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Species diversity, elevational distribution and reproductive

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Sarawak (Borneo)

Article · December 2007

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 Hamburg, Dezember 2007
Mitt. hamb. zool. Mus. Inst. Band 104 S. 141-174
ISSN 0072 9612

Species diversity, elevational distribution and

reproductive modes in an amphibian community at the
Matang Range, Sarawak (Borneo)

INDRANEIL DAS1, ANDRÉ JANKOWSKI2, MOHD. IQBAL B. MAKMOR3 &

ALEXANDER HAAS2

1
Institute of Biodiversity and Environmental Conservation, Universiti Malaysia Sarawak, 94300,
Kota Samarahan, Sarawak, Malaysia. E-mail: idas@ibec.unimas.my.
2
Biozentrum Grindel und Zoologisches Museum, Universität Hamburg, Martin–Luther–King–
Platz 3, 20146 Hamburg, Germany.
3
Department of Zoology, Faculty of Resource Science and Technology, Universiti Malaysia
Sarawak, 94300, Kota Samarahan, Sarawak, Malaysia.

ABSTRACT. – We present an inventory of the amphibian fauna of the Matang Range, including
Kubah National Park and the Matang Wildlife Centre areas of western Sarawak, East Malaysia
(Borneo). A total of 55 species is reported, including 53 Anura and two Gymnophiona. Relative
abundance and diversity of anuran amphibians was also examined along an elevational transect.
The transect was divided into four altitudinal bands of < 200 m, 201-400 m, 401-600 m, and
601-800 m elevations, all coinciding with mixed dipterocarp forests. Relative abundance and
species diversity decreased with elevation in general and was the highest below 200 m. Nine
distinct reproductive modes are shown by the fauna, the most common being Mode 1 (eggs and
feeding tadpoles in lentic water, with 12 species) and Mode 2 (eggs and feeding tadpoles in lotic
water, with 19 species). Variation in reproductive modes shown by the anuran amphibian fauna
at Matang is suspected to be the causal reason for the high regional species richness, helping
reduce competition via resource partitioning.

KEYWORDS: Amphibia, Matang Range, Sarawak, Malaysia, Borneo, biodiversity, species

turnover, elevation, reproductive modes.

Introduction
The Matang Range (Figs 1-2) rises about 22 km to the north-west of the city of Kuching,
and includes two protected areas– Kubah National Park (Park Headquarters at 01°36'41.7''N,
110°11'47.1''E) and the Matang Wildlife Centre (Park Headquarters at 01°36'33.8''N,
110°09'35.3''E). The summit of this massif, Gunung Serapi, is 911 m asl. The range is the
primary catchment area for Sungei (= river) Rayu, and consists of sand and mud
deposited alluvium, along fairly steep terrain, with streams and drains on the flanks, the
altitude under protected areas lying between 20-777 m asl. The summit region lies within
the jurisdiction of Telecom Malaysia. Vegetation represented includes mixed dipterocarp
forest and kerangas (Bornean heath forest). An account of the location, vegetation and
geology of the area can be found in HAZEBROEK & BIN A. MORSHIDI (2000).
142 I. DAS, A. JANKOWSKI, M. I. B. MAKMOR & A. HAAS

Fig. 1. Map of the Matang Range, Sarawak. Inset: Map of Borneo, showing location of Sarawak
and the Matang Range.

The earliest collection of amphibians from the region was made by EDWARD BARTLETT
(ca. 1836-1908), a former Curator of the Sarawak Museum, in 1895. Subsequent collections
in the last century were reported by SMITH (1925) and INGER (1966). Field collections
from Matang were also done in the 1990s by MASAFUMI MATSUI, from Kyoto University.
We present here an inventory of the amphibian fauna of the Matang Range, based on
field work that was conducted between 2004-2007. Collections were made along the summit
trail to Serapi and areas adjacent to the trail, along trails to the Sungei Rayu Waterfall,
transect within Sungei Rayu, transect within a stream close to the Sungei Rayu around
the Matang Wildlife Centre area, along Sungei Matang and in the “Frog Pond“ area
(including transects within two streams close to the “Frog Pond”). The Frog Pond is a
human excavated depression of 1.5-2 m depth, 25-40 m length, and 8-12 m width,
depending on season.
Altitudinal changes, correlated with changes in vegetation types, are known predictors
of amphibian distribution (e.g., LAKIM et al. 1999; INGER & STUEBING 1992; DANIELS
1992; NAVAS 2003). Additionally, the frequency of occurrence of certain microhabitats,
such as side pools which can be occupied by larval stages of some species, are reduced
with increase in stream gradient above 1,000 m. Nonetheless, montane regions may be
important in supporting endemics– INGER & STUEBING (1992) reported 30 species of
 Amphibians of Matang Range 143

Fig. 2. View of the Matang Range from Kampung Matang.

amphibians as restricted to montane regions of Borneo. Local precipitation (and

therefore humidity regimes) may also play an important role in the regional distribution
of amphibians. In a study of altitudinal distribution of amphibians of the Himalayas,
WALTNER (1973) found that eastern Himalaya which receives higher precipitation, has
greater species richness than the drier western Himalaya. He also noted that amphibian
species richness decrease with elevation, with a sharp turnover noticed at ca. 2,750 m.
Further south, in the Western Ghats of Peninsular India, DANIELS (1992) reported
turnover at ca. 1,000 m. NAVAS (2002) reported that herpetofaunal diversity along
elevational gradients is linked to physiological ecology and evolutionary physiology,
and biotic and abiotic factors, water availability, oxygen and carbon dioxide are possible
related ecological factors limiting regional diversity. The effect of ultraviolet radiation on
anuran amphibians on egg development is another factor which may constrain amphibians
at higher elevations (NAVAS 2006). Past anthropogenic activities may also shape current
amphibian assemblages on mountains. For instance, LAKIM et al. (1999) suggested that
habitat destruction and alteration through human activities have influenced regional species
richness at their study site on Gunung Kinabalu, Sabah (Borneo).
Specific objectives of our studies were preparing an inventory of the amphibians of
the Matang Range, determining abundance and diversity of amphibians along an
elevation transect on Gunung Serapi, to test for faunal change with elevation (since
faunal turnover has been recorded in previous studies in Borneo at elevations of ca.
1,000 m) and assess the diversity of reproductive types in the area, through analysis of
the reproductive modes (development type and oviposition site).

Materials and Methods

S p e c i e s a c c o u n t s a n d n o m e n c l a t u r e. – For each recorded species, to aid
future workers, we provide a short description of adults of each species (the status of knowledge
of larval stages is currently incomplete, not permitting us to provide descriptions of all species),
and whenever possible, a colour image from our extensive files of images, typically from the
Matang region. In case of images from extralimital areas, we have mentioned the locality in the
144 I. DAS, A. JANKOWSKI, M. I. B. MAKMOR & A. HAAS

captions. We follow the nomenclature proposed in the Amphibian Tree of Life (FROST et al.
2006), as updated in FROST (2007).
I n v e n t o r y t e c h n i q u e s. – Data of analysis of altitudinal distribution on Gunung
Serapi were collected during field work between December 2005 and January 2006. Other data on
inventories and reproductive diversity were collected between 2004-2007. Specimens were
collected during transect walks by day or night (= Visual Encounter Surveys), which are generally
useful for compiling species lists of the area (see CRUMP & SCOTT 1994). Specimens collected
were photographed in life, euthanised with chlorobutanol typically within 12 h of capture,
tissues (generally liver) preserved in 95% ethanol, the specimen fixed in 4% formalin and
subsequently washed after 4 days and stored in 70% ethanol. Individual specimens (except
tadpoles) were numbered using the field collectors’ number (see below), and will be eventually
deposited in the museums of the Faculty of Resource Science and Technology, UNIMAS and
Biozentrum Grindel und Zoologisches Museum, Universität Hamburg. Tadpoles were numbered
in lots and prefixed ID.BT (Indraneil Das.Bornean Tadpole collection). Additional specimens
were examined at the following systematic institutions (abbreviations after LEVITON et al. 1985;
1988 indicated with asterisk):
BMNH* – The Natural History Museum, London, U. K.,
FMNH* – Field Museum of Natural History, Chicago, U.S.A.,
SM* – Sarawak Museum, Kuching, Malaysia,
UNIMAS – Zoological Museum, Universiti Malaysia Sarawak, Kora Samarahan, Malaysia, and
USDZ* – (also, ZRC), Raffles Museum of Biodiversity Research, National University of
Singapore, Singapore.

Abbreviations of field collectors include:

AJA – ANDRÉ JANKOWSKI
ID – INDRANEIL DAS
MW – MARK WILKINSON
RFI – ROBERT FREDERICK INGER

In the following section, we refer to the nine main collecting and observation sites (stations).
Geographic coordinates are available for five of them. Coordinates were taken with a Garmin
VistaTM Geographic Positioning System (datum: WGS 84).:
1. Kubah National Park Headquarters: 01°36'22''N, 110°11'45''E
2. Stream to right (facing downhill) of 200 feet point on Summit Trail : 01°36'42''N, 110°11'45''E
3. Frog Pond, situated at ca. 1,000 feet point on Summit Trail: 01°36'24''N, 110°11'22''E
4. Sungei Rayu 300 m transect starting at ca. 680 feet: 01°35'50''N, 110°10'47''E
5. Matang Wildlife Centre Headquarters: 01°36'33''N, 110°09'35''E

E l e v a t i o n a l d i s t r i b u t i o n. – Elevational line transect method was applied, the

transect divided into four elevational bands of 200 m interval as follows: below 200 m, 201-400
m, 401-600 m and 601-800 m elevations. The 200 m intervals seemed to be the most suitable
scale considering the difference in temperature at each interval would be ca. 1o Celsius (PASCAL
1988). Each band is more or less 500 m in length and frogs were sampled 2-3 m of both sides of
the Summit Trail. Stream bank and forest species were not considered in this study. All sampling
efforts were done at night (between 1930-2300 h), as all local frog species are nocturnal and
capture rate are higher at night (INGER 1980). Since all samplings were carried out at night, there
is no bias introduced by the capture of frogs by day.
D a t a a n a l y s e s. – For statistical analysis, the Shannon-Weiner Index (PIELOU 1966) was
used, with the modified Divers programme to describe the heterogeneity. Originally written by
KREBS (1989), we used the Rarefract and Divers programmes as modified by LAMAN (2001a;
2001b). ZAR’S (1996) modified t-test was used to test for significance between altitudinal bands,
using the MS-Excel 2003.

Formulae used in the Zar’s (1996) modified t-test are:

t = (H’1-H’2)/SH’1-H’2
where,
SH’1 – H’2 = sqrt { S2H’1 + S2H’2}
 Amphibians of Matang Range 145

The variance for each H’ may be approximated by:

S2H’ = {(£ fi log2 fi – (£ fi log (fi))2 /n)} / n2

Degrees of freedom (d.f.) for this test is v, as:

v = (S2H’1+ S2H’2) / {(S2H’1)2/n1 + (S2H’2)2/n2}

Species Inventory
Bufonidae

Ansonia leptopus (GÜNTHER, 1872): Brown Slender Toad.

(Fig. 3a)

Bufo leptopus A. C. L. G. GÜNTHER. 1872. Proc. Zool. Soc. London 1872:598.

M a t e r i a l. – Summit Trail to Gunung Serapi, off Frog Pond (ID-8393; ID-8402; ID-8650;
ID-8750; ID-8854; ID-8865); Sungei Rayu (AJA-006), stream close to Sungei Rayu (pers. obs.,
AJA). The species was described based on types from Matang.

D e s c r i p t i o n. – SVL to 62 mm; habitus slender; limbs long and thin; tips of fingers
rounded, third and fourth toes free of web; tympanum visible externally; skin of dorsum
with numerous small rounded warts; dorsum brown, flanks slightly darker; and venter
grey with darker mottling.

Ansonia longidigita INGER, 1960: Long-fingered Slender Toad.

(Fig. 3b)

Ansonia longidigita R. F. INGER. 1960. Fieldiana Zool. 39:480; Figs 81C, 83B, 84A.

M a t e r i a l. – Summit Trail (AJA-033).

D e s c r i p t i o n. – SVL to 65 mm; habitus slender; tympanum visible externally;

fingers slender; finger tips rounded; first finger reaching disk of second; males with 2-4
rows of black spines under lower jaw, and four rows of large, spinose tubercles on
dorsum; tarsal ridge absent; one to two phalanges of third and fifth toes free of web;
dorsum dark brown to blackish-brown, with lighter spots; limbs with dark cross-bars;
and venter cream with darker mottling.

Ansonia minuta INGER, 1960: Minute Slender Toad.

(Fig. 3c)

Ansonia minuta R. F. INGER. 1960. Fieldiana Zool. 39:493; Figs 81A, 89, 90.

M a t e r i a l. – Stream to right of 200 m point on Summit Trail (ID-8419-8422; ID-8484;

ID-8862, AJA-002). Also Sungei Rayu (pers. obs., AJA).

D e s c r i p t i o n. – SVL to 28 mm; habitus slender; tips of fingers disk-like; tip of

finger I not reaching disk of finger II; toes III and V with two phalanges free of web;
tarsal ridge sharp; interorbital ridge absent; dorsum brown, tubercles yellow; limbs with
darker bands; and venter yellow.
146 I. DAS, A. JANKOWSKI, M. I. B. MAKMOR & A. HAAS

Fig. 3a. Brown Slender Toad (Ansonia leptopus); b, Long-fingered Slender Toad (Ansonia
longidigita); c, Fig. 5. Minute Slender Toad (Ansonia minuta); d, Spiny Slender Toad (Ansonia
spinulifer); e, Common Asian Toad (Duttaphrynus melanostictus; specimen from Kuching);
f, Crested Toad (Ingerophrynus divergens); g, Brown Tree Toad (Pedostibes hosii; specimen
from Mulu Natl. Park); h, Short-legged Dwarf Toad (Pelophryne signata).
 Amphibians of Matang Range 147

Ansonia spinulifer (MOCQUARD, 1890): Spiny Slender Toad.

(Fig. 3d)

Bufo spinulifer F. MOCQUARD. 1890. Nouv. Arch. Mus. Nat. Hist. Nat. Paris, Ser. 3, 2:160.

M a t e r i a l. – Summit Trail to Gunung Serapi, off Frog Pond (ID-8240; ID-8258–59; ID-8629;
ID-8867), Sungei Rayu and stream near Sungei Rayu (pers. obs., AJA) Also, SM D.a.3.2.5,
collected 1898 from “Matang“ by an unknown collector.

D e s c r i p t i o n. – SVL to 45 mm; habitus slender; limbs long and thin; tips of fingers
rounded; toes nearly unwebbed; tympanum visible; large, spinose tubercles on the
dorsum, which is black with yellowish-cream oval patch between shoulders; and venter
dark with pale spots and streaks.

Duttaphrynus melanostictus (SCHNEIDER, 1799): Common Asian Toad.

(Fig. 3e)

Bufo melanostictus J. G. SCHNEIDER. 1799. Hist. Amphib. Nat.:216.

M a t e r i a l. – None collected, but observed at the Kubah Park Headquarters. This species has
been recorded from “Mt. Matang“ by SMITH (1925) and INGER (1966:74).

D e s c r i p t i o n. – SVL to 115 mm; habitus stocky with warty skin; snout obtusely
pointed; tympanum distinct about ½ eye diameter; elongated parotoid gland; toes half-
webbed; black bony ridges from snout to eye and around upper orbit and tympanum;
dorsum greyish-brown, with orange and yellow markings during the breeding season;
and venter cream.

Ingerophrynus divergens (PETERS, 1871): Crested Toad.

(Fig. 3f)

Bufo divergens W. C. H. PETERS. 1871. Mber. Koninkl. Akad. Wiss. Berlin 1871:579.

M a t e r i a l. – Summit Trail to Gunung Serapi, off Frog Pond (ID-8247; ID-8630); on Summit
Trail to Gunung Serapi, ca. 400 feet (ID-8737). Stream close to Frog Pond, Waterfall Trail (pers.
obs., AJA). Also, “Matang“ (SM D.a.3.2.8– 4 specimens), collected 4.1912 by J. GEE FOOK.
“Mt. Matang“ cited by SMITH (1925).

D e s c r i p t i o n. – SVL to 55 mm, habitus stocky; a pair of ridges between eyes;

parotoid gland longer than wide, followed by rows of warts; fingers not fully webbed;
dorsum bright orange or yellow-brown, sometimes with yellow vertebral stripe; and
venter cream.

Pedostibes hosii (BOULENGER, 1892): Brown Tree Toad.

(Fig. 3g)

Nectophryne hosii G. A. BOULENGER. 1892. Proc. Zool. Soc. London 1892:508; Pl. 30; Fig.2.

M a t e r i a l. – Not collected. A calling group of males observed on the bank of Sungei Rayu on
148 I. DAS, A. JANKOWSKI, M. I. B. MAKMOR & A. HAAS

27 June 2006.
D e s c r i p t i o n. – SVL to 105 mm; habitus stocky; a bony ridge connects eyes to
parotoid gland; tympanum visible; toes (except fourth) fully webbed; fingers basally
webbed, finger tips expanded into pads; dorsum of males and some females unpatterned
brown, other females black, purple or greenish-olive, with bright yellow spots; and
venter cream or yellow.

Pelophryne guentheri (BOULENGER, 1882): Günther’s Dwarf Toad.

Nectophryne guentheri G. A. BOULENGER. 1882. Cat. Batr. Sal. Coll. Brit. Mus.:280; Pl. 18, Fig. 3

M a t e r i a l. – None collected by us. The holotype (BMNH 1947.2.19.28; ex-BMNH

72.2.19.27) was taken in “Matang“. A second specimen was collected during undergraduate
student field work in 31 August 2006, from a forest quadrat and deposited in the museum of
UNIMAS.

D e s c r i p t i o n. – SVL to 30 mm; habitus slender; head slightly depressed; limbs

slender; webbing on hand thick, reaching tip of first finger but leaving half of outer
fingers free; fifth toe webbed to disks; dorsum greenish-grey, with reddish-brown
tubercles; and venter cream with dark mottling.

Pelophryne signata (BOULENGER, 1894): Short-legged Dwarf Toad.

(Fig. 3h)

Nectophryne signata G. A. BOULENGER. 1894. Proc. Zool. Soc. London 1894:645; Pl. 40; Fig. 1.

M a t e r i a l. – Ridge behind Matang Wildlife Centre (ID-7159).

D e s c r i p t i o n. – SVL to 25 mm; habitus slender; webbing on hand thick, reaching

tip of first finger but leaving half of outer fingers free; webbing reach disks of first toe
only; parotoid gland absent; dorsum dark brown, small black spots or a light or dark
cruciform pattern, sometimes with a pale lateral stripe along flanks; and venter spotted
with brown or black.

Phrynoidis aspera (GRAVENHORST, 1829): River Toad.

(Fig. 4a)

Bufo asper J. L. C. GRAVENHORST. 1829. Delic. Mus. Zool. Vrat.: 58.

M a t e r i a l. – None collected, but the distinctive chorus of this species was heard on 4 August
2006 from the river near the head of the Waterfall Trail.

D e s c r i p t i o n. – SVL to 170 mm; habitus stocky; parotoid small, rounded; bony

ridges on forehead absent; tympanum small, indistinct; all toes (except fourth) webbed
to tips; dorsum grey to dark brown; throat, chest and sides and undersides of limbs with
small red and black spots; and rest of venter cream.
 Amphibians of Matang Range 149

Fig. 4a. River Toad (Phrynoidis aspera; specimen from Serian); b, Giant River Toad
(Phrynoidis juxtaspera; specimen from Mulu Natl. Park); c, Dwarf Mountain Frog (Ingerana
baluensis); d, Grass Frog (Fejervarya limnocharis); e, Kuhl’s Creek Frog (Limnonectes kuhlii);
f, Rivulet Frog (Limnonectes laticeps); g, Giant River Frog (Limnonectes leporinus); h, Malaysian
River Frog (Limnonectes malesianus);
150 I. DAS, A. JANKOWSKI, M. I. B. MAKMOR & A. HAAS

Phrynoidis juxtaspera (INGER, 1964): Giant River Toad.

(Fig. 4b)

Bufo juxtasper R. F. INGER. 1964. Fieldiana Zool. 44:154.

M a t e r i a l. – None collected but reported for the area by HAZEBROEK & BIN A. MORSHIDI
(2000:368).

D e s c r i p t i o n. – SVL to 215 mm; habitus stocky; head broad; bony crests on

forehead absent; tympanum visible; parotoid gland 2–4 times as long as wide; toes
(except fourth) webbed to tips; dorsum dark grey-brown to black, sometimes with
indistinct darker spots; and venter cream with dark mottling.

Ceratobatrachidae

Ingerana baluensis (BOULENGER, 1896): Dwarf Mountain Frog.

(Fig. 4c)

Cornufer baluensis G. A. BOULENGER. 1896. Ann. & Mag. nat. Hist. ser. 6 17:449.

M a t e r i a l. – Sungei Rayu (AJA-024).

D e s c r i p t i o n. – SVL to 40 mm; habitus squat; short head; fingers short, broadened

at tip; toes half webbed, with widened tips; hind limbs short; skin of dorsum and venter
with pebbly texture, more smooth on venter; dorsum brown to brownish-grey, with
scattered dark spots or a dark longitudinal band; chest and rest of venter pale grey or
cream and throat heavily pigmented with brown.

Dicroglossidae

Fejervarya limnocharis (GRAVENHORST, 1829): Grass Frog.

(Fig. 4d)

Rana limnocharis J. L. C. GRAVENHORST. 1829. Delic. Mus. Zool.Vratislav. 1:42.

M a t e r i a l. – >300 m asl, Summit Trail to Gunung Serapi (ID-8476); Summit Trail near Frog
Pond (AJA-018, AJA-020 and AJA-023; UNIMAS 147–48). Also, SM D.a.1.2.4 from
“Matang“, collected 6.1893 by EDWARD BARTLETT (cited in BARTLETT 1895), SMITH (1925)
and INGER (1966: 206).

D e s c r i p t i o n. – SVL to 60 mm; habitus moderately slender; snout narrow; dorsum

with broken longitudinal ridges; outer metatarsal tubercle absent; toes with pointed tips,
less than half webbed; dorsum brown to brownish-grey, some with a pale yellow
vertebral line; males with a black, M-shaped band across throat and venter cream.

Limnonectes kuhlii (TSCHUDI, 1838): Kuhl’s Creek Frog.

(Fig. 4e)

Rana kuhlii J. J. de TSCHUDI. 1838. Classif. Batr.: 40.

 Amphibians of Matang Range 151

M a t e r i a l. – Summit Trail to Gunung Serapi, off Frog Pond (ID-8205; ID-8255; ID-8257;
ID-8757; ID-8849; ID-8868); stream south of Frog Pond; Sungei Rayu; stream east of Sungei
Rayu (pers. obs., AJA). Kubah, without precise data (UNIMAS 013; UNIMAS 016; UNIMAS
158; UNIMAS 162–164; UNIMAS 166–167). Also, SM uncatalogued (ex-RFI 76383), from
“Sungei China, Matang“, collected 15.7.1956 by R. F. INGER. “Matang“ (in INGER, 1966:202).

D e s c r i p t i o n. – SVL to 69.0 mm; habitus robust; head enlarged in adult males, with
large, tooth-like processes on mandibles; limbs short; flanks of body and limbs with
rounded tubercles; toes fully webbed; tympanum hidden; dorsum brown with darker
blotches; a pale brown band from eye to jaws and one across snout and venter cream
except for the dark-spotted throat and chest.

Limnonectes laticeps (BOULENGER, 1882): Rivulet Frog.

(Fig. 4f)

Rana laticeps G. A. BOULENGER. 1882. Cat. Batr. Sal. British Mus.:20; Pl. 1; Fig. 1.

M a t e r i a l. – Along Summit trail to Gunung Serapi (ID-8190–95; ID-8530; ID-8643; ID-

8645; ID-8648; ID-8734–35; ID-8856–57; ID-8866); stream south of Frog Pond (pers. obs.,
AJA). Kubah, without precise data (UNIMAS 007; UNIMAS 015; UNIMAS 037; UNIMAS
132; UNIMAS 157; UNIMAS 159–161; UNIMAS 165).

D e s c r i p t i o n. – SVL to 35.7 mm; habitus robust; head enlarged in adult males, with
large, tooth-like processes on mandibles; limbs short; skin of dorsum with a corrugated
pattern; tympanum hidden; toes with one phalange free of web; dorsum brown with or
without darker blotches; a pale brown band from eye to jaws and one across snout and
venter cream.

Limnonectes leporinus (ANDERSSON, 1923): Giant River Frog.

(Fig. 4g)

Rana macrodon var. leporina L. G. ANDERSSON. 1923. Nyt. Mag. Nat. 61:123.

M a t e r i a l. – Frog Pond (ID-8179; ID-8184; ID-8260; ID-8526); > 300 m asl, Summit Trail
to Gunung Serapi (ID-8480); near Park HQ, Summit Trail to Gunung Serapi (AJA-042); stream
south of Frog Pond; Sungei Rayu (pers. obs., AJA). Kubah, without precise data (UNIMAS
066; UNIMAS 075; UNIMAS 102).

D e s c r i p t i o n. – SVL to 180 mm; habitus robust; snout pointed; limbs long and
powerful; short ridges on back; toes almost completely webbed; males much larger than
females, with lower jaws bearing an enlarged fang-like structure; dorsum grey; a dark
streak between eye and nostril marking the canthal ridge and limbs dark-banded.

Limnonectes malesianus (KIEW, 1984): Malaysian River Frog.

(Fig. 4h)

Rana malesiana B.-H. KIEW. 1984. Malayan Nat. J. 37:154.

M a t e r i a l. – Frog Pond (ID-8731); Stream south of Frog Pond (ID-8863); Waterfall Trail
close to Sungei Rayu (AJA-046).
152 I. DAS, A. JANKOWSKI, M. I. B. MAKMOR & A. HAAS

D e s c r i p t i o n. – SVL to 150 mm; habitus robust; toes 3/4 webbed and swollen at
tips; tympanum visible and tympanic region flattened; snout profile between eye and
nostril flat and smooth (no canthal ridge); dorsum reddish-brown to brownish-pink; in
many specimens snout set off in colour from dorsum by interorbital band and eye to
upper lip bands; a pale, thin vertebral stripe present in some individuals and venter
cream with dark mottling.

Limnonectes palavanensis (BOULENGER, 1894): Smooth Guardian Frog.

(Fig. 5a)

Rana palavanensis G. A. BOULENGER. 1894. Ann. & Mag. nat. Hist. ser. 6 14:85.

M a t e r i a l. – Off Frog Pond (ID-8263; ID-8634); Waterfall Trail (AJA-016); stream south of
the Frog Pond (pers. obs., AJA). Kubah, without precise data (UNIMAS 137–38; UNIMAS 154).

D e s c r i p t i o n. – SVL to 40 mm; habitus slender; hind limbs long; toes half webbed,
with slightly swollen tips; a V-shaped ridge of skin between shoulders; dorsum reddish
brown to chocolate brown; a dark interorbital bar and venter yellow.

Limnonectes paramacrodon (INGER, 1966): Masked Frog.

(Fig. 5b)

Rana paramacrodon R. F. INGER. 1966. Fieldiana Zoology 52:228; Fig. 47.

M a t e r i a l. – On trail to Sungei Rayu (ID-8636); stream south of the Frog Pond (pers. obs., AJA).

D e s c r i p t i o n. – SVL to 75 mm; habitus robust; snout rounded; tympanum and

supratympanic fold distinct; a few low tubercles on dorsum; toe tips distinctly widened
and webbed to tips, except toe IV, with one phalange free; a dark mask covers
tympanum and supratympanic fold; dorsum greyish-brown and venter yellow.

Occidozyga baluensis (BOULENGER, 1896): Seep Frog.

(Fig. 5c)

Oreobatrachus baluensis G. A. BOULENGER. 1896. Ann. & Mag. nat. Hist. ser. 6 17:401; Pl. 17.

M a t e r i a l. – None collected; cited in INGER (1966:240) from “Matang“.

D e s c r i p t i o n. – SVL to 35 mm; habitus stocky; vomerine teeth absent; toes 3/4

webbed; dorsum tan or brown and venter cream with black blotches.

Occidozyga laevis (GÜNTHER, 1859): Yellow-bellied Puddle Frog.

(Fig. 5d)

Oxyglossus laevis A. C. L. G. GÜNTHER. 1859. Cat. Batr. Sal. British Mus.:7; Pl. 1; Fig. A.

M a t e r i a l. – In shallow stream, close to Frog Pond, off Summit Trail (ID-8532; ID-8632;
ID-8729–30; AJA-030–32; AJA-044); Sungei Rayu trail (AJA-015 and AJA-019). Also,
“Matang“ (SM D.a.1.1), date and collector unknown; cited from “Matang“ in INGER (1966:243).
 Amphibians of Matang Range 153

Fig 5a. Smooth Guardian Frog (Limnonectes palavanensis); b, Masked Frog (Limnonectes
paramacrodon); c, Seep Frog (Occidozyga baluensis; specimen from Mulu Natl. Park);
d, Yellow-bellied Puddle Frog (Occidozyga laevis); e, Lowland Litter Frog (Leptobrachium
abbotti); f, Mjöberg’s Dwarf Litter Frog (Leptobrachella mjobergi); g, Sarawak Slender Litter
Frog (Leptolalax gracilis); h, Bornean Horned Toad (Megophrys nasuta).
154 I. DAS, A. JANKOWSKI, M. I. B. MAKMOR & A. HAAS

D e s c r i p t i o n. – SVL to 48 mm; habitus stocky; hind limbs short and fat; toes thick,
fully webbed, with rounded tips; a single tooth-like projection on lower jaw; dorsum
dark grey brown, sometimes with a pale vertebral stripe; and venter lemon yellow.

Megophryidae

Leptobrachium abbotti (COCHRAN, 1926): Lowland Litter Frog.

(Fig. 5e)

Megophrys abbottii D. COCHRAN. 1926. J. Washington Acad. Sci. 16:446.

M a t e r i a l. – Summit Trail to Gunung Serapi, off Frog Pond (ID-8404; ID-8647); > 300 m asl,
Summit Trail to Gunung Serapi (ID-8481); stream close to Frog Pond (AJA-052).

D e s c r i p t i o n. – SVL to 95 mm; habitus stocky; head wider than body; eyes large;
limbs short and weak; toes nearly free of webbing; skin of dorsum smooth; dorsum dark
brown, usually with a darker pattern on forehead; iris dark brown; and bulbus culi blue
beyond iris.

Leptobrachella mjobergi SMITH, 1925: Mjöberg’s Dwarf Litter Frog.

(Fig. 5f)

Leptobrachella mjöbergi M. A. SMITH. 1925. Sarawak Mus. J. 3:13; Figs A-D.

M a t e r i a l. – Summit Trail to Gunung Serapi, off Frog Pond (ID-8241); > 300 m asl, Summit
Trail to Gunung Serapi (ID-8482–83); Waterfall Trail and river (AJA-007, AJA-041 and AJA-047);
stream close to Frog Pond; stream near Sungei Rayu (pers. obs., AJA). Also, SM uncatalogued
(ex-RFI 77206) from “Sungei China, Matang“, collected 28.7.1956 by R. F. INGER, GAUN and
BIDAI.

D e s c r i p t i o n. – SVL to 23 mm; head narrower than body; digits free, their tips
swollen but with pointed disks; sides of body with elongated cream-coloured glands
between axilla and groin; supratympanic fold distinct; brown or grey-black, with several
black spots; a crossbar across eyes and another on shoulder; and venter cream.

Leptolalax gracilis (GÜNTHER, 1872): Sarawak Slender Litter Frog.

(Fig. 5g)

Leptobrachium gracile A. C. L. G. GÜNTHER. 1872. Proc. Zool. Soc. London 1872:598.

M a t e r i a l. – Near entrance to Kubah National Park, on Summit Trail (ID-8254); Summit Trail
to Gunung Serapi, off Frog Pond (ID-8397; ID-8401); Waterfall Trail (ID-8851; AJA-027–028);
Sungei Rayu; stream close to Sungay Rayu (pers. obs., AJA); Kubah, without precise data
(UNIMAS 128). Also, SM uncatalogued (ex-RFI 77218) from “Sungei China, Matang“, collected
28.7.1956 by R. F. INGER, GAUN and BIDAI.

D e s c r i p t i o n. – SVL to 50 mm; habitus slender; head longer than wide; tips of

fingers and toes rounded; toes basally webbed; dorsum dark greyish-blue or brown,
with black blotches and a black interorbital bar; venter cream with black spots; upper
surfaces of limbs are dark barred; and upper surface of forearm cream, but changeable
under stress.
 Amphibians of Matang Range 155

Megophrys nasuta (SCHLEGEL, 1858): Bornean Horned Toad.

(Fig. 5h)

Ceratophrys nasuta H. SCHLEGEL. 1858. Nat. Leercursus Geb. Koninkl. Militaire Akad.
Handl. beof. Dierk.:57;Pl. 4; Fig. 72.

M a t e r i a l. – Summit Trail to Gunung Serapi, off Frog Pond (ID-8362; ID-8848); Summit
Trail near Kubah National Park Headquarters; Waterfall Trail; stream close to Sungei Rayu (pers.
obs., AJA) Kubah, without precise data (UNIMAS 071). Also, “km 16, Matang“ (SM
uncatalogued, in jar no. 18, 26.6.1987, collected by R. AHMED) and “Matang“ (SM D.a.4.2.1.d),
collected 9.1891, by A. MC. D. GIBSON.

D e s c r i p t i o n. – SVL > 125 mm; habitus robust in females; large, dermal

appendages on tip of snout and upper eyelids; head wider than long; tympanum hidden;
tips of fingers and toes blunt; fingers free; toes basally webbed; skin with two pairs of
parallel dermal folds; dorsum brown, with a black subocular bar, limbs with narrow
crossbars; and throat brownish-black in males.

Microhylidae

Calluella brooksii (BOULENGER, 1904): Brookes’ Burrowing Frog

Colpoglossus Brooksii G. A. BOULENGER. 1904. Ann. & Mag. nat. Hist. Ser. 7, 13:42; Pl. 2.

M a t e r i a l. – None collected by us; reported from “Matang“ by INGER (1966:120).

D e s c r i p t i o n. – SVL to 60 mm; habitus robust; head short; limbs short, thick;

nostrils at tip of snout; toes basally webbed; dorsum brown with scattered black spots;
and venter cream or yellow.

Chaperina fusca MOCQUARD, 1892: Saffron-bellied Frog.

(Fig. 6a)

Chaperina fusca F. MOCQUARD. 1892. Mem. Soc. Zool. France 5:35.

M a t e r i a l. – Waterfall Trail (ID.BT-316); Kubah National Park Headquarters (pers. obs., AJA).

Description. – SVL to 26 mm; snout short; small flexible spine at elbow and at heels; toes
free of web; tips of fingers and toes slightly dilated; dorsum greenish- or bluish-black,
with darker patches and venter and undersurfaces of limbs with a black network
enclosing bright yellow spots.

Kalophrynus heterochirus (BOULENGER, 1900): Variable Sticky Frog.

(Fig. 6b)

Calophrynus heterochirus G. A. BOULENGER. 1900. Proc. Zool. Soc. London 1900:186;

Pl. 17; Fig. 4.

M a t e r i a l. – On trail to Sungei Rayu (ID-8527; ID-8749); near Frog Pond (AJA-035), along
Summit Trail; at ephemeral pond (pers. obs., AJA).
156 I. DAS, A. JANKOWSKI, M. I. B. MAKMOR & A. HAAS

D e s c r i p t i o n. – SVL to 33 mm; habitus slender; head narrow; snout pointed; toes

basally webbed; dorsum brownish-orange; groin with several dark-bordered cream
spots; and venter cream with darker markings.

Kalophrynus pleurostigma TSCHUDI, 1838: Red-sided Sticky Frog.

(Fig. 6c)

Kalophrynus pleurostigma J. J. DE TSCHUDI. 1838. Classif. Batr.:86.

M a t e r i a l. – Near stream, on Waterfall Trail (ID-8533); at stream close to Sungei Rayu

(AJA-051). Also, SM D.a.2.1.1.a from “foot of Matang“, collected 6.1893 by E. BARTLETT
(cited by BARTLETT 1895).

D e s c r i p t i o n. – SVL to 58 mm; habitus stout; snout pointed; outer fingers and

toes small; toes barely webbed; dorsal surface of females pebbly, that of males with tiny
spines; dorsum light or dark brown, usually with darker blotches, flanks brownish-orange;
a dark eye-like spot in inguinal region; throat and chest brown and rest of venter grey.

Metaphrynella sundana (PETERS, 1867): Bornean Tree Hole Frog.

(Fig. 6d)

Calophyla sundana W. C. H. PETERS. 1867. Mber. Koninkl. Akad. Wiss. Berlin 1867:35.

M a t e r i a l. – Waterfall Trail (ID-8534); throughout the National Park (acoustically

encountered by AJA).

D e s c r i p t i o n. – SVL to 25 mm; habitus robust; tips of fingers widened; inner meta-

tarsal tubercle low; toes webbed; dorsum and flanks with scattered tubercles; dorsum
light or dark brown or grey-black, with a darker mark in the middle of the back; dorsal
surface of fingers yellow to orange and venter grey.

Microhyla borneensis PARKER, 1926: Bornean Narrow-mouthed Frog.

(Fig. 6e)

Microhyla borneensis H. W. PARKER. 1926. Ann. & Mag. nat. Hist. ser. 10 2:473.

M a t e r i a l. – Frog Pond (ID-8196–97; ID-8250; ID-8508; ID-8529; ID-8640; ID-8738);

stream close to Frog Pond (pers. obs., AJA) Kubah, without precise data (UNIMAS 089). Also,
“Matang“ in SMITH (1925).

D e s c r i p t i o n. – SVL to 23 mm; habitus stocky, broad and depressed; head short,

triangular; snout slightly pointed; tympanum not visible; four fingers; toe tips slightly
widened; dorsum brownish-pink to grey, with a light-edged, dark brown patch on the
middle of the back; black blotches below eye and nares; black dots along flanks and
venter mottled with brown.

Microhyla sp.: Matang Narrow-mouthed Frog.

(Fig. 6f)

M a t e r i a l. – Summit Trail to Gunung Serapi, off Frog Pond (ID-8198–8201; ID-8243;

ID-8270; ID-8633; ID-8855; AJA-004; AJA-010; AJA-012-13; AJA-025; AJA-029; AJA-050).
 Amphibians of Matang Range 157

Fig. 6a. Saffron-bellied Frog (Chaperina fusca); b, Variable Sticky Frog (Kalophrynus
heterochirus); c, Red-sided Sticky Frog (Kalophrynus pleurostigma, specimen from Gg.
Santubong); d, Bornean Tree Hole Frog (Metaphrynella sundana); e, Bornean Narrow-mouthed
Frog (Microhyla borneensis); f, Matang Narrow-mouthed Frog (Microhyla sp.); g, Green Paddy
Frog (Hylarana erythraea); h, White-lipped Frog (Hylarana raniceps).
158 I. DAS, A. JANKOWSKI, M. I. B. MAKMOR & A. HAAS

R e m a r k s. – This represents a new species, whose formal description will be

published elsewhere.

Ranidae

Hylarana baramica (BOETTGER, 1901): Brown Marsh Frog.

(Fig. 7a)

Rana baramica O. BOETTGER. 1901. Abh. Senckenb. Naturforsch. Ges. 25:391.

M a t e r i a l. – Kubah, without precise data (UNIMAS 004). The distinctive calls of this
species were heard from a peat swamp at the Matang Wildlife Centre on 27 June 2006.

D e s c r i p t i o n. – SVL to 67 mm; habitus slender; head relatively large; eye promi-

nent; tympanum visible; toes less than half webbed; fingers long; upper lip with pale
spots; dorsum dark brown, with darker spots on the sides, granular to slightly warty in
texture; tympanum black, with a distinct pale centre; venter cream or pale brown and iris
golden with red-copper tint anterior and posterior to pupil.

Hylarana erythraea (SCHLEGEL, 1827): Green Paddy Frog.

(Fig. 6g)

Hyla erythraea H. SCHLEGEL. 1837. Abbild. Amph.: 27; Pl. 9; Fig. 3.

M a t e r i a l. – None collected by us, but one was taken from Frog Pond by a UNIMAS team,
and one photographed by AJA. Also Summit Trail near Headquarters (pers. obs., AJA).

D e s c r i p t i o n. – SVL to 75 mm; habitus robust; hind limbs long and muscular; toes
half webbed; skin of back smooth, with flat dorsolateral glandular ridges; dorsum bright
green with a broad cream-coloured dorsolateral bands and venter unpatterned cream.

Hylarana glandulosa (BOULENGER, 1882): Rough-sided Frog.

(Fig. 7b)

Rana glandulosa G. A. BOULENGER. 1882. Cat. Batr. Sal. British Mus.: 73; Pl. 8.

M a t e r i a l. – Frog Pond (ID-8180).

D e s c r i p t i o n. – SVL to 93 mm; habitus robust; head relatively large; skin granular

with warts; flanks with large, flat warts; tympanum large; a large gland on upper arm;
toes half webbed; upper lip wit pale spots; dorsum dark brown or brown with darker
spots; iris red; venter cream and iris red.

Hylarana laterimaculata (BARBOUR & NOBLE, 1916): Side-spotted Swamp Frog.

(Fig. 7c)

Rana laterimaculata T. BARBOUR & G. K. NOBLE. 1916. Proc. New England ZoQl. Club
6:21; Fig. 3

M a t e r i a l. – Waterfall Trail (AJA-040). Also heard and recorded from bank of shallow
stream up from Frog Pond and from a swampy area close to Sungei Rayu.
 Amphibians of Matang Range 159

D e s c r i p t i o n. – SVL to 47.5 mm; habitus slender; head relatively large; eyes promi-
nent; tympanum visible; toes less than half webbed; fingers long; dorsum dark brown,
with darker spots on the sides; tympanum black, lacking a pale centre; upper lip with a
continuous pale streak from below level of eyes to posterior edge of tympanum and
venter cream with dark mottlings.

Hylarana luctuosa (PETERS, 1871): Mahogany Frog.

(Fig. 7d)

Limnodytes luctuosus W. C. H. PETERS. 1871. Monatsber. Preuss. Akad. Wiss. Berlin 1871:579.

M a t e r i a l. – Frog Pond (ID-8181–83; ID-8390–92; ID-8637; ID-8646); Kubah, without

precise data (UNIMAS 099–100; UNIMAS 153). Also, “Matang“ (INGER 1966:208).

D e s c r i p t i o n. – SVL to 60 mm; habitus stout; head triangular; limbs short;

tympanum as large as eye; dorsum reddish-brown, flanks and venter greyish-black;
limbs, especially the hind limbs, dark with distinct narrow light grey bands.

Hylarana picturata (BOULENGER, 1920): Spotted Stream Frog.

(Fig. 7e)

Rana (Hylorana) picturata G. A. BOULENGER. 1920. Rec. Indian Mus. 20:179.

M a t e r i a l. – Frog Pond (ID-8189; ID-8271; ID-8403; ID-8644); Summit Trail; stream south
of Frog Pond; Sungei Rayu (pers. obs., AJA) Kubah, without precise data (UNIMAS 127;
UNIMAS 130).

D e s c r i p t i o n. – SVL to ca. 68 mm; tympanum visible; tips of digits slightly

swollen; toes over half webbed; dorsum and flanks dark brown to nearly black, with
yellow or yellowish-orange angular spots; a yellow-orange stripe starts at the snout
towards the eye and dissolves into spots behind eye and venter grey

Hylarana raniceps (PETERS, 1871): White-lipped Frog.

(Fig. 6h)

Polypedates raniceps W. C. H. PETERS. 1871. Monatsber. Preuss. Akad. Wiss. Berlin 1871:580.

M a t e r i a l. – Frog Pond (ID-8188; ID-8628; ID-8641; ID-8859); Sungei Rayu; stream east of
Sungei Rayu (pers. obs., AJA). Kubah, without precise data (UNIMAS 005–6; UNIMAS 028;
UNIMAS 031; UNIMAS 050). Also “Matang“ (INGER 1966:183: as Rana chalconota).

D e s c r i p t i o n. – SVL to 60 mm; habitus slender in males, relatively robust in adult

females; head pointed; toes nearly completely webbed; finger and toe tips with disks;
tympanum large; dorsum green, changeable to brown; a distinct white lip and ventral
surface of thighs reddish-pink.

Hylarana signata (GÜNTHER, 1872): Striped Stream Frog.

(Fig. 7f)

Polypedates signatus A. C. L. GÜNTHER. 1872. Proc. Zool. Soc. London 1872:600; Pl. 40; Fig. C.
160 I. DAS, A. JANKOWSKI, M. I. B. MAKMOR & A. HAAS

Fig. 7a. Brown Marsh Frog (Hylarana baramica; Mulu National Park) b, Rough-sided Frog
(Hylarana glandulosa; Mulu Natl. Park); c, Side-spotted Swamp Frog (Hylarana
laterimaculata); d, Mahogany Frog (Hylarana luctuosa); e, Spotted Stream Frog (Hylarana
picturata); f, Striped Stream Frog (Hylarana signata; specimen from Mulu Natl. Park);
g, Matang Stream Frog (Meristogenys jerboa); h, Black-spotted Rock Frog (Staurois guttatus).
 Amphibians of Matang Range 161

M a t e r i a l. – Summit Trail to Gunung Serapi, off Frog Pond (ID-8396; ID-8398; ID-8858).

D e s c r i p t i o n. – SVL to ca. 68 mm; tympanum visible; tips of digits slightly

swollen; toes over half webbed; dorsum and flanks dark brown to nearly black, with
yellow or orange spots, a red to orange stripe is starting from the snout and continues
to the end of the body without interruption, separating on both sides the dorsum from
the flanks; venter grey

Meristogenys jerboa (GÜNTHER, 1872): Matang Torrent Frog.

(Fig. 7g)

Hylorana jerboa A. C. L. G. GÜNTHER. 1872. Proc. Zool. Soc. London 1872:599.

M a t e r i a l. – Sungei Kubah, off Summit Trail to Gunung Serapi (ID-8238); Sungei Rayu
(AJA-003 and AJA-009); stream east of Sungei Rayu; Kubah, without precise data (UNIMAS 036;
UNIMAS 039; UNIMAS 168). Also, SM D.a.1.2.13 (no data); SM uncatalogued (ex-RFI 76534),
from “Sungei China, Matang“ collected 28.7.1956 by R. F. INGER. “Matang“ cited by INGER
(1966:266).

D e s c r i p t i o n. – SVL to 82 mm; habitus slender; tympanum diameter ½ or less eye

diameter; outer metatarsal tubercle present; toes fully webbed; posterior rim of
tympanum at level of angle of jaws in males; dorsum medium brown; limbs with dark
cross-bars and ventral surface of tibia either unpigmented or with a few scattered dark
pigments.

Staurois guttatus (GÜNTHER, 1858): Black-spotted Rock Frog.

(Fig. 7h)

Ixalus guttatus A. C. L. G. GÜNTHER. 1858. Cat. Batr. Sal. Brit. Mus.:76; Pl. 4, Fig. D.

M a t e r i a l. – Off Summit trail (ID-8846); Sungei Rayu (AJA-026); stream east of Sungei
Rayu Kubah, without precise data (UNIMAS 019; UNIMAS 026–27). Also, SM D.a.1.2.16.k
(6.1902) from “Matang“. Also listed for Matang by SMITH (1925) and INGER (1966: 250: as
Staurois natator).

D e s c r i p t i o n. – SVL to 55 mm; habitus slender; snout pointed; skin of dorsum

granualr with some rounded warts; toes fully webbed; tips of fingers and toes broad
disk-like, with white or cream margin; dorsum olive green to brown (especially in adult
females), with large black spots, changeable to olive-green when stressed; webbing of
toes turquoise blue and venter cream.

Rhacophoridae

Nyctixalus pictus (PETERS, 1871): Cinnamon Frog.

(Fig. 8a)

Ixalus pictus W. C. H. PETERS, 1871. Monatsber. Preuss. Akad. Wiss. Berlin 1871:580.

M a t e r i a l. – Summit Trail, above Frog Pond (ID-8864); Waterfall Trail (AJA-017); ephemeral
Pond, Stream east of Sungei Rayu (pers. obs., AJA).
162 I. DAS, A. JANKOWSKI, M. I. B. MAKMOR & A. HAAS

D e s c r i p t i o n. – SVL to 35 mm; habitus slender; hind limbs long; tympanum large,

visible; tips of fingers and toes expanded; toes half webbed; skin on forehead and back
rough; dorsum and forehead cinnamon, orange or red, white spotted; iris white in dorsal
third and venter cream.

Philautus tectus DRING, 1987: Obscure Bush Frog.

(Fig. 8b)

Philautus tectus J. C. M. DRING, 1987. Amphibia-Reptilia 8:30; Fig. 1.

M a t e r i a l. – Summit Trail at region of Frog Pond (AJA-011). Many observations by AJA,

always in association with Resam fern (Dicranopteris linearis).

D e s c r i p t i o n. – SVL to 27.3 mm; habitus slender; head narrow; lores deeply

concave; canthus prominent; hind limbs long; nuptial pads large; tips of fingers and
toes expanded into disks; toes half webbed; back weakly tuberculate; dorsum tan or
yellow, changeable under stress to mottled grey and brown; a light-edged dark inter-
orbital bar and venter greyish-cream.

Philautus sp.: Bush Frog.

(Fig. 8c)

M a t e r i a l. – On trail to Sungei Rayu (ID-8528).

R e m a r k s. – The systematic status of this species is currently under study.

Polypedates leucomystax (GRAVENHORST, 1829): Four-lined Tree Frog.

(Fig. 8d)

Hyla leucomystax J. L. C. GRAVENHORST. 1829. Delic. Mus. Zool.Vratislav. 1:26.

M a t e r i a l. – Frog Pond (ID-8186–87; ID-8651); > 300 m asl, Summit Trail to Gunung Serapi
(ID-8478–79). Also, SM D.a.1.3.1 from “Matang“, collected 9.1894 by A. MC. D. GIBSON.
“Matang“ listed by SMITH (1925).

D e s c r i p t i o n. – SVL to 75 mm; habitus slender in males, relatively robust in

females; snout profile rounded; fingers free of web; no fringe of skin along arm; no
tubercle on heel; skin smooth; colour of dorsum variable, ranging from yellow, olive to
brownish-yellow, with or without dark lines on dorsum and venter cream.

Polypedates macrotis (BOULENGER, 1891): Dark-eared Tree Frog.

(Fig. 8e)

Rhacophorus macrotis G. A. BOULENGER. 1891. Ann. & Mag. nat. Hist. ser. 6 14:282.

M a t e r i a l. – Frog Pond (ID-8185; ID-8636; ID-8649); > 300 m asl, Summit Trail to Gunung
Serapi (ID-8477); stream east of Sungei Rayu (pers. obs., AJA). Kubah, without precise data
(UNIMAS 110; UNIMAS 152).
 Amphibians of Matang Range 163

Fig. 8a. Cinnamon Frog (Nyctixalus pictus); b, Obscure Bush Frog (Philautus tectus); c, Bush
Frog (Philautus sp.); d, Four-lined Tree Frog (Polypedates leucomystax); e, Dark-eared Tree Frog
(Polypedates macrotis); f, File-eared Tree Frog (Polypedates otilophus); g, Frilled Tree Frog
(Rhacophorus appendiculatus); h, Gunung Gading Tree Frog (Rhacophorus gadingensis).
164 I. DAS, A. JANKOWSKI, M. I. B. MAKMOR & A. HAAS

D e s c r i p t i o n. – SVL to 90 mm; habitus slender in males, relatively robust in

females; eyes large; tips of fingers and toes disk-like; skin smooth; dorsum and forehead
dark brown; a dark brown band from nostril through eyes and across tympanum to mid-
flank and venter cream.

Polypedates otilophus (BOULENGER, 1893): File-eared Tree Frog.

(Fig. 8f)

Rhacophorus otilophus G. A. BOULENGER. 1893. Proc. Zool. Soc. London 1893: 527; Pl. 44.

M a t e r i a l. – Frog Pond (ID-8176–78; ID-8265); Kubah, without precise data (UNIMAS

085). Also, “Matang“ (SM D.a.1.3.6.f), 14.10.1893, collected by A. McD. Gibson. “Matang“ in
INGER (1966: 329).

D e s c r i p t i o n. – SVL to 97 mm; habitus robust; a bony ridge over tympanum; head

large, triangular; tips of fingers and toes disk-like; toes half webbed; a fleshy projection
at heel; dorsum and forehead bright yellow to yellowish brown, with thin dark lines and
venter cream.

Rhacophorus appendiculatus (GÜNTHER, 1859): Frilled Tree Frog.

(Fig. 8g)

Polypedates appendiculatus A. C. L. G. GÜNTHER. 1859. Cat. Batr. Sal. British Mus.: 79.

M a t e r i a l. – Frog Pond (ID-8635; ID-8639); a large calling group at the ephemeral Pond
(pers. obs., AJA).

D e s c r i p t i o n. – SVL to 50 mm; habitus slender; head triangular; males with a

conical snout-tip; females with a conical snout-tip; toes with some free webbing; a
narrow wavy fringe of skin along limbs; a series of serrated dermal projections ventrally
along mandible; dorsum variable and also changeable, ranging from olive grey to brown,
with darker markings; some frogs have a pink tinge on front edge of thighs and venter
cream with a yellow tinge.

Rhacophorus gadingensis DAS & HAAS, 2005: Gunung Gading Tree Frog.
(Fig. 8h)

Rhacophorus gadingensis I. DAS & A. HAAS. 2005. Raffles Bull. Zool. 53:258; Figs 1-2.

M a t e r i a l. – Summit Trail to Gunung Serapi, off Frog Pond (ID-8394) and Sungei Rayu
(AJA-049). New record for the Matang Range, and the first record from outside the type locality
(Gunung Gading).

D e s c r i p t i o n. – SVL to 29.5 mm; habitus slender; head wider than long; fingers
lacking nuptial pads; a dermal flap along forearm; spinose calcar present; supratympanic
fold weak; dorsum brown, with a dark interorbital bar; iris golden with copper tint
around pupil; dark brown blotches on dorsum and flanks with blue blotches and venter
yellow-cream.
 Amphibians of Matang Range 165

Rhacophorus kajau DRING, 1984: White-eared Tree Frog.

(Fig. 9a)

Rhacophorus kajau J. C. M. DRING. 1984. Amphibia-Reptilia 4: 112; Fig. 3.

M a t e r i a l. – ca. 100 m from Chalet, on Summit Trail to Gunung Serapi, off Frog Pond
(ID-8264; ID-8522; ID-8642; ID-8852; AJA-021). Also at the stream south of the Frog Pond
(pers. obs., AJA).

D e s c r i p t i o n. – SVL to 22.6 mm; habitus slender; webbing on fingers rudimentary;

narrow crenulated dermal ridge along outer edges of hands, feet, tarsus and postcloacal
region; vomerine teeth absent; dorsum leaf green, with sharp transition to cream at mid-
flank; tiny white spots present on dorsum; white spots on lips, tympanum and flanks
and venter cream.

Rhacophorus nigropalmatus BOULENGER. 1895: Wallace’s Flying Frog.

(Fig. 9b)

Rhacophorus nigropalmatus G. A. BOULENGER. 1895. Ann. & Mag. Nat. Hist., Ser. 6, 16:170.

M a t e r i a l. – Frog Pond (ID-8733; UNIMAS 082).

D e s c r i p t i o n. – SVL to 100 mm; habitus robust; snout rounded; tips of fingers and
toes disk-like, fully webbed; fleshy projection on heel; dorsum bright green, with white
spots; webbing black margined with yellow; iris pale yellow venter yellowish-cream.

Rhacophorus pardalis GÜNTHER, 1859: Harlequin Tree Frog.

(Fig. 9c)

Rhacophorus pardalis A. C. L. G. GÜNTHER. 1859. Cat. Batr. Sal. British Mus.: 83; Pl. 6; Fig. D.

M a t e r i a l. – Frog Pond (ID-8175; ID-8399–8400; ID-8850); stream south of Frog Pond;

ephemeral pond (pers. obs., AJA). Kubah, without precise data (UNIMAS 079).

D e s c r i p t i o n. – SVL 75 mm; habitus robust; snout rounded; tips of fingers and

toes disk-like; fingers and toes fully webbed; fleshy flap at heel; forehead and dorsum
red, pink or reddish brown, with darker markings; webbing orange to red; iris tan around
pupil, lighting up to gold in the periphery and venter yellow with red areas.

Ichthyophiidae

Caudacaecilia asplenia (TAYLOR, 1965): Broad-striped Caecilian.

(Fig. 9d)

Ichthyophis asplenius E. H. TAYLOR. 1965. Univ. Kansas Sci. Bull. 46:278.

M a t e r i a l. – One specimen, an adult, was collected from a sluggish stream within Matang
Wildlife Centre, within a patch of kerangas forest (ZRC 1.10835, ex-MW 3789).
166 I. DAS, A. JANKOWSKI, M. I. B. MAKMOR & A. HAAS

Fig. 9a. White-eared Tree Frog (Rhacophorus kajau); b, Wallace’s Flying Frog (Rhacophorus
nigropalmatus); c, Harlequin Tree Frog (Rhacophorus pardalis); d, Broad-striped Caecilian
(Caudacaecilia asplenia).

D e s c r i p t i o n. – SVL to 204 mm; length 23-26 times body width; total annuli 247-270;
grooves and folds do not cross back except in posterior half of body; splenial teeth
absent in adults; dorsum bluish-grey, with a yellow lateral stripe.

Ichthyophis biangularis TAYLOR, 1965: Angular Caecilian

Ichthyophis biangularis E. H. TAYLOR. 1965. Univ. Kansas Sci. Bull. 46:272.

M a t e r i a l. – No specimens were collected by us. “Matang“ is the type locality of this

species (BMNH 72.2.19.59A; holotype), although further details, such as altitudinal distribution
and precise locality, remain unknown.

D e s c r i p t i o n. – SVL to 253 mm; length 26 times body width; primary and

secondary annuli form angles pointed forward dorsally; total annuli 330-333; annuli on
tail 10; dorsum blackish-slate with a yellow lateral stripe.

Results and Discussion

S p e c i e s d i v e r s i t y. – Currently, 55 species of amphibians (53 species of Anura
and two of Gymnophiona) are recorded from the Matang Range (Table 1). This
represents about a third of the Bornean amphibian fauna (DAS & HAAS 2005). Two
amphibian species, Pelophryne guentheri and Ichthyophis biangularis, are endemic to
this mountain massif, and two additional species (Microhyla and Philautus) are new to
science and will be formally described elsewhere.
 Amphibians of Matang Range 167

Table 1. Species list and reproductive modes shown by the amphibian community at the Matang
Range, Sarawak, Malaysia. Definitions in CRUMP (1974, modified by DUELLMAN & TRUEB, 1986).
Sl. Species Reproductive Mode
____________________________________________________________________
Bufonidae
1. Ansonia leptopus 2 – egg and feeding larvae, lotic waters
2. Ansonia longidigita 2 – egg and feeding larvae, lotic waters
3. Ansonia minuta 2 – egg and feeding larvae, lotic waters
4. Ansonia spinulifer 2 – egg and feeding larvae, lotic waters
5. Duttaphrynus melanostictus 1 – egg and feeding larvae, lentic waters
6. Ingerophrynus divergens 1 – egg and feeding larvae, lentic waters
7. Pedostibes hosii 2 – egg and feeding larvae, lotic waters
8. Pelophryne guentheri ? (possibly Mode 6)
9. Pelophryne signata 6
10. Phrynoidis aspera 2 – egg and feeding larvae, lotic waters
11. Phrynoidis juxtaspera 2 – egg and feeding larvae, lotic waters

Ceratobatrachidae
12. Ingerana baluensis ?

Dicroglossidae
13. Fejervarya limnocharis 1 – egg and feeding larvae, lentic waters
14. Limnonectes kuhlii 2 – egg and feeding larvae, lotic waters
15. Limnonectes laticeps 1 or 2 – egg and feeding larvae, lotic or lentic waters
16. Limnonectes leporinus 3 – eggs in constructed basin, stream
17. Limnonectes malesianus 2 – egg and feeding larvae, lotic waters
18. Limnonectes palavanensis 14 – terrestrial eggs, larval transport, feeding larvae
19. Limnonectes paramacrodon ? (possibly Mode 2)
20. Occidozyga baluensis 1 – egg and feeding larvae, lentic waters
21. Occidozyga laevis 1 – egg and feeding larvae, lentic waters

Megophryidae
22. Leptobrachium abbotti 2 – egg and feeding larvae, lotic waters
23. Leptobrachella mjobergi 2 – egg and feeding larvae, lotic waters
24. Leptolalax gracilis 2 – egg and feeding larvae, lotic waters
25. Megophrys nasuta 2 – egg and feeding larvae, lotic waters

Microhylidae
26. Calluella brooksii ?
27. Chaperina fusca 4 – phytothelm, feeding larvae
28. Kalophrynus heterochirus ? (possibly Mode 4)
29. Kalophrynus pleurostigma 4 – phytothelm, feeding larvae
30. Metaphrynella sundana ? – phytothelm, feeding unclear
31. Microhyla borneensis 1 – egg and feeding larvae, lentic waters
32. Microhyla sp. 6 – phytothelm, endotrophic larva

Ranidae
33. Hylarana baramica ? (possibly Mode 1)
34. Hylarana erythraea 1 – egg and feeding larvae, lentic waters
35. Hylarana glandulosa 1 – egg and feeding larvae, lentic waters
36. Hylarana laterimaculata ? (possibly Mode 1)
37. Hylarana luctuosa 1 – egg and feeding larvae, lentic waters
38. Hylarana picturata 2 – egg and feeding larvae, lotic waters
39. Hylarana raniceps 1 or 2 – egg and feeding larvae, lotic or lentic waters
40. Hylarana signata 2 – egg and feeding larvae, lotic waters
41. Meristogenys jerboa 2 – egg and feeding larvae, lotic waters
42. Staurois guttatus 2 – egg and feeding larvae, lotic waters
168 I. DAS, A. JANKOWSKI, M. I. B. MAKMOR & A. HAAS

Rhacophoridae
43. Nyctixalus pictus 4 – phytothelm, feeding larvae
44. Philautus tectus 17 – direct development
45. Philautus sp. ? (possibly Mode 17 – direct development)
46. Polypedates leucomystax 8 – foam on pond, feeding tadpoles in pond
47. Polypedates macrotis 23– arboreal foam nest, feeding tadpoles in pond
48. Polypedates otilophus 23 – arboreal foam nest, feeding tadpoles in pond
49. Rhacophorus appendiculatus ? – feeding tadpoles in pond
50. Rhacophorus gadingensis 23 – arboreal foam nest, feeding tadpoles in stream
51. Rhacophorus kajau 18 – arboreal nest, feeding tadpoles in pond/stagnant stream
52. Rhacophorus nigropalmatus 23 – arboreal foam nest, feeding tadpoles in pond
53. Rhacophorus pardalis 23 – arboreal foam nest, feeding tadpoles in pond

Ichthyophiidae
54. Caudacaecilia asplenia ?
55. Ichthyophis biangularis ?

Direct comparisons of species richness across sites in Borneo are difficult, given the
size difference in areas that have been sampled and the unequal time expended in such
inventory activities, not to mention the rather unequal experience of different field
collectors and timing (and hence, local weather conditions) of collection. Nonetheless,
such comparisons can be useful in understanding the amphibian richness of sites for
conservation purposes, such as inventories of protected and other geographical areas.
INGER & VORIS (1993) examined species richness of riparian frogs from 18 localities in
Sabah and Sarawak, and found total species counts in the range 11-27. Matang therefore
has a relatively large riparian assemblage – 25 species. Non-riparian components of the
fauna also contribute significantly to the overall species count, and at Matang, included
species with endotrophic tadpoles (those of the genus Pelophryne and one species of
Microhyla) and direct-developers (genus Philautus).
E l e v a t i o n a l d i s t r i b u t i o n. – A total of 100 individuals comprising 15
species from five families of amphibians were collected from Gunung Serapi, from 201 to
800 m above sea level. Combined with the earlier collections, a total of 22 species were
recorded below 200 m elevation. These numbers indicate that mixed dipterocarp forests
of Gunung Serapi has a diverse amphibian fauna.
Table 2 shows diversity indices based on the amphibian distribution data recorded
for the three altitudinal bands (201-400 m, 401-600 m and 601-800 m). A noteworthy result
is that frog diversity decreased with elevation. ZAR (1996)’s modified t-test shows that
species diversity between altitudinal band 2 (201-400 m) and altitudinal band 4 (601-800 m)
are significantly different (Table 2). A total of 19 species of anurans have been recorded
from altitudinal band 1 (below 200 m). This zone showed the highest species richness
compared to the rest of the three altitudinal bands. Of these, 11 were also recorded from
higher elevations. Species not recorded beyond 200 m include three species of
Bufonidae (Ansonia spinulifer, Ingerophrynus divergens and Duttaphrynus
melanosticus), two species of Megophryidae (Leptolalax gracilis and Megophrys
nasuta), two species of Ranidae (Hylarana raniceps and Staurois guttatus), and one
species of Microhylidae (Metaphrynella sundana). The highest number of individuals
of all frogs were also found at altitudinal band 2 (201-400 m), with 58 individuals. With
 Amphibians of Matang Range 169

Table 2. Summary of diversity indices based on the data recorded for the three altitudinal bands
sampled on Gunung Serapi, Matang Range, Sarawak.
_______________________________________________________________________________________________________________________
Diversity Indices Altitudinal band 2 Altitudinal band 3 Altitudinal band 4
(201–400) m (401–600) m (601–800) m
_______________________________________________________________________________________________________________________
Simpson’s 0.994 0.692 0.533
Shannon-Weiner (H’) 3.807 1.585 1.000
_________________________________________________________________________________________________________________________

increase in elevations, there is a distinct decrease in the abundance of individuals with

fewer individuals captured at altitudinal band 3 (27 individuals) and the lowest numbers
of individuals were recorded at altitudinal band 4 (15 individuals) (Fig. 10), similar to
species richness values. Band 4 had the lowest species richness value, with just two
species recorded between 601-800 m (Fig. 10).
Not all species recorded from the Range were found in the elevational transects,
which may be due to a combination of factors, such as time constraints (more effort
expended in encounter surveys in forested areas), the focus of the transect along a
relatively open area and small sampling volume (2-3 m of the transect) and the exclusion
of stream banks and leaf litter from most of the area covered during the transect
sampling. The most abundant species, representing the bulk of the transect capture,
belong to Limnonectes laticeps, with 52 individuals, and was recorded at all four
elevational bands. At the study site, this species seems to have taken the opportunity
of the road-side rainwater drains to breed. In other areas, it has been recorded to breed
on the banks of small and intermittent streams.

Fig. 10. Relationship between amphibian diversity, amphibian species richness, and elevation on
Gunung Serapi, Matang Range, Sarawak.
170 I. DAS, A. JANKOWSKI, M. I. B. MAKMOR & A. HAAS

Table 3. Summary of results of Zar’s (1996) modified t-tests, comparing species diversity
between altitudinal bands on Gunung Serapi, Matang Range, Sarawak.
_________________________________________________________________________________________________________________________
Altitudinal bands H’ x
H’ y
t cal.
t crit.
Conclusion
_________________________________________________________________________________________________________________________
2 vs 3 3.807 1.585 -0.4752 -1.96 Not significant
2 vs 4 3.807 1.000 -7.80503 -2.000 Significant
3 vs 4 1.585 1.000 -1.0269 -2.021 Not significant
_________________________________________________________________________________________________________________________

Species richness decreased with elevation on Gunung Serapi, similar to the findings
by DANIELS (1992) in the Western Ghats, India, and by many others, which highlight
that portions of hills of low to medium elevation are more likely to have higher diversity
of amphibians than higher elevations. According to NAVAS (2003), the most important
factor that control amphibian species diversity is humidity. In a study of the montane
amphibian fauna of north-western Borneo, INGER & STUEBING (1992) found that
environmental changes with altitude affect amphibians. Changes in vegetation alter the
temperature, soil moisture, and the abundance of particular breeding sites. Restriction of
the variety of potential breeding sites and microhabitats at higher elevations are
therefore linked to the decrease in species richness with elevation. HEYER (1967) also
showed close correlation between vegetation types and herpetofaunal assemblages.
However, since all altitudinal bands in this study coincided with mixed dipterocarp
forest, vegetional changes cannot be attributed to the changes in species richness and
diversity observed. At the Serapi site, there were fewer streams and water resources at
higher elevations, and most major streams and rivers are found < 400 m asl. A large pond
located at altitudinal band 2 (ca. 269 m asl) is utilized for breeding by at least seven
species (including Microhyla borneensis, Hylarana raniceps, Hylarana luctuosa,
Polypedates leucomystax, Polypedates macrotis, P. otilophus and Rhacophorus pardalis).
Other species recorded at the Range and likely to breed include Hylarana erythraea,
Limnonectes palavanensis, Limnonectes paramacrodon, Rhacophorus appendiculatus,
R. nigropalmatus, Caudacaecilia asplenia and Ichthyophis biangularis.
Thus, higher elevations on Serapi have fewer potential breeding sites, effecting
species diversity and richness. Additionally, elevational gains are concurrent with
reduced land area and typically, reduced habitat complexity (HEATWOLE 1982). Another
contributing factor leading to higher species richness and diversity of amphibians at
lower elevations is the rapid run-off of rainfall, caused by the steepness of the terrain
(INGER & STUEBING 1992).
Notwithstanding relatively low species richness and diversity values, montane regions,
whose existence is threatened by anthropogenic factors (see BROOKS et al. 1999 and
references therein) are important in supporting high levels of genetic divergence
(ROBERTS et al. 2006), endemic species (DAS 2006), novel ecological processes
(MORRISON & HERO 2003), and altitudinal species turnover is considered more important
in regional species richness, rather than geographic distribution (VASUDEVAN et al. 2006).
R e p r o d u c t i v e m o d e s. – CRUMP (1974) studied a New World species-rich site
(Santa Cecilia in Ecuador, with 81 amphibian species) and characterized the anuran
amphibian fauna in terms of several quantitative reproductive variables that many
subsequent workers have used. DUELLMAN & TRUEB (1986) revised the scheme of
 Amphibians of Matang Range 171

classification, and recognised 29 reproductive modes shown by anurans worldwide.

Reproductive data for the amphibian fauna of Matang are incomplete but permit a
preliminary characterisation of the reproductive modes seen in the assemblage (Table 1).
Nine distinct reproductive modes are shown by the fauna. The most common is
Reproductive Modes 2 (eggs and feeding tadpoles in lotic water, with 19 species).
Reproductive Mode 1 (eggs and feeding tadpoles in lentic water, with 12 species) was
next common. Few species reproduced in more than one microhabitat. Examples include
Hylarana raniceps and Limnonectes laticeps which were found breeding in both lentic
and lotic habitats (hence, both Modes 1 and 2). In the latter species, the occupancy of
lentic habitats by the larval stages may occur during locally dry periods as it inhabits
ephemeral streams or mountain brooks.
Some correlations between Reproductive Modes and recognised lineages are evident.
For instance, members of the Bufonidae predominantly show Mode 2; Dicroglossidae
mostly Mode 1 or 2; Megophryidae, Mode 2; Ranidae, Modes 1 or 2; and Rhacophoridae,
Mode 23. A major divergence from Mode 23 (arborial foam nest; feeding tadpoles) is
shown by Rhacophorus kajau, which does not produce foam, but retains its eggs in a
coat of jelly, applied to leaves overhanging water. Other significant divergences from
the typical mode in the Rhacophoridae are shown by Nyctixalus pictus (Mode 4– eggs
and feeding larvae in tree holes or aerial plants, i.e., phytothelmata), Philautus tectus
(Mode 17– eggs hatch into froglets, with no free-swimming larval stage) and
Rhacophorus gadingensis (Mode 9– foam nest on leaf and feeding tadpoles in stream).
The causal factors for the great diversity of reproductive modes known in the living
anurans worldwide (for example, DUELLMAN 1992) has been attributed to both
phylogenetic diversity and ecological processes, particularly, resource partitioning
(CRUMP 1974), and the latter factor may be one of the mechanisms permitting high
species richness of amphibians at the Matang Range of Sarawak.

ACKNOWLEDGEMENTS. Permission and facilities to conduct field work in Sarawak was

provided by DATUK CHEONG EK CHOON, Director and Controller of National Parks and Nature
Reserves, Sarawak Forestry Corporation, permit numbers 5/2004, dated 06.07.2004; 9/2005,
dated 18.8.2005 and 38/2006, dated 3.8.2006. We are also grateful to LIM CHAN KOON and
OSWALD BRAKEN TISEN for advice on permitting and logistics and especially to the staff of
Kubah National Park and Matang Wildlife Centre for permitting us to work in areas under their
jurisdiction, and for their advice on localities and other help. In particular, we would like to name
MOHIDIN RAJULI, Park Manager and SUZIANI BINTI SULAIMAN, Conservation Officer at Kubah
National Park and SAIP SULONG, Park Manager, Matang Wildlife Centre, for their support. We
thank our respective institutions, the Institute of Biodiversity and Environmental Conservation,
Universiti Malaysia Sarawak, and Hamburg University, for supporting our researches on the
herpetology of south-east Asia.
Financial support for ‘Inventory and biodiversity of the frog fauna of East Malaysia (Borneo)
with emphasis on their larval forms (Amphibia: Anura)’ was received from Volkswagen-Stiftung,
Germany (grant number: 1/79 405). In the field, we thank our colleagues and students for
assistance in collecting and preparing specimens and providing help in the field, including DAVID
GOWER, GENEVIEVE V. A. GEE, LIM CHAN KOON, SAIBAL SENGUPTA, LILY ANAK SIR, JEET
SUKUMARAN, SANDRA JAMES TINGSOM AND NORSHAM BINTI YAAKOB. Special thanks to JEET
SUKUMARAN for preparing the maps in Fig. 1. We thank CHARLIE J. LAMAN for statistical
advice. Material from the BMNH, SM and UNIMAS could be examined, for which we are
grateful to C. J. MCCARTHY, C. M. LEH and FATIMAH ABANG. Finally, we thank GENEVIEVE
GEE and especially TZI MING LEONG for comments on the manuscript.
172 I. DAS, A. JANKOWSKI, M. I. B. MAKMOR & A. HAAS

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 174 I. DAS, A. JANKOWSKI, M. I. B. MAKMOR & A. HAAS

APPENDIX 1
Output data of computations on ZAR’S (1996) modified t-tests.

Altitudinal band
4 (601–800 m) 3 (401–600 m) 2 (201–400 m )
fi (total) 15 27 58
fi x Log (fi) 13.49485002 29.50581425 52.48791132
fi x Log(power 2) (fi) 12.44279533 33.93328622 61.69014846
n (log(n)) 17.64136889 38.64682163 102.2788236
fi x (Log (fi))2 182.1109771 870.5930745 2754.980835
n2 225 729 3364

H1 0.276434591
S2 H1 0.001342505
(S2 H1)2 1.80232E-06
(S2 H1)2/n 1.20155E-07
(S2 H1+S2 H2) 0.003659512
(S2 H1+S2 H2)2 1.3392E-05

H3 0.858464005
S2 H3 0.004218335
(S2 H3)2 1.77944E-05
(S2 H3)2/n 3.06799E-07
(S2 H2+S2 H3) 1.197019834
(S2 H2+S2 H3)2 1.432856484

H2 0.338555829
S2 H2 0.002317007
(S2 H2)2 5.36852E-06
(S2 H2)2/n 1.98834E-07
(S2 H1+S2 H3) 0.00556084
(S2 H1+S2 H3)2 3.09229E-05

Received: 01 October 2007; accepted: 30 October 2007.

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