Phylogenetic relationships of geckos of the Hemiphyllodactylus harterti group, a new species from Penang Island, Peninsular Malaysia, and a likely case of true cryptic speciation
Author
Cobos, Anthony
Author
Grismer, L. Lee
Author
Wood, Perry L.
Author
Quah, Evan S. H.
Author
Anuar, Shahrul
Author
Muin, Mohd Abdul
text
Zootaxa
2016
4107
3
367
380
journal article
39157
10.11646/zootaxa.4107.3.5
2aa0be33-190d-44be-ac3a-7ee41d4cd706
1175-5326
265560
7791AAD7-2F1D-47EB-88A7-1B97C765BFFC
Hemiphyllodactylus cicak
sp. nov.
Penang Island Slender
Gekko Cicak Kerdil
Pulau
Pinang
Figs. 3
,
4
Holotype
. Adult male (
LSUCH
11762) collected by Evan Quah and L. Lee Grismer on
5 May 2014
at
600 m
elevation from the living room of the old Ban Hin Lee Guest House on Penang Hill,
Pulau
Pinang, Peninsular
Malaysia
(
5°25’23.14”N
,
100°16’19.79”E
) at approximately 1800 hrs during a heavy thunderstorm.
Paratypes
.
Three adult females (
LSUHC
11763–65) are associated with the same collection data as the
holotype
.
FIGURE 3
. Male Holotype (LSUCH 11762) of
Hemiphyllodactylus cicak
sp. nov.
from Penang Hill, Penang Island, Penang, Peninsular Malaysia.
Diagnosis.
Hemiphyllodactylus cicak
sp. nov.
can be separated from all other species of
Hemiphyllodactylus
by possessing the unique combination of characters, having a maximum SVL of
31.4 mm
; three circumnasal scales; one or two scales between supranasals (=postrostrals); eight supralabials; eight infralabials; 10 or 11 longitudinally arranged ventral scales at midbody contained within one eye diameter; a series of 42 pore-bearing precloacal-femoral scales in the male; lamellar formula on hand 2–3–3–2; lamellar formula on foot 2–3–3–3; dorsal body pattern consisting of dark, transverse, paravertebral blotches coupled with white speckles; dark pre- and postorbital stripes; faint postorbital stripe extends along the flanks to the hind limbs. These characters, and other diagnostic characters, are scored across all species of
Hemiphyllodactylus
within the
harterti
group listed in
Table 3
.
Description of
holotype
. Adult male; head sub-triangular in dorsal profile, depressed, distinct from neck; lores and interorbital regions flat; rostrum moderate in length (NarEye/HeadL 0.23); prefrontal region flat to weakly concave; canthus rostralis smoothly rounded, barely discernable; snout moderate, rounded in dorsal profile; eye large; ear opening round, small; eye to ear distance greater than diameter of eye; rostral wider than high, partially divided dorsally, bordered posteriorly by small supranasals; one internasal (= postnasal); external nares bordered anteriorly by rostral, dorsally by supranasal, posteriorly by two postnasals, ventrally by first supralabial (= circumnasals 3R,L); 8 (R,L) square supralabials tapering to below posterior margin of orbit; 8 (R,L) square infralabials tapering to below posterior margin of orbit; scales of rostrum, lores, top of head, and occiput small, granular, those of rostrum largest and slightly raised; dorsal superciliaries flat, rectangular, subimbricate; mental triangular, bordered laterally by first infralabials and posteriorly by two large postmentals; each postmental bordered laterally by a single sublabial; one row of smaller scales extending transversely from juncture of second and third infralabials and contacting mental; seven chin scales; gular scales small, subimbricate, grading posteriorly into slightly larger, subimbricate, throat and pectoral scales which grade into slightly larger, subimbricate ventrals.
TABLE 3.
Diagnοstic characters οf the species in the
Hemiphyllodactylus harterti
grοup.
harterti
|
tehtarik
|
larutensis
|
bintik
|
titiwangsaensis
|
cicak
sp. nov.
|
max SvL 39 |
40.4 |
52.2 |
36.6 |
52.2 |
31.4 |
chin scales 6‾8 |
8 |
6‾10 |
7 |
7‾11 |
7 |
pοstmentals distinctly enlarged (1) οr nοt (0) 1 |
1 |
1 |
1 |
1 |
1 |
circumnasal scales 2‾5 |
5 |
3 ‾5 |
5 |
3 ‾4 |
3 |
scales between supranasals 3 ‾4 |
3 |
3 |
3 |
2 ‾4 |
1 ‾2 |
supralabial scales 10‾11 |
11 |
9,10 |
11 |
5‾8 |
8 |
infralabial scales 10‾11 |
10 |
7‾10 |
12 |
5‾8 |
8 |
dοrsal scales 14‾19 |
18 |
13‾20 |
17 |
17 |
16‾18 |
ventral scales 6‾14 |
12 |
7‾13 |
7 |
8 |
10‾11 |
lamellar fοrmula οn hand 3‾3‾3‾3 |
3‾3‾3‾3 |
⁄ |
2‾4‾4‾3 |
3‾4‾4‾4 |
2‾3‾3‾2 |
lamellar fοrmula οn fοοt 3‾3‾4‾3 |
3‾4‾5‾4 |
⁄ |
3‾4‾4‾4 |
4‾5‾5‾5 |
2‾3‾3‾3 |
subdigital lamellae οn first finger 3 |
5 |
3,4 |
4 |
2 |
2 |
subdigital lamellae οn first tοes 4 |
5 |
3‾5 |
5 |
2 |
2 |
Preclοacal and femοral pοres series separate (1) οr cοntinuοus (0) 0 |
nοne |
0 |
⁄ |
0 |
0 |
Preclοacal and femοral pοres 42‾45 |
0 |
27‾36 |
⁄ |
26‾32 |
42 |
clοacal spurs 1,2 |
3 |
2,3 |
1 |
2‾3 |
2 |
subcaudals enlarged, plate-like (1) οr nοt (0) 0 |
⁄ |
0 |
0 |
0 |
0 |
dark pοstοrbital stripe present (1) οr absent (0) 1 |
1 |
1 |
1 |
1 |
1 |
light pοstοcular οr trunk spοts (1) οr absent (0) 1 |
0 |
1 |
0 |
1 |
1 |
dark dοrsοlateral stripe οn trunk (1) οr absent (0) 1,0 |
0 |
0 |
0 |
0 |
0 |
dοrsal pattern unicοlοr (1) οr nοt (0) 0 |
1 |
1 |
0 |
0 |
0 |
dark dοrsal transverse blοtches (1) οr nοt (0) 0 |
0 |
0 |
0 |
1 |
0 |
lοngitudinal series οf white (1) οr yellοw οr red (0) dοrsal spοts 0 |
0 |
0 |
0 |
1 |
1 |
pοstsacral mark lacking anteriοr arms (1) οr arms present (0) 0 |
1 |
1 |
0 |
1 |
1 |
caecum pigmented (1) οr nοt (0) 1 |
0 |
0 |
1 |
0 |
0 |
gοnads pigmented (1) οr nοt (0) 0 |
0 |
0 |
0 |
0 |
0 |
Τrunk⁄SvL 0.48‾0.53 |
0.55 |
0.46‾0.51 |
0.49 |
0.43‾0.48 |
0.47‾0.50 |
HeadL⁄SvL 0.22‾0.24 |
0.2 |
0.21‾0.24 |
0.23 |
0.15‾0.27 |
0.25 |
HeadW⁄SvL 0.16‾0.18 |
0.16 |
0.15‾0.17 |
0.18 |
0.15‾0.19 |
0.16‾0.17 |
HeadW⁄HeadL 0.65‾0.85 |
0.8 |
0.63‾0.73 |
0.8 |
0.63‾0.71 |
0.63‾0.67 |
SnEye⁄HeadL 0.41‾0.48 |
0.47 |
0.39‾0.51 |
0.41 |
0.34‾0.38 |
0.35 |
NarEye⁄HeadL 0.28‾0.33 |
0.4 |
0.27‾0.36 |
0.33 |
0.23‾0.29 |
0.22‾0.23 |
EyeD⁄HeadL 0.22‾0.30 |
0.28 |
0.22‾0.28 |
0.24 |
0.21‾0.24 |
0.22 |
SnW⁄HeadL 0.15‾0.22 |
0.16 |
0.11‾0.15 |
0.17 |
0.14‾0.21 |
0.16 |
EyeD⁄NarEye 0.81‾1.00 |
0.72 |
0.66‾0.90 |
0.74 |
0.84‾0.94 |
0.95‾1 |
Snv⁄HeadW 0.20 ‾0.32 |
0.2 |
0.18 ‾0.21 |
0.21 |
0.21 ‾0.31 |
0.19 |
Body somewhat elongate (Trunk/SVL 0.49), dorsoventrally compressed; ventrolateral folds absent; dorsal scales small, granular, 17 scales at midbody contained within one eye diameter; ventral scales, flat, subimbricate much larger than dorsal scales, ten scales contained within one eye diameter; enlarged, precloacal scales; 42 porebearing femoral and precloacal scales; forelimbs short, slender in stature, covered with flat, subimbricate scales dorsally and ventrally; palmar scales flat, subimbricate; all digits except digit I well-developed; digit I vestigial, clawless; distal, subdigital lamellae of digits II–V undivided, angular and U-shaped; lamellae proximal to these transversely expanded; lamellar formula of digits
II–V 2–3
–3–2 (R,L); two transversely expanded lamellae on digit I; claws on digits II–V well developed, unsheathed; distal portions of digits strongly curved, terminal joint free, arising from central portion of lamellar pad; hind limbs short, more robust than forelimbs, covered with flat, juxtaposed scales dorsally and by larger, flat subimbricate scales ventrally; plantar scales low, flat, subimbricate; all digits except digit I well-developed; digit I vestigial, clawless; distal, subdigital lamellae of digits II–V undivided, angular and U-shaped; lamellae proximal to these transversely expanded; lamellar formula of digits
II–V 2–3
–3–3 (R,L); two transversely expanded lamellae on digit I; claws on digits II–V well developed, unsheathed; distal portions of digits strongly curved, terminal joint free, arising from central portion of lamellar pad; tail rectangular in cross-section. Morphometric data are presented in table 4.
TABLE 4.
Mensural and meristic data from the type series of
Hemiphyllodactylus cicak
sp. nov.
from the LSUHC collection.
11762 |
11763 |
11764 |
11765 |
Sex |
m |
f |
f |
f |
Chin
Scales
|
7 |
7 |
7 |
7 |
Postmentals distinctly enlarged (1) or not (0) |
1 |
1 |
1 |
1 |
Circumnasal scales |
3 |
3 |
3 |
3 |
Scales between supranasals |
1 |
1 |
1 |
2 |
Supralabial scales |
8 |
8 |
8 |
8 |
Infralabial scales |
8 |
8 |
8 |
8 |
Dorsal scales |
17 |
16 |
16 |
18 |
Ventral scales |
10 |
10 |
10 |
11 |
Lamellar formula on hand |
2–3–3–2 |
2–3–3–2 |
2–3–3–2 |
2–3–3–2 |
Lamellar formula on foot |
2–3–3–3 |
2–3–3–3 |
2–3–3–3 |
2–3–3–3 |
Subdigital lamellae on first finger |
2 |
2 |
2 |
2 |
Subdigital lamellae on first toes |
2 |
2 |
2 |
2 |
precloacal and femoral pore series separate (0) or continuous (1) |
0 |
/ |
/ |
/ |
# precloacal and femoral pores |
42 |
/ |
/ |
/ |
dark postorbital strip present (0) or absent (1) |
0 |
0 |
0 |
0 |
Number of cloacal spurs |
2 |
1 |
1 |
1 |
Subcaudals enlarged, plate like (1) or not (0) |
0 |
0 |
0 |
0 |
Dorsal pattern unicolor (1) or not (0) |
0 |
1 |
1 |
0 |
Dark dorsal transverse blotches (1) or not (0) |
0 |
0 |
0 |
0 |
Snout Vent Length |
31.4 |
28.9 |
25.4 |
28.5 |
Trunk Length |
15.6 |
14.04 |
11.9 |
14.5 |
Head Length |
7.7 |
6.8 |
6.3 |
6.5 |
Head Width |
4.9 |
4.7 |
4.2 |
4.6 |
Eye Diameter |
1.7 |
1.6 |
1.4 |
1.6 |
Snout Eye Length |
2.8 |
2.7 |
2.2 |
2.7 |
Nar Eye Internarial Width (SnW) |
1.8 |
1.4 |
1.4 |
1.6 |
FIGURE 4.
Ventral view of the type series
Hemiphyllodactylus cicak
sp. nov.
Coloration before preservation (
Fig. 3
)
. Top of head, body, and limbs beige with black spots and white speckles; dark pre- and postorbital and paired, paroccipital stripes present; dorsum overlain with paired, dark, slightly offset, squarish, paravertebral markings; ground color of the dorsum on the anterior portion of the tail gray while the ground color of the rest of the tail light beige; large, dark, lateral markings on anterior portion of tail that fade toward the posterior; flanks and dorsal surfaces of limbs darkly mottled; ventral surfaces of head, neck, body, and limbs whitish, semi –transparent with greyish brown speckling especially along the sides of the body; subcaudal region orange, especially bright on the underside of the tail (
Fig. 4
).
Distribution
.
Hemiphyllodactylus cicak
sp. nov.
is known only from the
type
locality of Penang Hill, Penang, Peninsular
Malaysia
(
Fig. 1
).
Natural history.
All specimens of the
type
series were collected at night in the Ban Hin Lee Guest House during a heavy downpour. This species occurs in syntopy with various other species of geckos in the guesthouse those being
Hemiphyllodactylus
typus
Bleeker
,
Gehyra mutilata
Wiegmann
,
Gekko monarchus
Schlegel
, and
Hemidactylus frenatus
Duméril, Bibron.
LSUHC 11764 was a gravid female. An additional gravid female (LSUHC 12488 not part of the
type
series) was collected late at night crossing a road near the guest house bordered by hill dipterocarp forest on the
30 July 2015
. This indicates the reproductive biology of this species extends from at least early May through late July.
Etymology
. This specific epithet “
cicak
” is the Malay word for lizard.
Variation (
Fig. 4
).
The general color patterns of the
paratypes
closely match that of the
holotype
. LSUCH 11763 is an adult female that has a partially regenerated tail with a soft gray color. Differences in scales counts are presented in
Table 4
.
Comparisons.
The molecular analysis indicates that
Hemiphyllodactylus cicak
sp. nov.
is embedded within the
harterti
group. It can be distinguished from all other species in that group by having a SVL of
31.4 mm
, which is smaller than all other species in the group (
Table 3
).
Hemiphyllodactylus cicak
sp. nov.
has a manual lamellar formula of 2–3–3–2 as opposed to 3–3–3–
3 in
H. harterti
, 2–4–4–
3 in
H. tehtarik
, or 3–4–4–
4 in
H. bintik
and
H. titiwangsaensis
,
respectively, and a pedal lamellar formula of 2–3–3–3 as opposed to 3–3–4–
3 in
H. harterti
, 3–4–5–
4 in
H. tehtarik
, 3–4–4–
4 in
H. bintik
, or 4–5–5–
5 in
H. titiwangsaensis
, respectively. It also differs from all other species of the
harterti
group by having only one or two scales between the supranasals as opposed to three, four, or five.
Hemiphyllodactylus cicak
sp. nov.
has eight supralabial scales, whereas
H
.
harterti
has 10 or 11,
H. tehtarik
has 11,
H. larutensis
has nine or 10,
bintik
has 11, and
H. titiwangsaensis
has 5–8.
Hemiphyllodactylus cicak
sp. nov.
can be distinguished from
H. tehtarik
and
H. bintik
by having light postocular and trunk spots.
Hemiphyllodactylus cicak
sp. nov
.
lacks the dark transverse blotches present in
H. titiwangsaensis
, and lacks the unicolor dorsal pattern of
H. tehtarik
and
H. larutensis
. Additionally, the new species can be differentiated from
H. titiwangsaensis
and
H. larutensis
by having a precloacal and femoral pores series of 42, as opposed to 26–32 or
27–36 in
these species, respectively. Although
H. cicak
sp. nov
.
is most closely related to
H. bintik
and
H. harterti
it has two lamellae on its first finger as opposed
H. harterti
which has three and
H. bintik
which has four. A similar difference can be seen on digit one of the foot where
H. harterti
has four lamellae and
H. bintik
has five, whereas the new species has two. There is an uncorrected pairwise sequence divergence of approximately 8.07% between
H. harterti
and
H. bintik
.
Previously,
Grismer
et al.
(2013)
noted that a divergence of at least 5.0% in
Hemiphyllodactylus
was consistent with discrete, diagnostic, morphological differences delimiting species boundaries within gekkonids in general.