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New Species of Narrow-Mouthed Frog (Amphibia: Anura: Microhylidae; Genus
Kaloula) from the Mountains of Southern Luzon and Polillo Islands, Philippines
Author(s): Arvin C. Diesmos, Rafe M. Brown, Angel C. Alcala
Source: Copeia, 2002(4):1037-1051.
Published By: The American Society of Ichthyologists and Herpetologists
DOI: http://dx.doi.org/10.1643/0045-8511(2002)002[1037:NSONMF]2.0.CO;2
URL: http://www.bioone.org/doi/full/10.1643/0045-8511%282002%29002%5B1037%3ANSONMF
%5D2.0.CO%3B2
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q
2002 by the American Society of Ichthyologists and Herpetologists
Copeia, 2002(4), pp. 1037–1051
New Species of Narrow-Mouthed Frog (Amphibia: Anura: Microhylidae;
Genus Kaloula) from the Mountains of Southern Luzon and Polillo
Islands, Philippines
A
RVIN
C. D
IESMOS
,R
AFE
M. B
ROWN
,
AND
A
NGEL
C. A
LCALA
We describe a new species of narrow-mouthed frog of the genus Kaloula from the
volcanic mountains of southern Luzon Island (Mt. Banahao, Mt. Isarog, and Mt.
Mayon) and adjacent Polillo Island, Philippines. On the volcanoes of southern Lu-
zon, the new species is found in habitats ranging from small dry stream beds to
stationary pools of rivers in mid- to upper montane primary forest. On Polillo Is-
land, the new species has been found near quiet streams in selectively logged pri-
mary forest and second growth near sea level. The new species presumably is allied
to Kaloula rigida (a forest species endemic to northern Luzon Island) and to Kaloula
picta (a more widespread Philippine endemic that is found in a variety of habitats)
as indicated by possession of narrow disks on the digits and by the presence of
supernumerary tubercles on the palmar surface of the manus. It differs from these
species by its smaller body size, reduction of webbing on toes, absence or extreme
reduction of outer metacarpal tubercles, and by characteristics of the advertisement
call.
Isang ba´gong uriı´n ng palaka´ng kı´tirang-bibig (Amphibia: Anura: Microhylidae;
genus Kaloula) mula sa bu´lubundu´kin ng Pulong Luzon at Pulong Polillo, Pilipinas
(Lagom).—Inilalarawan namin ang isang ba´gong uriı´n ng palaka´ng kı´tirang-bibig ng
angka´ng Kaloula mula sa mabulka´ng bulu´ bundukin ng katimu´gang Luzon (Bk. Ba-
nahao, Bk. Isarog, at Bk. Mayon) at sa mga pulo´ ng Polillo, Pilipinas. Sa mga bulkan
ng katimu´gang Luzon, ang bagong uriı´n ay matatagpuan sa mga pa´nirahang abo´t
mula sa mga maliliit na patay na sapa´ hanggang sa mga tining na layo´ n ng mga ilog
sa may kalagitnaan hanggang sa ilayang gubat o orihinal na tampo´ k-gubat. Ang isang
populasyon ay nakilala rin mula sa mga mababang hayon ng mga pa´ nirahang-gubat
sa pulo´ ng Polillo. Ano bagong uriı´n ay kaanib ng Kaloula rigida (isang uriı´ng-gubat
na taa´l sa Pulong Luzon) at ng Kaloula picta (higit na palasa´ k na uriı´ng ta´al sa
Pilipanas at matatagpuan sa mga iba’t-ibang uri ng pa´nirahan) ba´tay sa ipinapakita
nitong pag-aangkin ng mga makikitid na disk sa mga daliri at ng kawalan nito ng
mga butlig sa sa´latang pa´lad sa kama´y nito´ . Ita ay naiiba sa mga na´banggit na uriı´n
dahil sa pangangatawan nitong may kaliitan bawa´s na pa´likpikan sa mga dulong
daliri, at sa maraming katangian nito sa usaping taga´l at ispektrum ng kanyang hud-
ya´t-panta´wag.
A
MONG the first reports of microhylids in
the Philippines were accounts of species
eventually referred to the genus Kaloula Gray,
1831. These included the descriptions of Plectro-
pus pictus Dume´ ril and Bibron, 1841 (
5
Kaloula
picta) and Hylaedactylus conjunctus Peters, 1863
(
5
Kaloula conjuncta), both from the island of
Luzon. Taylor (1920, 1922a,b) later recognized
six Philippine species of Kaloula (Kaloula baleata,
K. conjuncta,Kaloula kalingensis,Kaloula negrosen-
sis,K. picta, and Kaloula rigida). In his mono-
graphic treatment of Philippine Amphibia, In-
ger (1954) considered K. kalingensis (from Lu-
zon Island) and K. negrosensis (Negros Island)
to be subspecies of K. baleata and K. conjuncta,
respectively. He also described two new subspe-
cies of K. conjuncta:Kaloula conjuncta meridionalis
of Mindanao Island and Kaloula conjuncta stickeli
of Leyte Island. In total, Inger (1954; see also
Alcala and Brown, 1998) recognized four Phil-
ippine Kaloula species within two unnamed,
nonphylogenetic species groups. The first
group (characterized by the presence of super-
numerary tubercles) consisted of K. conjuncta
(with four subspecies, Kaloula conjuncta conjunc-
ta,K. c. meridionalis,Kaloula conjuncta negrosensis,
and K. c. stickeli), K. picta, and K. rigida. The
second group (supernumerar y tubercles ab-
sent) was represented by K. baleata (with sub-
species Kaloula baleata baleata from Java, Bor-
neo, and Palawan and Kaloula baleata kalingensis
from Luzon).
More recently, Ross and Gonzales (1991) el-
evated K. kalingensis to the status of a full species
1038 COPEIA, 2002, NO. 4
(see also Brown et al., 1996, 2000a; Diesmos,
1998) and described a new Philippine species
(K. kokacii) from Catanduanes Island. In addi-
tion to Catanduanes, Kaloula kokacii has recently
also been collected from Camarines Norte, Al-
bay, and Sorsogon provinces of the Bicol Pen-
insula, southern Luzon (RMB and ACD, un-
publ. data). This distinct species should be con-
sidered part of Inger’s (1954) second species
group by virtue of its close phenotypic similarity
to K. kalingensis (Ross and Gonzales, 1991). In
this paper, we describe a fourth species, presum-
ably allied to the first species group (K. picta,K.
rigida, and possibly K. conjuncta) of Philippine
Kaloula.
M
ATERIALS AND
M
ETHODS
We examined specimens deposited at the Cal-
ifornia Academy of Sciences (CAS), the Philip-
pine National Museum (PNM), the Texas Nat-
ural History Collection of the Texas Memorial
Museum (TNHC), and the Field Museum of
Natural History (FMNH).
To minimize interobserver error (Lee, 1982;
Hayek et al., 2001), only measurements taken
by ACD (with needle point dial calipers, to the
nearest 0.1 mm) were used in statistical analyses
and the description. Morphometric characters
(defined by Matsui, 1984; Heyer et al., 1990)
included snout–vent length (SVL), head length
(HL), head width (HW), snout length (SL), in-
terorbital distance (IOD), internarial distance
(IND), eye–nostril distance (END), eye–tympa-
num distance (ETD), horizontal diameter of
eye (ED), horizontal diameter of tympanum
(TD), manus length (ML), pes length (PL), ra-
dius length (RL) femur length (FL), tibia
length (TBL), tarsus length (TSL), third finger
length from proximal edge of basal tubercle
(Fin3L), third finger disk width (Fin3DW),
third finger penultimate phalanx width
(Fin3PPW); forth toe length (Toe4L), and forth
toe disk width (Toe4DW). Other qualitative
morphological characters examined include
color pattern, degree of terminal digital disk ex-
pansion, structure of tubercles on hands and
feet, extent of ventral adhesive glands in males,
and degree of webbing between fingers and
toes.
Calls of the new species, K. kalingensis,K. c.
conjuncta, and K. picta were recorded between
25 C and 27 C with either a Sony
y
WM DC6
Professional Walkman with a Sennheiser
y
ME80 condenser microphone (equipped with
K3U power module) or a Sony
y
TCM-453V
voice-activated recorder. Calls were recorded at
distances ranging from 0.5–2.0 m and ambient
temperatures were recorded immediately after
calls were recorded. Calls were digitized and an-
alyzed using Soundedit
q
(vers. 16 Users’ Guide,
Macromedia, Inc., San Francisco, CA, 1995) and
Canary
q
(vers. 2.0 Users’ Manual, R. A. Charif,
S. Mitchell, and C. W. Clark, Canary Cornell
Laboratory of Ornithology, Ithaca, NY, 1996)
software installed on a Macintosh computer. We
examined oscillograms (waveforms), audiospec-
trograms (sonograms), and results of the Fast
Fourier Transformation (frequency spectrum)
for a variety of temporal and spectral characters.
R
ESULTS
Kaloula walteri sp. nov.
Figures 1–3
Holotype.—PNM 6725 (field no. ACD 168), an
adult male collected 5 May 1995 at 1700 h by
A. C. Diesmos and R. A. N. Altamirano at an
elevation of 950 m on the southeast slope of Mt.
Banahao, Barangay Lalo, Municipality of Taya-
bas, Quezon Province, Luzon Island, Philip-
pines (14
8
4
9
N, 121
8
29
9
E).
Paratopotypes.—Two adult males; PNM 6662
(ACD 166) and PNM 6724 (ACD 167), collected
with the holotype on the same date.
Other paratypes.—Fourteen males; CAS 211741–
42 (ACD 810–811) and TNHC 59666, all adult
males collected July 1995 by R. E. Fernandez at
an elevation of 700 m on the eastern slope of
Mt. Banahao, Sitio Barod, Barangay Palola, Mu-
nicipality of Lucban, Quezon Province; PNM
6663, collected 10 April 1996 by A. C. Alcala at
650 m in Solsogin Creek on Mt. Banahao, Ba-
rangay Samil, Municipality of Lucban, Quezon
Province; PNM 6664–6665 and TNHC 59667,
TNHC 60116, collected at the same locality on
9 September 1995 by A. C. Diesmos; PNM 6763–
6767, collected 21 April, 1995 by R. E. Fernan-
dez on Mt. Banahao, Municipality of Tayabas,
Quezon Province; TNHC 61117, adult male col-
lected 28 Aug 2001 by R. M. Brown, R. E. Fer-
nandez and V. Yngente at an elevation of 675
m, southeast slope of Mt. Banahao, Barangay
Lalo, Municipality of Tayabas, Quezon Province.
Referred specimens.—PNM 6726, an adult female
from 990 m on the east slope of Mt. Isarog, Ba-
rangay Hiwacloy, Municipality of Goa, Camari-
nes Sur Province, southern Luzon Island, Phil-
ippines (13
8
39
9
N, 123
8
23
9
E); PNM 5294 and
5307, from 650 m elevation, Mt. Mayon, Baran-
gay Buang, Municipality of Buang, Albay Prov-
ince; PNM 6723 adult male from 25 m Polillo
1039DIESMOS ET AL.—NEW PHILIPPINE KALOULA
Fig. 1. Color pattern variation in live Kaloula walteri, photographed on Mt. Banahao, Luzon Island. Light
phase (above, tan dorsum with dark marbling) and dark phase (below, dark gray with very dark brown marbling
both from the same population in the Municipality of Tayabas in 1998 (see Fig. 4). These specimens were not
collected; photos: RMB.
Watershed Reserve, Municipality of Polillo,
Quezon Province, Polillo Island.
Etymology.—The new species is named in honor
of Walter C. Brown in recognition of his im-
mense contributions to our knowledge of the
systematics, zoogeography, and ecology of Phil-
ippine amphibians and reptiles. Suggested
Common Name: Walter’s narrow-mouthed frog.
Diagnosis.—A small species of Kaloula, similar in
appearance to K. picta and K. rigida but differ-
ing by a smaller body size of males (SVL
5
24.5–
31.5 mm; vs 38.1–51.2 in K. picta and 33.1–43.9
in K. rigida; Appendix 1) by the absence (in four
specimens) or presence (eight specimens) of
only a faint, very reduced, rounded outer meta-
tarsal tubercle (vs tubercles invariably present,
very distinct, enlarged and pointed), by the ab-
sence of lateral glandular flank ridges (vs pres-
ence of a faint glandular ridge or tubercle row
stretching from eye to axilla), and by the pres-
ence of two subarticular tubercles of the fourth
toe in 70% of the specimens (vs three in K. picta
and in 80% of K. rigida specimens examined).
The new species also differs from K. picta by the
presence of an inner metatarsal tubercle short-
er (vs equal to or longer) than the first toe and
1040 COPEIA, 2002, NO. 4
Fig. 3. Dorsal (above) and lateral (below) views of
head of holotype of Kaloula walteri (male PNM 6725;
head length
5
7.6 mm).
←
Fig. 2. Palmar view of right manus (above; ML 8.1
mm) and plantar view of right pes (below; PL 12.9
mm) of male holotype (PNM 6725) of Kaloula walteri.
by its unusual advertisement call, consisting of
a soft chain of single snapping pulses (vs long,
loud, low-frequency trilled croaks; Table 1).
Kaloula walteri differs from K. kalingensis,K.
kokacii,K. baleata,K. c. conjuncta,K. c. negrosensis,
K. c. meridionalis, and K. c. stickeli by nonexpan-
sion of terminal digital disks (vs terminal disks
moderately to widely expanded). The new spe-
cies also differs from K. kalingensis,K. kokacii,
and K. baleata by the presence of stratified (light
above, dark below midline) coloration on the
posterior surface of the thigh and flanks (vs col-
or homogeneous or mottled), absence (vs pres-
ence) of a light pericloacal ring (reduced to
light patches on either side of the vent in some
K. baleata), and presence (vs absence) of super-
1041DIESMOS ET AL.—NEW PHILIPPINE KALOULA
T
ABLE
1. C
OMPARISONS OF
Q
UALITATIVE AND
Q
UANTITATIVE
D
IAGNOSTIC
C
ALL
C
HARACTERS IN
F
OUR
S
PECIES OF
P
HILIPPINE
Kaloula: Kaloula walteri, Kaloula picta, Kaloula
kalingensis,
AND
Kaloula conjuncta conjuncta.
Character K. walteri
(26 C) K. picta
(25 C) K. kalingensis
(25–27 C) K. c. conjuncta
(24 C)
Overall call structure slow snapping
pulse train
trilled call
(many brief pulses)
single tonal frequency
sweep
repeated croaks
Note/call
Call rate
Call duration
Pulse rate
44–198
2.2/sec
20–90 sec
0.002.2/ms
32–37
1.2/sec
290 ms
0.15/ms
1
1–3/min
155–200 ms
—
30–180
1–2/min
30–50 sec
6.1–6.3/ms
Pulse duration
Pulse rise time
Interpulse interval
14–17 ms
2.0–2.2 ms
390–496 ms
4.5–7.9 ms
0.9–1.1 ms
5.8–6.2 ms
—
—
—
27.1–48.1 ms
0.5–1.0 ms
111.1–141.1
Dominant frequency peak
5
2.2 kHz
(2nd harmonic)
peak
5
2.0 kHz
(2nd harmonic)
peak
5
2.2–1.8 kHz sweep
(3rd–4th harmonic)
peak
5
2.3–2.4 kHz
# of component frequencies
Dom. frequency modulated?
5–7
no
4
no
7–8
yes
3–5
no
numerary tubercles at the base of each digit of
the manus. Kaloula walteri also differs from K.
kokacii and K. baleata by a much smaller body
size (Appendix 1) and by its unique advertise-
ment call (Table 1). The new species also differs
from K. c. conjuncta,K. c. negrosensis,K. c. stickeli,
and K. c. meridionalis by the absence of dorsal
tubercles or limitation of tubercles to the sacral
region (vs tubercles distributed over the entire
dorsum), limitation of the ventral epidermal ad-
hesive glands to the sternal region (vs glands
covering anterior two-thirds to three quarters of
venter), and limitation of webbing of pes to the
level of the proximal subarticular tubercles of
all digits in males (vs webbing extending to at
least the level of the distal subarticular tuber-
cle). The new species also differs from K. c. ne-
grosensis,K. c. stickeli, and K. c. meridionalis by a
markedly smaller body size (Appendix 1).
Description of holotype.—A mature male; habitus
rotund; head proportionally small and not dis-
tinct from neck, length 26% of SVL, 81% of
head width; snout slightly pointed beyond lower
jaw, its tip square in dorsal aspect, pointed and
ventrally sloping in lateral aspect; eyes protrud-
ing laterally beyond silhouette of head in dorsal
aspect, protruding only slightly beyond dorsal
surface of head in lateral aspect; pupil subcir-
cular; interorbital region flat; eye diameter 59%
of interorbital distance; canthus rostralis con-
cave and swollen; loreal region sloping, mod-
erately concave; eye diameter 61% of snout
length; eye diameter 73% of eye–narial dis-
tance; labial region not flared or swollen; nos-
trils barely protuberant; eye–nostril distance
much longer than distance from nostril to tip
of snout; internarial region slightly convex; tym-
panum visible under skin, not exposed, its an-
nulus 82% of eye diameter; dorsal edge of tym-
panum partially concealed by supratympanic
fold extending from posterior corner of eye to
supra-axillary region; tongue oval, elongate,
lacking posterior notch; choanae round, very
small, situated at anterolateral edge of palate,
separated by a distance equal to or slightly larg-
er than their diameter, not obscured by palatal
shelf; dentigerous process of vomer evident but
vomerine teeth apparently absent; vocal slits
large, just posterior to rictus.
Skin of dorsal surfaces of body, head, and
limbs smooth; ventral surfaces of trunk and
limbs smooth; ventral surface of throat slightly
granulated; loose skin overlying median subgu-
lar vocal sac forms slight sternal fold; ventral
epidermal adhesive gland occupies sternal re-
gion and area immediately posterior; limbs, ma-
nus and pes well developed, manus length 63%
1042 COPEIA, 2002, NO. 4
of pes length; pes length 97% of tibia length;
tibia 95% of femur length; tibia length 46% of
snout–vent length; digits of manus slender, ter-
minal disks slightly expanded and swollen; de-
creasing digit length (longest to shortest when
adpressed) 3, 2
5
4, 1; one subarticular tuber-
cles on digits I–II, two under digits III–IV; su-
pernumerary tubercles present at the base of
each figure; thenar (inner metacarpal) tubercle
oval, elongate, smaller than subarticular tuber-
cle of digits; palmar (outer metacarpal) tubercle
large, flat, nearly divided; nuptial pads absent,
forearms not hypertrophied.
Tarsus smooth, lacking folds; plantar surfaces
of pes smooth, with well-developed subarticular
tubercles; decreasing digit length (longest to
shortest when adpressed) 4, 3, 5, 2, 1; one sub-
articular tubercle on digits I–II; two on digit III;
three on digit IV; minute outer metatarsal tu-
bercle barely evident; inner metatarsal tubercle
prominent, with sharp outer edge, about two-
thirds length of toe I; digits of pes almost com-
pletely free of web; fleshy basal toe webbing
barely evident, and extending to the level of the
proximal subarticular tubercles among digits
III–V but not extending to the level of these
tubercles among digits I–III.
Measurements of holotype (in millimeters).—SVL
29.2; HL 7.6; HW 9.4; SL 3.6; IND 2.6; IOD 3.7;
ED 2.2; END 1.6; TD 1.8; ETD 0.54; FL 14.0;
TBL 13.3; TSL 7.5; RL 6.5; Fin3L 4.5; Fin3DW
0.9; Fin3PPW 0.8; Toe4L 7.0; Toe4DW 0.8; PL
12.9; ML 8.1.
Color in preservative.—Dorsum generally dark
brown; marbled with very dark brown, darker
central region and black blotches in interorbit-
al, interscapular, sacral, and supracloacal re-
gions; lateral surfaces of body with stratified col-
or pattern of dark lavender below and dark red-
dish brown above; separated by thin pale line
that runs from the nares, across the canthus and
lateral edge of palpebrum, diagonally across the
flanks and posteriorly to groin; similar color
pattern (dark below, light above) on anterior
and posterior surfaces of the femur separated
by a light tan line; lateral surfaces of rostrum,
tympanic region, and supraaxial regions dark
lavender to charcoal gray; dorsal surfaces of
forelimbs dark brown with charcoal gray blotch
on joint of humerus and forearm; two distinct
dark charcoal gray blotches on forearm; dorsal
surfaces of digits dark brown with charcoal gray
and tan flecks; dorsal surfaces of femur with
one dark brown to charcoal gray crossbar con-
tinuous (when specimens are in a resting posi-
tion) with similar bars on tibial segments; tarsi
light brown, with small dark brown blotches and
solid very dark brown on the posterior (supra-
plantar) surfaces; three small black spots on
skin overlying sacral region; venter and ventral
surfaces of limbs reddish brown with distinct
pale yellowish cream spots with irregular bor-
ders; epidermal adherence gland visible as ligh-
ter orangish-brown patch in the sternal region;
gular region black; palmar surface of the manus
dark brown; plantar surface of the pes black;
webbing at base of toes gray with small black
spots; dorsal surfaces of terminal digital disks
dark brown, ventral surfaces light cream.
Coloration in life.—(from field notes of ACD)
Dorsum reddish brown with chocolate brown
marbling to dark gray with very dark brown
marbling; stratified lateral color pattern sepa-
rated by bright thin white lines; venter light yel-
lowish brown with bright white spots; throat
very dark brown to black; groin and flanks with
minute white flecks; iris metallic brown.
Variation.—Our sample consists of 21 males and
one female. Based on this modest sample, no
obvious color pattern or body size differences
exist between the sexes. Intersexual variation in
degree of webbing of the foot (documented in
other Philippine Kaloula; R. Crombie, pers.
comm.) is not apparent given our small sample.
Four specimens lack outer metatarsal tuber-
cles (CAS 211741, PNM 6662, 6724–25); in the
remainder, these tubercles are reduced and
barely perceptible under high magnification.
Dorsal ground color reddish brown, gray
brown, dark lavender, or light gray; dorsum of
17 specimens generally marbled, with darker
central regions composed of dark brown to
charcoal gray blotches in interorbital, interscap-
ular, sacral, and supraanal regions; remaining
four specimens are homogeneous charcoal gray,
dark lavender, or reddish brown; specimens pos-
sessing marbled dorsal pattern also possess lat-
erally stratified color patterns of dark lavender
to charcoal gray below and reddish brown, gray
brown, or light gray above thin pale lateral line;
three specimens lack dorsal marbled color pat-
tern and also lack laterally stratified color pat-
terns; these four specimens (Mt. Isarog: PNM
6726; Mt. Banahao: TNHC 59666, 61117; CAS
211742) possess stratified color patterns on pos-
terior segments of hindlimbs; tarsi reddish
brown to dark lavender, with small dark brown
to black spots or solid dark lavender to charcoal
gray coloration on the posterior (supra-plantar)
surfaces; eight specimens with 2–12 (mean
5
6.8
6
3.7 SD) black spots on skin overlying sa-
cral region; venter and ventral surfaces of limbs
1043DIESMOS ET AL.—NEW PHILIPPINE KALOULA
Fig. 4. Distribution of Kaloula walteri on Mt. Ba-
nahao (Banahaw) at the extreme southern end of the
Sierra Madre mountain range (stippled) of Luzon Is-
land (and Polillo Island; left) and details of type lo-
cality in the Municipality of Tayabas (inset; right).
Known collection localities include (1) Municipality
of Lucban, Mt. Banahao, (2) Municipality of Tayabas
(
5
type locality), Mt. Banahao, (3) Watershed Re-
serve Area, Polillo Island, (4) Mt. Isarog, and (5) Mt.
Mayon.
Fig. 5. Typical appearance of habitat at type lo-
cality (Municipality of Tayabas, Mt. Banahao, Luzon
Island, Philippines) of Kaloula walteri: stream bed with
stationary pools (above) and small pool with over-
hanging vegetation where the first specimens were
collected (below).
light reddish to orangish brown with distinct
pale yellowish to white spots; one specimen
(PNM 6664) with very pale gray venter with
cream spots; males with very dark gray to black
throat; female with reddish brown throat. The
Polillo Island specimen (PNM 6723) is some-
what lighter in coloration than others but oth-
erwise is indistinguishable.
Measurements of males (in millimeters; n
5
21).—
SVL 24.5–32.3 (mean
5
29.1 mm
6
2.0 SD);
HL 6.4–9.2 (mean
5
7.9
6
0.7); HW 7.4–10.5
(mean
5
9.3
6
1.0); SL 3.0–3.7 (mean
5
3.4
6
0.3); IND 1.9–2.6 (mean
5
2.3
6
0.2); IOD 2.6–
3.9 (mean
5
3.2
6
0.4); ED 1.7–2.7 (mean
5
2.3
6
0.3); END 1.4–1.9 (mean
5
1.6
6
0.2);
TD 1.2–1.9 (mean
5
1.6
6
0.2); ETD 0.4–0.8
(mean
5
0.6
6
0.1); FL 11.9–14.8 (mean
5
13.4
6
0.9); TBL 11.3–14.3 (mean
5
12.8
6
0.9);
TSL 6.7–8.5 (mean
5
7.5
6
0.4); RL 6.0–7.4
(mean
5
6.7
6
0.5); Fin3L 3.6–5.0 (mean
5
4.2
6
0.4); Fin3DW 0.5–0.9 (mean
5
0.8
6
0.1);
Fin3PPW 0.4–0.8 (mean
5
0.8
6
0.1); Toe4L
6.3–7.9 (mean
5
7.0
6
0.5); Toe4DW 0.6–1.0
(mean
5
0.8
6
0.1); PL 11.9–14.8 (mean
5
13.4
6
0.9); ML 7.3–9.0 (mean
5
8.2
6
0.5).
Measurements of females.—SVL 31.4; HL 7.6; HW
9.8; SL 3.1; IND 2.1; IOD 2.6; ED 2.7; END 1.7;
TD 1.2; ETD 0.6; FL 13.7; TBL 13.5; TSL 7.8;
RL 7.4; Fin3L 4.4; Fin3DW 0.8; Fin3PPW 0.7;
Toe4L 7.6; Toe4DW 0.8; PL 13.5; ML 8.4.
Distribution.—The new species is known from
Mt. Banahao (S. central Luzon Isl.), Mt. Isarog
and Mt. Mayon (of the Bicol Peninsula; P
HI
-
VOLCS
, 1990; Diesmos, 1998) and from Polillo
Isl. (Fig. 4).
Ecology, habitat, and life history.—Specimens of K.
walteri have been observed or collected in ripar-
ian habitats ranging from lowland dipterocarp
forest to midmontane closed-canopy rain forest
(sensu Whitmore, 1984). On Mt. Banahao, the
new species has been observed in habitats at el-
evations of 650–950 m ranging from small (
5
3
m wide) dry stream beds to larger rivers (
$
6
m wide) during the dry season (obser ved in
March and April) when water levels decline and
large stationary pools form (Fig. 5). The single
specimen from Mt. Isarog was collected at an
elevation of 990 m, and Mt. Mayon specimens
were collected at 650 m. The specimen from
Polillo was collected at approximately 25 m
above sea level in well-regenerated secondary
forest. Specimens at this locality have been
heard calling near clearings and even on edges
of agricultural areas adjacent to forest (V.
Yngente, pers. comm.).
Kaloula walteri is a burrowing species; when
1044 COPEIA, 2002, NO. 4
Fig. 6. The advertisement call of Kaloula walteri.
(A) Fast Fourier Transformation (FFT; frequency
spectrum: relative amplitude vs frequency, in kHz),
(B) expanded waveform (relative amplitude vs time
in ms), (C) audiospectrogram (sonogram: frequency
in kHz vs time in ms), and (D) oscillogram (relative
amplitude vs time in ms). The FFT was calculated at
the second pulse in this figure.
disturbed, two individuals were observed mak-
ing vigorous digging motions of the hind limbs
in attempt to burrow backward into gravel
stream beds near small stationary pools (ACD,
pers. obs.). Males call from under rocks, beside
small pools, and under vegetation or debris.
Calling males were observed in small choruses
of 2–6 individuals per aggregation. Chorus ac-
tivity was arranged in bouts of calling and si-
lence, usually with a single male initiating a cho-
rus, to be joined by 1–5 individuals; the chorus
persists for 30–90 sec and rapidly declines after
which a period of 60–180 sec persists until a sin-
gle male initiates a new bout of calling. During
vocalization, initial pulses are produced without
inflation of the vocal sac. After several pulses,
vocal sacs extend fully and subsequent pulses
are produced by oscillation of the vocal sac, be-
tween partially and fully inflated. Frogs were not
observed nor heard during nights immediately
following rain when streams were rapidly run-
ning and loudly cascading. Calling males were
only recorded when streams had declined to a
point when stationary pools were present and
ambient noise levels had decreased. On several
instances, choruses were heard in late after-
noon hours (1430–1800 h). Despite extensive
searches at the type locality and several nights
attending choruses, females of K. walteri have
not been collected or observed on Mt. Bana-
hao. The single specimen from Mt. Isarog that
we refer to this species is a female with mature
oviducts but without maturing eggs. We have no
definitive data on the site of egg deposition or
the nature of larval development, but pools of
stagnant water near calling male K. walteri con-
tained small dark brown tadpoles (not collect-
ed) that may either belong to this species or the
sympatric K. conjuncta conjuncta.
Other sympatric species of anurans include
Limnonectes macrocephalus,Limnonectes woodwort-
hi,Rana luzonensis,Rana similis,Philautus surdus,
Platymantis banahao,Platymantis corrugatus,Platy-
mantis dorsalis,Platymantis luzonensis,Platymantis
montanus,Platymantis indeprensus, and Platyman-
tis pseudodorsalis. Other amphibians known from
Mt. Banahao (Diesmos, 1998; unpubl. data),
that we have not found at the specific type lo-
cality, include Occidozyga laevis,Rana erythraea,
Polypedates leucomystax,Rhacophorus bimaculatus,
Rhacophorus pardalis, and two additional undes-
cribed species of Platymantis. On Mt. Isarog, the
new species also is sympatric with Platymantis isa-
rog (Brown et al., 1997), and an undescribed
species of Platymantis (unpubl. data) and on Po-
lillo Island, Rhacophorus appendiculatus,Platyman-
tis polillensis,Platymantis sp. (of uncertain taxo-
nomic status, possibly conspecific with P. luzo-
nensis) and Rana (
5
Fejervar ya)vittigera also
have been recorded sympatrically (Hampson,
1999a,b; ACD and RMB, pers. obs.).
Descriptions of advertisement calls of Kaloula walteri
and some related species.—In this section we pro-
vide brief descriptions of the advertisement calls
of Kaloula walteri,K. picta,K. kalingensis, and K.
c. conjuncta, and we comment on the calls of K.
rigida and K. kokacii. Because at present there
are so few quality recordings of these species,
detailed quantitative analyses of these calls are
not possible. We expect that further studies will
require reconsideration of the advertisement
calls of these species because the known range
of interspecific variation likely will expand with
the collection of further data and the clarifica-
tion of species boundaries (see Discussion).
Nevertheless, a comparison of the gross struc-
ture of the calls is useful at present and contrib-
utes to the diagnosis of the new species.
The advertisement call of K. walteri (recorded
at the type locality) is presented in Figure 6. We
have recorded K. walteri on five occasions on
Mt. Banahao and descriptions of the species’
calls on Polillo Isl. (courtesy of V. Yngente, pers.
comm.) qualitatively match those at the type lo-
cality. To the human ear, the call sounds like
‘‘tuk...tuk...tuk...,’’ resembling the sound
produced by snapping fingers or slowly knock-
ing together two fist-sized stones. This resem-
blance is so strong that we have initiated chorus
activity by tapping together stones on several oc-
casions.
The call consists of series of slowly delivered
1045DIESMOS ET AL.—NEW PHILIPPINE KALOULA
Fig. 7. The advertisement call of Kaloula picta.
Parts A–D and definitions as in Figure 6. The FFT was
calculated at midcall (210 ms).
snapping notes (
5
pulses) delivered intermit-
tently, with no discernible temporal structure
(i.e., each note consists of just a single pulse)
and no modulation of pulse amplitude within
the call. Call length varies from 20–90 (mean
5
65.8
6
28.6 SD; n
5
5) sec and intercall interval
ranged from 60–180 sec (
5
127.5
6
50.4 SD; n
5
4).
The following description is based on one
high-quality call recording (length of 30 sec),
recorded by ACD at the type locality on 6 May,
1995 (2200 h; 25.5 C). Pulse rate [(total num-
ber of pulses
2
1)/time from beginning of first
pulse to beginning of last] for 30 sec of contin-
uous calling is 2.2 pulses/sec, and interpulse in-
terval ranges from 390 to 496 ms. Pulse dura-
tion varies from 14 to 17 ms and consists of a
rapid rise (2.0–2.2 ms) in amplitude and a
slightly longer decline to ambient sound levels
(3.7–4.1 ms). Although the distinction of clear
harmonics was difficult because of the transient
nature of the pulse, Fast Fourier Transforma-
tion (FFT) did allow for some generalizations
to be made with respect to the component fre-
quencies of the call. Each pulse had a range of
distinct frequency components, with marked
energy in the fundamental frequency and dis-
tinct energy peaks at four apparent multiples
(harmonics) of the fundamental. The funda-
mental frequency peaked at 1.2 kHz, the em-
phasized frequency peaked at 2.2–2.3 kHz, and
subsequent frequency components occurred at
3.6–3.8, 4.7–4.9, 6.0–6.3, and 7.7–7.9 kHz re-
spectively. Thus, although each pulse spanned a
frequency range of 0.11 Hz to 7.9 kHz, the ma-
jority of the energy in the call was located be-
tween 1.3 and 2.8 (peak frequency
5
2.2–2.3)
kHz.
Although calls of K. rigida are not currently
available for analysis, we have heard this species
in the type locality (Barangay Balbalan, Munic-
ipality of Balbalan, Kalinga Prov., Luzon Isl.). Its
call is a moderately rapid train of brief pulses
and sounds to the human ear like a small en-
gine idling.
Kaloula picta: This species calls in large cho-
ruses (sometimes hundreds of individuals) from
flooded rice fields and puddles in disturbed ar-
eas following heavy rains. Although the call
structure of K. picta (recorded in Antique Prov-
ince, Panay island; 25 C; 1–2 m; Fig. 7) is dras-
tically different from K. walteri, some similarities
in individual pulses warrant discussion. The call
of K. picta sounds to the human ear like a drawn
out ‘‘whhaaaak, whhaaaak, whhaaaak.’’ In cho-
ruses of many individuals of different body sizes
(and, thus, calls of different dominant frequen-
cies), nearby individuals alternate while calling,
which produces the onomatopoeic sound ‘‘ko-
kak, ko-kak, ko-kak’’ (‘‘Ko-kak,’’ or a similar var-
iation, is the common name for ‘‘frog’’ in many
Philippine provinces). Overall call structure (22
calls analyzed for one individual) is quite differ-
ent from that of K. walteri, with calls consisting
of 32–37 distinct pulses arranged in a trilled
croak (Fig. 7), delivered at 1.2 calls/sec (call
rate calculated as in pulse rate, above). Pulse
rate within each call is 0.15 pulses/ms and pulse
duration varies from 4.5 to 7.9 ms. Amplitude
modulation of the entire call occurs as a gradual
increase in pulse amplitude from 20–100% rel-
ative amplitude during the first 135 ms (
5
14–
16 pulses) a leveling off of amplitude at appar-
ent maximum for 120 ms (
5
14–16 pulses),
then a more rapid decline in amplitude to am-
bient levels over a period of 40–45 ms (
5
4–6
pulses). Only at the beginning (between first
10) and end (between last 10 pulses) of the call
are there discernible interpulse intervals, these
lasting from 5.8 to 6.2 ms.
As in K. walteri, individual pulses consist of a
rapid (0.9–1.1 ms) onset and increase to peak
amplitude, then a more gradual (4.5–5.7 ms)
decline to ambient levels. The Fast Fourier
Transformation revealed that the vast majority
of the energy of the call is packaged between
two frequency intervals, the first (fundamental)
between 0.6 and 1.7 kHz (peak frequency ap-
proximately 1.2 kHz) and the second (domi-
nant) between 1.7 and 3.8 kHz (peak of 2.2
kHz). Although the FFT (Fig. 7) did not resolve
higher frequency components of the call, the
audiospectrogram revealed the presence of at
least two faint harmonic bands at roughly 6 and
8 kHz, respectively. Thus, although the call
structure of K. picta and K. walteri appeared very
different (K. picta calls consist of 32–37 rapidly
1046 COPEIA, 2002, NO. 4
Fig. 8. The advertisement call of Kaloula kalingen-
sis. Parts A–D and definitions as in Figure 6. The FFT
was calculated at 200 ms and background noise evi-
dent in call is wind and rain.
Fig. 9. The advertisement call of Kaloula conjuncta
conjuncta. Parts A–D and definitions as in Figure 6.
Shown are the alternating pulse trains of two calling
males. The FFT was calculated in the middle of the
first pulse in this figure.
delivered, amplitude modulated pulses and K.
walteri calls consist of a series of slowly delivered
pulses with no amplitude modulation detected
across pulses) individual pulses were remarkably
similar and short in duration, with most energy
in the emphasized frequency (the second har-
monic) of around 2.1–2.3 kHz.
Kaloula kalingensis: The advertisement call of
K. kalingensis (recorded in the type locality: Bal-
balan, Kalinga-Apayao Province; 25–27 C; 0.5 m;
Fig. 8) are drastically different in components,
sound, and structure from the other three spe-
cies considered here. The call is a single loud
note and varies in the way it sounds to the hu-
man ear, ranging from ‘‘Bwop!’’ to ‘‘Eeow!’’ Ka-
loula kalingensis calls in loud tonal notes with a
slight decreasing frequency sweep. This species
calls from tree holes in forest trees and occa-
sionally from bamboo stands in slightly dis-
turbed areas adjacent to forested regions. Single
note calls are delivered at a rate (not quanti-
fied), qualitatively described by the recorder
(RMB) as 1–3/min. The waveform (Fig. 8) de-
picts an extremely rapid rise from ambient to
near peak amplitude in 1.8 ms; the rise from
ambient to peak amplitude for the call’s entirety
occurs from 28 to 37 ms. The call duration for
eight calls recorded over 15 min for one indi-
vidual ranges from 155 to 250 ms and exhibits
erratic amplitude modulation with no apparent
temporal pattern (Fig. 8). Call structure in-
cludes a number of harmonically related fre-
quency sweeps, with most energy in the forth
(emphasized) harmonic. The dominant fre-
quency swept in a shallow concave fashion from
2.2 to 1.8 kHz over a period of 50 to 60 ms. The
Fast Fourier Transformation (just after call on-
set, i.e., 200 ms) revealed four distinct frequen-
cy components harmonically relating to the
forth (dominant) frequency, and then possibly
three or four less distinct component harmon-
ics. The first four distinct frequency compo-
nents had peak frequencies of approximately
0.5, 0.9, 1.4, and 1.9 kHz. Less distinct frequen-
cy peaks were detected at 2.6, 3.3, and 3.8 kHz,
respectively. The similar species Kaloula kokacii
(Ross and Gonzales, 1991) has recently been
collected on the Bicol Peninsula of Luzon Is-
land (unpubl. data and R. Sison, pers. comm.)
and the advertisement call has been heard (by
ACD and RMB) but not recorded. The calls of
this species are similar to those of K. kalingensis
in that they consist of loud descending tonal
honks. They differ by a much more rapid call-
ing rate, and sound to the human ear like
‘‘Auk. . . auk. . . auk. . . auk. . . auk’’ (pers. obs.).
Kaloula conjuncta conjuncta: This species calls
singly or in small choruses (
#
10 individuals)
in quiet pools of accumulated rainwater at the
beginning of the rainy season ( July–August) in
montane forested regions on S. Luzon. We have
also observed large choruses in flooded fields
(adjacent to forest) following heavy rains in Al-
bay Province, S. Luzon. We recorded one pair
calling together with pulses alternating for 90
sec (Fig. 9). The advertisement call of K. c. con-
juncta (recorded on Mt. Banahao; 24–26 C)
consists of a long train of croaking pulses pro-
duced continuously for 30–180 sec (mean
5
95.5
6
68 SD). To the human ear, the call
sounds like a continuous series of grunts:
‘‘Errr...rrr...Errr....’’ Pulse duration varied
from 27.1–33.4 ms (mean
5
30.8
6
2.0 SD; n
5
25) for one individual and 34.3–48.1 ms
(mean
5
40.8
6
4.3; n
5
25) for another. In-
terpulse intervals ranged from 133.4–141.1
1047DIESMOS ET AL.—NEW PHILIPPINE KALOULA
(mean
5
136.3
6
2.9 SD; n
5
24) and 111.1–
140.3 ms (mean
5
129.3
6
6.5; n
5
24), re-
spectively. Pulse rate for these two same individ-
uals was 6.1 pulses/sec and 6.3 pulses/sec for
35 and 50 sec of continuous calling. Individual
pulses were amplitude modulated, with 4–6 sub-
pulses (Fig. 9). Relative amplitude of each sub-
pulse increased throughout the pulse with the
first subpulse approximately 60% of maximum
amplitude observed in the ultimate or penulti-
mate subpulse (Fig. 9). Each subpulse consisted
of an instantaneous (0.5–1 ms) rise to peak am-
plitude and then a slower (3–4 ms) decline to
near ambient levels before the onset of the next
subpulse. The Fast Fourier Transformation in-
dicates that the dominant frequency lies be-
tween 2.3 and 2.4 kHz and is flanked above (to
4 kHz) and below (to 1 kHz) by a near sym-
metrical range of component frequencies.
D
ISCUSSION
Local folklore in the municipality of Tayabas
at the base of Mt. Banahao maintains that
troupes of macaques descend from the forest at
night, reportedly to play in river beds after dark.
Descriptions of their behavior by resident hunt-
ers and Tayabas mountaineers includes anec-
dotes of monkeys tapping rocks together in co-
ordinated group efforts at night ( J. Redor, pers.
comm.). We assume that the distinctive call of
K. walteri is the basis of this belief. Frequent
sightings of macaques in and around the type
locality ( J. Redor, pers. comm., and pers. obs.)
and the unique sound of the advertisement call
supports this interpretation and suggest how
the association of these might be the basis of
this common belief.
The description of K. walteri brings the num-
ber of Kaloula species currently recognized in
the Philippines to seven. More than half of the
known species of the genus Kaloula occur in the
Philippines (Frost, 1985, 2000; Duellman,
1993), and the recent descriptions of K. kokacii
and K. walteri further emphasize the level of di-
versity and endemism in Philippine microhylids.
Nevertheless, we consider seven species to be a
clear underestimation of Kaloula species diver-
sity in the archipelago, and we note the follow-
ing unresolved taxonomic issues. First, the sta-
tus of Inger’s (1954) subspecies of K. conjuncta
needs to be fully reconsidered within the con-
text of a lineage-based species concept (sensu
Frost and Hillis, 1990; de Queiroz, 1998, 1999).
Most of Inger’s (1954) subspecies of Kaloula are
characterized by nonintegrating diagnostic
characters of external morphology, advertise-
ment call, ecology, body size, and insular allop-
atry on separate Pleistocene aggregate island
platforms (sensu Heaney, 1986; see also Brown
and Diesmos, 2001), suggesting that some sub-
species may be full evolutionary lineages worthy
of specific rank (Frost and Hillis, 1990; de Quei-
roz, 1998, 1999). Second, preliminary work with
newly collected material (ACD and RMB, un-
publ. data) suggests that patterns of montane
endemism on Luzon may require the further
subdivision of both K. c. conjuncta and K. kalin-
gensis. Third, the taxonomic status of K.‘‘balea-
ta’’ populations on Palawan Island is poorly de-
fined and based on only a few specimens in ma-
jor museum collections. We doubt that this for-
est species is conspecific with true K. baleata
frequently encountered in disturbed and agri-
cultural areas of the Sunda Shelf islands of Java,
Bali, Sumatra, and Borneo (Inger, 1966; Iskan-
dar, 1998; F. Yuwono, pers. comm.). Fourth, In-
ger (1954) remarked on the patterns of varia-
tion in K. picta that, at the time of his review,
did not, in his opinion, warrant taxonomic re-
vision. Reconsideration of these populations
with additional data may reveal patterns of is-
land endemism and evidence of diversification
that would require taxonomic recognition.
Fifth, Gaulke (2001a–c) and Ferner et al. (2001)
have noted the presence of a population related
to K. kalingensis on Panay Island and recent sur-
veys suggest that Negros Island may support a
related population as well (C. N. Dolino, ACD,
and RMB, pers. obs.); we doubt that the Visayan
Aggregate Island Complex population is con-
specific with K. kalingensis of Luzon (for discus-
sion, see Brown and Diesmos, 2001), and we
suspect that these Visayan populations repre-
sent an additional undescribed species of Kalou-
la. Sixth, unpublished data suggest the presence
of at least two additional undescribed species of
Kaloula that appear to be residents of lesser in-
sular centers of Philippine endemism (Heaney,
1986; Brown and Diesmos, 2001) and may soon
be recognized as single, small island endemics.
Finally, disjunct distribution of frogs referred to
K. rigida (which have been collected in the Cen-
tral Cordillera and the Sierra Madre mountain
ranges) requires careful consideration before
definite statements can be made regarding the
true identity of these allopatric populations. All
of these additional new species definitions await
a comprehensive taxonomic review of Philip-
pine Kaloula already in progress (ACD and
RMB, unpubl. data).
The curious disjunct distribution of K. walteri
on Mt. Banahao, Mt. Isarog, Mt. Mayon and Po-
lillo Island (Fig. 4) implies that the new species
is likely to be found between these localities if
suitable forested habitat still exists. Accordingly,
1048 COPEIA, 2002, NO. 4
future surveys should sample montane habitats
on Mt. Labo, Mt. Cadig, Mt. Malinao, and other
forested areas of southern Quezon, Camarines
Norte, Camarines Sur and Albay Provinces
(PHIVOLOCS, 1990). Other recent studies
have demonstrated the presence of consider-
able levels of montane endemism on the moun-
tains of southern Luzon (Brown et al., 1996,
2000b; Diesmos, 1998).
The results of this study contribute to a grow-
ing appreciation of the diversity (Brown et al.,
1996, 2000a, 2000b) and zoogeographic com-
plexity (Brown and Alcala, 1970; Auffenberg,
1988; Diesmos, 1998) of the amphibian and rep-
tile fauna of Luzon island (see also Ross and
Gonzales, 1991; Brown et al., 1999; Alcala et al.,
1998). The full appreciation of that diversity is
challenged by at least partial continuing accep-
tance of a taxonomy based on antiquated spe-
cies concepts (Inger, 1954; Alcala and Brown,
1998; Brown and Diesmos, 2001), a lack of sur-
vey data for the mountains of Luzon (Brown et
al., 1996, 2000b), the near complete and con-
tinuing removal of Luzon’s primary forests
(Collins et al., 1991; Kummer, 1992; Heaney
and Regalado, 1998) and by current restrictions
placed on biodiversity specialists working in the
Philippines (La Vin˜a et al., 1997). We suspect
that current taxonomy reflects a gross under-
estimation of the diversity and endemism of
Philippine amphibians and that this ignorance
regarding the magnitude of Philippine biodi-
versity in part contributes to the destructive ex-
ploitation of the country’s remaining rain for-
ests. Great strides could be made toward the
conservation of Philippine biodiversity by im-
posing effective limits on further deforestation,
empowering local communities to sustainably
manage natural resources and by lifting current
bureaucratic obstructions to biodiversity re-
search.
S
PECIMENS
E
XAMINED
Kaloula cf baleata: Philippines, Palawan Island,
Palawan Prov., about 1.5 km south-southwest of
Iwahig: CAS-SU 21770; Cagayan Island, Palawan
Prov.: CAS 157186.
Kaloula kalingensis: Philippines, Luzon Island,
Kalinga Prov., Municipality of Balbalan: CAS
61462 (Holotype); Kalinga Prov., Municipality
of Balbalan, Barangay Sadsadan, Mt. Nagtud:
TNHC 59646, 59647; Quezon Prov., Municipal-
ity of Lucban, northeast slope of Mt. Banahao:
CAS 201203 PNM 5063; Quezon Prov., Munici-
pality of Dolores, Barangay Kinabuhayan, Mt.
San Cristobal: TNHC 59645; Zambales Prov.,
Municipality of Masinloc, Barangay Coto, south
slope of Mt. Apoy: PNM 2417.
Kaloula kokacii: Philippines, Luzon Island,
Sorsogon Prov., Municipality of Irosin, Mt. Bu-
lusan: TNHC 59648, PNM 5062; Camarines
Norte Prov., San Lorenzo Ruiz, Mt. Labo: PNM
2137; Catanduanes Isl., Catanduanes Prov., Mu-
nicipality of Gigmoto, Borodan summit: PNM
2043 (Holotype).
Kaloula conjuncta conjuncta: Philippines, Poli-
llo Island, Burdeos: CAS 62445–62447; Luzon
Island, Laguna Prov., Municipality of Los Ban˜os,
Barangay Batong Malaki, University of the Phil-
ippines, College of Forestry campus: TNHC
59626; Quezon Prov., Municipality of Tayabas,
Barangay Camaysa, Mt. Banahao: TNHC 59627,
59628.
Kaloula conjuncta meridionalis: Philippines,
Mindanao Island, Surigao Prov.: CAS-SU 16166–
16168; ‘‘Mindanao Institute of Technology’’:
CAS 122190; ‘‘North Mindanao’’: CAS 66208;
Lanao Prov., Dansalan: CAS-SU 14307; Davao
del Sur Prov., Municipality of Toril, Barangay
Upper Baracatan, Sitio San Roque: TNHC
59629–59643.
Kaloula conjuncta negrosensis: Philippines, Ne-
gros Island, Negros Oriental Prov., Dumaguete:
CAS-SU 14168–14173; Negros Oriental Prov., 5
km west of Municipality of Valencia, ridge north
of Maite River: CAS-SU 15773–15775, 16195–
16198, 16200–16209; Negros Oriental Prov.,
pool at Camp Tas-Tubig: 139242, 139243,
139246–139249; Panay Island, Antique Prov.,
Municipality of Valderrama, Barangay Lublub,
base of Mt. Baloy: TNHC 56340–46; Panay Is-
land, Antique Prov., Municipality of Culasi, Ba-
rangay Alojipan: PNM 2555.
Kaloula conjuncta stickeli: Philippines, Leyte Is-
land, Leyte Prov. Municipality of Carigara, Ca-
rigara City: FMNH 60786 (holotype).
Kaloula picta: Philippines, Luzon Island, Nue-
va Vizcaya Prov., Municipality of Bayombong:
CAS-SU 9842, 9848, 9850; Zambales Prov., vicin-
ity of Subic Bay: CAS 15631; Batangas Prov.,
Camp McGrath: CAS 15731; Laguna Province,
Municipality of Los Ban˜os: CAS 61056, 61057;
Rizal Prov., Manila: CAS 61634–61638, 61653;
Polillo Island: CAS 62257; Cebu Island, 1 km
west of Tisa, Guianoran River: CAS-SU 16175,
16181–82, 16184, 16187–88; Samar Island, West-
ern Samar Prov., Calbayog City, Barangay Ranis:
TNHC 54982; Polillo Island, Quezon Province,
Municipality of Polillo, Barangay Sibucan, Sitio
Tambangin: TNHC 54983, 54984; Bohol Island,
Bohol Province, Municipality of Bilar, Barangay
Riverside: TNHC 56368–56375, 56430–56432;
Cebu Island, Cebu Prov., Municipality of Cebu
City, Barangay Talamban, Sitio Pulang Bato:
1049DIESMOS ET AL.—NEW PHILIPPINE KALOULA
TNHC 56468; Mindanao Island, Davao del Sur
Prov., Municipality of Toril, Barangay Upper Ba-
racatan, Sitio San Roque: TNHC 59649–59665,
59871; Davao City, Municipality of Calinan, Ma-
lagos Eagle Station: TNHC 59870.
Kaloula rigida: Philippines, Luzon Island, Ka-
linga Prov., Municipality of Balbalan: CAS
61461, 61466, 61467, 61469, 61470, 61473,
61474, 62549, 62550, (Paratypes); Barangay Sad-
sadan, Mt. Nagtud: TNHC 59644; Cagayan
Prov., Municipality of Baggao, Via Barrio: PNM
5471.
Kaloula walteri: See holotype, paratypes, and
referred specimens sections.
A
CKNOWLEDGMENTS
We thank the Protected Areas and Wildlife
Bureau of the Philippine Department of Envi-
ronment and Natural Resources for facilitating
collecting and export permits necessary for
field portions of this and related studies. Some
collections and data contributing to the descrip-
tion of K. walteri were taken as part of the re-
quirements fulfilling ACD’s master’s degree
program from the University of the Philippines
at Los Ban˜os. ACD’s early fieldwork on Mt. Ba-
nahao was made enjoyable by the company and
comic relief of N. Mallari and R. Altamirano.
We thank R. Altamirano, N. Mallari, J. Sedlock,
R. Fernandez, J. Redor, G. Gaytano, M. Pedre-
gosa, J. Ibanez, G. Bueser, M. Duya, R. Quiber,
M. Amor, P. Villanueva, R. Riso, J. Rosell, C. Tor-
no, J. Gonzales, A. Dans, W. Reyes, M. Leonida,
R. Maranan, R. Lagat, S. Rivera, W. Faller, and
B. Obmerga for various assistance associated
with fieldwork on Mt. Banahao. A. Manamtam,
D. Balete, and L. Afuang access provided the
female specimen from Mt. Isarog and we thank
K. Hampson and V. Yngente for sharing obser-
vations of Kaloula populations from Polillo Is-
land. Visits to the Tayabas and Lucban areas of
Mt. Banahao were facilitated by J. Redor, G.
Gaytano, Kabanahaw, Inc., Tayabas Mountain-
eers, Luntiang Alyansa Para sa Bundok Bana-
haw, and J. Jaro, and W. Faller of the Southern
Luzon Polytechnic College. Financial support
for some fieldwork was provided by the Society
of Systematic Biologists, the U.S. National Sci-
ence Foundation, the Society for the Study of
Amphibians and Reptiles, the American Society
of Ichthyologists and Herpetologists, The Ex-
plorers Club, the Section of Integrative Biology
and Texas Memorial Museum of the University
of Texas at Austin. In particular, we thank J. Re-
dor for his consistent support of our research
in Tayabas, for assistance with the tagalog ab-
stract presented in this paper, and for his efforts
in documenting Tayabas folklore. For the gen-
erous loans of specimens and provision of work-
ing space during museum visits we thank R. Dre-
wes, J. Vindum, M. Koo, K. Wiseman, C. Read-
ing and L. Henwood (CAS), R. Sison (PNM), A.
Resetar, H. Voris, R. Inger (FMNH), J. Rosales
and D. Cannatella (TNHC). Financial support
for visits (by ACD and RMB) to CAS was gen-
erously provided by the Charles Stearns Fellow-
ships and support for ACD’s visit to TMM was
facilitated by E. Theriot and D. Cannatella
(TNHC). Critical reviews of the manuscript
were provided by R. Crombie, E. Moriarty, R.
Inger, G. Pauly, and an anonymous reviewer.
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A
NGELO
K
ING
C
ENTER FOR
R
ESEARCH AND
E
NVI
-
RONMENT
M
ANAGEMENT
;M
ARINE
L
ABORATORY
,
S
ILLIMAN
U
NIVERSITY
,B
ANTAYAN
,D
UMAGUETE
C
ITY
,N
EGROS
O
RIENTAL
,P
HILIPPINES
;
AND
(RMB) S
ECTION OF
I
NTEGRATIVE
B
IOLOGY AND
T
EXAS
M
EMORIAL
M
USEUM
,U
NIVERSITY OF
T
EXAS
,A
USTIN
,T
EXAS
78712–1064. E-mail:
(ACD) kaloula@i-manila.com.ph; (RMB)
rafe@mail.utexas.edu; and (ACA) suakcrem@
philwebinc.com. Send reprint requests to
RMB at Texas address. Submitted: 11 July
2001. Accepted: 13 May 2002. Section editor:
C. Guyer.
1051DIESMOS ET AL.—NEW PHILIPPINE KALOULA
A
PPENDIX
1. S
UMMARY OF THE
D
ISTRIBUTION
(
1
,P
RESENT
;
2
,A
BSENT
)
OF
D
IAGNOSTIC
C
HARACTER
S
TATES IN
P
HILIPPINE
M
EMBERS OF THE
G
ENUS
Kaloula.
Character walteri rigida picta conjuncta negrosensis meridionalis stickeli kalingensis kokacii baleata
SVL (m)
(f)
24.5–31.5
31.4
33.1–43.9
40.3–56.1
38.1–51.2
34.7–56.1
26.6–30.1
28.1–33.1
29.8–35.6
39.6–45.3
31.9–44.2
38.7–48.5
41.7–44.4 24.2–32.9
29.0–39.3
37.0–39.1
42.6–44.3
51.1–61.2
45.1–60.2
expanded digital disks
dorsal tubercles
black sacral spots
inner metatarsal tubercle
red axilla spots
web reaching disk of fourth toe
2
2
,
1
2
,
1
1
2
2
2
1
1
1
2
2
2
1
2
,
1
1
2
2
1
2
,
1
2
1
2
1
1
1
2
1
2
1
1
1
2
,
1
1
2
1
1
1
2
1
2
2
1
2
2
2
2
,
1
2
1
2
2
,
1
2
2
2
1
2
2
2
1
2
outer metatarsal tubercle ver y faint
or absent
prominent,
pointed
prominent,
pointed
weak weak weak weak weak distinct weak
subarticular tubercles on fourth toe 3 (30%) 3 (80%),
2 (20%)
3 2223223
lateral glandular ridge or tubercle rows
glandular venter
supernumerary tubercles
2
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
2
2
2
2
2
2
2
2
2
stratified color on posterior thigh
stratified lateral flank coloration
perianal ring
1
1
2
1
1
2
2
,
1
1
2
1
1
2
1
1
2
1
1
2
1
1
2
2
2
1
2
2
1
2
2
1
