VIEWS: 142 PAGES: 16

									Herpetologica, 64(3), 2008, 305–320
E 2008 by The Herpetologists’ League, Inc.

                           NORTHERN PHILIPPINES
       RAFE M. BROWN1,4, CARL H. OLIVEROS1,2, CAMERON D. SILER1,                       AND   ARVIN C. DIESMOS3
      Natural History Museum and Biodiversity Research Center, Department of Ecology and Evolutionary Biology,
                               The University of Kansas, Lawrence, KS 66045-7561, USA
         ISLA Biodiversity Conservation; 9 Bougainvillea St, Manuela Subdivision, Las Pinas City 1741 Philippines
            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
      CORRESPONDENCE: e-mail,                         gonads or by scoring of prominent secondary

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
Distal femoral          2            ?3             +             2                                     2                 2                  2                  2              2           2         2
   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

Internasals              1            1             1            1–2                                     1               1–2                 1                1                1            1       1–3
Scales contacting        5            5             5             5                                      5                5                  5                 5–6             5            5       5–6
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
Paravertebrals in 192–226           175         173–191       171–203                               180–195           175–195            175–207            178–200        155–170       158–179   91–102
Ventrals in AGD      67–85           64          64–74          5761                                 58–63             63–66              65–74              58–62          54–58        66–72     60–64
     Defined as all enlarged/differentiated supralabials, counted posteriorly to the point at which scales were no longer differentiated.
     Taylor (1922) reported 80 preanofemorals; Brown and Alcala (1974) reported 70–72; we count 82 enlarged scales in the pore-bearing series
     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.
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)
      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 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.

To top