; 13f 4m; 6f 5m; 4f 4m; 8f 3m; 5f 3m; 2f 9m; 12f male SV:L 111.8–117.9 91.0–96.7 56.2–80.7 55.0–88.2 62.7–89.2 89.7–104.7 82.0–92.1 57.2–65.7 99.2–119.9 120.1–166.1 female SVL 85.5–106.9 49.3 91.0–96.7 40.6–69.7 68.2–70.9 58.6–72.5 79.7–87.9 78.0–88.0 44.5–61.8 88.2–117.1 119.2–144.1 Vertebral light tri- light circular indistinct dark transverse dark thin light + dark dark paired white circular dark paired light + dark rust colored coloration lobed blotches transverse spot rows transverse vertebral blotches spots spots inverted spots bars dark bands bands blotches V-shaped Supralabials1 13–15 12 12–13 11–13 11–13 11–14 11–13 15–16 12–14 11–13 12–14 Preanofemorals 58–74 822 74–80 31–40 52–66 71–84 52–66 36–42 64–70 20–24 12–20 Preanal & + (75%) ?3 2 2 2 + (33%) + (28%) 2 + (39%) 2 2 femoral series interrupted Distal femoral 2 ?3 + 2 2 2 2 2 2 2 2 pore-bearing patch present Toe IV scansors 15–18 16 14–16 13–15 12–14 12–13 16–19 17–19 16–19 18–22 17–20 Dorsal protuberant convex convex protuberant protuberant convex protuberant protuberant conical absent conical Tubercles to conical to conical HERPETOLOGICA Internasals 1 1 1 1–2 1 1–2 1 1 1 1 1–3 contacting rostral Scales contacting 5 5 5 5 5 5 5 5–6 5 5 5–6 nostril Midbody ventrals 38–42 40 35–40 38–44 40–47 37–42 41–50 42–48 38–43 30–36 30–35 Midbody Dorsals 107–132 100 88–103 96–112 102–125 102–108 123–135 112–127 114–121 92–104 94–106 Midbody 18–22 15 15–17 16–20 16–20 12–15 12–18 10–16 10–20 — 10–12 tubercle rows Vertebral 29–33 18 17–24 18–23 17–26 18–24 19–28 17–25 23–27 — 18–22 tubercles In AGD Paravertebrals in 192–226 175 173–191 171–203 180–195 175–195 175–207 178–200 155–170 158–179 91–102 AGD Ventrals in AGD 67–85 64 64–74 5761 58–63 63–66 65–74 58–62 54–58 66–72 60–64 1 Defined as all enlarged/differentiated supralabials, counted posteriorly to the point at which scales were no longer differentiated. 2 Taylor (1922) reported 80 preanofemorals; Brown and Alcala (1974) reported 70–72; we count 82 enlarged scales in the pore-bearing series 3 This character cannot be confidently assessed in the immature female holotype (CAS 60526) due to the absence of pores in the preanofemoral series of females. 313 314 HERPETOLOGICA [Vol. 64, No. 3 TABLE 2.—Continuous morphometric variation in the type series of G. crombota. Data are presented for adults only (subadults and juveniles excluded). See Materials and Methods for character definitions. Entries are means 6 1 SD (range in parentheses). Character Males (n 5 4) Females (n 5 9) SVL 101.1 6 16.2 (85.5–117.9) 98.7 6 8.4 (85.1–106.9) TL1 101.7 6 18.7 (80.5–115.7) 83.0 6 8.6 (71.9–94.0) HL 27.7 6 3.6 (24.5–31.1) 26.7 6 1.9 (23.5–28.1) HW 21.2 6 3.3 (18.3–24.6) 20.2 6 1.7 (17.7–21.8) HD 11.6 6 2.0 (9.6–13.5) 11.2 6 1.2 (10.0–12.7) SNL 12.7 6 1.7 (11.2–14.6) 12.7 6 0.7 (11.3–13.1) ED 5.9 6 0.6 (5.3–6.7) 5.1 6 0.6 (4.0–5.7) END 9.9 6 1.5 (8.6–11.5) 9.3 6 0.6 (8.4–10.2) AO 2.5 6 0.7 (1.8–3.2) 2.0 6 0.3 (1.7–2.4) IND 3.3 6 0.9 (2.5–4.2) 3.5 6 0.3 (3.1–3.9) IOD 4.7 6 0.3 (4.4–5.2) 5.0 6 0.4 (4.5–5.6) AGD 48.1 6 8.8 (40.0–58.1) 47.2 6 3.7 (42.3–52.0) FL 22.8 6 3.8 (19.4–26.1) 21.8 6 2.4 (17.8–23.6) TBL 17.4 6 3.8 (14.9–19.9) 17.0 6 1.4 (14.8–18.2) TIL 6.0 6 1.0 (4.8–7.2) 5.5 6 0.7 (4.3–6.3) TIVL 11.6 6 1.8 (9.9–13.2) 10.6 6 0.7 (9.7–11.9) TW 9.8 6 1.0 (8.4–10.8) 9.0 6 1.2 (7.5–10.6) TD 7.3 6 1.0 (6.4–8.2) 7.3 6 0.8 (6.3–8.2) 1 Includes data on regenerated tails (4m; 7f). gecko. The new species is further distin- 46.9% of HL; dorsal surfaces of head rela- guished from G. athymus by the presence of tively homogeneous, with only slightly pro- dorsal body tubercles (absent from G. athymus) nounced concave postnasal, prefrontal, inter- and from G. porosus and G. romblon by the oribital, and parietal depressions; auricular presence of sharply conical dorsal tubercles (vs. opening large, round, angles slightly latero- tubercles only slightly convex in these species). posteriorly from beneath temporal swellings Finally, the presence of distinct transverse light on either side of head; tympanum very deeply cream-colored tri-lobed bars across the dorsum sunken; orbit large, bordered by only slightly appears to distinguish G. crombota from the distinct supraorbital crest; eye large, pupil highly variable suite of color patterns exhibited vertical, margin wavy (Fig. 3A); AO 49.1% of by other Philippine species of Gekko. These and ED; limbs relatively long and slender; femoral other differences among Philippine Gekko segments of hind limbs thick, bulky; TBL species are summarized in Table 1. 16.9% of SVL, 76.2% of FL. Description of holotype.—Adult male in Rostral large, subrectangular, not quite excellent condition (Figs. 3–4), with a small twice as broad as high, with two dorsomedial incision in the sternal region (portion of liver depressions between raised posterodorsal removed for tissue specimen), and hemipenes projections that form the anterolaterally-pro- partially everted. SVL 118.0 mm; habitus jecting edge of the nares and suture anteriorly robust, limbs well-developed, relatively slen- with the supranasals; nostril surrounded by der; tail relatively long; margins of limbs rostral, the first labial, a single lower postnasal, smooth, lacking cutaneous flaps or dermal an enlarged upper postnasal, and an enlarged, folds; a thin adipose line (cutaneous fold) round, convex supranasal; supranasals sepa- running along ventrolateral margin of trunk. rated on either side by a single large Head wide, characterized by extremely internasal; supranasals and internasal followed hypertrophied temporal and adductor and posteriorly by a pair of slightly enlarged temporal musculature; noticeably broader posterosupranasals on each side; supranasals (1.2 times) body at widest point; snout and posterosupranasals each separated by a subtriangular, rounded at tip in dorsal and single minute median scale; scales immedi- lateral aspect (Fig. 3A,B,C); HW 79.1% of HL ately posterior to posterosupranasals only and 20.8% of SVL; SNL 59.3% of HW and slightly enlarged. September 2008] HERPETOLOGICA 315 Total number of differentiated supralabials blade-like ridges) preorbital scales (Fig. 3A, 12/12 (L/R; 9/8–9 to center of eye), bordered B); 38/42 circumorbitals in total, differentiat- dorsally by one row of slightly differentiated ed into the following distinct regions: (1) snout scales; total number of differentiated minute precircumorbitals, (2) enlarged, flat, infralabials 11/11 (8–9/9 to center of eye), squarish circumoribtals dorsoanterior to orbit bordered ventrally by one row of enlarged (7 on left, 8 on right), (3) transverse elongation scales and 3 rows of only slightly differentiated and modification into fringe-like points (spiny chin scales; mental triangular; mental and first ciliaria, 11 on left, 9 on right) across three infralabials on left (first four on right) dorsoposterior margin of orbit, gradually greatly enlarged and wrapping onto ventral reducing to (4) minute postcircumorbitals; a surfaces of chin, nearly twice the size of total of 35 interorbital scales (straight line individual infralabials 4–11; mental followed distance from center of each eye, across both by a pair of slender, elongate postmentals; eyelids). postmentals bordered posterolaterally by a Axilla–groin distance 49.2% of SVL; undif- secondary pair, approximately one half the ferentiated dorsal body scales round to length of first pair, and a tertiary pair of lateral irregularly octagonal, nonimbricate, relatively postmentals, one quarter the length of prima- homogeneous in size, convex; dorsals sharply ry postmentals; postmental scale series bor- transition to imbricate ventrals along the dered posteriorly by a single series of slightly ventrolateral adipose fold; dorsals lack sur- enlarged scales; followed immediately by a rounding interstitial granules but are inter- sharp transition to nondifferentiated chin and spersed with 18 irregularly transverse rows (33 gular scales; postrictal scales slightly enlarged, paravertebral rows) of highly enlarged and 2 or 3 times the size of gular scales; remainder protuberant, to strongly conical dorsal body of undifferentiated gular scales very small, tubercles; each dorsal tubercle with a raised, round, nonimbricate, juxtaposed (Fig. 3C). thorn- to keel-like posteriorly-projecting Dorsal cephalic scales highly heterogeneous point; each tubercle surrounded by a circle and varied in shape, disposition, and distribu- of very slightly enlarged adjacent dorsals; 109 tion; scales of rostrum enlarged, round, oval to transverse midbody dorsals; 202 paraverte- subrectangular, and convex to conical; post- brals between midpoints of limb insertions; 42 nasal, prefrontal, and interorbital depressions transversely arranged ventrals; scales on possess noticeably smaller scales; palpebral dorsal surfaces of limbs larger than dorsals, scales heterogeneous, with some scales as with interspersed enlarged tubercles extend- small as adjacent interorbital region and ing down limbs and terminating at the dorsal others as large and raised as rostral scales; surfaces of hands and feet; enlarged patches undifferentiated posterior head scales granu- of distinct imbricate scales present on wrist, lar, flat to irregularly convex, reducing in size anterior (preaxial) surface of upper arm and posteriorly, interspersed with numerous thigh, on knee, and on distal ventral surface of slightly enlarged conical tubercules, and hind limb; scales on dorsal surfaces of hands making a smooth transition through nuchal and feet similar to dorsal limb scales (but region to small, juxtaposed, flat trunk scales lacking tubercles); ventral body scales flat, interspersed with enlarged, sharply conical cycloid, strongly imbricate, much larger than body tubercles; throat and chin scales small, lateral or dorsal body scales, largest at mid- juxtaposed, nonimbricate; gular and pectoral ventral line. regions with enlarged cycloid, imbricate Seventy-four dimpled pore-bearing scales scales, continuing to increase in size through (Fig. 4B) in a near-continuous preanofemoral ventral abdomen, becoming very enlarged and series (18 preanals on left, 19 on right; strongly imbricate. separated from femorals by a single undiffer- Ornamental cephalic scalation includes entiated scale; 19 femorals on left; 18 on right) numerous conical tubercles on posterolateral each punctured by pore bearing dark orange, portions of head (temporal, supratympanic, exudate, arranged in a wavy, widely obtuse, and postrictal regions) and a short curved inverted ‘‘V’’ formation and continuing to just series of 4–5 enlarged, sharply conical (with before the patellar region; preanal pores 2–3 316 HERPETOLOGICA [Vol. 64, No. 3 times the diameter of femoral pores; preanals subcaudals enlarged, plate-like, 3–4 rows per situated atop a substantial preanal bulge that annulations; subcausals widely expanded to folds over into precloacal region in preserved cover majority of ventral surface of tail, or split specimen but was erect and protuberant in into a pair of subcaudals along posterior life; preanals preceded by five similarly margin of each annulation. enlarged but non-dimpled scale rows; pre- Variation.—Ranges of selected diagnostic anals followed by five enlarged scales rows, meristic characters are presented in Table 1. roughly forming a triangular patch of scales Morphometric variation in mensural charac- before vent; femoral series lacks preceding or ters of adult paratypes is presented in Table 2. following enlarged scale rows; scales latero- The type series contains two large mature posterior to preanofemoral series (i.e., along males, two small but presumably mature ventroposterior surfaces of hind limb) reduce males (with swollen preanal buldges and in size sharply to minute scales of the hemipenal tail base swellings), nine mature posterior edge of the hind limb. females (all gravid, with large white oblong Digits moderately expanded and covered on eggs visible through skin of posteroventral palmar/plantar surfaces by bowed, unnotched, body wall) and eight juveniles/subadults, four undivided scansors (Fig. 4A); digits lack in- of which are males, with hemipenes partially terdigital webbing; subdigital scansors of everted. manus: 11/12, 10/11, 14/15, 14/16, and 13/12 Coloration of holotype in ethanol.—Dorsal on left/right digits I–V respectively; pes: 13/12, ground coloration of head, body, tail, and 11/12, 16/15, 15/15, and 15/14 on left/right dorsal surfaces of limbs medium gray with digits I–V respectively; subdigital scansors of scattered indistinct light gray blotches and manus and pes bordered proximally (on black flecks; light tri-lobed bars traverse the palmar and plantar surfaces) by 1–4 slightly body in nuchal region, above limb insertions, enlarged scales that form a near-continuous and across torso to base of tail; anterior series with enlarged scansors; all digits clawed, margin of tri-lobed light gray dorsal bars but first (inner) claw greatly reduced; remain- bordered by faintly darker pigmentation; ing terminal claw-bearing phalanges com- posterior edge of light gray transverse bars pressed, with large recurved claws, not rising colored as dorsal trunk ground coloration. free at distal end until they extend beyond Dorsal and lateral surfaces of head similar dilated hyperextensible portion of digit. to dorsal ground coloration, but with distinct Tail base bordered by a single, greatly light gray spots; a light cream bar extends enlarged conical postcloacal spur on each side posteriorly from the orbit; palpebra dark gray; of vent; postcloacal swellings pronounced; rostral and supralabials medium gray; infra- hemipenes incompletely everted, their distal labials very light gray. structures not evident; tail long, 98.0% of Limbs colored as torso, lacking transverse SVL; tail not depressed, subcylindrical, divid- banding; dorsal surfaces of hands and feet ed into distinct fracture planes/autotomy light gray; digits light gray with dark gray grooves (5 whorls or annulations); distal tail surfaces of expanded distal portions; tail portions not original; tail with clear autotomy medium gray with dark gray bands corre- scar and distally regenerated portion; 11 sponding to two caudal annuli; regenerated caudal annulations before autotomy scar portion of tail flat gray. (22.2 mm), 12–13 annulations estimated in Ventral head, neck, and torso light cream autotomized portion based on length ventral surfaces of limbs slightly darker; (47.1 mm), for an estimated total possible ventral surfaces of digits (scansors) dark gray; annulation count of 25–26; TD (not including preanofemoal region white with orange pore basal postcloacal swelling) 75.9% of TW; exudate; ventral surfaces of tail medium gray dorsal tail (following description based on (both original and regenerated portions). original portions of tail) as heavily adorned as Coloration of holotype in life.—(from pho- dorsum with tubercles; caudal tubercles tographs of holotype before preservation; concentrated along posterior edge of caudal Fig. 2A–D) Dorsal ground coloration dark annulation; caudals similar in size to dorsals; purplish-gray to yellowish-brown, with indis- September 2008] HERPETOLOGICA 317 tinct dark gray to black blotches; dorsum with Juveniles are patterned more brightly than six light cream tri-lobed bars traversing the adults, with more intensely contrasting light axilla-groin region, each bordered anteriorly and dark dorsal and tail coloration. In some by an accompanying thin transverse wavy juveniles (KU 304821) the tail banding black band (the darkest above insertion of contrast is nearly extreme black and white. hind limbs) and posteriorly by normal trunk Juveniles (KU 304814, 304821, 304829 ground coloration. 304832) also possess cream spots on the Dorsal nuchal region and posterior portions dorsal surfaces of articulations between adja- of head very similar to trunk coloration but cent phalanges. This gives the fingers and toes with distinct, round, cream-colored spots; a light and dark banded appearance. This similar cream bars radiate out from the orbit; pattern is faint in the subadult paratype (KU postrictal region flat gray; labial scales pur- 304807) and absent in all adults. plish-gray with cream spots on every third Ventral coloration is nearly invariant, with labial scale; darker black blotches and flecks the exception that some specimens have very congregate on snout, interorbital region, and dark gray to black ventral tail surfaces (e.g., parietal region; infralabial region and chin KU 304809, 304826, 304849) while others gray to light gray; snout scale purplish gray; have very light gray ventral tail surfaces that gular region light gray to brownish-tan. are only slightly darker than ventral body Dorsal surfaces of limbs light gray with coloration (e.g., KU 304845, 304847). Only in numerous dark brown flecks; dorsal surfaces two juveniles is the black and white transverse of digits dark gray with slightly lighter claws; banded caudal pigmentation strong enough to dorsal and lateral portions of tail banded wrap around on to the ventral surface of the alternating dark gray and cream (correspond- tail (KU 304814, 304821). ing to tail annuli); distal autotomy regrowth Distribution and natural history.—The new dark brown. species is known only from Babuyan Claro Ventral body and limbs yellow with scat- Island where it was collected low (,3 m) on tered gray and dark brown flecks; preano- trunks and buttresses in primary dipterocarp femoral region bright yellow with dark orange forest at low elevations, close to the island’s pores; palmar and plantar surfaces of manus coast. Given the island’s volcanic origin and and pes yellowish with light gray subdigital geological history of isolation (Marini et al., scansors; ventral tail cream with brown 2005; McDermott et al., 1993) we do not transverse bars, and solid brown ventral expect G. crombota to be distributed on other coloration distal to autotomy scar and subse- islands in the Babuyan Island group, the quent regrowth. Batanes Island group, or mainland Luzon. Color variation.—Our sample of four adult Our impression is that the new species is very males, a subadult male, two juvenile males, common at the type locality. No vocalizations nine adult females, and five juveniles of by this species were heard during our brief undetermined sex exhibits moderate color visits to the type locality. The only other variation. The adult male paratype (KU gekkonids encountered during our three night 304825) and two female paratypes (KU stay on Babuyan Claro were Hemidactylus 304809, 304836) have darker dorsal coloration frenatus, Cyrtodactylus philippinicus, and than that of the holotype, with very dark gray Luperosaurus macgregori (see Discussion in anterior margins of the light transverse bars. Brown et al., 2007). Two females (KU 304826, 3004849) have Etymology.—The specific epithet crombota much darker dorsal ground coloration than is a group of letters derived from the names do any of the other specimens. In these Crombie and Ota, and treated as a noun in specimens, the tri-lobed light gray transverse apposition. We employ the amalgam crombota bars take on a more striking, contrasting to jointly honor Ronald Crombie and Hide- appearance. Three adult female paratypes toshi Ota in recognition of their collaborative (KU 304845, 304847, 304851) have a light to survey efforts in the Babuyan Islands (Ota and medium gray, nearly patterenless dorsum, Crombie, 1989; Ota and Ross, 1994) and their with only faint vestiges of transverse banding. continued work on the gekkonid fauna of this 318 HERPETOLOGICA [Vol. 64, No. 3 archipelago (H. Ota and R. Crombie, unpub- Sarangani, Maranduque, Coron, Busuanga, lished data). Suggested common name: Ba- Burias, Ticao, Semira, Semirara, Maestre de buyan Claro Gecko. Campo, Cuyo, Basilan, Jolo, Tawi-Tawi, and many other similarly small, isolated landmass- DISCUSSION es. Fourth, it is clear that comprehensive efforts to survey gekkonid fauna of the The description of Gekko crombota brings Philippines would do well to target isolated the total number of endemic Philippine Gekko species to eight (ten total, when G. hokouensis limestone karst areas and natural caves. Such is excluded and the nonendemic G. mon- habitats are not only patchily distributed (and archus and G. gecko are included). We are thus can be expected to have promoted certain that this number represents an under- evolutionary divergence via isolation) but are estimate of species diversity and we enumer- also heavily imperiled and increasingly under ate the following series of unresolved taxo- threat from overexploitation by humans for a nomic issues that need to be addressed before variety of natural resources (Clements et al., a full realization of Philippine gekkonid 2006). The discovery of G. ernstkelleri in an diversity can be achieved. First, the Babuyans isolated limestone outcop on Panay Island (C. and Batanes island groups require additional D. Siler, personal observations; Roesler et al., survey work before we can be reasonably 2006) demonstrates that hidden gekkonid certain that the total gekkonid fauna is known. diversity awaits field herpetologists willing to We are aware of at least four additional, target long-overlookecd limestone habitats. undescribed Gekko species in the Babuyans Similarly, G. gigante (endemic to the lime- alone and we suspect that several additional stone landbridge Gigante island group; W. undescribed species in the Babuyans and Brown and Alcala, 1978; R. Brown and Alcala, Batanes await discovery on small isolated 2000) is proof that geological isolation (e.g., islands surrounded by deep water. Second, landmasses separated by deep water) may not although morphologically distinct species are be necessary to promote gekkonid diversifica- still being described from other parts of the tion if limestone habitats have been isolated archipelago (Roesler et al., 2006), the majority over geological timescales. of the Philippines’ more subtle or possibly Finally, numerous isolated mountain ranges cryptic species diversity undoubtedly mas- on larger islands (Luzon, Palawan, Samar- querades under the widespread species Gekko Leyte, Mindanao) can be expected to support mindorensis and Gekko monarchus. Prelimi- as yet undocumented gekkonid diversity. nary molecular sequence data (C. Siler, A. Recent discoveries from the Sierra Madre Diesmos, and R. Brown, unpublished data) mountain range of Luzon (Brown et al., suggest that these taxa may be comprised of 2000b, 2007; A. Diesmos, unpublished data) numerous cryptic, unrecognized evolutionary suggest that these remaining forested regions lineages, and potentially are worthy of specific all warrant extensive biodiversity surveys. rank. At a minimum, the major Pleistocene Whatever the circumstances, we are certain Aggregate Island Complexes of the Philip- that the diversity of Philippine Gekko species pines (Brown and Diesmos, 2002; Brown et is substantially underestimated. Given the al., 2007; Gaulke et al., 2007) might each be undeniable fact that destructive exploitation expected to harbor endemic species diversity of Southeast Asian forests is partly related to in both of these species complexes. Third, an ignorance of their biodiversity, it is crucial numerous deep water islands (e.g., not that future faunal inventories throughout the connected to adjacent islands during the last Philippines pay careful attention to habitats glaciations; Brown and Diesmos, 2002) have known to harbor endemic gekkonids. We not been exhaustively surveyed for herpeto- would not be surprised if the eventual number fauna and are thus ripe for the potential of known endemic Philippine Gekko species discovery of additional Gekko species. Islands were to double in the very near future. is this category are are Lubang, Camiguin Acknowledgments.—For the loans of specimens or Norte, Camiguin Sur, Calayan, Dalupiri, assistance while visiting museum collections, we thank Fuga, Masbate, Siquijor, Dinagat, Siargao, the following individuals and their respective institutions September 2008] HERPETOLOGICA 319 (museum abbreviations follow Leviton et al., 1985): A. BROWN, W. C., AND A. C. ALCALA. 1978. Philippine Lizards Resetar, M. Kearney, and H. Voris (FMNH); J. Vindum, of the Family Gekkonidae. Silliman University Press, R. Drewes, and A. Leviton (CAS); J. Simmons and L. Dumaguete City, Philippines. Trueb (KU), A. Wynn, R. Wilson, R. Heyer, and K. de CLEMENTS, R., N. SODHI, M. SCHILTHUIZEN, AND P. K. L. Queiroz (USNM) and R. Sison (PNM). Support for NG. 2006. Limestone karsts of Southeast Asia: Imper- fieldwork was provided by the University of Kansas, and iled arks of biodiversity. Bioscience 56:733–742. the National Science Foundation. The Stearns Fellowship DE QUEIROZ, K. 1998. The general lineage concept of of the California Academy of Sciences provided support species, species criteria, and the process of speciation. that allowed RMB and ACD to undertake multiple visits Pp. 57–75. In D. J. Howard and S. H. Berlocher (Eds.), to CAS. We thank the Department of the Environment Endless Forms: Species and Speciation. Oxford Uni- and Natural Resources, and the Protected Areas and versity Press, New York, New York, U.S.A. Wildlife Bureau (especially C. Custodio, T. M. Lim, and DE QUEIROZ, K. 1999. The general lineage concept of A. Tagtag), for facilitating research and export permits for species and the defining properties of the species this and related studies, KU IACUC for approving category. Pp. 49–89. In R. A. Wilson (Ed.), Species: research protocols, and M. Pedregosa, and M. A. Reyes, New Interdisciplinary Essays. Massachusetts Institute N. Antoque, B. Fernandez, and J. Fernandez for untiring of Technology Press, Cambridge, Massachusetts, assistance in the field. We also thank Municipal DENR U.S.A. authorities of Calayan, Cagayan Province for logistical GAULKE, M., H. ROESLER, AND R. M. BROWN. 2007. A new support. Thanks are due to M. Garfield for scientific species of Luperosaurus (Squamata: Gekkonidae) from illustration and J. Weghorst, A. Bauer, and an anonymous Panay Island, Philippines, with comments on the reviewer for comments on previous versions of the taxonomic status of Luperosaurus cumingii (Gray, manuscript. 1845). Copeia 2007:413–425. LEVITON, A. E., R. H. GIBBS, JR., E. HEAL, AND C. E. DAWSON. 1985. Standards in herpetology and ichthyol- LITERATURE CITED ogy: Part I. Standard symbolic codes for institutional BROWN, R. M. 1999. New species of parachute gecko resource collections in herpetology and ichthyology. (Squamata; Gekkonidae; Genus Ptychozoon) from Copeia 1985:802–821. northeastern Thailand and Central Vietnam. Copeia MANTHEY, U., AND W. GROSSMAN. 1997. Amphibien und 1999:990–1001. Reptilien Sudostasiens. Natur und Tier-Verlag, Berlin, BROWN, R. M., AND A. C. ALCALA. 2000. Geckos, cave Germany. frogs, and small land-bridge islands in the Visayan sea. MARINI, J.-C., C. CHAUVEL, AND R. C. MAURY. 2005. Hf Haring Ibon 2:19–22. isotope compositions of northern Luzon lavas suggest BROWN, R. M., AND A. C. DIESMOS. 2002. Application of involvement of pelagic sediments in their source. lineage-based species concepts to oceanic island frog Contributions to Mineralogy and Petrology populations: the effects of differing taxonomic philos- 149:216–232. ophies on the estimation of Philippine biodiversity. The MCDERMOTT, F., M. J. DEFANT, C. J. HAWKESWORTH, R. C. Silliman Journal 42:133–162. MAURY, AND J. L. JORON. 1993. Isotope and trace BROWN, R. M., AND S. I. GUTTMAN. 2002. Phylogenetic element for three component mixing in the genesis of systematics of the Rana signata complex of Philippine north Luzon lavas (Philippines). Contributions to and Bornean stream frogs: reconsideration of Huxley’s Mineralogy and Petrology 113:9–23. modification of Wallace’s Line at the Oriental-Austra- OTA, H., AND R. I. CROMBIE. 1989. A new lizard of the lian faunal zone interface. Biological Journal of the genus Lepidodactylus (Reptilia: Gekkonidae) from Linnean Society 76:393–461. Batan island, Philippines. Proceedings of the Biological BROWN, R. M., A. C. DIESMOS, AND M. V. DUYA. 2007. A Society of Washington 102:559–567. new Luperosaurus (Squamata: Gekkonidae) from the OTA, H., AND C. A. ROSS. 1994. Four new species of Sierra Madre of Luzon Island, Philippines. Raffles Lycodon (Serpentes: Colubridae) from the northern Bulletin of Zoology 55:167–174. Philippines. Copeia 1994:159–174. BROWN, R. M., J. W. FERNER, AND A. C. DIESMOS. 1997. OTA, H. K.-Y. LUE, S.-H. CHEN, AND W. C. BROWN. 1989. Definition of the Philippine Parachute Gecko, Ptycho- Taxonomic status of the Taiwanese Gekko, with zoon intermedium Taylor 1915 (Reptilia: Lacertilia: comments on the synonymy of Luperosaurus amissus Gekkonidae): redescription, designation of a neotype, Taylor. Journal of Herpetology 23:76–78. and comparisons with related species. Herpetologica ROESLER, H., C. D. SILER, R. M. BROWN, A. D. 53:357–373. DEMEGILLO, AND M. GAULKE. 2006. Gekko ernstkelleri BROWN, R. M., J. SUPRIATNA, AND H. OTA. 2000a. Discovery sp. n.—a new gekkonid lizard from Panay Island, of a new species of Luperosaurus (Squamata; Gekko- Philippines. Salamandra 42:197–211. nidae) from Sulawesi, with a phylogenetic analysis of RUSSELL, A. P. 1979. A new species of Luperosaurus the genus and comments on the status of L. serrati- (Gekkonidae) with comments on the genus. Herpeto- caudus. Copeia 2000:191–209. logica 35:282–288. BROWN, R. M., J. A. MCGUIRE, J. W. FERNER, N. TAYLOR, E. H. 1922a. The lizards of the Philippine Islands. ICARANGAL, JR., AND R. S. KENNEDY. 2000b. Amphibians Philippine Bureau of Science, Manila, Philippines. and reptiles of Luzon Island, II: Preliminary report on TAYLOR, E. H. 1922b. Additions to the herpetological the herpetofauna of Aurora Memorial National Park, fauna of the Philippine Islands, I. Philippine Journal of Philippines. Hamadryad 25:175–195. Science 21:161–206. 320 HERPETOLOGICA [Vol. 64, No. 3 WERMUTH, H. 1965. Gekkonidae, Pygopodidae, Xantusi- ILOILO PROVINCE, Municipality of Carles: CAS 124866–67 dae. Pp. 154–155. In R. Mertens, W. Hennig, and H. (Paratypes); Barangay Asloman: KU 305138–40. Wermuth (Eds.), Das Tierreich, Vol. 80. Walter de Gruyter and Co., Berlin, Germany. Gekko hokouensis.—(1) ‘‘Philippines’’ FMNH 17812 WILEY, E. O. 1978. The evolutionary species concept (Luperosaurus amissus holotype). reconsidered. Systematic Zoology 21:17–26. Gekko mindorensis.—(56) NEGROS ISLAND, NEGROS ORIENTAL PROVINCE, Himangpangon Cave, Manjayod: .Accepted: 3 June 2008 CAS-SU 28656–60; GUIMARAS ISLAND, GUIMARAS .Associate Editor: Christopher Raxworthy PROVINCE, Municipality of Buenavista, Barangay Old Poblacion: KU 302721, 302725; NEGROS ISLAND, NEGROS OCCIDENTAL PROVINCE, Municipality of Cauayan, APPENDIX I Barangay Camalandaan: 302722–24; MASBATE IS- LAND, MASBATE PROVINCE, Municipality of Mandaon, Comparative Material Examined Barangay Poblacion: 302726–28; PANAY ISLAND, CAPIZ All specimens examined are from the Philippines. PROVINCE, Municipality of Pilar, Barangay Natividad: Numbers in parentheses indicate the number of speci- 302729–32; LUBANG ISLAND: OCCIDENTAL MINDORO mens examined for each species and museum abbrevia- PROVINCE, Municipality of Lubang, Barangay Vigo: KU tions follow Leviton et al. (1985). 303913–16, 303917–951. Gekko athymus.—(7) PALAWAN ISLAND, PALAWAN Gekko monarchus.—(3) PALAWAN ISLAND, PALA- PROVINCE, ca. 10 km WSW of Iwahig: CAS 137677; ca. 8– WAN PROVINCE, ca. 1.5 km. W.S.W. of Iwahig: CAS-SU 9 km S. of Balico: CAS-SU 23119 (holotype); ca. 20 km 28416; ca. 5 km SSE of Iwahig: CAS-SU 28496; ca. 7 km SW of Iwahig: CAS-SU 23121 (paratype); Municipality of WNW of Iwahig: CAS-SU 28554. ˜ Brooke’s Point, Barangay Samarinana; Mt. Mantalingahan: Gekko palawanensis.—(25) PALAWAN ISLAND, PA- KU 309331–34. LAWAN PROVINCE, 7 km WNW of Iwahig: CAS 17318; 8 km Gekko ernstkelleri.—(10) PANAY ISLAND, ANTIQUE W of Iwahig: CAS 17319; ca. 9 km W of Iwahig: CAS PROVINCE, Municipality of Pandan, Barangay Duyong, 17320–22; KU 30948, 309171, 309279–95, 309468. Duyong Hillside (5 ‘‘Mt. Lihidian’’), 300 m.a.sl.: PNM Gekko porosus.—(4) BATANE ISLAND, BATANES 9152–54; KU 300196–202. PROVINCE, 3 km ENE of Basco Town: USNM 266519, Gekko gecko.—(13) LUBANG ISLAND, OCCIDENTAL 291387; Mahatao: USNM 266517; ITBAYAT ISLAND: MINDORO PROVINCE, Municipality of Lubang, Barangay CAS 60526 (holotype). Paraiso: KU 303960–72. Gekko romblon.—(12) SIBUYAN ISLAND, ROMBLON Gekko gigante.—(8) SOUTH GIGANTE ISLAND, PROVINCE, Taclobo Barrio: CAS 139180–82 (paratypes); ILOILO PROVINCE, Municipality of Carles, Tantangan: CAS ROMBLON ISLAND, ROMBLON PROVINCE, Municipality 124315–17 (paratypes); NORTH GIGANTE ISLAND, of Romblon, Barangay Li-O: KU 302736–42, 303977–78.
Pages to are hidden for
"ANEW GEKKO FROM THEBABUYAN ISLANDS, NORTHERN PHILIPPINES"Please download to view full document