AMPHIBIANS & REPTILES

of
MOUNT KINABALU
(North Borneo)

Rana decorata MOCQUARD, 1890

Calamaria lateralis MOCQUARD, 1890

Pelturagonia cephalum MOCQUARD, 1890

after MOCQUARD 1890

AMPHIBIANS & REPTILES
of
MOUNT KINABALU
(North Borneo)

Rudolf Malkmus, Ulrich Manthey, Gernot Vogel,

Peter Hoffmann & Joachim Kosuch

with 384 photographs

by

C. Brühl, C.L. Chan, I. Das, W. Denzer, T. Eltz, R. Gerhardt, A. Götzke,

W. Grossmann, T. Hikida, W. Hödl, D. Knowles, U. Kuch, M. Kunkel, B. Leideritz,
K.E. Linsenmair, W. Malkmus, S. Manthey, M. Maronde, M. Matsui, J.C. Murphy,
A. Nöllert, M. Pfeiffer, H. Sauer, M. Schroth, R.B. Stuebing,
T. Ulber, M. Veith, S. Wilson, and the authors
Frontcover: Backcover:
Philautus aurantium © R. Malkmus Trimeresurus cf. sumatranus © K.E. Linsenmair

Address of the authors

Rudolf Malkmus, Schulstr. 4, D-97859 Wiesthal, Germany

(Introduction, Mount Kinabalu - the surrounding environment, Amphibia (with exception of the family Ranidae))

Dipl. Ing. Ulrich Manthey, Kindelbergweg 15, D-12249 Berlin, Germany; Email: manthey.sseah@t-online.de
(Sauria (lizards) and Testudines (turtles, tortoises, and terrapins))

Dr Gernod Vogel, Im Sand 3, D-69115 Heidelberg, Germany; Email: Gernot.Vogel@t-online.de

(Serpentes (snakes))

Dipl. Biol. Peter Hoffmann, Im Oberdorf 41, D-79292 Pfaffenweiler, Germany; Email: dorin.hoffmann@t-online.de
Joachim Kosuch, Im Plänzert 39, D-53474 Neuenahr-Ahrweiler, Germany; Email: joachim@falco.biologie.uni-mainz.de
(Amphibia, family Ranidae)

Translation from the German manuscript (with some exeptions) by Herprint International CC,
P.O.Box 14241, Bredell 1623, South Africa

ISBN 3-904144-83-9
© 2002 A.R.G. Gantner Verlag Kommanditgesellschaft, FL 9491 Ruggell

Distributed by
Koeltz Scientific Books, Herrnwaldstr. 6, D-61462 Königstein / Germany
Fax: (+49) 6174 937240 Email: koeltz@t-online.de Phone: (+49) 617493720
Internet: http://www.koeltz.com

Layout by Sibylle Manthey, Berlin / Germany

Design by Ulrich Manthey, Berlin / Germany
Printed by Fuldaer Verlagsanstalt, D-36037 Fulda / Germany
 Contents

Contents

Preface .......................................................................................................................................... 7
Acknowledgements ................................................................................................................... 8
How to use this book ................................................................................................................ 9
Map of Borneo .......................................................................................................................... 11
Introduction ............................................................................................................................... 12

Mount Kinabalu - the surrounding environment ......................................................... 13

Topography, Geology, Geomorphology ................................................................................ 13
Zoogeography ........................................................................................................................ 15
Climate ................................................................................................................................... 16
Vegetation .............................................................................................................................. 20
Tropical lowland vegetation zone ( to 900 m a.s.l.) ............................................................. 21
Tropical lower montane vegetation zone (900-1800 m a.s.l.) ............................................. 22
Tropical upper montane vegetation zone (1800-3000 m a.s.l.) ............................................ 23
Tropical subalpine vegetation zone (3000-3700 m a.s.l.) .................................................... 25
Fauna ..................................................................................................................................... 28
Herpetofauna ......................................................................................................................... 32
Human beings in the vicinity of Mount Kinabalu ................................................................ 38
A retrospective glance on herpetological research in Borneo and the area of Mt. Kinabalu ..... 40

Amphibia (amphibians) ......................................................................................................... 46

Anura (tailless amphibians or frogs) .................................................................................. 46
Important traits for the identification of frogs and their larvae ............................................ 48
Morphology ........................................................................................................................... 49
Habitats of tadpoles ............................................................................................................... 53
Larval feeding types .............................................................................................................. 54
Bioacoustics ........................................................................................................................... 56
Reproductive strategies ......................................................................................................... 58
Key to the families of Bornean frogs .................................................................................... 60
Preliminary key to the larvae of Bornean frog families ....................................................... 61
Bufonidae .......................................................................................................................... 62
Megophryidae ................................................................................................................... 88
Microhylidae ..................................................................................................................... 115
Ranidae .............................................................................................................................. 133
Rhacophoridae .................................................................................................................. 175
Gymnophiona .......................................................................................................................... 218
Ichthyophiidae ................................................................................................................... 218

5
Contents

Reptilia (reptiles) ...................................................................................................................... 220

Squamata (scaly reptiles) ...................................................................................................... 220
Sauria (lizards) ................................................................................................................... 221
Distinguishing features in lizards ..................................................................................... 222
Key to the families of Bornean lizards .............................................................................. 223
Agamidae ...................................................................................................................... 224
Anguidae ....................................................................................................................... 250
Gekkonidae ................................................................................................................... 251
Scincidae ....................................................................................................................... 269
Varanidae ...................................................................................................................... 293
Serpentes (snakes) ............................................................................................................... 297
Distinguishing features in snakes ..................................................................................... 298
Key to the families of Bornean snakes ............................................................................. 299
Colubridae ..................................................................................................................... 300
Crotalidae ...................................................................................................................... 373
Cylindrophiidae ............................................................................................................ 380
Elapidae ........................................................................................................................ 382
Pythonidae .................................................................................................................... 389
Typhlopidae .................................................................................................................. 392
Xenopeltidae ................................................................................................................. 393
Testudines (turtles, tortoises, and terrapins) .................................................................... 395
Distinguishing features in turtles .......................................................................................... 395
Key to the families of Bornean turtles .................................................................................. 395
Bataguridae ....................................................................................................................... 396
Trionychidae ...................................................................................................................... 399
Glossary ....................................................................................................................................... 401
Bibliography .............................................................................................................................. 403
Index ............................................................................................................................................. 417

6
 Preface

Preface
Compared with previous times, the substantially improved possibilities for travelling have enabled an
increasingly large number of people to reach other countries or even distant continents. Amongst these
are a number who, be it at home or outdoors, spend their spare time with intense studies of certain groups
of animals and/or plants. Small by comparison are the numbers of those who are able to study as a
profession subjects such as behaviour, systematics and evolution, or the geographical distribution and
ecology of animals and plants. For many of these, and other nature lovers, journeys to poorly known
zoogeographical zones will only be sensible if they are able to recognise at least a portion of the biologi-
cal species living there. Enabling this is one of the main goals of this book. The authors have dealt in the
main chapters of their treatise with all species of amphibians and reptiles currently known to exist on
Mount Kinabalu. Rising to 4,095 m Mount Kinabalu is the highest mountain between the Himalayas and
the alpine mountain ranges of New Guinea. It is situated in the north of the tropical island of Borneo, in
the Malayan state of Sabah, and is marked by a variety of ecological conditions which, in turn, have
created an extraordinary diversity of species. The visitor from central Europe will be surprised to learn
that the Kinabalu massif is home to 77 species of amphibians and 112 species of reptiles, which equals
the species diversity of the entire European continent. The present book is by no means a compilation of
existing literature published by other authors. Rather all authors except one have been to the Kinabalu
region themselves and have partly worked there for several months. They have therefore been able to
combine the results of their own studies with an in-depth analysis of the literature to create a treatise
original in character. The more than 30 pages of introductory chapters contain valuable information that
may be interesting not only to the herpetologist. The attentive reader will be particularly pleased to find:

• The thorough analysis of the relevant primary literature with synonymy lists of scientific names,
something which was, unfortunately, insufficiently attended to in many books of a similar nature
published during the past decades;
• A wealth of information, not yet otherwise published, on the behaviour and ecology of the individual
species;
• Identification keys for all herpetological taxa known to occur in the Kinabalu region;
• Excellent colour photographs of live representatives of almost all species;
• Oscillograms and audiospectrograms of the calls of many amphibians.

In order to make the book accessible to an international readership, it has not been published in the
original language of the authors, but in English. “Amphibians and Reptiles of Mount Kinabalu” places a
tool in the hands of amateur researchers as well as professional herpetologists which will not only elimi-
nate many identification problems in the field, but which can also be used for the identification of photo-
graphs and preserved material or that of recorded amphibian calls. I am convinced that, despite the recent
publication of a considerable number of checklists and some books more orientated towards the general
public on the amphibians and reptiles of this region, this book will find a justified place in the libraries of
many nature lovers. Those interested in the herpetofauna of Southeast Asia will receive it with particular
appreciation.

Berlin, January 2002

Dr. sc. Rainer Günther
Museum für Naturkunde der Humboldt-Universität zu Berlin

7
Acknowledgements

Acknowledgements

We wish to acknowledge the help of a number of people during the preparatory stages of this book. Many
people contributed information, technical assistance and organisatorial help. Many thanks to all of you:
Markus Arning, John H. Beaman, Walter Brodt, Julian Dring, Helmut Fleißner, Uwe Fritz, Klaus Kittel,
Klaus Kuhn, Peter Lenk, Nicola Lutzmann, Bernhard Malkmus, Mark-Oliver Rödel, Andreas Schlüter,
and Barbara Sein.

We are grateful to Kraig Adler, Carsten Brühl, Chan Chew Lun, Indraneil Das, Wolfgang Denzer,
Thomas Eltz, Ronald Gerhardt, Armin Götzke, Wolfgang Grossmann, Tsutomu Hikida, Walter Hödl,
David Knowles, Ulrich Kuch, Michael Kunkel, Bernd Leideritz, Karl E. Linsenmair, Walter Malk-
mus, Sibylle Manthey, Manfred Maronde, Masafumi Matsui, John C. Murphy, Andreas Nöllert, Martin
Pfeiffer, Horst Sauer, Martin Schroth, Robert B. Stuebing, Thomas Ulber, Michael Veith, and Steve
Wilson for the permission to use their photographs.

Several of our colleagues reviewed one or more sections of this book. In particular, we wish to thank
Rafe Brown, Indraneil Das, Patrick David, Wolfgang Denzer, and Jimmy McGuire.
We also acknowledge the help of Patrick David, Robert F. Inger and Van Wallach regarding taxonomical
problems.

To curatorial staff of the following institutions we owe a debt for loaning us specimens under their care:
Museum für Naturkunde der Humboldt-Universität zu Berlin (ZMB), Berlin (Rainer Günther and Detlef
Langer); British Museum of National History (BM), London (E. Nicholas Arnold and Colin McCarthy);
Field Museum of Natural History (FMNH), Chicago (Harold K. Voris and Alan Resetar); Zoologisches
Forschungsinstitut und Museum A. Koenig (ZFMK), Bonn (Wolfgang Böhme and Wolfgang Bischoff);
Forschungsinstitut Senckenberg (SMF), Frankfurt/M (Gunther Köhler); Instituut voor Taxonomische
Zoölogie (ZMA), Amsterdam (L. van Tuijl); Muséum National d’Histoire Naturelle (MNHN), Paris
(Annemarie Ohler); The Raffles Museum of Biodiversity Research, National University of Singapore
(USDZ/ZRC), Singapore (Kelvin Lim, Peter Ng and C.M. Yang); Kyoto University Zoological Collec-
tion (KUZ), Kyoto (Masafumi Matsui, Tom Hikida and Hidetoshi Ota (Okinawa)); Nationaal
Natuurhistorisch Museum (RMNH), Leiden (Marinus S. Hoogmoed and H.A.J. in den Bosch).

For the preparation of drawings we would like to thank Vera Heinrich, Jan Hnizdo, and Petr Kliment.
Maklarin Lakim, Francis Liew, Klaus Riede, Anna Wong, and Paul Yambun made our stay at Kinabalu
Park very pleasant and provided help in many ways.

We are indebted to Rainer Günther (ZMB) for presenting the preface.

Finally our heartfelt thanks to our families for their patience and support in many ways.
Last but not the least, we would like to thank Sven Koeltz, our publisher, for keeping a positive attitude
towards this publication during all stages of its production.

8
 How to use this book

How to use this book

This book is a comprehensive guide to the reptiles and amphibians occurring on Mt. Kinabalu treating all
species recorded from the area and including all related taxa known to occur on Borneo. Inclusion of all
taxa serves the purpose to identify also species, which are not yet recorded from Mt. Kinabalu. EBENHARD
& SJÖGREN (1984) listed three species of snakes for Mt. Kinabalu, omitted here, viz.: Calamaria lowi
BOULENGER, 1887, Ptyas carinatus (GÜNTHER, 1858) and Cerberus rynchops (SCHNEIDER, 1799). Since no
sources were provided for these records, the three species have been ignored in the present work. DAS &
ISMAIL (2001) listed additional lizard species for Mt. Kinabalu that have not been included here as the
compilation is still under revision (pers. comm. DAS).

After a general introduction to the orders of amphibians (Anura, Gymnophiona) and reptiles (Squamata,
Testudines) occurring on Mt. Kinabalu supported by numerous illustrations, explanation of terms, for-
mulae, and analyses individual families are discussed in detail. Taxa are introduced alphabetically irre-
spective of any taxonomical or phylogenetical relationship.

With regard to illustrations preference was given to colour photographs that were taken on-site in the
area of Mt. Kinabalu, even if this meant to accept an inferior photographic quality. While the photo-
graphs primarily serve to illustrate the general appearance of a species, additional drawings are provided
for specific features (e.g. oral fields in tadpoles, details of pholidosis in reptiles).

Genera are differentiated via dichotome keys based solely on morphological criteria. Additional data
useful for unambiguous identification can be found in the species account. Discussion of an individual
genus includes a table of distribution for all Bornean species and a key to these species. All taxa recorded
from Mt. Kinabalu so far are given in boldface.

Species accounts are standardized providing the following information: the valid scientific name, refer-
ence of the original description including the locus typicus (type locality; origin of the type specimen), a
chresonymy list (as of end of 2000), a diagnosis, description of external features as well as ecological
and ethological observations, and distribution.

Scientific denomination follows trinomial classification. The basic unit of zoological classification is the
species. Its scientific name is composed of two names (binomial), that of a subspecies of three names
(trinomial). The first name is the generic name, the second the species name, and the possible third the
subspecies name. The taxonomic concept of subspecies is subject to controversial debates and we have
therefore limited information on subspecies to unambiguous characters.

Synonymy/Chresonymy lists are limited to those references, which refer to an occurrence of the
species on Mt. Kinabalu. Species names and relevant synonymies are given with the respective authors’
names, year of the publication, and page number(s); complete references can be found in the bibliogra-
phy.

Diagnosis is a compilation of generally valid data and serves to distinguish a taxon from other taxa on
generic, specific or sub specific level. In case of a monotypical genus, the relevant taxon is differentiated
from similar Bornean taxa.

9
How to use this book

Description is based on own examinations and/or the standard publications for the region, e. g. DE ROOIJ
(1915, 1917), INGER (1966), and MANTHEY & GROSSMANN (1997). A descriptive summary of adult external
features is accompanied by data on ontogenetic development or developmental stages where appropriate
(e. g. amphibian larvae). Measurements relate to standardized methods described in the introductory
chapters and are given in millimetres.

Ecology/Ethology summarizes data on general behaviour, habitat preferences, vertical distribution, popu-
lation density, feeding, predators and antipredator behaviour. It should be noted that there is a painful
lack of ecological and ethological information on many of the species.

Distribution compiles the entire geographical distribution of a species as well as its distribution on
Borneo (for localities see map on page 11). Distribution maps are limited to those species for which a
larger number of localities are known. Locality data relate to our own studies, those taken from the
literature are marked differently.

Mt. Kinabalu contains listing of all known localities on Mt. Kinabalu (> 500 m a.s.l.) where a taxon has
been recorded. Apart from a few unidentified places, all localities are shown on the maps on page 200,
203 and 204. Their spellings are very much inconsistent in commonly used maps, and we largely follow
the spellings used on a draft map kindly made available to us by BEAMAN. It is unfortunate that older
records usually do not include altitudinal information. In many cases the altitude of the actual collecting
site is likely to vary substantially from that of the stated locality as only the nearest known location is
given in publications.

Species determination of specimens housed in the collection of Kinabalu Park (HQ-Collection) has been
accepted without further verification.

Literature used in the introductory chapters and further readings are marked by an asterisk (*) in the
bibliography section of this book.

10
11
Introduction

Introduction
Covering an area of 739 204 square kilometres, the
biggest of the Sunda Islands, Borneo, lies in the
central tropics. As far as size is concerned, it ranks
third among the largest islands of the world, after
Greenland and New Guinea.
It has more than a thousand years of varied history,
during which Indian, Chinese and - from the 16th
century onwards - European tradesmen and colo-
nial powers strived for territory, resources, and com-
modities. It was not before the end of World War II
that the territories of Borneo became independent.
* In 1949, the formerly Dutch part of Borneo be-
came Indonesian under the name of Kalimantan
(535 834 km²).
* In 1963, the formerly British northern part of
Borneo joined the Malaysian Federacy; the prov-
inces Sabah (73 620 km²) and Sarawak 123 985
km²) were established.
* In 1984, the formerly British protectorate Brunei
(5 765 km²) gained independence. The economic
dominance Brunei exerted in the region lead Euro-
peans to using its name in gradual linguistic distor-
tions (Brunei ® Baruney® Borneo) to signify the
whole island. This name is still in common use after WHITEHEAD (1893)
nowadays.
Gunung (= Mount) Kinabalu, 4095 m, is the high- Fig. II Mt. Kinabalu from near Melankup.
est summit of the entire Malay Archipelago and is
situated in the far north of Borneo, in the province
of Sabah. Apart from the summit of Makita/ during the first decade after its first ascent, only 53
Kalimantan (2987 m) it towers more than 1500 m people climbed the mountain, whilst nowadays tens
above all the other peaks of the island. of thousands annually follow their tracks.
In former times indigenous people regarded the In spite of this mass rush, the mountain has not
mountain as a place where they expected the souls lost a bit of its aura. Nobody manages to elude the
of their deceased to be gathered in a village of the magic and majesty radiated by the granite giant
dead (Aki Nabalu), which was strictly off limits. emerging from the rain forest. The elevated lan-
From the mid-19th century onwards, the mountain guage used in films and leaflets to entice the tour-
has been radiating a virtually magic attraction for ist to Sabah, the land of the “Sacred Mountain”,
botanists and zoologists due to its unrivalled di- and to evoke the paradisaical wonders which await
versity of species. the visitor shows quite unmistakingly: the former
Almost instantly after having been incorporated realm of the dead has undergone a metamorphosis
into the Malaysian territory, it gained the status of to a national sanctuary. In biological terms, Mt.
a protected Park, and even the capital of Sabah, the Kinabalu has remained what it used to be in the
former Jesselton, derives its name from the great last century: a great centre of biological diversity
peak: Kota Kinabalu. For tourists it has been an which is unparalleled - not only in the Sunda area,
unrivalled attraction from the very beginning: but also worldwide.

Mount Kinabalu - the surrounding environment
In order to understand the distribution of specific
species at any place on the earth, it is necessary to
have a clear idea about the fundamental abiotic and
biotic conditions of the respective area under ex-
amination. The valid criteria are:
* the geological and geomorphological conditions
* the zoogeographical circumstances
* the recent climatic conditions
* the structure of the vegetation cover
* the anthropogeneous impact on the environment
The following chapters try to delineate an over-
view of these basic conditions for the area under
inspection.
Topography, Geology, Geomorphology
Mount Kinabalu, with 4095 m the highest mountain
between the Himalayas and New Guinea, is located
at the northern tip of Borneo (5°57'-6°18’N, 116°28'-
Fig. 1 Mt. Kinabalu in the very early morning, view from the Twin Bed Cabins.
Mount Kinabalu - the surrounding environment
116°45’E). The central Kinabalu massif forms the
shape of a horseshoe open to the north. The uninter-
rupted rocky cliff facing south sends two ridges to
the north separated by the 1500 m deep Low’s Gully
(the western ridge already descends rapidly after
2 kilometres, while the eastern one stretches far to
the north, reaches once more an altitude of 2576 m
in the summit of Tambuyukon and descends in the
range of Mt. Templar-Mandalon) and continues to
the east down to Poring jointed into various lower
ridges to the north and to the south.
The massif is one of the most important water catch-
ments in Sabah and the source of many streams
(e.g. Sg. Kinaram and Tuaran flow to the north,
Wariu, Panataran, and Kadamaian to the west,
Liwago, Mesilau, Luidan, Langanan, Kinapassan,
and Kapuakan to the east). Together with their, in
part temporary, tributaries and crossing through all
altitudes they represent just about every type of
habitat one may expect in tropical montane run-
ning waters.
© M. Maronde
13
 Mount Kinabalu - the surrounding environment
© R. Malkmus
Fig. 2 Rock area polished by ice during the last glacial, with nunataks in the background (northern plateau of Mt.
Kinabalu.
The area examined comprises in principle the en-
tire mountain massif above 500 m above sea-level,
but, for reasons of inaccessibility, is largely re-
stricted to localities around Poring, Headquarters
(inclusive of the Summit Trail), Mesilau, Sayap,
Kiau, and Bundu Tuhan. About 80% of the area lie
within the boundaries of the Kinabalu Park.
The crystalline basement of northern Borneo is
covered with a thick sequence of marine sediments
which developed in the Tertiary period. During
the middle of the late Miocene period (some 15
million years ago), these layers of shale, siltstone,
and sandstone were folded and uplifted to the
mountain ranges of the Crockers and Trusmadi.
During the Pliocene period (some 10 to 5 million
years ago), a subvolcanic intrusion was exposed
uplifting the overlying layers at the northern end
of the Crockers. The central summit plateau - today
called Mount Kinabalu - consists of a granitic
centre (hornblende adamellite), covering a surface
of 150 km², and an elevation of 2000 and 3000 m,
surrounded with a ring of porphyritic and
ultrabasic rocks (serpentine). These are followed
14
by tertiary sediments at lower elevations. On
the southeastern slopes (Pinosuk Plateau) up to
140 m thick Pleistocene layers of gravels are de-
posited.
During the last Pleistocene period, the summit re-
gion was covered by an ice-cap, about 5 km² in
extent, reaching down to about 3000 m. This gla-
ciation has left distinctive landforms not only in
the summit area (polished and plucked surfaces by
the erosional action of the ice on the plateau,
nunataks, U-shaped gullies, moraine material with
subangular boulders at the end of such gullies and
near Pakka Cave, cirques on the summit), but the
geomorphology of the lower slopes, situated in the
periglacial zone, was stamped likewise (deposits
by blockstreams and mudflows on Pinosuk, forms
of valleys and canyons).
The agents of the recent, chiefly chemical, weath-
ering under tropical conditions (high temperatures
and humidity) can be studied on the denuded sum-
mit area (exfoliation of slabs of rocks) and espe-
cially in regions deforested by man (partially up to
1800 m) with considerable soil erosion.

Zoogeography
Together with the islands of Palawan, Sumatra,
Java, and the Malay Peninsula, Borneo forms part
of the so-called Sunda Plate, a southeastern spur
of the Asian continental shelf. Between Borneo and
mainland Southeast Asia, large parts of the Sunda
Plate were exposed for much of the Tertiary. It was
not before the Pliocene period that a shallow shelf
sea separated the continent from the Sunda Islands.
The Quaternary then was a period of repeated al-
ternation of colder (glacial) and warmer (intergla-
cial) spaces of time. During the glacials a substan-
tial portion of the water was bound terrestrially as
glacial ice causing eustatic sea level changes world-
wide (> 100 m max.). Under these conditions large
areas of land of the Sunda Shelf were re-exposed
and connected Borneo with the Asian continent.
Distinct indications of this intermittent Quaternary
falling-dry are given by submarine channels. They
indicate the courses of former rivers at times when
the platform was above sea level. Corresponding
to the alternation of colder and warmer periods,
these land connections were formed and broken
several times.
Fig. III The area of Sundaland (surrounded by a bro-
ken line) exposed at times of lowest sea level (dark shade)
during the last glacial with ancient river system (sub-
marine channels).
partly after PIJA 1980
Mount Kinabalu - the surrounding environment
Situated east of the Sunda platform, and reaching
depths of several thousand metres in places, the
Makassar Straits separates Borneo and Sulawesi.
The latest connection between these two islands
existed in the middle Eocene, more than 40 mil-
lion years ago. During the Tertiary, arcs of volcanic
islands were important as potential land bridges and
island hopping routes for organisms to and from
the Philippines, Borneo, and Sulawesi.
The climate of the Sunda region during the glacials
was probably 2-4°C cooler and more arid than it is
today while the interglacials had a warm and hu-
mid climate.
These paleo-climatogeological conditions manifest
themselves in the existing zoogeographical situa-
tions. They help us to explain many disjunct plant
and animal distribution patterns and the composi-
tion of species in the different geographical regions.
It is significant that a major zoogeographical bound-
ary separates Borneo and Sulawesi: the Wallace’s
Line. The islands west of this boundary - thus in-
cluding Borneo - form part of the Oriental Region,
those to the east of it belong to the Wallacea, a tran-
sitional region between Orientalis and Australis.
The relationship of the herpetofauna of Borneo is
significantly closer to the Asian mainland and other
Sunda islands than to Sulawesi despite its shorter
geographical distance to the latter.
This is proven by facts such as 70% of the snake
and 43% of the lizard species of Borneo also oc-
curring on the Malay Peninsula whereas only 25
and 19%, respectively, exist also east of the
Wallace’s Line. Nearly all non-endemic species of
Bornean frogs (ca. 35%) are also found on the
mainland, but only 7% also on Sulawesi (of which
70% are frogs settling in cultivated areas which are
very likely to have been introduced by man).
The large diversity of species of amphibians and
reptiles on Borneo in general and on Mt. Kinabalu
in particular can be based for a large part on the
following factors:
• Most of the species of animals and plants show
a tendency to increase their diversity the nearer
to the equator they occur. Borneo is situated in
the central tropics within the species-rich Ori-
ental Region. The amphibians and reptiles na-
tive to this region furthermore include numer-
ous genera which are known for their rapid
15
Mount Kinabalu - the surrounding environment
speciation (e.g. Ansonia, Meristogenys, Philau-
tus, Calamaria, Sphenomorphus).
• The repeated connection and separation of
Borneo to, respectively from, the mainland dur-
ing the Quaternary favoured an exchange of
faunae in one situation and speciation processes
in the other resulting in ca. 25% of the snakes,
45% of the lizards, and 67% of the frogs being
endemic to the island. If the hypothesis presented
by BRANDON-JONES (1996) were applicable, how-
ever, saying that during the last two glacials
“Asian rainforests were reduced by drought to a
few scattered pockets of distribution”, a migra-
tion of the almost exclusively sylvicolous
herpetofauna of this region over the landbridge
could be fathomed only with great difficulty.
• Studies of the biogeography of islands have
shown that small islands support fewer species
than do larger islands of similar habitats (areal
effect), and that isolated islands support fewer
species than islands close to the mainland.
Borneo is one of the largest islands in the world,
and together with other islands, lies close to the
Asian continent.
• By displacing competing generalists, a multi-
tude of niche-specific species have developed,
which is a characteristic trait of tropical rain for-
ests. This process is powered by a wide variety
of interactions (e.g. symbiosis, parasitism, prey-
predator relationship, mimicry, communication
mechanics, etc.), but also by the isolation ef-
fects of mosaic-like distribution patterns of
sparse populations found in many species.
• Due to a climate which becomes proportionally
unfavourable with increasing altitude, along with
a reduction of resources, and a limitation of suit-
able spawning possibilities, the number of am-
phibian species decreases at increasing altitudes.
At the same time, the tendency to form localized
montane endemites increases. The altitudinal
structure of Mt. Kinabalu induces that its habi-
tats above 1500-2000 m function like ecological
islands. These habitats are grouped in plant-so-
ciologically defined altitudinal zones (comp.
chapter “Vegetation”). It was mainly their climatic
differentiation which necessitated particular ad-
aptations from the plants and animals invading
them and so favoured speciation processes.
16
The geological age of Mt. Kinabalu is high enough
to enable many genera to have reached consider-
able degrees of differentiation through adaptive
radiations manifesting themselves in the form of
localized endemites which have adapted to certain
ecological niches within certain altitudinal zones
(e.g. Ansonia guibei, Ansonia fuliginea, Kalo-
phrynus baluensis, Leptobrachium gunungense,
Philautus saueri, Leptolalax maurus, Calamaria
schmidti, etc.). Recognizing these species as local-
ized endemites, however, requires a considerable
amount of caution due to the fact that our knowl-
edge of the distribution of the animals is still very
limited. As the average temperatures were several
degrees lower than today during the Pleistocene,
some of the forms classified as localized endemites
may turn out to be in fact glacial relicts which pre-
viously inhabited also lower elevations, but were
then forced to retreat to higher altitudes when the
temperatures rose. Today they occur only in
montane isolates scattered widely over the entire
islands.
Climate
“The Marei Parei spur offered a fine position
for a sanitarium, at any height between 4 000 ft
and 5000 ft. Our tent was pitched at about 4 700 ft,
and we found that the thermometer marked 75° in
the mid-day shade, 56° at six a.m., and 63° at six p.m.
This would be a delightful climate in a well-built
house.”
Spencer St. John,
Life in the Forests of the Far East, 1862
Borneo maintains a tropical rain forest climate up to
altitudes of above 1000 metres. It is characterized
by an annual precipitation in excess of 2000 mm, no
dry periods for several months, day temperatures
of +27°C, +22-23°C at night, and a relative humid-
ity of usually above 90%, and thus lies in the range
of the monsoon weather pattern of tropical Asia.
The constantly present equatorial low pressure area
dominating the Indo-Pacific region, known as the
Inter-Tropical Convergence Zone, shifts by follow-
ing the position of the sun to the south in winter
and to the north in summer, triggering the north-

Fig. IV Monthly precipitation at the Headquarters of Mt. Kinabalu in the years 1987-1990 (after MALKMUS 1996b)
east monsoon (November/December through
March) and the southwest monsoon (June through
September), respectively, with their heavy zenith
rains. Substantial precipitation is, however, ob-
served throughout the year in all parts of Borneo
so that the monsoons merely cause it to intensify
in certain seasons.
Notwithstanding this situation, extended dry peri-
ods are observed aperiodically with monthly
precipitations of less than 30 mm. Cyclic changes
in the atmospheric circulation (amplification of the
southeast passats) in the form of the so-called El
Niño Southern Oscillation often result in months-
long dry periods at irregular intervals (e.g. in 1977,
1979, 1982/83, 1992, 1997/98) exposing the ecosys-
tem rain forest to substantial stress and favouring
catastrophic forest fires. For example, in 1972/73
a prolonged drought disturbed all woody plants of
the upper montane forest in some areas with shal-
low soils on Mt. Kinabalu.
Mount Kinabalu - the surrounding environment
The general climate may be altered significantly
in places due to the individual topography. Out-
side prolonged rainy or dry periods, a very charac-
teristic daily weather pattern can be observed on
Mt. Kinabalu:
The first hours of daylight are marked by a clear or
minorly overcast sky. Depending on the intensity
of sunlight and wind, humidity drops from 90-95%
to 40-70% outside the forest and in the crowns of
the trees, while it oscillates very little within the
forest. Air saturated with moisture ascends the
slopes of the mountain following its relief. As the
temperatures drop by an average of 0.55 to 0.65°C
per 100 m increased altitude, the cooling water
vapour condenses. The condensation level during
the morning hours normally lies between 2000 and
3000 metres. This results in the formation of a ring
of fog and clouds up to 3000 m altitude, covering
the entire mountain down to about 1500 m during
the course of the afternoon. Heavy rain showers,
17
 Mount Kinabalu - the surrounding environment
© A. Nöllert

Fig. 3 A typical view of Mt. Kinabalu in the morning when humid air is condensing above 2000 m.

often in the form of thunder storms, usually set in slopes so that insolation and wind exposition are
from between 2.00 and 4.00 p.m. often lasting up highly variable factors. For example, entirely dis-
into the early night hours (i.e. 7.00 to 9.00 p.m.). similar climatic conditions may be found when
While the annual pattern of the temperatures
is relatively balanced, pronounced oscilla-
tions are discernible during the course of a Fig. V Oscillations of temperature at various altitudes on
Mt. Kinabalu (measured in August/September 1986).
day. They are least apparent in the cloudy
modified after KÜRSCHNER
moss forest and most distinct above the tree
1990
line. Corresponding oscillations are found in
the relative humidity. The summit region is
known to experience occasional snow falls
and frost at night.
The visitor from central Europe finds a Sepilok/Sandakan
rather agreeable climate between 1400 and
1600 m which was appraised already by Poring

the pioneer explorers of Mt. Kinabalu in Park Headquarters

the 19th century (see quote at the begin-
ning of this chapter). This climatic situa-
tion is subject to numerous localized aber- Kamborangoh
rations. Isotherms and isohyets show no
parallel alignment anywhere on the moun-
Panar Laban Huts
tain as a result of the very differentiated
structure of the relief (in the form of gorges,
canyons, outcrops, edges, and ridges) and
the varying exposition and inclines of the hrs

18

comparing an exposed ridge with a stream that
flows in a protected gorge, even if both are situ-
ated at the same elevation and only a few hundred
metres apart. It is significant that it is exactly these
gorges which represent the upper altitudinal distri-
bution limits of many species of frogs.
The topographically induced network of microcli-
matic niches experiences another fine-tuning of
its differentiation by the vegetation cover. Below
3200 m there is only one natural type of plant for-
mation on Mt. Kinabalu, i.e. forest. Depending
on the height of its crown canopy, significant ab-
errations of the individual climatic components
may be observed within vertical distances of as
little as 10 to 60 metres. The climate immediately
above the leaf litter up to about 2 m is marked by
being very constant as far as temperatures are
concerned (daily variation usually less than 5°C).
Due to the ineffectiveness of air currents, the
ground level is constantly covered by air which is
saturated with moisture; the relative humidity thus
measures more than 95% most of the day, with a
minimum of 80-90% between 12.00 a.m. and 3.00
p.m. It is only when the dry winds accompanying
the El Niño weather phenomenon take effect that
the humidity may decrease to less than 60% for
prolonged periods of time.
The partly many-tiered crown canopy blocks out
the sun so that merely 1 to 3% of the sunlight reach
the forest floor although the intensity may vary
Fig. VI Calling activity of frog species on Sg. Liwago (1450 m) in August 1990. A – after one week of drought
(rel. humidity < 60%). B – after two hours of thunderstorm between 3.00 and 5.00 p.m. (rel. humidity > 95%).
A B
Mount Kinabalu - the surrounding environment
locally during the course of the day due to the shift-
ing of sun-exposed patches.
Entirely different climatic conditions are found at
the level of the crowns since they are highly ex-
posed to wind and insolation. The result is that in-
dividual climatic components are subject to sub-
stantial oscillations during the course of a day, with
temperatures varying by more than 10°C, and hu-
midity by 50 to 60%.
Between the floor and the crown level, the climatic
components experience alterations of all kinds, so
that it is just to speak of a vertical tiering of cli-
mates according to the tiering of the rain forest it-
self, in other words a “storey climate”.
Climatic peculiarities, lastly, result from temporary
gaps in the forest caused by wind storms and slides
of the soil, and in the immediate vicinity of streams.
With amphibians and reptiles being poikilothermic
animals, their activity is limited to a relatively nar-
row bandwidth of climatic preconditions. The
knowledge of the many-faceted climatic conditions
in their biotopes is therefore an essential tool for
the understanding of their often insular niche seg-
regation in a larger biosphere, their activity
rhythms, and numerous other adaptive strategies.
Figure VI demonstrates this principle clearly by
showing calling activities of frogs on the same site
during various weather conditions, while fig. VIII
on page 26 shows the influence of the vertical dis-
tribution.
19
 Mount Kinabalu - the surrounding environment

Every visitor to the mountain cannot but note how

Vegetation the physiognomy of the forest changes the higher
he ascends. The mountain is virtually covered by
stacked rings of vegetation zones with each having
“The people of Sabah possess on this fa-
a specific composition of species. These altitudinal
mous mountain what I believe is the richest
vegetation zones are the response by the flora to the
and most remarkable assemblage of plants in
altitudinally changing climatic and edaphic condi-
the world.”
tions. Notwithstanding this there is no strict demar-
E.J.H. Corner, 1978
cation between one zone and the subsequent one,
but a gradual transition. While Indo-Malayan taxa
With the exception of the slopes of the Kadamaian dominate the lower elevations, the moderate altitudes
valley, the flanks of Mt. Kinabalu above 500 m used are home to mainly East Asian ones, and Austral-
to be covered by virgin rain forest until as recently Antarctic floral elements reign above 2800 m.
as the early 20th century. These montane forests
are home to the largest variety of plant species in
the entire Malesian region as the perma-moist Fig. 5 Coelogyne ragioferens, a beautiful orchid.
Southeast Asian tropics are referred to in the sci-
ence of phytogeography. Besides hundreds of up-
per montane, subalpine and alpine endemisms (no-
tably neoendemisms in the vegetation facies on
ultramafic substrate), it includes truly spectacular
plants such as Rafflesia which parasites on vines
(Tetrastigma), sixteen species of carnivorous pit-
cher plants (Nepenthes), twenty-five species of
Rhododendron, and not less than 610 species of
ferns, 350 of mosses, and 750 of orchids. BEAMAN
& BEAMAN (1990) estimated some 4000 species of
vascular plants to occur on Mt. Kinabalu.

Fig. 4 Flower of Rafflesia pricei, Mamut Valley. On

Mt. Kinabalu Rafflesia is common round the Mamut Val-
ley at 900-1500 m a.s.l.
© R. Malkmus

© M. Schroth

20

Fig. 6 Sg. Kipungit, near Poring Hot Springs (550 m).
Tropical lowland vegetation zone
(to 900 m a.s.l.)
Climate: Levelled, moist-warm; average annual
temperature +26-27°C; precipitation throughout the
year (2000-3000 mm).
Soil: Yellow latosol with a thin layer of humus in
the form of mould or mould-like compost as a re-
sult of a rapid rotting and mineralization of leaf
litter.
Vegetation: Multi-layered dipterocarp rain forest
Upper storey: 30-50 m high tree giants (Shorea,
Dipterocarpus, Hopea, Vatica), moderately evident
epiphytism.
Middle storey: Dense, closed crown canopy 20 m
above the ground (Annonaceae, Meliaceae,
Myrtaceae, Sapotaceae), bamboo, lianas.
Bottom storey: Lacunate, 8-10 m high under-
brush; vines, bamboo and rattan thickets; plank
roots; the floor is covered with a closed layer of
leaf litter.
Mount Kinabalu - the surrounding environment
© A. Nöllert
Water: Mountain streams with moderate to strong
currents and occasional cascades; pronounced tidal
changes relative to the weather.
Structures: stream bed sediments dominated by
coarse gravel, forming an extensive scattered pat-
tern (hyporheic interstitial area); sand and gravel
banks; rocks and fallen logs which protrude above
the water surface; permanently or temporarily con-
nected rocky pools with fine sediment and deposits
of leaf litter. Accounting for the strong current and
shade (usually only 2-5% of the sunlight reach the
water surface), no higher aquatic plants and few al-
gae can develop. The water of these mountain
streams is in general very rich in oxygen, poor in
dissociating salts, alkalines, and acids as a result of
the filter performance of the fungal plexus which is
associated with the root systems of the trees; pH 6
to 7; water temperatures rather stable (+17 to 20°C).
Ephemeral rain pools or puddles and phyto-
thelmes (water bodies fit for vegetation): limited
to cavities in trunks or limbs of trees, tree buttresses,
and lianas, as well as bamboo internodes, funnels in
plank roots, and in the axils of leaves (e.g. Pandanus).
21
 Mount Kinabalu - the surrounding environment

Tropical lower montane vegetation zone Soil: Micropodzolic soils with a layer of raw hu-
mus whose thickness increases with increasing al-
(900-1800 m a.s.l.) titude.
Vegetation
Climate: Levelled, moderately moist-warm; aver- 900-1300 m: Transition from submontane
age annual temperature +18-24°C; precipitation dipterocarp forest to lower mixed Fagacea forest
throughout the year (> 2500 mm). (30-35 m).
1300-1800 m: Mixed Fa-
gacea forest (oak-chest-
Fig.7 Sg. Liwago (1500 m) nut forest). Reduction of
the crown height to 15-18
m; very large variety of
species (Lauraceae, Thea-
ceae, Myrtaceae, Elaeo-
carpaceae), first conifers
(Dacrydium, Podocar-
pus, Agathis); the reduced
height of the canopy per-
mits a higher intensity of
light on the floor result-
ing in a thicker under-
brush. Phorophytes (car-
rier trees for epiphytes)
are thickly covered with
mosses, ferns, and or-
chids; the first cutaneous
ferns (Hymenophyllum)
and terrestrial mosses ap-
pear pointing towards the
presence of climatic ele-
ments typical for mossy
forests.
Water: Usually fast-
flowing, cascadeous, cold
(+14 to 17°C) mountain
streams with the struc-
tural traits of submon-
tane streams. The bank-
ing vegetation includes
thickets of shrubs (Gun-
nera, Impatiens, Bego-
nia, Pitospatha) and Rat-
tan (Calamus, Daemono-
rops).
Phytothelmes: Dead pit-
© R. Malkmus

chers of Nepenthes and

leaf axils, while cavities
in trees and lianas are rare.

22

Fig. VII Daily oscillations of air and water temperatures at the confluence of Sg. Silau-Silau and Sg. Liwago
(ca. 1400 m a.s.l.) in August 1994 (after MALKMUS 1996c).
Tropical upper montane vegetation zone
(1800-3000 m a.s.l.)
Climate: Levelled, cool-moist; average annual tem-
perature +10-13°C; precipitation throughout the
year (> 4000 mm); daily, long-persistent cloud lay-
ers; extremely high atmospheric humidity.
Soil: Very acid brown earths, podzols and peaty
gley soils.
Vegetation: Cloudy moss forest. An 8-12 m high
forest of conifers, Ericaceae, Myrtaceae, Fagaceae,
tree ferns (Cyathea) and Bambusa, with dense
underbrush. The trees are usually stunted and
deeply packed with epiphytes such as mosses (es-
Mount Kinabalu - the surrounding environment
pecially Bazzanis, Plagiochila, Mastigophora),
liverworts (Lepidozia), lichens (Usnea), orchids (e.g.
Dendrochilum, Coelogyne, Bulbophyllum), and club
mosses (Lycopodiatae). Many species absorb water
from the fog (“fog-stripping”). Since the plaster of
mosses releases water only very slowly, the coating
of the phorophytes never dries up completely. Waft-
ing mists turn the stunted forest with its heavy load
of epiphytes into a bizarre, spooky environment
which accounts for its name “elfin forest”.
While the mossy forest rises up to 3000 m altitude on
the southeastern side, it is replaced along the Summit
Trail between 2600 and 3000 m by ultrabasic rock
(serpentine) with highly demineralized, variously
moist soil which carries a small-crown forest (< 8 m)
23
Mount Kinabalu - the surrounding environment

Fig. 8 Usnea lichens

© S. Manthey

Fig. 9 Nepenthes lowii

© M. Schroth

Fig. 10 Nepenthes villosa

© M. Maronde
© R. Malkmus

Fig. 11 Mossy forest, along the summit trail.

with colonies of few leading species (Podocarpus, torrential streams with many cascades which often
Dacrydium, Myrica, Leptospermum) which is fall dry during prolonged dry periods above 2500 m.
marked by a substantially reduced epiphytism. Phytothelmes: Pitchers of Nepenthes are very com-
Water: Almost exclusively very rapidly flowing to mon; cavities in trees are rare.

24
 Mount Kinabalu - the surrounding environment

© R. Malkmus
Fig. 12 Tropical alpine dwarf scrub.

Tropical subalpine vegetation zone

(3000-3700 m a.s.l.)

Climate: Distinct fluctuations in the temperature

during the course of the day (often > 10°C); cool;
average annual temperature +8-10°C; precipitation
throughout the year (no data available); extreme

winds with chilling and drying effects common; rela-
tive humidity varying between 20 and 100%.
Soil: 3000-3200 m: A layer of acid brown podzo-
lic soils and peaty podzols. 3200-3700 m: Accu-
mulations of podzolic soil limited to eroded grooves
and rifts between large granite boulders (skeleton
soils).
Vegetation: Low-growing (< 5 m), very dense colo-
nies of hard-leafed plants (Leptospermum, Schima,
Rhododendron, Myrica, Daphniphyllum, Phyllo-
clades, Podocarpus)) which form the tree-line al-
ready between 3200 and 3400 m for edaphic rea-
sons (the potential tree-line in the Indo-Australian
region lies at 3700-3800 m, Mt. Wilhelm on New
Guinea being an example). Species of the same gen-
© S. Manthey
Fig. 13 Leptospermum recurvum
era colonize crevices and rock fissures which are
filled with humus up to the summit zone (4100 m
a.s.l.), scattered over an alpine rock desert (tropical
alpine dwarf-scrub).
Water: The water levels of the streams are directly
related to the amount of precipitation; water-filled
rock basins are occasionally found.
Phytothelmes: Pitchers of Nepenthes villosa be-
low 3200 m.

25
Mount Kinabalu - the surrounding environment

Fig. VIII Number of amphibian species in the differ-

ent altitudinal vegetation zones on Mt. Kinabalu.

Fig. VIII While average temperatures decrease continually with increasing altitude, frequency and quantity of
precipitation increase. The altitudinal distribution of amphibians and reptiles is influenced to a significant extent by
both factors: the diversity of species of reptiles decreases dramatically with increasing altitude; although number-
ing fewer in species, amphibians are better adapted to lower temperatures and moisture and therefore extend their
ranges a 1000 m higher into the montane regions.

26
 Mount Kinabalu - the surrounding environment

Fig. IX Number of reptile species in the different altitudinal vegetation zones on Mt. Kinabalu.

27
 Mount Kinabalu - the surrounding environment
© U. Manthey
© S. Manthey
© U. Manthey
From top to base
Fig. 14 Tragulus javanicus, the Smaller Mouse Deer and
Callosciurus prevostii (Fig. 15), the White-striped Tree
Squirrel, appear to be common in the lowland zone. The
Palm Civet Paradoxurus hermaphroditus (Fig. 16) was
also observed up to 1800 m and higher.
28
Fauna
“The first impression which one per-
ceives in the tropical forests is that almost no
animal life can be found.”
A.R. Wallace
The Malay Archipelago, 1869
The diversity of faunal species of the Kinabalu
massif is in no way inferior to that of its flora. It is
a pure forest fauna of the East Oriental region with
a high content of endemisms. On the other hand
tropical rain forests are characterized by a quanti-
tatively unproportional ratio between plant and
animal biomasses with an average of 1000 tons of
plant mass to only 200 kilograms of animal mass.
The fact that the number of animal species is not-
withstanding substantially higher than that of plants
is explained by the large diversity of invertebrates
which amount to little in weight. Most of the spe-
cies are furthermore rather rare so that a high spe-
cies diversity (richness) goes along with a low
population density (evenness).
The spatial niche segregation of the individual spe-
cies - a large percentage of the latter is confined to
the canopy - is so perfectly adapted to the habitat
that every observer who enters a tropical rain for-
est for the first time (and who would not do so with
the expectation to encounter an unlimited species
richness?) will surely be disappointed and agree
with WALLACE’s statement quoted above.
The following is an inventory of what has been re-
corded to date from Mt. Kinabalu:
110 species of mammals, e.g. Orang Utan (Pongo
pygmaeus), Red Leaf Monkey (Presbytis rubicun-
da), Grey Gibbon (Hylobates moloch), Slow Lori
(Nycticebus coucang), Tarsier (Tarsius bancanus),
Flying Lemur (Cynocephalus variegatus), Pango-
lin (Manis javanica), Porcupine (Trichys fascicu-
lata), Malay Bear (Helarctos malayanus), Leop-
ard Cat (Felis bengalensis), Mouse Deer (Tragulus
sp.), Barking Deer (Muntiacus sp.), 22 species of
bats (Chiroptera), 26 squirrels (Sciuridae), 7 tree
shrews (Tupaiidae), and 7 civets (Viverridae).
65% of the mammal species endemic to Borneo
occur on Mt. Kinabalu.
 Mount Kinabalu - the surrounding environment

306 species of birds, e.g. Great Argus Pheasant

(Argusianus argus), 6 species of hornbills (Buce-
rotidae), 11 woodpeckers (Picidae), 19 cockoos
(Cuculidae), 3 pittas (Pittidae), 17 bulbuls (Pycno-
notidae), 8 broadbills (Eurylaimidae), 13 sunbirds
and spiderhunters (Nectarinidae), 5 drongos (Dicru-

From top to base

Fig. 17 Argusianus argus, a common lowland resident
is more heard than seen. The Rhinoceros Hornbill,

© M. Schroth
Buceros rhinoceros (Fig. 18) is rearly seen so near. It
was observed on Mt. Kinabalu up to 1750 m.

© U. Manthey
From top to base
Fig. 19 An unknown spider.
Fig. 20 Phlebogryllacris venosa from the Headquarters.
© S. Manthey

ridae), 29 laughing thrushes (Timaliidae), and 9

barbets (Capitonidae).
48% of bird species endemic to Borneo have been
recorded from Mt. Kinabalu.
40 species of fish; predominantly species which
have adapted to fast-flowing waters by the devel-
opment of adhesive organs, e.g. catfishes (Sisoridae,
Bagridae) and gastromyzontids (Gastromyzon-
tidae).
Only a low percentage of the legions of arthropods
has as yet been described: crustaceans (Crustacea),
millipeds (Diplopoda), centipeds (Chilopoda), spi-
ders (Arachnida) and espacially insects (Insecta).
© U. Manthey

Insects are represented in thousands of species in

many orders: brilliant odonates (Odonata), butter-
flies (Lepidoptera; e.g. Papilionidae, Pieridae,

29
 Mount Kinabalu - the surrounding environment

© M. Schroth
© M. Schroth

© M. Schroth
© W. Malkmus

From top to base

Fig. 23 Pseudomyagrus waterhousei
Fig. 24 Brahmidia hearseyi
Fig. 25 Cethosia hypsea
© R. Malkmus

Danaidae, Lycaenidae; Sphingidae, Geometridae,

From top to base
Fig. 21 Pyrops intricata
Fig. 22 A newly emerged cicada.
Drepanidae) and beetles (Coleoptera; e.g. Ceram-
bycidae, Lucanidae, Buprestidae, Elateridae,
Passalidae, Curculionidae), bizarre stick insects
(Phasmida), crickets (Orthoptera) and bugs (Hemi-
ptera); very diverse, in many cases colonial and

30

therefore contributing more individuals than any
other order of insects are termites (Isoptera), ants
and bees (Hymenoptera).
Most species occupy certain ecological niches -
defined as that part of the environment which a
species actually exploits - and partly represent
highly specialized morphological and ethological
adaptations as a result of complex co-evolutionary
developments of animals and plants, or of differ-
ent animal species. They are fixed components of
the rain forest and cannot survive without it. The
highly advanced degree of differentiation of the
adaptations also provides a clear indication of the
old age of the Malayan rain forests as a stable eco-
system.
As far as amphibians and reptiles are concerned,
other animals are of vital importance as predators
and prey. Unfortunately that the prey spectra of the
species occurring on Mt. Kinabalu are as poorly
known as their predators.
The major part of the food of metamorphosed am-
phibians is composed of arthropods and their lar-
vae, worms, and molluscs. Larger species, such as
Limnonectes leporina, may also prey upon smaller
vertebrates, including members of their own spe-
cies, and crabs. Most frogs are broad-spectrum car-
nivores (dietary generalists); dietary specialists are
rare. Microhylids, for example, feed mainly on ants
and termites.
Predators of amphibians are particularly numer-
ous among small mammals (e.g. civets, rats, ot-
ters), birds (owls, hawks, thrushes), and snakes
(Amphiesma flavifrons, Amphiesma sarawacense,
Dendrelaphis pictus, Rhabdophis chrysargos,
Xenochrophis trianguligerus), but also include
large spiders, centipedes, and crabs. The carnivo-
rous larvae of dragonflies (Anisoptera) and aquatic
beetles, hygrophilous snakes, and fish prey upon
tadpoles living in the mountain streams and some
may also ingest frog eggs. Nothing is known, on
the other hand, about the predators of anuran lar-
vae which live in stagnant pools. Potential dan-
gers might come in the form of terrestrial crabs,
larvae of small dragonflies, pond skaters (Baptista
sp.), the larvae of mosquitos (Toxorhynchites sp.)
and young, aquatic snakes. Due to their substan-
tial sizes and physical powers, snakes (e.g. spe-
cies of the genera, Python and Ptyas) play impor-
Mount Kinabalu - the surrounding environment
© U. Manthey
Fig. 26 Dendrelaphis pictus eats a frog of the family
Ranidae (photograph was not taken from Borneo).
tant roles as predators among vertebrates. Many
of the climbing species (e.g. Boiga, Dendrelaphis,
Gonyosoma, Trimeresurus) prey mainly upon
birds and plunder nests. Smaller arboreal (e.g.
geckos, agamids, scincids), and fossorial species
(e.g. Calamaria, Typhlops, Sphenomorphus) feed
mainly on arthropods and worms; the snail-eat-
ing snakes of the genera Pareas and Internatus
are largely specialized in preying upon molluscs
and Gongylosoma upon spiders.
Predators of reptiles are predominantly found
among predatory small mammals, wild boars, and
birds (e.g. hawks, cockoos, thrushes, pittas). Lizards
are the favourite food of many snakes, the smaller
ones being often captured by large hunting spiders.
Besides the entirely unresearched parasites, the
numerous predators are a major controlling factor
of the populations of amphibians and reptiles.
Being carnivorous, amphibians and reptiles are sec-
ondary consumers of other secondary and primary
(vegetarians) consumers with amphibians usually
occupying a midfield position in the food chains
as second, third, and fourth level consumers. This
is also true for many reptiles, although larger spe-
cies may also be top predators in many of food
chains, within the food web.
31
Mount Kinabalu - the surrounding environment
Herpetofauna
As of 2000, 77 species of amphibians and 112 of
reptiles are known from Mt. Kinabalu. Besides
some species settling in cultivated areas which oc-
cur in the marginal zones (e.g. Kaloula pulchra,
Polypedates leucomystax, Rana erythraea, Rana
nicobariensis, Fejervarya limnocharis, Gehyra
mutilata, Hemidactylus frenatus, Hemidactylus
garnotii), the herpetofauna of the Kinabalu massif
is composed mainly of forms particular to sub-
montane and montane forests. These inhabit either
trees at various storeys, the leaf-littered forest floor,
they may live subterrestrially, or have adapted to
habitats dominated by mountain streams.
Many species spend their entire life in the branch-
work of trees and may only descend to, or near, the
ground for a short period of time to reproduce; oth-
ers are ground-dwellers, which occasionally can
be found in the lower vegetation stratum. While
some species (e.g. Chaperina fusca, Rhacophorus
nigropalmatus) undertake extensive journeys
through the forests, others inhabit rather small
niches which they do not leave for years. The ver-
tical distribution of some may stretch over more
than 1300 m in altitude (e.g. Ansonia fuliginea,
Ansonia longidigita, Pelophryne misera, Cyrtodac-
tylus baluensis, Phoxophrys cephalum, Amphiesma
sarawacense), while others are only known from
certain, well-delimited localities (e.g. Ansonia
guibei, Leptolalax maurus).
The adaptations to a life on the trees are particu-
larly manifold. Arboreal reptiles have developed
specific organs which aid climbing, such as
scansory pads (Gekkonidae), strong, curved, and
pointed claws (Gekkonidae, Agamidae), keels be-
tween the ventral and lateral scales (Serpentes: e. g.
Dendrelaphis, Chrysopelea), or prehensile tails
(Serpentes: Trimeresurus). Frogs, on the other hand,
may have adhesive, glandular surfaces on their ven-
tral sides and the pads of their finger and toe tips
(Rhacophorids possess a cartilaginous joint be-
tween the subterminal and terminal joints which
allows the pad to be placed onto the surface at a
right angle).
To be able to bridge larger distances between
branches and tree trunks, frogs employ various leap-
32
ing techniques which are aided by particularly long
hind limbs. Snakes have solved this problem by
employing a longitudinal muscle contraction which
results in a stiffening of the body axis. Some spe-
cies have acquired highly specialized adaptations,
such as the ability to reduce the sinking speed of a
free fall by paragliding as a result of an active en-
largement of the body surface thus enabling them
to bridge larger distances in an oblique, controlled
gliding flight. This may be achieved by a spread-
ing of the ribs (flying snake: Chrysopelea), which
may also be extremely enlarged and interconnected
by skin membranes (patagia) (flying dragons:
Draco), by employing appendicular lateral skin
fringes like sails (flying geckos: Ptychozoon), or
unproportionally large digital webbings (flying
frogs: Rhacophorus). WALLACE (1869) was the first
to describe and illustrate a “flying” frog (Rhaco-
phorus nigropalmatus) from Sarawak, a species
also present on Mt. Kinabalu.
Rain forests are marked by a distinct stratification
of animal communities in relation to its zonation.
The various genera of scincid lizards (Scincidae)
may serve as an illustration of this fact, with
Sphenomorphus and Mabuya living mainly on the
forest floor, Apterygodon on tree trunks, and
Lamprolepis in the canopy. In the case of agamid
lizards, the circumference of the tree trunks leads
to another type of segregation, with Draco distinctly
favouring the mighty trunks of dipterocarps,
Fig. X Rhacophorus nigropalmatus after WALLACE 1869.

Gonocephalus trunks with a smaller diameter (usu-
ally < 30 cm in diameter), and Phoxophrys and
Aphaniotis slim trunks and bushes. Frogs of the
genus Megophrys spend their lives on the ground,
Philautus amoenus prefers levels between 10 and
200 cm above the ground in the mossy forests, and
Philautus bunitus occurs at 3 metres plus on trees.
Due to a lack of research we know next to nothing
about the herpetofauna of the canopy.
In contrast to the arboreal species, numerous
ground-dwelling inhabitants exhibit particular ad-
aptations to a shady, moist environment. Roots,
fallen logs in all stages of rot, rock flakes, and an
all-covering layer of leaf litter offer a highly dif-
ferentiated range of niches. Besides caecilians (i.e.
Ichthyophiidae) and terrestrial frogs (Lepto-
brachium, Megophrys, Kalophrynus), the leaf lit-
ter community includes numerous fossorial to semi-
fossorial snakes (e.g. Bungarus, Calamaria, Cylin-
drophis, Calliophis, Pseudorabdion, Oreocalamus,
Typhlops), forest skinks (Sphenomorphus), worm
skinks (Brachymeles), and glass lizards (Ophi-
saurus). Blind snakes (Typhlopidae) and dwarf
snakes (Calamaria) exhibit typical adaptations to
a burrowing-boring way of life in the forest floor
with the head being reinforced and compact with a
reduced scutellation and reduced eyes (the blind
snakes have eyes covered by scales), no distinct
slimming of the neck, a short, spine-like, pointed
tail suitable for anchoring, and a very smooth
scutellation of the body. Mabuya and Spheno-
morphus actively utilize shifting sun patches on the
forest floor for their thermoregulation.
Frogs of the genera Ansonia, Huia, Limnonectes,
Meristogenys, Rana, Staurois, and keeled skinks
(Tropidophorus) strongly depend on mountain
streams; some snakes (e.g. Amphiesma flavifrons,
Xenochrophis trianguligerus) and the turtle
Dogania subplana are hygrophilous. The larvae of
71% of the anuran species of Mt. Kinabalu are
known, 56% of which are inhabitants of mountain
streams, the remainder develops in ponds and
phytothelms. The species of the genus Philautus,
which are known for their eggs to develop into
froglets within a shell of jelly, deposit their eggs
terrestrially at sufficiently moist sites (cushions of
moss, rock crevices, beneath bark, and in dead
pitchers of Nepenthes). Some ranids (Limnonectes
Mount Kinabalu - the surrounding environment
© R. Malkmus
Fig. 27 Semiadult Rhacophorus everetti macroscelis,
as example for lichen mimesis.
finchi, L. leporina, L. palavanensis) engage them-
selves in parental care for their developing off-
spring. Tadpoles living in mountain streams are
usually adapted to a life in certain ecological niches
with some possessing highly specialized adhesive
organs to withstand strong currents.
A remarkable feature in the appearances of amphib-
ians and reptiles is the high content of shape and
colour elements which correspond with plant struc-
tures (phytomimesis). In this respect some repre-
sentatives of the families Megophryidae, Bufoni-
dae, and Microhylidae imitate dead leaves (inclu-
sive of the leaf’s ribs and imperfections caused by
phyllophagous insects), while lichen and bark struc-
tures are found in the body coloration of various
frogs (Rhacophorus everetti, Ansonia hanitschi) and
reptiles (Draco, Phoxophrys, Ptychozoon), while
again others resemble twigs and lianas (Ahaetulla,
Dendrelaphis, Pareas, Psammodynastes, Trimere-
surus). By their shapes and colour patterns, frogs
of the genera Ansonia, Meristogenys, and Staurois
blend perfectly into their surroundings of rocks in
the splashing zone of small waterfalls which are
partly overgrown with algae. Some Amphibians and
reptiles further enhance these cryptic effects by
remaining motionlessly attached to the surface
when they feel they might have been discovered.
Many species (e.g. of the genera Philautus, Rhaco-
phorus, Bronchocela, Phoxophrys, Draco) further-
more possess the ability to rapidly alter their col-
orations. This physiological colour change is hor-
mone-driven and may be triggered by changes in
33
Mount Kinabalu - the surrounding environment

Fig. XI Numbers of species in anuran families and genera present on Borneo and Mt. Kinabalu.

34
 Mount Kinabalu - the surrounding environment

Fig. XII Numbers of species in saurian families and genera present on Borneo and Mt. Kinabalu.

35
Mount Kinabalu - the surrounding environment

Fig. XIII Numbers of species in snake families and genera present on Borneo and Mt. Kinabalu.

36

the surrounding temperatures, the intensity of light,
the level of humidity, but also by excitement. These
colour changes do, however, not only serve the
purpose of camouflage, but also play roles in com-
munication and thermoregulation. A perfect exam-
ple for the function mentioned last is Philautus
aurantium which possesses guanophores in its skin
which reflect sunlight. It enables the frog to avoid
overheating by turning almost white in response to
being exposed to sunlight while resting on a leaf
during the day. Aposematic, brightly contrasting
warning colours, as known from Neotropical
anurans (e.g. Atelopus, Dendrobates) are almost
entirely absent in the species of Southeast Asia.
The change of climate-induced ecological actuali-
ties, particularly between altitudes of 1200 and
1400 m, and between 1700 and 1900 m, respectively,
corresponds not only with changes in the composi-
tion of the flora, but also with those of the fauna.
Numerous species of amphibians and reptiles reach
their maximum, respectively minimum, vertical dis-
tribution limits at these two altitudinal levels. The
overlapping existence of highland and lowland forms
in the area between 1200 and 1400 m causes the
slight increase of number of species, though tenden-
tious decreasing with increasing altitude. The often-
observed congruences of these distributional rela-
tions in species of various genera underlines the dis-
tributional ecological importance of these bounda-
ries. A drastic reduction of the specific diversity be-
comes evident at elevations above 1800 m, in the
transitory zone to the mossy forests, so that the state-
ment by INGER (1978), “One of the disappointing
things about Kinabalu’s frog fauna is that it shows
so little altitudinal stratification.”, clearly demon-
strates the need for fundamental revisions. This state-
ment was very obviously based on an insufficiently
large pool of information on the distribution of the
amphibians in this area at that time.
While our knowledge of the vertical distribution
of reptiles on Mt. Kinabalu is grossly insufficient
(only a handful, partly doubtful records exist for
altogether five species from above 1800 m of which
the records of Rhabdophis murudensis and Cyrto-
dactylus baluensis from 2500 m currently represent
the maximum of the vertical distribution of rep-
tiles from Borneo), at least some of the more vocal
amphibians provide us with more acceptable in-
Mount Kinabalu - the surrounding environment
formation on their altitudinal distribution. Many
genera (e.g. Leptolalax, Leptobrachium, Mego-
phrys, Kalophrynus, Ansonia, Philautus) appear on
Mt. Kinabalu in the form of species groups, with
the distribution of the individual species being lim-
ited to certain elevations. It requires only an ascent
up the mountain at night and the personal experi-
ence of just how sharply the individual vocaliza-
tions supersede one another and can be heard
sympatrically only in narrow overlapping zones,
to eliminate any doubts as to the existence of such
abrupt vertical distribution boundaries.
Figures VIII and IX (p. 26 and p. 27) shows clearly
(with the exception of the area between 1200 and
1500 m) that the number of species diminishes
gradually with increasing altitude. Various reasons
are accountable for this. A particularly important
factor - especially as far as reptiles are concerned -
is of course the climate whose supportive qualities
for poikilothermic species of tropical areas dete-
riorate rapidly with increasing altitude (lower tem-
peratures, increasing precipitation, frequency of
cloud cover). With increasing altitude the follow-
ing changes are observed:
• a reduction of the variety of ecological niches
which is largest in the dipterocarp forests;
• a reduction of the availability of food with the
specific diversity and density of potential prey
(mainly invertebrates) being subject to substan-
tial limitation with increasing altitude;
• a reduction of potential spawning sites. As a re-
sult of the topography, stagnant water bodies are
absent and the streams above 2000 m usually have
steep gradients. Their water levels often fluctuate
between drying up completely and extreme high
water. With the exception of Ansonia fuliginea,
whose larvae are unknown, none of those species
of frogs are found above 2200/2300 m which de-
posit their spawn in streams.
For reasons already discussed in chapter
“Zoogeography”, the occurrence of endemic am-
phibians and reptiles on Mt. Kinabalu correlates
positively with the altitudinal level between 1400
and 1900 metres. It must be said, however, that their
stati as endemites can at best be considered tenta-
tive as the totality of the species inhabiting the moun-
tains of Borneo is not sufficiently known. In fact,
many species previously considered endemites of
37
 Mount Kinabalu - the surrounding environment
Mt. Kinabalu have since also been recorded from
other mountains of the island, examples being
Leptobrachella baluensis, Megophrys baluensis (?),
Occidozyga baluensis, Sphenomorphorus kinaba-
luensis, Meristogenys kinabaluensis, Pelophryne
misera, Cyrtodactylus baluensis, Oreocalamus
hanitschi, etc.
At present, 13% of the amphibian species of Mt.
Kinabalu are considered endemic to the moun-
tain, i.e. Leptolalax arayai, Leptolalax maurus,
Leptobrachium gunungense, Megophrys koba-
yashii, Ansonia fuliginea, Ansonia guibei, Kalo-
phrynus baluensis, Philautus amoenus, and
Philautus saueri. All of them have as to date only
been found above 1400/1500 metres. 7 % of the
reptiles of Mt. Kinabalu are currently regarded en-
demic, i.e. the lizards Hypsicalotes kinabaluensis,
Complicitus nigrigularis, Ptychozoon rhacophorus,
Sphenomorphus tenuiculus, and the snakes,
Calamaria schmidti, Hydrablabes praefrontalis, and
Trimeresurus malcolmi. Since some of these spe-
cies descend as far down as to Poring, their stati as
endemites must be considered doubtful.
Fig. 28 Vegetable plantations between Bundu Tuhan and Kundasang.
© R. Malkmus
38
Human beings in the vicinity of
Mount Kinabalu
“Enough has been discovered for it to be
realized that for many animals the rainforest
habitat is essential and that they cannot live in
secondary forest.”
Whitmore, 1988
“The loss of biological diversity is the
great tragedy of our age - I consider it to be the
single most important and lasting of our spe-
cies on this planet.”
Davis B. Wake, 1994
Discoveries in the Niah Caves, Sarawak, prove that
there has been human life on Borneo for at least
40000 years. The population density, however, was
at such a low level until the last century, that around
1900 more than 90% of the island were still cov-
ered with a complete and coherent primary forest.

Fig. 29 Woods detroyed by fire (Wariu Valley, March 1998).
Although SHELFORD (1916) had already complained
about an extensive destruction of the forests in the
vicinity of settlements, the real decline of the rain
forests did not start until the end the World War II.
By 1957 the island had already lost 25% of its for-
est. In 1953, Sabah, where every citizen could claim
land as long as he was willing to clear and culti-
vate it, was still covered with forests to an extent
of 86%. In 1983 there were only 63% left, in 1990
only 41%. The time has passed in which one could
still write like MJÖBERG (1928): “The gigantic is-
land of Borneo is clothed from head to foot, as it
were, in green virgin forests.”.
How did this dramatical decline come about?
The most important reasons are:
* Drastic increase of population. This leads to a
continually growing need for new areas to settle
in, which automatically implies infrastructural
measures and woodland clearances.
* Slash-and-burn as well as shifting cultivation by
farmers; during some years with prolonged
droughts in relation to El Niño Southern Oscil-
Mount Kinabalu - the surrounding environment
© R. Malkmus
lation events between 1982 and 1987 as well as
in 1997/98 this kind of agriculture caused fires
of devastating extent; a total area of about 5-6
million ha. had to be sacrificed to these fires in
the course of just the last ten years.
* Highly developed industrial nations (especially
Japan) bought permissions for extensive logging
for a period stretching far beyond the turn of the
millennium.
* Agro-industrially organised, large concerns pro-
mote the monocultural cultivation of oil palm
trees and rubber plants on the cleared soil.
These developments did not spare the forests of Mt.
Kinabalu, as well. The surroundings of the moun-
tain are inhabited by the Dusun people. Tradition-
ally, they practise shifting cultivation (economy of
changing land based on slash-and-burn cultivation:
after a period of a 1-2 years’ cultivation of a soil
extremely lacking nutrients there follow fallow pe-
riods of various lengths), but restricted their eco-
nomic actions to the glens of Kadmaian and the
vicinity of Ranau, for the mountain itself was a
39
Mount Kinabalu - the surrounding environment
religious taboo. It was not before the middle of the
20th century, when the belief in the mountain gods
had deteriorated to a mere myth, that extensive
forested areas facing south were cleared - up to an
altitude of 1400 metres. Between Kundasang and
Ranau extensive vegetable plantations were estab-
lished; in 1958 a jeep track was built between the
west coast and Ranau which was extended in 1972
to the so-called Transsabah Highway connecting
Kota Kinabalu and Sandakan. In 1964, 711 km² of
the Kinabalu massif were declared a protected Park
under the administration of Sabah Parks; moun-
tain lodges (in between 3200 and 3800 m) and a
visitors’ centre (Headquarters, at 1524 m) were es-
tablished, followed by additional ones at Poring
(500 m), close to “Hot Springs”, in 1967, and in
Mesilau (1860 m) in 1998.
Eventually, 25.5 km² were sacrificed to enable the
establishment of a copper mine in the vicinity of the
Sungei Mamut source. In order to compensate for
the loss and to silence critical voices, the park was
enlarged to the north into the Templer region. This
region covers an area of 93 km², but large parts of it
consist of substituted forests on logged-over land.
In 1983, one of the big fires raged in the eastern
foothills, and it was only due to sheer coincidence
that it did not reach the buttresses of the mountain
itself. The omnipresent tampering with fire contin-
ues to be common practice around the mountain and
is not liable to any restrictions. In 1984, the park
boundaries were redefined and the protected area
was reduced by 26 km². Two areas of a particularly
rich biodiversity were sacrificed up to an altitude of
about 1800 m during these reductions, the rain for-
ests of Bukit Hampuan and of the Pinosuk plain.
Temperate fruit and flower farms, a dairy farm, and
a golf course were established on the deforested
aeras. When in 1989 first steps were taken to re-
integrate Bukit Hampuan into the park, a fire de-
stroyed the major part of the remaining forests.
Within the boundaries of the park as well, there
are deficiencies hard to understand, e.g. the intro-
duction of a sewage into Sg. Silau Silau, water sup-
ply taken from the upper parts of Sg. Silau Silau,
Sg. Liwago and its tributaries. During persistent
periods of drought this leads to a partial running
dry of entire stretches of the rivers and brooks, the
overdimensional new road at Mesilau with ditches
40
for rainwater (40 cm deep) running alongside with-
out the slightest support for animals to crawl back
up. These ditches form deadly pits for many of them
(in March 1998 we found 34 withered Ansonia and
Meristogenys and three Sphenomorphus within a
stretch of about 100 metres).
Travelling from the Headquarters down to Ranau,
respectively from Sayap down the Wariu valley, is
an experience of a devastatingly harsh scenic con-
trast. Mountain forests still untouched and pure a
second ago abruptly give way to a chaotic view with-
out any gradual transition; the traveller is confronted
with the entire range of unthought-of methods of
overfelling and overexploitation: all over the plaxe
randomly dispersed huts of corrugated iron, slopes
eroded by mud avalanches, bare ridges, residues of
secondary forests crumbled to charcoal, simmering
rubbish dumps, polluted brooks with an increased
water runoff and increased sedimentation rates, and
the churned-up areas of copper mines.
Ignorance, lack of organisation, unlimited pursuit
of profit, and the hunger for land amongst an in-
cessantly growing population have contributed to
the gradual shrinkage of the belt of a protecting
buffer zone around the mountain. But the time of
patchwork tailoring is drawing to an end: it is not
possible anymore to burn away forest in the south
and add its territory in the north, for one is running
out of wood resources there as well. In the mean-
time, the mountain’s substance has become totally
exposed; its overboarding biological treasures are
forced to recede to a highly fragile and extremely
jeopardised insular territory.
A retrospective glance on
herpetological research in Borneo and
the area of Mt. Kinabalu
“The collector is far too prone to kill at
sight every animal he captures; he is usually a
bird of passage, and has not the time to devote
to the patient and difficult observation of an
animal’s behaviour and habits of life.”
R.W. Shelford,
A Naturalist in Borneo, 1916

Until the beginning of the 19th century the Nether-
lands did not have a firm grip on more than a cou-
ple of forts along the coast. Ventures into the inte-
rior of the islands, into the realm of cannibals, were
looked upon as suicidal. Not before the time be-
tween 1820 and 1850 did the “Natuurkundige
Comissie” organize research excursions, which
invaded the central part of the island (e.g. by Hor-
ner, Gaffron, Müller, Schwaner). The first zoologi-
cally important collection (of mammals and birds)
was established on behalf of the Rijks Museum van
Natuurlijke Historie at Leiden. The herpetofauna
did not attract any attention for quite a long time.
The great snake atlas by SCHLEGEL (1837) does not
mention more than one species as far as Borneo is
concerned, Hydrophis pelamis. SCHLEGEL (1848-
54), GRAY (1845, 1859), and GÜNTHER (1859, 1864)
then described for the first time the amphibians and
reptiles of Borneo stored in museum collections at
Leiden and London. Shorter herpetological works
were published by BLEEKER between 1857 and
1860, and EDELING in 1865. In 1885, FISCHER de-
scribed a small collection made by Grabowsky, in
1892, MOCQUARD classified one by Chaper. The first
check list of the amphibians and reptiles of Bor-
neo (GÜNTHER 1872) comprised already 157 spe-
cies. Since by now more than a hundred years no
significant progress has been accomplished as to
the knowledge about the distribution and ecology
of the herpetofauna of Kalimantan, which suffers
from a lack of research anyway.
The scientific research in the nowadays Malaysian
part of Borneo has undergone a totally different
development. Its beginning is inextricably inter-
twined with the reign of the “White Radjas” of
the Brooke dynasty, who resided at Kuching/
Sarawak. Scientifically interested Sir James
Brooke was of paramount importance in this re-
spect. He established his own collection, founded
the Sarawak Museum at Kuching and roused in-
terest in undertaking trips into the interior amongst
naturalists and civil servants of his own adminis-
tration by providing generous support. Without
doubt, A.R. Wallace, who toured Sarawak in 1856,
was the most well-known amongst them. During
this time the first publication dealing with local
herpetological data (MOTLEY & DILLWYN 1855) oc-
curred.
Mount Kinabalu - the surrounding environment
Under the rule of James Brooke, the “Sacred Moun-
tain” became an object of scientific research. His
colonial minister Hugh LOW reached the summit
plateau for the first time on the 11th of March 1851,
and he repeated the ascent together with Spencer
St. John in 1858. The latter vividly depicts the ex-
pedition in his book, “Life in the Forests of the Far
East (1862)”. The trials and tribulations, which
lasted for weeks, did not mainly consist of the
physical challenge of the ascent itself, but rather
of the problems with establishing oneself at the base
camp (the Dusun village Kiau) and with the re-
cruitment of porters, who had to be convinced to
invade the sacred aura of the mountain. The topog-
raphy only allows one route of ascent, which was
also taken by all subsequent expeditions. Still to-
day the Summit Trail sticks to this route from 1800
m onwards, i.e. from Kiau (914 m) passing Lumu
Lumu (1676 m), Kamborangoh (2195 m) and Pakka
cave (3109 m), up to the region of the summit.
New route up Mount Kinabalu Mesilau trail
promises more challenges
Source: The Star, Friday, January 8, 1999
KOTA KINABALU
Trekkers intending to make their way up Mount Kina-
balu, whether for the first or umpteenth time, can now
try out a new route which is also more challenging
than the existing trail which has been used more than
30 years. The new route, called the Mesilau Summit
Trail, is located on the lesser-known eastern face of
the mountain. It starts at the recently-opened Mesilau
Nature Resort ..... near a restaurant at the resort.
The new 6 km route ends at the Layang Layang hut
and hikers will have to trek for another 2 km before
reaching Laban Rata ......the new trail will trek through
a montane forest and pass the Mesilau waterfall.
..... “After 2 km at an altitude of 3000 to 4000 m, the
trail goes downhill through a valley ..., hikers can see
a huge Agathis damara tree with its unique branches
and roots,”.... After passing through a number of
streams and about 2.5 km into the journey, a 45 m-high
waterfall can be seen. Then for another 2 km, the path
snakes up a steep and narrow hill, the toughest part of
the route. ...after this section, the trail levels for 1.5 km
and pitcher plants abound on both sides of the path. ...
the trail then takes hikers through a rocky slope before
they reach the Layang Layang hut at an altitude of
2702 m.
41
Mount Kinabalu - the surrounding environment
Fig. XIV The summit route of the expeditions between 1851 and 1930.
It has to be pointed out already at this stage that
these altitudes, which were applied to later mate-
rial, should be given a closer look before being
taken over. This is especially valid as to “Kambo-
rangoh, 2195 m”. In spite of intensive research,
many species (e.g. Ansonia longidigita, Kalophry-
nus pleurostigma, Cyrtodactylus baluensis, Sphe-
nomorphus kinabaluensis) which were collected by
Chasen and Pendlebury in 1929 and attributed by
SMITH (1931) to this location could not be found at
this altitude. Presumably they were collected some-
where alongside the track between Lumu Lumu and
Kamborangoh.
While the first attempts to climb the summit pla-
teau are to be treated as rather sportive events with-
out any scientific fruits worth mentioning, the or-
nithologist J. Whitehead did not only manage to
climb the highest point of the plateau (Low’s Peak,
1888) for the first time, but also to earn scientific
42
modified after
PENDLEBURY & CHASEN 1932
merits; to Whitehead “belongs the credit of the
first extensive exploration of the mountain and to
him we owe our first real insight into the zoologi-
cal treasures” (MOULTON 1915). In 1893, WHITE-
HEAD’s brilliant work “The Exploration of Kina-
balu North Borneo” was published. It was a deci-
sive contribution to the formation of a new aura,
the aura of an unfathomable biological treasure.
During the following decades Mt. Kinabalu be-
came the place of pilgrimages for botanists and
zoologists alike, but never did a herpetologist oc-
cur amongst them. The herpetological section of
Whitehead’s collection was not edited by him him-
self but by BOULENGER (1887) and M OCQUARD
(1890). Their list of species presents the first sci-
entifically recorded amphibians and reptiles in the
Kinabalu region.
During the following 40 years civil servants of the
Sarawak and Raffles Museums brought minor

collections from Mt. Kinabalu every now and again,
which were described by herpetologists who were
not acquainted with the country by own experience;
examples are the systematic descriptions of the col-
lections of A.H. Everett by G.A. BOULENGER (1893,
1896), of R. Hanitsch by G.A. BOULENGER (1899),
of J.A. Griswold by A. LOVERIDGE (1938), and of the
biggest one, comprising 600 specimens, of F.N. Cha-
sen and H.M. Pendlebury by M.A. SMITH (1931).
Due to the habit of considering the collector when
creating the names of new species, we frequently
encounter in them the names of scientists and of
civil servants working as curators who dealt with
fields of research totally different from herpetology
(e.g. Tropidophorus beccarii, Ovophis chaseni,
Pedostibes everetti, Oreocalamus hanitschi, Cala-
maria griswoldi, Philautus mjobergi, Meristogenys
whiteheadi, etc.).
Since the mid-fifties of the 20th century the research
into the herpetofauna of Sarawak and Sabah is very
much intertwined with the personality of R.F. Inger,
now the Emeritus Curator of the Amphibians and
Reptiles at the Museum of Natural History, Chi-
cago. His classical work “The Systematics and
Zoogeography of the Amphibia of Borneo (1966)”
comprises the entire knowledge about the amphib-
ians of the island available at that time. It also
shows, however, how fragmentary the knowledge
was back then. While already a high percentage of
the descriptions of species is based on very old ma-
Fig. 30 G.A. Boulenger Fig. 31 M.A. Smith
Archiv: K. Adler
Mount Kinabalu - the surrounding environment
terial, data about ecology and ethology can be found
only in fragments or are totally missing. This is the
result of a research methodology predominant dur-
ing more than a hundred years, which is character-
ized quite concisely by Shelford, as quoted in the
heading of this chapter. As to the data concerning
distribution, INGER himself remarked: “Analysis of
the distribution of amphibians within Borneo at this
time is more likely to reflect the distribution of col-
lectors than of frogs.”.
During the subsequent years INGER wrote (partly in
co-operation with other authors like BACON, DRING,
EMERSON, FROGNER, GRITIS, HAILE, HOSMER, KING,
LLOYD, TAN, VORIS, and WALKER), a number of trea-
tises on population dynamics, the biology of re-
production, and the settling in ecological niches
by various species for the first time. Apart from
that he published many taxonomical papers describ-
ing numerous new species; he wrote particularly
recommendable works about the systematics and
ecology of tadpoles.
Further descriptions of new amphibians were pub-
lished by ISKANDAR (1978), DRING (1983a, 1983b,
1984), MALKMUS (1992b, 1996d), MATSUI (1986,
1997), MALKMUS & RIEDE (1996a), MALKMUS &
MATSUI (1997), and MANTHEY & STEIOF (1998). Due
to these new insights and data the number of known
and described species has increased from 93 (INGER
1966) to 152 (Anura from 91 to 146, Ichthyophiidae
from 2 to 6) amongst them Barbourula kalimanta-
Fig. 32 R.F. Inger
Archiv: Forschungsinstitut Senckenberg
43
Mount Kinabalu - the surrounding environment
nensis by ISKANDAR, 1978, a member of a family
not yet verified for Borneo until then.
Comments on the ecomorphology, ethology, activ-
ity rhythms, and the bioacoustics of specific species
were published by DENZER (1994b), DRING (1983a,
b, 1987), EMERSON (1981), HARDING (1982), HOFF-
MANN (1995c, 1998c, 2000), L AKIM , Y AMBUN &
AIRAME (1999), M ALKMUS (1989, 1992a, 1993,
1994a, 1994b, 1995a, 1996a, b, e, 1997, 1998),
MALKMUS & RIEDE (1996a, b), MATSUI (1979a, 1996),
MATSUI, WU & YONG (1993), MATSUI, YONG, ARAYA
& HAMID (1996), and WONG (1994).
Research on reptiles still lingers on a rather low level
compared with that on amphibians. The latest total
survey is provided by the work of N. DE ROOIJ (1915,
1917), which encompasses the whole Indo-Malayan
territory. MANTHEY & GROSSMANN (1997) published
a volume, which comprises the whole variety of all
known genera of the Southeast Asian herpetofauna;
its impressive gallery of brilliant colour photographs
adds to its scientific merits. Amongst the more re-
cent revisions of the description of families, genera
and species, INGER’s works (partly in co-operation
with GREENBERG, HOSMER, LEVITON, and MARX) oc-
cupy once again a dominant position, besides those
of BACON (1967), HIKIDA (1982, 1990), GÜNTHER &
MANTHEY (1995), MANTHEY (1991) MANTHEY &
Fig. XV Graphic illustration of new records of amphibian and reptile species in the Mt. Kinabalu region
(> 500 m a.s.l.) during the individual decades since the beginning of observations.
44
GROSSMANN (1997), MANTHEY & DENZER (2000),
MUSTERS (1983), OTA & HIKIDA (1991) and WALLACH
& GÜNTHER (1998).
HAILE (1958) and STUEBING (1991) published check
lists of the snakes of Borneo, INGER & TAN (1996)
did the same with regard to frogs, the lizards of Sabah
were listed by TAN (1993), the amphibians and rep-
tiles of Brunei by DAS (1995), the amphibians and
reptiles of Mt. Kinabalu by MANTHEY (1983) and
MALKMUS (1994b), the amphibians of Kinabalu by
INGER, STUEBING & TAN (1996) and INGER, TAN &
YAMBUN (2000). It is pleasant and satisfying to note
that recently five books about Bornean amphibians
and reptiles, written in a popular scientific vein, were
published (INGER & STUEBING 1989, 1997, INGER &
TAN 1996, STUEBING & INGER 1999, LIM & DAS 1999).
Around the middle of the 20th century Mt. Kina-
balu lost its formerly dominant position among the
research territories of Borneo. This was not exclu-
sively due to the fact that one was tempted to be-
lieve that after a whole generation of scientists’ in-
tensity of work its zoology and vegetation were suf-
ficiently explored; another important role played the
fact that an exacerbated infrastructure and the use
and deployment of helicopters made the remotest
areas accessible (especially the central highland of
Sarawak, the mountainous regions between Gunung

Mulu - Murud - Lumaku and Crocker, the Danum
Valley in the eastern part of Sabah, etc.). The expan-
sion of the examination area brought about the in-
sight that many species, which were believed to be
endemic, turned out to inhabit other areas of the is-
land as well (cf. INGER 1978, INGER & STUEBING 1992).
This was, however, nothing new to ornithologists
(MJÖBERG 1928).
Surprisingly, investigations undertaken since 1984
lead to the discovery of numerous new species of
amphibians and reptiles, all of which still have to be
treated as endemic species. This demonstrates that
the investigation of the herpetofauna of Mt. Kinabalu
is far from approaching a final conclusion.
Figure XV demonstrate the historical development
of coming to know the range of species at Mount
Kinabalu. Scientists doing field work regularly
brought back collected material from their travels.
Most of the herpetological material is looked after
in the following collections in museums throughout
the world:
The British Museum of Natural History, London
(BM); Field Museum of Natural History, Chicago
(FMNH); Kyoto University Zoological Collection
(KUZ); Museum of Comparative Zoology, Cam-
bridge/Mass. (MCZ); Muséum National d’Histoire
Naturelle, Paris (MHNP); Naturhistorisches Museum
Wien (NHMW); Naturhistorisches Museum Basel
(NMB); Osaka Museum of Natural History, Osaka
(OMNH); Sabah Parks-Field Museum, Headquarter/
Kinabalu Park; Sarawak Museum, Kuching (SMK);
Forschungsinstitut Senckenberg, Frankfurt/Main
(SMF); Raffles Museum of Biodiversity Research,
National University of Singapore (USDZ/ZRC);
Sabah State Museum, Kota Kinabalu (SSM); Coll.
Universiti Kebangsaan Malaysia, Sabah Univ. Cam-
pus (UKMS); United States national Museum, Smith-
sonian Inst., Washington (USNM); Zoologisches
Forschungsinstitut und Museum A. Koenig, Bonn
(ZFMK); Museum für Naturkunde der Humboldt-
Universität zu Berlin (ZMB).
One could still agree with Inger’s comment on am-
phibians, dating from 1978, although a moderate
modification seems recommendable: “Almost none
of the information on natural history comes from
information made at Kinabalu. There is much to
be done.”
Mount Kinabalu - the surrounding environment
This quotation does not only delineate a defi-
ciency, but also functions as an appellation to
draft a programme of future research activities.
It is also an appellation to get going. Especially
three fields should form the centre of investiga-
tion:
* Systematics: a whole variety of systematic prob-
lems are to be tackled in this context. This should
not be attempted on a merely morphological ba-
sis, but also by including and considering ecologi-
cal, ethological, and molecular data. Naturally,
some uncertainties will not yield to clarification,
as the products of a systematic approach are artifi-
cial and - like WILSON (1997) adequately pointed
out: “science to a high degree, but a bit of art as
well”.
* Distribution: nearly all the data about places where
animals were found is restricted to the close envi-
ronment of Poring, the Headquarters, Mesilau,
Sayap, Bundu Tuhan, and Kiau; apart from the
summit trail there are no data concerning the areas
above 2000 metres. All the data refer to the lower
stratum of the forest (below 5 metres). Future in-
vestigations should include areas outside this very
limited scope of observation, which barely covers
approximately five percent of the total space. Al-
most nothing is known about the equitability (rela-
tive density and distribution) of the various spe-
cies.
* Natural History: we barely know anything about
the natural history of most of the species (includ-
ing the larvae). There is a lamentable lack of stud-
ies research dealing with ecology and ethology
(e.g. reproduction, acoustics, population dynam-
ics, activity rhythms, settlement in niches, etc.).
As a crucial step forward in investigating the
herpetofauna of Mt. Kinabalu, Sabah Parks have
initiated a program in 1991. They work together
with INGER, who monitors the population dynam-
ics of selected species at three sites at different lev-
els of elevation.
This book attempts to sum up the present state of
knowledge about the amphibians and reptiles of
Mount Kinabalu. It soon will become evident that
the number of unsolved problems is vast. No doubt:
“There is much to be done”!
45
 Rhacophoridae

Amphibia (amphibians)
Some 350 million years ago, during the Devon period, the first
amphibians developed from archaic fishes eventually reaching their
evolutionary peak between the periods of Carbon and Triassic. They

us
were the first vertebrates to conquer firm land. Their close tie to a

km
al
M
moist environment and the fact that, in most of the species, the early

R.
©
ontogenetic phases necessitate water for the gill-bearing larvae still
give a clear indication of their origin.
The recent amphibians are poikilothermic vertebrates with a nude skin
which is kept moist by excretions from mucous glands. With the
exception of the gymnophionians (caecilians) amphibians possess two
pairs of limbs. The skull is flattened and articulated by the atlas (first
neck vertebra) to the spine, the sternum (breastbone) is not connected
to the ribs. The gymnophionians lack a sternum as well as shoulder
and pelvic girdles. They breathe through lungs, via the mucous
membranes of the mouth cavity, and through the moist skin.
About 5000 species of amphibians are known, which are grouped in
three orders, i.e.:
Tailed amphibians (Urodela or Caudata)
Tailless amphibians (Anura or Salientia)
Caecilians (Gymnophiona or Apoda)
Borneo is home to tailless amphibians and caecilians whereas tailed
amphibians are absent, just as the latter are missing in the entire Sunda
Archipelago.

Anura (tailless amphibians or frogs)

Frogs are marked by very obvious external traits, i.e. a stout build of Ranidae
the body, no tail, two pairs of limbs of which the hind pair is usually
stronger and longer, four fingers and five toes, and large eyes and a
very large mouth in most of the species.
Their ontogenetic development involves a larval stage in the form of
gill-bearing, tailed tadpoles which either develop free-swimming or
within an egg to frogs (metamorphosis).
Anurans are represented on Borneo by 6 families comprising 30 genera
th

and 146 species (as of 2000). The ongoing process of discovering new
ro
ch
.S

species, and the ever-increasing knowledge gathered by means of

M
©

research in comparative anatomy, morphology, ethology, and by genetic

and phylogenetic analyses demonstrate, however, that the systematics
of the amphibians of Southeast Asia still holds an abundance of
unanswered questions and uncertainties. For the purpose of this book
we follow here the classification of INGER & TAN (1996) who for their
part followed DUBOIS (1987, 1992), DUELLMAN (1993), and DUELLMAN
& TRUEB (1986) with reservations (comp. INGER 1996).

Background: Sg. Silau-Silau © M. Maronde

 Systematic of Bornean
higher taxa of anurans

Suborder Archaeobatrachia
Superfamily Discoglossoidea
Family Bombinatoridae
Genus Barbourula
Superfamily Pelobatoidea
Family Megophryidae
Genera Leptobrachella
Leptobrachium
Leptolalax
Megophrys
©
U.
M
Suborder Neobatrachia
Bufonidae ant
hey Superfamily Bufonoidea
Family Bufonidae
Genera Ansonia
Bufo
Leptophryne
Pedostibes
Pelophryne
Superfamily Microhyloidea
Family Microhylidae
Genera Caluella
Chaperina
Gastrophrynoides
Kalophrynus
Kaloula
Metaphrynella
Microhyla
Superfamily Ranoidea
Family Ranidae
Genera Fejervarya
Hoplobatrachus
Huia
Ingerana
Limnonectes
Meristogenys
© R. Malkmus Occidozyga Microhylidae
Rana
Staurois
Family Rhacophoridae
Genera Nyctixalus
Philautus
Polypedates
©

Rhacophorus
H

Megophryidae
.S
au

Theloderma
er
Anura
Important traits for the identification
of frogs and their larvae
According to MAYR (1975) the members of a spe-
cies form a genetic and ecological unit and a re-
productive community. The phenotype, which
presents itself in the form of never absolutely iden-
tical individuals, is a summary of traits which re-
sults from both endogenous (genotype) and ex-
ogenous (biotic and abiotic influences of the en-
vironment) factors. The selective pressures exerted
by the latter may have favourable, eliminating, or
varying effects on the development of individual
traits. Such traits, or groups of traits, show a more
or less pronounced consistency specific to indi-
vidual taxa and are therefore of importance for
the classification of an animal.
• The species as a genetic unit: the analysis of
the genotype is the task of the laboratory bi-
ologist. As far as the frogs of Southeast Asia
are concerned, relevant work has just begun.
• The species as an ecological unit: it mani-
fests itself in many-faceted interactions with
the animated and unanimated environment,
in particular by behavioural patterns typical
for the species or genus, and by physiologi-
cal adaptations. It requires only a single
apomorphic feature to prevent populations
from producing fertile progeny. Such differ-
ences may, for example, be variations in the
aposematic behaviour, communication, ac-
tivity rhythms, social structure, choice of
ecological niche and calling sites, etc. As far
as Mt. Kinabalu is concerned, there are sta-
tionary bank-dwellers along the streams (e.g.
species of the genera Leptolalax, Ansonia,
Meristogenys, Huia, Staurois, Rana),
ground-dwellers of the forests which never
(e.g. Pelophryne, Kalophrynus), or only for
the purpose of reproduction visit streams
(e.g. Leptobrachium, Megophrys), or arbo-
real species which never (e.g. Metaphry-
nella, Nyctixalus, Philautus), or also only for
the purpose of reproduction visit streams
(e.g. Pedostibes, Polypedates, Rhaco-
phorus).
48
• The species as a reproductive community: be-
havioural patterns which can be linked to re-
production are to a large extent ritualized and
particular specific to a species . This serves to
ensure a reproductive separation among closely
related, sympatric species. Ritualized behav-
ioural parameters can be found in all interac-
tions related to reproduction. Oviposition and
volume and appearance of clutches may as well
provide information as to which species or ge-
nus we are dealing with. Diurnal, small frogs,
for example, which show a “flagging” behav-
iour with stretched-out hind limbs are certain
to belong to the genus Staurois; if we encoun-
ter a free-hanging foam nest or one which floats
on the water surface, we can be sure it were
Polypedates or Rhacophorus which deposited
them there. The advertisement calls often offer
the key for distinguishing species which can-
not be separated, or are difficult to differenti-
ate, by morphological traits (e.g. species of
Philautus or Leptobrachium montanum vs. Lep-
tobrachium gunungense).
As valuable as biochemical and ethological data
are for the identification of species, they usually
require the employment of costly instrumentation,
large amounts of time, or depend on the coinci-
dental concurrence of certain factors. The herpe-
tologist in the field will therefore make use of the
immediately available traits of the phenotype (ex-
ternal morphology) for the identification of a spe-
cies in most of the cases.
Many species of Bornean frogs can be identified
without doubt on the basis of morphological traits
only. It is only for highly variable and phenotypi-
cally very similar forms (e.g. in genera such as
Ansonia, Leptobrachium, Meristogenys, Micro-
hyla, Pelophryne, Philautus) that even the expe-
rienced systematist is faced with difficulties. As a
consequence the identification keys for these gen-
era are unsatisfactory.
It is important for the user of this book to know the
meaning of the terms used in the species accounts
and how the metric details were taken, and the same
applies to the definition of the call characteristics.
Only by this knowledge will it be possible to verify
and compare them with own data.

Morphology
Body shape and size of a frog usually suggests an
affiliation to the family, and often points to a cer-
tain genus already. In order to subsequently de-
termine the respective genus and species, a series
of traits are of great importance, such as the shapes
of the snout and pupils, the shape, relative size,
and position of the tympanum, and certain skin
structures (skin surface smooth, granular, rippled,
warty; size, shape, and arrangement of glandular
raises, skin ridges and appendages; meristically
highly variable groups of keratinized spikes on
the lower jaw, chest, and fingers).
Metric ratios and comparative data of two body
dimensions are often much more important than
indications of linear sizes (SVL of adult speci-
mens) as they are not only relevant for the identi-
fication of a species, but also for that of the re-
spective sex.
Here, we restrict ourselves largely to the follow-
ing indices:
• tibia length/snout-vent-length (TL/SVL);
• comparison head length – head width;
• distance of the anterior eye margin from nostril;
Fig. XVI Place of measurements in anurans.
Anura
• distance of the posterior eye margin to tympa-
num;
• horizontal diameter of eye – horizontal diame-
ter of tympanum;
• inter-ocular distance – inter-narial distance;
• lengths of fingers and toes; and the position of
the tibio-tarsal articulation when the hind leg is
plied forward to the body, etc.
The presence, respectively absence, of mandibu-
lar and vomerine teeth, presence or absence of
nuptial pads in males, and the morphology of the
tongue may also be of importance.
The arrangement and shape of the tubercles on the
lower side of the hands and feet are often of great
taxonomic relevance, and so are the shapes of the
finger and toe tips and the extent of the interdigital
webbings. To describe the latter we will use a for-
mula in accordance with GLAW & VENCES (1994):
“Fingers and toes are numbered. Each toe or finger
is followed by the numbers of the phalanges, which
are not webbed (in parentheses). Only the visible
phalanges, which are bordered by the subarticular
tubercles, are considered. ‘i’ means inner side, ‘e’
means outer (exterior) side of a finger or toe. So,
‘4i(1)’ means ‘at the inner side of the fourth toe
there is one unwebbed phalange’.”
Fig. XVII Anuran palate.
49
Anura

Fig. XIX Underside of the hand of a frog.

Fig. XVIII External morphological characters of frogs and toads.

Fig. XX A section through the tip of the digit of a treefrog. Fig. XXI Underside of the foot of a frog.

Webbing formula: Fully webbed; Webbing formula: 4i(2) 4e(1); Full formula: 1(0,5) 2i(1)2e(0.5)
4i(0) 4e(0) 4i(2): 2 phalanges of the inner side 3i(1)3e(1) 4i(2)4e(2) 5(0)
i=inner side e=exterior side of the fourth toe are unwebbed we use a simplified version:
of phalanges 4e(1): 1 phalange of the exterior side 1(0,5) 2(1/0.5) 3(1/1) 4(2/2) 5(0)
of the fourth toe is unwebbed
Fig. XXII Webbing formula (after GLAW & VENCES 1994) for describing the extension of webbing.

50

Fig. XXIII Shape of snout of Bornean frogs.
truncate, blunt
(e.g. species of
Ansonia, Pedostibes)
round
(e.g. species of Leptolalax,
Meristogenys, Occidozyga,
Rana)
sloping
(e.g. species of
Polypedates, Rana)
protruding
(e.g. species of Ansonia,
Bufo, Kalophrynus)
angular
(e.g. species of
Leptobrachium, Rhacophorus)
In some cases tip of snout with a dermal projec-
tion, e.g. triangular projection (Megophrys nasuta),
conical tip (females of Philautus petersi).
Next to the habitus, it is often the colouration and
the colour pattern which give a species its typical
appearance (e.g. in Leptolalax pictus, Pedostibes
everetti, Chaperina fusca, Rana signata, or Nyc-
tixalus pictus). The pigmentation of the iris is un-
mistakable in some forms: Leptobrachium monta-
num has, for example, a white, and Leptobrachium
abbotti a bluish eye background, Rhacophorus
angulirostris possesses a blue upper margin of the
iris, Leptolalax arayai shows a coloured segment
covering ca. 120° of the iris, while a network of
blood vessels covers the eyes of Leptolalax pictus
and Philautus bunitus. Some species exhibit a
marked sexual dichromatism (e.g. Philautus
amoenus, Rhacophorus angulirostris, Rhacophorus
everetti).
Referring anatomic research of frogs, details of the
skull, the shoulder (pectoral) girdle (arciferous or
Anura
Fig. XXIV Pectoral girdles
(cartilaginous areas are stippled).
A firmisternal pectoral girdle (epicoracoid carti-
lages fused in the midline) have species of the fami-
lies Microhylidae, Ranidae and Rhacophoridae.
An arciferal pectoral girdle have species of the
families Bufonidae and Megophryidae.
firmisternal), and details of the vertebrae are of
great importance for determining a specimen’s sys-
tematic position.
The larvae of anurans (tadpoles) can also be iden-
tified by species-specific external criteria. Major
morphological traits include the position of the
spiracle, the anal tube, the eyes, nostrils, the shape
and extent of the caudal fin, morphometric rela-
tions, colourations and colour patterns. However,
the most important group of character for identifi-
cation of tadpoles are those associated with the oral
field (shape and extent of marginal oral papillae,
the number of rows of labial teeth, and that of the
keratinized beak). The rows of labial teeth are again
described in a formula which was originally pro-
posed by ALTIG (1970). It must be pointed out,
though, that the proportions of the body of a tad-
pole may vary with both the individual and age as
51
Anura

Fig. XXV Embryonic and larval development of a frog (substantially simplified after GOSNER 1969).

Fig. XXVI Lateral view of a tadpole.

Fig. XXVII Structures of the oral disc of a tadpole.

5(3-5) / 3 5(2-5) / 3(1-2)

Teeth row formula after ALTIG 1970.

52
 Anura

Habitats of tadpoles

The type of a water body and its altitudinal loca-

tion can also assist in the attempt to determine tad-
poles as the larvae of the individual anuran species
occupy in general very specific niches. The origin
MALKMUS of a tadpole is therefore valuable information for
Fig. XXVIII Tadpole of Leptobrachium gunungense the taxonomist. INGER (1985) distinguished 9 of
from Sg. Tibabar (1850 m), showing system of lateral- those microhabitats (“m” in species accounts) for
line organs. running waters and 7 for stagnant ones.

can the structures of the oral field (e.g. in Lepto-
brachium, Meristogenys). If visible, the arrange-
ment of the lateral organ, and the position of glands,
may be of importance for the identification of a
species.
The significance of larval morphology for the sys-
tematics can be demonstrated by a look at the Amo-
lops-group where the genus Amolops was split up
into the genera Amolops, Huia, and Meristogenys
on the basis of morphological differences in their
tadpoles (YANG 1991) and larval characters were
crucial for determining the evolutionary relation-
ships among the taxa.
1. Streams
a. Torrent: surface of water broken, occasionally
foaming, bed of rocks (diameter > 5 cm) and
boulders, current strong (e.g. Leptobrachella,
Meristogenys)
b. Riffle: surface of water in shallow waves but
never foaming, bed of gravel, current moderate
(e.g. Leptolalax, Ansonia, Rhacophorus cyano-
punctatus)
c. Shingle area: surface of water smooth, bed of
flattened rocks (3-10 cm), current weak to mod-
erate (e.g. Rana chalconota, Megophrys nasuta)
d. Leaf drift: areas with deep accumulations of dead
leaves, drifts up to 1.5 m deep and 10 m long,

Fig. XXIX Microhabitats in a mountain brook (in part. after MALKMUS 1999).

typical microhabitat for larvae with ventral sucker disk like e.g.
Meristogenys

rocks, exposed to the current

cascade

interstices of gravel interstices of gravel

barrier of leaf drift, caused by leaf drift
rocks or branches and trunks
fallen into the stream

typical microhabitat for larvae with cup-like expanded lips like of Leptolalax
and funnel-shaped lips like of Megophrys

53
Anura
current weak (e.g. Leptobrachium montanum,
Rana signata, Rhacophorus gauni)
e. Open pool: ponded parts of streams, bottom silt
to sand and gravel (e.g. Leptobrachium monta-
num, Megophrys nasuta)
f. Side pool: an embayment off the main channel,
partially or completely cut off from the current,
bottom fine to coarse sand, usually covered by
dead leaves, current weak or absent (e.g. Rana
signata, Leptobrachium montanum, Bufo diver-
gens)
g. Pothole on rock bank: water-filled depression on
rocky bank, bottom bed rock, often with dead
leaves (e.g. Rana signata, Rana chalconota)
h. Seepage area: trickle of water less than 2 cm
deep, usually emerging from a steep clay bank
(e.g. Limnonectes kuhli)
i. Pool of intermittent stream: up to 0.5 m in maxi-
mum surface dimension, depth up to 5 cm, bot-
tom fine silt, sand, and pea-gravel, current only
after moderate to heavy rain (e.g. Limnonectes
kuhli, Rana chalconota)
2. Microhabitats away from streams
a. Seepage area: film of water less than 1 cm deep
emerging from a slope of clay and small rocks,
current negligible (e.g. Limnonectes kuhli).
b. Rain pool: rain-filled depression of variable size
and depth, bottom usually clay, often with dead
leaves (e.g. Kalophrynus pleurostigma, Rana
finchi, Polypedates otilophus, Microhyla).
c. Animal wallow: rain-pool modified by the ac-
tion of mammals, bottom fine silt, length more
than 1 m, depth up to 15 cm (e.g. like in 2b)
d. Log cavity: mainly open on upper surface or side
of a log, surface up to 15 × 20 cm, depth up to
10 cm (e.g. Chaperina fusca, Nyctixalus pictus,
Microhyla borneensis).
e. Hole in living surface root: small holes up to
18 cm in diameter and 10 cm in depth, height
above ground 5-16 cm (e.g. Nyctixalus pictus).
f. Buttress tank: cavity formed by anastomosis of
two or more buttresses, surface up to 30 × 40 cm,
depth 30 cm, height above ground up to 4 m.
g. Hole in trunk or large branch: surface area up to
20 × 30 cm, depth 30 cm, height above ground
>0.5 m (e.g. Chaperina fusca, Metaphrynella
sundana, Nyctixalus pictus).
54
h. Water-filled phytothelmous cavities in bamboo
internodes, leaf axils (Pandanus), natural cups
in litter (fruit husks, palmtree spathe, fallen
leaves) and dead Nepenthes pitchers: very re-
duced water content (e.g. Leptophryne borbo-
nica, Metaphrynella sundana, Philautus saueri)
Larval feeding types
As adaptations to their individual habitats and
partly in order to be able to make use of certain
food resources, anuran larvae have often devel-
oped substantial modifications to their body struc-
tures (e.g. shape of the body, development of cau-
dal musculature and fins, position and structures
of the mouth area). In principle two categories are
distinguished, i.e. endotrophic and exotrophic lar-
vae. Endotrophic larvae gain nutrition for their
development from parental sources (yolk): these
froglets may either hatch from egg capsules (Phi-
lautus) or are free-swimming, nidicolous, non-
feeding tadpoles (Kalophrynus, Pelophryne). All
the other species of Bornean genera have tadpoles
of the exotrophic type, i.e. they are free-swim-
ming, feeding larvae.
INGER (1986) distinguished 5 larval feeding types:
1) Bottom suspension feeders: in microhabitats away
from streams and in stream areas with weak cur-
rents where fine suspension and leaf drift cover
the bottom. Tadpoles of this type (generalized
pond-type) show no morphological specialization
(e.g. Bufo [fig. 39], Rana, Polypedates [fig. 40]).
2) Mid-water suspension feeders: in microhabitats
away from streams (rain pools, pig wallows,
potholes); larvae with large branchial baskets
with dense ruffles, lacking beaks and denticles
(e.g. Chaperina [fig. 41], Microhyla [fig. 42]).
3) Obligate benthic feeders: in streams with mod-
erate to strong currents; tadpoles feed on the
epilithic film on rocks to which they cling. They
have cup-like oral disks (e.g. Rhacophorus),
sucker-type oral disks (e.g. Ansonia [fig. 43]),
or abdominal suckers (e.g. Huia [fig. 44], Meris-
togenys); larvae are rheophilous, with elongated,
depressed bodies, long muscular tails, low fins,
and small, non-bulging eyes.
 Anura

4) Macrophagous feeders: larvae inhabit a wide larval feeding type 3

range of microhabitats in streams; they are com-
mon in leaf drifts and in interstices of gravel on
the stream bottom (e.g. Leptobrachella, Lepto-
brachium [fig. 45], Leptolalax [fig. 46]); larvae
with heavy, sharp beaks.
5) Surface film feeders: larvae dwell in the entire

© W. Denzer
range of stream microhabitats from riffles to pot-
holes; they suspend particles of floating material
from the surface film by means of their upturned
funnel mouths (e.g. Megophrys [fig. 47]).
Larval feeding type 3
larval feeding type 3
Larval feeding type 1
© U. Manthey

© U. Manthey
Larval feeding type 1 Larval feeding type 4
© U. Manthey

© U. Manthey
Larval feeding type 2 Larval feeding type 4

© M. Schroth
© U. Manthey

Larval feeding type 5

Larval feeding type 2
© U. Manthey

© H. Sauer

From top to base: Tadpoles of the genera From top to base: Tadpoles of the genera
Fig. 39 Bufo Fig. 40 Polypedates Fig. 41 Chaperina Fig. 43 Ansonia Fig. 44 Huia Fig. 45 Leptobrachium
Fig. 42 Microhyla. Fig. 46 Leptolalax Fig. 47 Megophrys.

55
Anura

All these types of larvae are represented on Mt.

Kinabalu although the bottom suspension feeders,
which are typical elements of lowland waters, fea-
ture much less prominent than the rheophilous
benthic feeders. Due to geomorphological and hy-
drological reasons, the number of water bodies
which would allow a successful development of
amphibian larvae is very limited above 2000 m on
Mt. Kinabalu; here, endotrophic forms dominate
(e.g. Philautus species).
The realization of an identification key to the lar-
vae is impossible at present since only 50-60% of

the amphibians of Borneo are known by their tad-
poles (comp. I NGER 1985, 1992a, b, I NGER &
WASSERSUG 1990). We must therefore be content
with a key to the families here.
Bioacoustics
On reaching maturity the males of most anuran
species are able to vocalize. Most species develop
- partly in both sexes - several types of vocalizations
(advertisement, release, distress calls, etc.). Among
these, the advertisement calls of the males are of
exceptional importance with regard to biological
and systematic sciences.
© P. Hoffmann
Fig. 48 A calling male of Leptolalax pictus from Sg.
Wariu, Sayap, 1000 m.
Listening to nightly concerts of frog voices is cer-
tainly one of the most memorable experiences dur-
ing a visit to a rainforest. The volume of these
concerts is again and again surprising. It is mainly
due to the resonant properties of the vocal sacs
(sac-like extrusions of the bottom of the oral cav-
ity) which vary in size and position, but are gen-

Fig. XXX Circadian calling activity of some anurans on Mt. Kinabalu (after MALKMUS 1996b).

maximum calling activity limited calling activity

56
 Anura

Which part of an acoustic signal must be considered

the “call” is often a question of the definition chosen.
In a series of signals largely made up of about equal
pulses, each signal considered a note and the entire sig-
nal series a call. (e.g. Philautus aurantium).

• note means a pulse group (n)

Fig. XXXI Physical call characteristics.
• note duration means the time from the beginning of
the first to the end of the last pulse in a note
• internote interval means the time from the end of
one note to the beginning of the next note (i)
• note repetition rate means the number of notes per
second
• p: pulse duration (ms)
• ip: interpulse interval (ms)
• pulse rate means the number of pulses per second

erally positioned in the throat region. The sounds
are actually produced by the channelling of air
back and forth between the lungs and the oral cav-
ity passing past vocal chords which oscillate and
produce sounds which are then amplified by the
vocal sac.
The biological significance of the advertisement
call lies in the identification of the own species,
the localization of the caller, and the stimulation
of the sexual partner, thus serving a reproductive
isolation mechanism. In conjunction with aggres-
sion calls the advertisement call simultaneously
functions as an acoustic marker for an occupied
territory and the maintenance of the same, thus
regulating population densities. In order to prevent
confusion closely related species which occur

Fig. XXXII Graphic illustration of acoustic signals using Philautus mjobergi as an example.

1 Sonagram (graphic illustration of an acoustic signal illustrating its course of time, its composition of frequen-
cies, and the relative intensity of the individual frequency components. The horizontal axis indicates time, the
vertical one frequencies).
2 Oscillogram (serves to interpret structural traits recognizable in a sonagram).
3 Enlarged section of an oscillogram showing a high resolution of structural elements in order to perform de-
tailed analysis of the call/note structure.

57
Anura
sympatrically have developed more distinct call
structures than those with an allopatric distribution
(e.g. in species of the genera Leptobrachium and
Philautus). But it is not always the male which ac-
tively attempts to attract a sexual partner. For ex-
ample, female Polypedates leucomystax ready to
mate advertise by producing single croaking sounds
and a rhythmical drumming with their toes (seis-
mic communication) for minutes on end.
On Mt. Kinabalu we are met with spatially firmly
established calling colonies (e.g. of Megophrys
baluensis, Metaphrynella sundana, Staurois tuberi-
linguis, Ansonia longidigitata, Philautus auran-
tium), those with calling densities which are un-
equally distributed over larger areas (e.g. Leptobra-
chium montanum, Kalophrynus baluensis, Philau-
tus bunitus and P. pictus), and species which pro-
duce singular calls at large intervals (e.g. Meristo-
genys orphocnemis, Rhacophorus angulirostris).
There are di- and nocturnal callers.
Some voices give evidence of an adaptation to the
acoustic background of their individual habitats. The
dwellers of mountain streams for example produce
sharp clicking, whip-cracking, or chirping signals
in order to rise above the sound level of their biotope
(rushing of water) and facilitate communication.
The fact that advertisement calls are species-spe-
cific is of immense value for the field herpetologist
since especially morphologically very similar spe-
cies are typically distinctly differentiated with re-
gard to their bioacoustics. It also helps to gain an
overview of the local distribution patterns of the
individual species.
Until about 40 years ago, the description of adver-
tisement calls was limited to onomatopoetic descrip-
tions such as a short, whip-cracking “cheep”-sound
for Meristogenys or a burring “errrrr...” for Philautus
petersi, or by using musical notes. Obviously these
methods of illustration are very subjective and little
informative. In order to record the physical call pa-
rameter (call length, frequency structure, intensity,
repetition rate of the impulses, length of intervals,
etc.) objectively, the calls are recorded on tape and,
with the aid of computers, displayed in the form of
sonagrams and oscillograms thus rendering them fit
for comparisons. Sonagrams provide a basis to
analyze sound levels, relative volumes, and the tim-
ing of acoustic signals.
58
When comparing analyses of voices it must be
taken into consideration that factors such as hu-
midity, temperature, light, and stimulation by the
given acoustic environment, but also the volume
produced by the individual, may have substantial
effects on certain call parameters (particularly vol-
ume, frequency, speed of the pulse rate).
To facilitate acoustic communication within the
acoustically heavily polluted rain forest the indi-
vidual species choose certain frequency bands. Just
like radio stations use various carrier frequencies,
frogs occupy various acoustic “channels”, in other
words acoustic niches within the overall back-
ground noise of their habitat.
Reproductive strategies
In a worldwide survey, DUELLMAN & TRUEB (1986)
distinguished altogether 29 different reproductive
modes which differ in the choice of oviposition sites
and type of development of the larvae. No less than
10 of these modes have been recorded for frogs on
Mt. Kinabalu:
I. Eggs aquatic
A. Eggs deposited in water
01. Eggs and feeding tadpoles in lentic water (e.g.
Chaperina, Kaloula, Occidozyga, Rana, Bufo)
02. Eggs and feeding tadpoles in lotic water
(e.g. Leptobrachella, Leptolalax, Ansonia,
Meristogenys, Huia, Staurois, Rana)
03. Eggs and feeding tadpoles in water in tree
holes or aerial plants (e.g. Metaphrynella)
04. Eggs and non-feeding tadpoles in water-
filled depressions (e.g. Kalophrynus)
05. Eggs and non-feeding tadpoles in water in
tree holes or aerial plants (e.g. Kalophry-
nus, Pelophryne, Philautus)
B. Eggs in foam nest
06. Foam nest on pond; feeding tadpoles in
pond (e.g. Polypedates, Rhacophorus)
II. Eggs terrestrial or arboreal
C. Eggs on ground or in burrows
07. Eggs hatch into feeding tadpoles that are
carried to water by an adult (e.g. Limno-
nectes finchi)
08. Eggs hatch into froglets (e.g. Philautus)

© P. Hoffmann
© R. Malkmus
D. Eggs arboreal
09. Eggs hatch into tadpoles that drop into wa-
ter-filled cavities in trees (e.g. Nyctixalus
pictus)
10. Eggs hatch into froglets (e.g. Philautus)
E. Eggs in foam nest
11. Nest arboreal; hatching tadpoles drop into
ponds or streams (e.g. Polypedates, Rhaco-
phorus); foam nests were found not over-
hanging water, but the tadpoles would hatch
during rain showers and then be washed to
a water body.
Anura
© S. Manthey
© M. Matsui
Above from top to base
Fig. 49 A typical foam nest of Polypedates leucomystax.
Fig. 50 Eggs of a Philautus species in a depression of a
dry fern.
Left from top to base: Fig. 51 Tadpoles of Chaperina
fusca in a bambusinternodium.
Fig. 52 In waterfilled tree holes live tadpoles of some
species (e.g. Metaphrynella).
Like the choice of oviposition, the details of the
individual clutches (e.g. arrangement, number,
size, and pigmentation of the eggs) are indicative
of genera and species. Unfortunately, only a rela-
tively small percentage of species are known by
their clutches. Species dwelling the upper montane
zone tend to produce smaller clutches with larger
eggs (e.g. Leptolalax, Pelophryne, Philautus).
Whether all or at least some of the anurans at the
various altitudinal levels are reproductively active
throughout the year, is still unknown. As far as
Borneo is concerned, respective information is only
available for lowland rain forests (I NGER &
GREENBERG 1963, INGER & BACON 1968). They show
a circannual reproduction for individual species.
59
Anura

Key to the families of Bornean frogs

(in parts after MANTHEY & GROSSMANN 1997)

A
B

1 Teeth present in upper jaw (A) ......................... 2

• Teeth absent in upper jaw ................................. 7

2 An intercalary cartilage between the 2 terminal

phalanges (B); 1st finger opposed to remaining D
fingers ....................................... Rhacophoridae C
• No intercalary cartilage; first finger not opposed
to remaining fingers ........................................... 3

3 Pupil vertical (C) ........................ Megophryidae

• Pupil horizontal (D) or round ........................... 4

4 Vomerine teeth present (E) ............................... 5

• Vomerine teeth absent .... Ranidae (Occidozyga) E
F
5 Inner metatarsal tubercle shovel-like (F) ............
.................................. Microhylidae (Calluella)
• Inner metatarsal tubercle not shovel-like or ab-
sent .................................................................... 6

6 Fingers fully webbed (G) ......... Bombinatoridae

• Fingers without webbing or rudimentary webbing
........................... Ranidae (except Occidozyga)

7 Skin dry, warty; parotoids (H) usually present ... G

........................................................... Bufonidae H
• Skin moist, smooth or rough, not warty; parotoids
absent ............ Microhylidae (except Calluella)

60
 Anura

Preliminary key to the larvae of Bornean frog families

(in parts after MANTHEY & GROSSMANN 1997)

A B
1 Free-swimming larvae ....................................... 2
• Direct development within the egg .....................
.............................. Rhacophoridae (Philautus)
C
2 Spiracle lateral (A) or ventral (B) ..................... 3
• Spiracle not visible, only 1 row of teeth (C) .......
.................................... Bufonidae (Pelophryne)
3 Spiracle lateral (A) ............................................ 4
• Spiracle ventral (B), no keratinized jaws, tooth- D
less ............................................... Microhylidae F
4 Oral disc directed ventral (D) ........................... 6 E
• Oral disc directed terminal (E), without rows of
teeth ................................................................... 5
5 Lips funnel-shaped (F) ......................................... H
.............................. Megophryidae (Megophrys)
• Lower lip horseshoe-shaped (G) ......... Ranidae G
(Occidozyga)
6 Marginal papillae of the mouth surround the entire
oral disc (H) .......... Megophryidae (with teeth:
Leptobrachium; without teeth: Leptobrachella;
with or without teeth: Leptolalax)
I K
• Upper lip without or only with lateral marginal
papillae (I) ......................................................... 7
7 Anal tube medial (K) ........................................ 8
• Anal tube lateral ..... Ranidae (Hoplobatrachus,
Limnonectes), Rhacophoridae (Nyctixalus,
Rhacophorus, Theloderma) M
L
8 Abdominal sucker (L), more than 2 rows of teeth
on the upper lip ....................................................
......................... Ranidae (Huia, Meristogenys)
• With or without oral sucker (M), 2 rows of teeth
on the upper lip ................................................. 9
9 3 rows of teeth on the lower lip ......... Bufonidae
(keratinized upper jaw absent or divided (N): N O
Ansonia; keratinized upper jaw continuous (O):
Bufo, Leptophryne, Pedostibes)
• At least 6 rows of teeth on the lower lip .............
............................................ Ranidae (Staurois)

61
Bufonidae

Bufonidae
Bufonids are cosmopolitan in temperate and tropical regions (except for Madagascar and the Australo-
Papuan region).
Oriental bufonids have an arciferous pectoral girdle, the sacrum has dilated diapophyses, an omosternum is
absent, a bony sternum present. There are 5-8 holochordal, procoelous presacral vertebrae. The maxillae
and premaxillae are edentate. Most species have dry, thick, glandular skins, often with pustular warts. The
pupil is horizontal. Mating takes place in an axillary amplexus.
Most bufonids produce egg clusters in string form (eggs normally pigmented) and deposit them in water.
Their tadpoles are free-swimming, aquatic, have beaks and denticles; the spiracle is sinistral, the anal tube
medial (except in Pelophryne). Some genera have torrent-adapted, oro-adhesive tadpoles with large, ex-
panded oral disks and suckers (Ansonia). The species of Pelophryne have non-feeding larvae which survive
entirely on yolk.
Bornean bufonids occur in various habitats, some are terrestrial (Bufo), others arboreal (Pedostibes) or both,
living partly on the ground and partly on low vegetation (Ansonia, Pelophryne); Pseudobufo is aquatic.
Nearly all species are distributed in primary and secondary forests, only a few appear in areas modified by
man. On Mt. Kinabalu, bufonids can be found over a wide altitudinal range. Of the three frog species
known to exceed an elevation of 3000 m a.s.l., two are bufonids (Ansonia fuliginea, Pelophryne misera).
The family is represented on Borneo by 6 genera and 29 species, of which 5 genera and 12 species occur on
Mt. Kinabalu.

Key to the Bornean genera of Bufonidae (after INGER 1966, MANTHEY & GROSSMANN 1997)
1 Tympanum visible (A) ...................................... 2 A
B
• Tympanum not visible (in part) ....... Pedostibes
2 Parotoids present (B) ........................................ 4
• No parotoid glands ............................................ 3
C
3 Toes fully webbed (C), body and limbs robust;
adults > 70 mm ............................... Pseudobufo
• Toes not fully webbed, body and limbs slender; D E
adults < 65 mm ..................................... Ansonia
4 At most one phalange of first finger projecting from
fleshy web (D); adults < 40 mm ....... Pelophryne
• two phalanges of first finger projecting free of
palm or web, hands without fleshy web .......... 5
5 Finger tips rounded, not dilated ....................... 6
• Finger tips dilated into truncate disks (E) ...........
............................................ (in part) Pedostibes
6 Underside of foot with a large elongated tubercle
near bases of each toe; body and limbs slender; length
of femur and tibia > SVL (F) .......... Leptophryne
• No such tubercles; body and limbs robust; length F G
of femur and tibia < SVL (G) .................... Bufo

62

Genus Ansonia STOLICZKA, 1870
Members of the genus Ansonia are known from
southern India, Thailand, the Malay Peninsula, Bor-
neo, and Mindanao (Philippines).
Bornean species are small to medium-sized, brown
to black toads (SVL < 65 mm), with a dorsolaterally
weakly flattened, elongate body with slender, long
legs. Tympanum distinct (except in A. albomacu-
lata); in most species rostrum projects over the
mouth; nostrils above symphysis, large eyes (di-
ameter ³ snout length), pupil horizontal. Tips of
fingers and toes mostly rounded, some species with
truncate disks. Finger lengths 3>4>2>1, toe lengths
4>5>3>2>1; fingers webbed at bases, webbing of
toes varies with the species. Skin rough, usually
warty; parotoid glands absent. Adult males with
subgular vocal sac (except in A. guibei), nuptial
pads and mandibular spines present.
Ova rather large and unpigmented. Tadpoles small
(usually < 20 mm) and adapted to a benthic life in
flowing water with a streamlined body (except in
A. leptopus), and a cup-like, sucker-type oral disk;
two upper and three lower rows of labial teeth,
upper beak divided, lower beak divided only in a
few species (e.g. A. guibei). One species (Ansonia
sp.) with an abdominal sucker. Body coloration
usually black.
These toads live in forest streams with strong cur-
rents (except A. leptopus). Bornean species inhabit
primary and old secondary forests in hilly to
montane areas. They can be found on the forest
Tab. 1 Distribution of Ansonia-species on Borneo.
Bufonidae
floor and on leaves in the lower stratum of veg-
etation. On Mt. Kinabalu, this genus has a dis-
tinct altitudinal stratification with A. fuliginea reach-
ing the maximum vertical distribution (3480 m)
of all amphibians of the Sunda Islands. Some spe-
cies are local endemics, restricted to very limited
areas (A. fuliginea, A. guibei).
Eleven species are known from Borneo, six of
them occur on Mt. Kinabalu.
Remarks: There is some doubt about the occur-
rence of A. albomaculata and A. leptopus on Mt.
Kinabalu. INGER (1985) described larval Ansonia
from Sg. Kipungit which he originally believed to
be A. albomaculata. In a later paper (INGER 1992b),
however, he distanced himself from this view by
leaving the specific identity of these larvae open
and referring to them as Ansonia “cruciform” due
to their dorsal pattern. On the other hand, A. albo-
maculata is listed again to occur on Sg. Kipungit
in the check list of the amphibians of Mt. Kinabalu
(INGER, STUEBING & TAN 1996), while the works by
INGER & TAN (1996a) and INGER & STUEBING (1997)
do not mention it for Sabah, but only for Brunei,
Sarawak, and Kalimantan.
Another larva, which could as yet not be assigned
to a species, was described by INGER (1992b) from
East Mesilau (1460 m) and from Sarawak. Like A.
guibei, it has both jaws divided and is therefore
referred to as Ansonia “beaks divided”. In a new
checklist of INGER et al. (2000) A. albomaculata,
A. leptopus and even A. minuta are mentioned for
Kinabalu Park.
63
Bufonidae
Preliminary key to Bornean species of Ansonia
1 3rd and 5th toe webbed to tips; skin flap on vent
present ..................................................... guibei
• 3rd and 5th toe not webbed to tips; no skin flap on
vent ................................................................... 2
2 1st finger does not reach disk of 2nd when fingers
adpressed .......................................................... 3
• 1st finger reaches disk of 2nd when fingers adpres-
sed .................................................................... 9
3 Sharp tarsal ridge present ................................ 4
• No tarsal ridge ................................................. 5
4 A white spot below eye; a whitish band from eye
to arm .......................................... albomaculata
• Without such pattern ............................. minuta
5 Tips of outer fingers dilated into spatulate disks
........................................................................... 6
• Tips of outer fingers rounded .......................... 8
Ansonia fuliginea (MOCQUARD, 1890)
Bufo fuligineus MOCQUARD, 1890: 158 – Type locality: North
Borneo
Nectophryne altitudinis SMITH, 1931: 14, pl. 1: fig. 1 – Type
locality: “Pakka, alt. 10,200 feet”, Mount Kina Balu, Sabah
(syn. after INGER 1966: 102; INGER & TAN 1996a: 554; PEND-
LEBURY & CHASEN 1932: 21
Pedostibes altitudinis, BARBOUR 1938: 192
Ansonia altitudinis, INGER & STUEBING 1992: 41-51
Ansonia fuliginea, INGER 1966: 102, 1978: 312; MATSUI 1979: 314;
MANTHEY 1983: 21; MATSUI, HIKIDA & NAMBU 1985: 152; MALK-
MUS 1988c: 176; INGER & DRING 1988: 463; INGER & STUEBING
1992: 41-51; MALKMUS 1992a: 106; INGER, STUEBING & TAN
1996: 362; INGER & TAN 1996a: 554; INGER et al. 2000: 16
Diagnosis
A medium-sized, moderately stocky species, with
a vertically truncate snout-tip in profile; tip of 1st
finger not reaching base of swollen tip of 2nd; nup-
tial pad composed of a few keratinized spines; re-
stricted to the high mountain region.
Description
Males up to 36 mm, females up to 44 mm; TL/SVL
0.38-0.41; head as wide as long; canthus rostralis
64
6 Disk of 3rd finger subequal to diameter of tympa-
num ....................................................... latidisca
• Disk of 3rd finger much narrower than tympanum
........................................................................... 7
7 Distance tympanum - mouth half or less than half
the distance nostril - mouth; flattened habitus ..
............................................................ platysoma
• Distance tympanum - mouth more than half dis-
tance nostril - mouth .......................... hanitschi
8 Snout projecting, oblique in profile .... torrentis
• Snout vertical in profile ..................... fuliginea
9 Dorsal warts low, rounded .................... leptopus
• Dorsal warts elevated, spinose ...................... 10
10 No interscapular spot ...................... longidigita
• An oval interscapular spot present; tips of fin -
gers dilated ........................................ spinulifer
sharp; lores oblique, a little concave; tympanum half
diameter of eye; tips of fingers and toes slightly
swollen; fingers webbed at their bases; webbing of
toes 1(0.5), 2(1/0.5), 3(2/1.5), 4(3/3), 5(2); a weak
outer metatarsal tubercle; mandibular spines black,
arranged in a double row; skin dorsally and laterally
with low, rounded warts, sometimes with melanic
spinules; lower sides of body granular.
Brown above, eyelids and snout darker; flanks and
lower sides black; legs light brown below; abdomen
with yellow or whitish spots; iris blackish with tiny
yellow spots.
Ecology/Ethology
Nearly nothing is known on the life history of this
toad. It is restricted to the high mountain region
(> 1500 m a.s.l.), to the upper and subalpine forest
zone and beyond the forest line. We observed a speci-
men active at night on the forest floor in the rocky
region with very dense vegetation near Pakka Cave
(3050 m). At the same site PENDLEBURY & CHASEN
(1932) had found a toad, “the stomach of this speci-
men contained fragments of beetles and small
crustacea”.

Fig. 53 Ansonia fuliginea from Pakka, 3050 m.
Distribution
Ansonia fuliginea is only known from Mt. Kina-
balu: Marei Parei; Kamborangoh; Layang Layang;
Pakka; Panar Laban (1500-3480 m); Sg. Mesilau
(East); Mentaki Ridge (1800-2000 m).
Ansonia guibei INGER, 1966
Ansonia guibei INGER, 1966: 104 – Type locality: Mesilau Cave
camp, 1800 m, Mount Kina Balu, Sabah
Ansonia guibei, INGER 1978: 312; MANTHEY 1983: 21; INGER &
DRING 1988: 463; MALKMUS 1988c: 176; INGER & STUEBING 1992:
41-51; MALKMUS 1994b: 221, 1994c: 87; INGER, STUEBING &
TAN 1996: 362; INGER & TAN 1996a: 554; LAKIM et al. 1999: 33;
MALKMUS & KOSUCH 2000: 121-124 ; INGER et al. 2000: 16
Diagnosis
A medium-sized species of Ansonia with long
limbs; snout vertical in profile; first finger reach-
ing base of tip of 2nd finger; toe webbing reaching
Bufonidae
© R. Malkmus
tips of 1st, 2nd, 3rd, and 5th toe; flaps of skin flanking
vent; no vocal sac. Restricted to high montane re-
gions.
Description
Males up to 32 mm, females up to 34 mm; TL/SVL
0.47-0.56; head as wide as long; canthus rostralis
sharp; lores weakly concave; tympanum one third
of eye diameter; tips of fingers and toes swollen;
fingers free of webs; a large, round palmar tuber-
cle; toe webbing 1(0), 2(0/0), 3(0/0), 4(2/2), 5(0);
small inner and outer metatarsal tubercles; skin
above with many round warts, sometimes with dark
spinules; flanks and ventrum granular; mandibular
spines black, arranged in one row.
Dorsally and laterally dark brown, sometimes with
lighter regions; yellowish brown crossbars on hind
limbs; knees and upper arms lightened; belly black-
ish with whitish granules; chest and throat greyish
65
 Bufonidae
© R. Malkmus

Fig. 54 A pair of Ansonia guibei from Sg. Mesilau East, 1850 m.

rosy; toe webbing greyish brown; nuptial pads
black; iris golden brown with fine black network.
Tadpoles with very broadly rounded snout; oral disk
wider than widest part of body; upper and lower
beaks divided; labial teeth dense, formula 2/3; tail
tapering to a rounded tip in the last third; tail length
64% of total length (23.8 mm at st 29); head and
Fig. XXXIII Lateral and dorsal view of the tadpole of
Ansonia guibei.
body uniformly black dorsally, circumlaterally
lighter; caudal muscle dark brown to black; fins
transparent with dark margins.
Ecology/Ethology
Ansonia guibei is restricted to the high montane pri-
mary forest of the Mesilau valley. Here it lives on
boulders protruding from the water and areas near
the banks of the rapidly flowing mountain stream. It
is nocturnal. When threatened it may also leap into

Fig. 55 Tadpole of Ansonia guibei.

© H. Sauer

1 mm
R. Malkmus

66

very rapidly flowing water, even into cascades, dive,
and hide in rock cavities below the water surface or
near the bank.
In mid-August we observed a female which car-
ried a male, attached to it in axillar amplexus, to
the stream. The male produced frequent series of
impulses with high-pitched, metallic clicking and
cracking sounds.
Large numbers of larval A. guibei were found in
the Sg. Mesilau (1800-1950 m) in the month of
March where they preferably resided on large boul-
ders in moderate to strong currents (m: 1a, b, c).
They were at about the same level of development
(st 25-30) indicating that reproduction follows an
annual rhythm.
The prey spectrum of A. guibei is wide with a large
variety of arthropods being accepted: cicadas,
pentastomids, small grasshoppers, plantlice, aphids,
beetles, small butterflies, spiders, mites, hairless
caterpillars, and beetle larvae.
Distribution
The species is known only from the eastern and
western Sg. Mesilau, between 1800 and 1950 m
a.s.l., on Mt. Kinabalu.
Fig. 56 Ansonia hanitschi from Sg. Silau-Silau.
Bufonidae
Ansonia hanitschi INGER, 1960
Ansonia hanitschi INGER, 1960b: 484 – Type locality: Mount
Kina Balu, Sabah
Bufo (Ansonia) penangensis, (not Ansonia penangensis
STOLICZKA, 1870) MOCQUARD 1890: 160
Bufo penangensis, (not Ansonia penangensis STOLICZKA, 1870)
HANITSCH 1900a: 74; VAN KAMPEN 1923: 75
Bufo leptopus, (not Bufo leptopus GÜNTHER, 1872) SMITH 1931:
13 (part); INGER 1956: 395 (part)
Ansonia minuta, (not Ansonia minuta INGER, 1960) MANTHEY
& DENZER 1982: 12; MANTHEY 1983: 21
Ansonia albomaculata, (not Ansonia albomaculata I NGER,
1960) MALKMUS 1991a: 28
Ansonia cf. albomaculata, MALKMUS 1992a: 108, 1994b: 222
Ansonia hanitschi, INGER 1966: 106, 1978: 312; MATSUI 1979:
314; MANTHEY 1983: 21; MALKMUS 1985: 7, 1987: 278, 1988c:
176; INGER & DRING 1988: 463; MALKMUS 1989: 184, 1991:
28, 1992a: 106; INGER & STUEBING 1992: 41-51; WONG 1994:
25-37; MALKMUS 1994b: 221, 1994c: 89; HOFFMANN 1995a: 9;
INGER, STUEBING & TAN 1996: 362; INGER & TAN 1996a: 554;
INGER & TAN 1996b: 73; MALKMUS 1996b: 20-26, 1996c: 281;
MANTHEY & GROSSMANN 1997: 28; HOFFMANN 1998b: 85; LAKIM
et al. 1999: 33; INGER et al. 2000: 16
Diagnosis
A medium-sized, moderately slender highland spe-
cies with long legs; snout oblique, sloping in pro-
file; 1st finger not reaching disk of 2nd.
© A. Nöllert
67
 Bufonidae

From top to base Description

Fig. 57 Dorsal pattern of a female of Ansonia hanitschi
from Sg. Silau-Silau.
Fig. 58 Belly of two females of Ansonia hanitschi.
Males up to 32 mm, female up to 37 mm; TL/SVL
0.45-0.57; head as broad as long; canthus rostralis
sharp; lores vertical; tympanum half eye diameter,
close to the eye; tips of fingers and toes swollen
into spatulate disks; rudimentary webbing at base
of fingers; weak outer palmar tubercle; toe web-
bing 1(0), 2(0/0-0.5), 3 (0.5-1/1), 4(1.5-2/1.5-2),
5(1.5-2); 2 long metatarsal tubercles; body and
limbs warty above; elongated warts dorsolaterally;
belly and chest coarsely, throat finely granular;
1 or 2 rows of yellowish mandibular spines.
Brown to greenish grey above, with oval light spots
on back and light bars on limbs; sometimes with an
extensive dark pattern on back; belly mottled with
brown or grey and yellow; chest and throat brown
flesh-coloured; nuptial pads dark brown; iris yellow-
ish grey to reddish golden with fine dark network.
Tadpoles black and teardrop-shaped, measuring
20 mm in total length (HERRMANN & ULBER 1992);
further description not published.

Ecology/Ethology
This species occurs along the edges of rocky
streams in montane primary forests (1000-1850 m
© R. Malkmus

a.s.l.). A. hanitschi is a particularly agile species

that frequents a variety of microhabitats (terrestrial,
riparian, arboreal). It usually tries to escape by
jumping into the stream
where it dives to seek
cover beneath rocks or
leaves, or crosses over to
the opposite bank. Cap-
tured specimens occa-
sionally arch their bodies
upward like the european
Bombina bombina (“Un-
ken reflex”), emit a sting-
ing scent, and give off a
high-pitched, sharply me-
tallic sounding distress
call (“tchigg”) which is
sometimes followed by a
series of clicking sounds.
Representatives of this
© R. Malkmus

species were found both

during the day and at
night, although calling

68
 Bufonidae

males were only encountered during the day (be-

tween 6.00 a.m. and 6.00 p.m.) perched on the
banks of streams, rock formations, sand and gravel
banks, and on emerged rocks in sprayzones. In con-
trast to most of the other anurans their calling ac-
tivity is hardly affected even by prolonged periods
of drought. Although territories are marked acous-
tically, aggressive behaviour may be witnessed oc-
casionally. In this regard it was observed that one
of two males, which were perched opposite each

© M. Maronde
other at a distance of only 1 cm and called
antiphonically, suddenly raised on its hind limbs,
grabbed the head of its opponent and pressed to its
chest for about 30 seconds. A captive pair produced Fig. 59 Male of Ansonia hanitschi.
strings comprising 45-73 eggs on rocks submerged
in water during the day (HERRMANN & ULBER 1992).
Tadpoles live in rocky streams with moderate to git/Poring (500 m), INGER, STUEBING & TAN (1996)
strong currents (m: 1a, b, c). for “Layang Layang” (2600 m), but both these
A. hanitschi can often be found resting or foraging records are doubtful, because of its normal altitudinal
on the leaves of low (up to 2 m) shrubs near streams range.
during the night. Captive specimens
accepted ants, flies, pentastomids,
earthworms, spiders, and the larvae Fig. XXXIV Sonagram and oscillograms of Ansonia hanitschi
of beetles and butterflies. (Sg. Liwago, 1500 m; 3.00 p.m. +21°C)

Call: (+21°C) A frequently repeated

note series which is very much remi-
niscent of the chirping of a cricket.
Each series takes 6-10 s and is com-
posed of equal notes with a note rep-
etition rate of 6 per sec. Each note
(duration 140-150 ms) comprises a
group of 3 pulsed single notes (du-
ration 21-50 ms) which are repeated
at intervals of 25-35 ms; frequency
5.4 to 6 kHz.

Distribution
A. hanitschi is only known from sev-
eral mountain ranges in western
Sabah and northern Sarawak. Mt.
Kinabalu: The species is very com-
mon in the Silau-Silau/Liwago region
(1400-1850 m); Kadamaian Falls
(1600 m); Mesilau Cave; Sayap; Sg.
Kinateki; HANITSCH (1900a) recorded
it from “2100 and 4200 ft.”; WONG
(1994) mentioned it for Sg. Kipun-

69
 Bufonidae
Ansonia longidigita INGER, 1960
Ansonia longidigita longidigita INGER, 1960b: 480 – Type
locality: “at 4,200 feet on Mount Kina Balu, North Borneo”
Bufo leptopus, (not Bufo leptopus GÜNTHER, 1872) BOULENGER
1887b: 95; MOCQUARD 1890: 159; WHITEHEAD 1893: 121;
BARTLETT 1894: 204; HANITSCH 1900a: 74; VAN KAMPEN 1923:
75; SMITH 1931: 13 (part);
Ansonia leptopus, (not Bufo leptopus GÜNTHER, 1872) INGER
1956: 395 (part)
Ansonia longidigita longidigita, INGER 1966: 111, 1978: 312
Ansonia longidigita, MANTHEY 1982: 21; INGER & DRING 1988:
463; MALKMUS 1988c: 176, 1989: 184, 1992a: 106; INGER
1992b: 228; INGER & STUEBING 1992: 41-51; WONG 1994:
25-37; MALKMUS 1996c: 281; HOFFMANN 1995a: 9; INGER &
TAN 1996a: 554; HOFFMANN 1998b: 85; LAKIM et al. 1999:
32; INGER et al. 2000: 16
Ansonia sp., INGER 1985: 20
Diagnosis
A moderately large to large, slender-bodied Ansonia
with an oblique, projected snout; fingers long and
Fig. 60 Female of Ansonia longidigita from Sayap, Sg. Lumotok Kecil, 1100 m.
© P. Hoffmann
70
slender; tip of 1st finger reaching base of swollen
tip of 2nd finger.
Description
Males up to 50 mm, females up to 70 mm; TL/SL
0.53-0.61; head slightly wider than long; canthus
rostralis sharply angular; lores vertical; tympanum
two fifths of eye diameter; tips of fingers and toes
swollen; fingers without webs; a large round pal-
mar tubercle; toe webbing 1(0), 2(0/0), 3(1.5/1-1.5),
4(2.5-3/2.5-3), 5(0.5-1); 2 low, round to oval meta-
tarsal tubercles; limbs with small, elevated,
spinose warts dorsally and dorsolaterally; throat
finely granular, other ventral surfaces coarsely tu-
bercular; 2-4 rows of brown to black mandibular
spines.
Dark brown above, with small irregular light spots
and single brick-red warts; obscure light cross-
bars on limbs; throat brownish, chest and belly
yellowish with brown or black spots; nuptial pads
 Bufonidae

© P. Hoffmann
© M. Schroth

Fig. 61 Tadpole of Ansonia longidigita from Sg. Fig. 62 Male of Ansonia longidigita from Kundasang,
Kemantis. Sg. Kinesepian, 1500 m.

yellowish brown; iris light golden brown, with a (duration 36-44 ms) consisting of 2-3 pulses (dura-
fine black network. tion 7-33 ms). The internote interval takes 31-36 ms,
Tadpoles teardrop-shaped; snout broadly roun- the dominant frequency is 3-4 kHz with a harmonic
ded; eyes dorsolateral, oral disk ventral; labial at 10.8 kHz.
teeth rows formula 2/3; upper beak
divided by a large gap; tail tapering
to a pointed tip; body and caudal Fig. XXXV Sonagram and oscillograms of Ansonia longidigita
muscle dark blackish brown; total (Sg. Langanan, 800 m; 7.30 p.m. + 22°C)
length 20-25 mm.

Ecology/Ethology
This toad is distributed over a wide
altitudinal range (150 to 2100 m
a.s.l.) in primary and old secondary
forests. It is active at night, and by
day stays hidden beneath logs,
leaves, and in crevices of rocks. Its
breeding sites are clear, swift, rocky
streams. The males call at night from
spray-moistened rocks, gravel bars,
and from the leaves and twigs of low
vegetation (up to 50 cm) forming
calling choruses.
Tadpoles live on the bottom of
streams with moderate to strong cur-
rents in primary forests (m: 1a, b, c).

Call: (+22°C) The long note series

(0.9-10.5 s) is composed of long,
high-pitched trills consisting of
4-135 notes at a note repetition rate
of 12.5-15.9 per second, each note

71
 Bufonidae
Distribution
Ansonia longidigita is known from Sabah, Brunei,
and Sarawak. Mt. Kinabalu: On Mt. Kinabalu, the
species has an extensive altitudinal range with a
recorded maximum of 2100 m, but it is less com-
mon above 1000 m a.s.l.
Sg. Panataran, 4800 ft. (WHITEHEAD 1893); Keno-
kok, Kiau, Sg. Kadamaian; Sg. Liwago/Silau-Silau/
Tibabar, Kamborangoh (SMITH 1931, as A. leptopus,
2100 m); Sg. Mesilau; Sayap: Sg. Kemantis, Sg.
Wariu, Sg. Lumotok Kecil; Sg. Luidan, near Bundu
Tuhan (1100 m); Poring: Sg. Langanan (very com-
mon between 650 and 850 m a.s.l.).
Fig. 63 Female of Ansonia cf. platysoma from Sayap, Sg. Wariu.
© M. Schroth
72
Ansonia platysoma INGER, 1960
Ansonia platysoma INGER, 1960b: 487 – Type locality: Luidan
River near Bundu Tuhan at 3,300 feet on Mount Kina Balu,
North Borneo
Ansonia platysoma, INGER 1966: 102, 1978: 312; MANTHEY
1983: 21; INGER & DRING 1988: 463; MALKMUS 1988c: 176;
INGER & STUEBING 1992: 25-37; MANTHEY 1994b: 222; INGER,
STUEBING & T AN 1996: 362; INGER & TAN 1996a: 554;
MALKMUS 1996c: 281; HOFFMANN 1995a: 9
Diagnosis
A small, very slender species with a distinctly de-
pressed body and head; snout strongly projected,
 Bufonidae

sloping in profile; 1st finger very short,

not reaching disk of 2nd; tips of outer fin-
gers spatulate; mandibular spines absent.

Description
Males and females 20-25 mm; TL/SVL
0.51-0.58; head longer than wide; can-
thus rostralis sharp; lores vertical; tym-
panum very close to the eye, half the lat-
ter’s diameter; fingers slender with a ru-
dimentary web between first 2 fingers;
a low, round outer palmar tubercle, an
inner one at base of 1st finger; tips of toes
swollen; toe webbing 1(0), 2(0/0.5-1),
3(1/1), 4(3/3), 5(1.5); 2 low metatarsal
tubercles; skin above and laterally with
low warts, spinose on limbs; belly and
chest coarsely granular, throat finely so;
males without mandibular spines.

© P. Hoffmann
Dark brown above and laterally, some-
times with yellowish dots; limbs with
light crossbars; below blackish, throat
and belly with small whitish spots; nup- Fig. 64 Ansonia platysoma from Poring, Sg. Kipungit II, 650 m.
tial pads blackish brown.

Ecology/Ethology Distribution
Little is known of this species. It was observed in Only a few sites are known for this species from Sabah
primary forest (150-1375 m a.s.l.). Males call and Sarawak. Mt. Kinabalu: Sayap: Sg. Kemantis,
throughout the day with considerable persistence Sg. Wariu, Sg. Lumotok Besar, Sg. Minodtuhan; Sg.
from rocks which may be sprayed by water in, or Luidan, near Bundu Tuhan; Poring: Sg. Kipungit
from the banks of, forest streams, usually in the (600 m), Sg. Langanan (900-950 m).
vicinity of small cascades. At
night, A. platysoma can be found
on the leaves of low vegetation Fig. XXXVI Sonagram and oscillogram of Ansonia platysoma
(up to 50 cm high), sometimes (Sg. Wariu, 1000 m; 2.00 p.m. +24°C)
also at some distance from the
streams.

Call: A series of high-pitched

trills, similar to a cricket song.
It usually consists of more than
100 weakly pulsed notes (dura-
tion 10-15 ms, internote interval
100-150 ms; the interval be-
tween the last two notes of a note
series is distinctly longer; note
repetition rate 5-7 per sec) with
a frequency of 7.8 to 8.2 kHz.

73
 Bufonidae

Ansonia spinulifer (MOCQUARD, 1890) dorsolateral warts large, elevated, juxtaposed, with
strong spines; characteristic light oval patch be-
tween shoulders.
Bufo spinulifer MOCQUARD, 1890: 160 – Type locality: Kina
Balu
Ansonia leptopus, (not Bufo leptopus GÜNTHER, 1872) INGER Description
1966: 110, 1978: 212; MALKMUS 1988d: 7, 1989: 282 Males up to 41 mm, females up to 50 mm; TL/SVL
Ansonia spinulifer, BOULENGER 1894a: 81; VAN KAMPEN 1923: 0.48-0.53; head broader than long; canthus rostralis
79; INGER & DRING 1988: 463; INGER 1992b: 229; INGER & sharp; lores vertical; tympanum two fifths of eye
STUEBING 1992: 41-51; WONG 1994: 25-37; HOFFMANN 1995a:
diameter; tips of fingers dilated into small round
9; INGER, STUEBING & TAN 1996: 362; INGER & TAN 1996a:
555; MALKMUS 1996c: 282; LAKIM et al. 1999: 32; INGER et disks, tips of toes a little smaller; fingers free of
al. (2000): 16 webbings; a large palmar tubercle present; toe web-
bing limited to the bases, rudimentary; 2 metatar-
Diagnosis sal tubercles; dorsal skin see Diagnosis above, be-
A medium-sized lowland species; snout project- low granular; males with 2-3 rows of dark man-
ing, oblique in profile; 1st finger reaching base of dibular spines.
tip of 2nd; toes almost free of webbings; dorsal and Dark brown to black above, with a light oval or
diamond-shaped spot
between shoulders; a
Fig. 65 Ansonia spinulifer from Poring, 550 m. light streak on groin,
and light crossbars on
limbs; the light areas
sometimes intermixed
with brick-red warts;
side of head light
brown; below blackish
with white dots, some-
times forming a net-
work; nuptial pads dark
brown; iris golden
brown.
Tadpoles teardrop-
shaped; snout very
broadly rounded; body
widest in plane before
eyes; eyes dorsal; oral
disk as wide as body; la-
bial teeth rows formula
2/3; upper beak divided
by a large gap; tail ta-
pering in the posterior
third to a rounded tip;
uniformly black; total
length up to 20 mm.

Ecology/Ethology
© R. Malkmus

A. spinulifer occurs in
primary forest up to
1000 m altitude. It is

74

after INGER 1992b
Fig. XXXVII Tadpole of Ansonia spinulifer.
nocturnal and has been observed on the forest floor
and in the neighbourhood of streams. Males call
from the top of rocks or from low vegetation, but
do not form calling aggregations.
Tadpoles live in forest streams with swift currents
(m: 1b, c), clinging to smooth rocks and feeding
on the organic film (algae) there. Occasionally, tad-
Genus Bufo LAURENTI, 1768
This cosmopolitan genus is the only Bornean toad
genus containing species outside the Oriental trop-
ics. None of the five Bornean species of Bufo is
restricted to this island, three of them also occur
on the Malay Peninsula.
The Bornean species are medium-sized to large
toads, with rather short limbs, stocky shapes, thick
warty skins, and large parotoid glands behind the
eyes. Tips of fingers and toes rounded, not dilated;
no webbing between fingers, all Bornean species
with distinct tympanae. The males with a median
Tab. 2 Distribution of Bufo-species on Borneo.
Bufonidae
after INGER 1992b
Fig. XXXVIII Oral disc of the tadpole of Ansonia
spinulifer.
poles can be found ascending vertical, water-
splashed cliffs up to heights of 5 cm.
Distribution
Known only from a few sites in Sabah and Sarawak.
Mt. Kinabalu: Sayap: Sg. Kemantis (1000 m); Por-
ing: Sg. Langanan, Sg. Kipungit, Bat Caves.
subgular vocal sac and nuptial pads; mating in ax-
illary amplexus.
The species of Bufo lay egg strings with numer-
ous, pigmented ova in stagnant or slowly flowing
water. Their larvae are of a generalised type, with
spheroidal bodies, keratinized beaks, and deep fins.
These toads are terrestrial, dwelling in both pri-
mary and secondary forests as well as clearing and
plantations; Bufo juxtasper can be found in agri-
cultural land, B. melanostictus in villages and
towns.
On Mt. Kinabalu, this genus is rarely encountered at
elevations above 1000 m a.s.l. Five species are known
from Borneo, two of them occur on Mt. Kinabalu.
75
 Bufonidae

Key to the Bornean species of Bufo (based on INGER 1966)

1 A pair of crests between eyes and parietal region
.......................................................................... 2
• If crests between eyes present, they do not
continue back onto the parietal region ............. 3
2 Parotoids long, sharply raised, their maximum
widths at most 1/5 of their lengths ......................
..................................................... quadriporcatus
• Parotoids triangular or oval in shape, their
maximum widths at least 2/5 their lengths .........
............................................................ divergens
3 A bony ridge curving around the eye to the
parotoid gland .............................. melanostictus
• No such ridge .................................................... 4
4 Parotoid gland elongated, much longer than eye
in diameter ........................................... juxtasper
• Parotoid gland round or triangular, about the size
of the eye .................................................... asper

Fig. 66 Bufo divergens, Mulu NP, Sarawak. Bufo divergens PETERS, 1871

Bufo divergens P ETERS , 1871: 579 – Type

locality: Sarawak
Bufo biporcatus, (not Bufo biporcatus GRA-
VENHORST, 1829) MOCQUARD 1890: 158; VAN
KAMPEN 1923: 88; SMITH 1925b: 30, 1931:
30
Bufo biporcatus biporcatus, INGER 1956: 394
Bufo biporcatus divergens, INGER 1966: 60,
1978: 312; MANTHEY 1983: 20
Bufo divergens, INGER , STUEBING & TAN 1996:
362; INGER & TAN 1996: 555

Diagnosis
A relatively small-sized, stocky toad;
snout with a small median bulge at
the tip, projecting, and oblique in pro-
file; a pair of narrow bony crests be-
tween the eyes; toes webbed over half
their lengths; 1st finger longer than 2nd.

Description
Males up to 45 mm, females up to
65 mm; TL/SVL 0.32-0.48; head
wider than long; canthus rostralis
sharp, lores vertical; tympanum about
© M. Schroth

3/5 the diameter of the eye; tips of

fingers and toes blunt, not swollen;
supernumerary metacarpal tubercles

76

present; toe lengths 4>3>5>2>1; toe webbing 1(1),
2(1/1), 3(1.5/1.5), 4(3/3-3.5), 5(1.5); inner meta-
tarsal tubercles oval; a smaller, rounded outer one;
skin above and laterally with many small, conical
spiny warts; a row of enlarged warts between eye
and groin; a row of spinose tubercles on inner edge
of tarsus; venter coarsely granular; parotoids tri-
angular to oval.
Reddish brown above, often with dark spots; dark
interorbital marking; limbs with dark crossbars;
nuptial pads black; below yellow or light brown,
with or without dark mottling; iris golden brown
with a dark network.
Tadpoles: Head-body portion of tadpoles oval,
slightly flattened above; eyes and nostrils dorsal,
oral disk ventral, subterminal; no expanded lips (in
contrast to the larvae of B. juxtasper); labial teeth
rows formula 2/3. Head-body portion and caudal
muscle dark brown to black; venter without pig-
mentation; dorsal fin dusky, ventral fin only along
the base so; total length 15-17 mm.
Ecology/Ethology
This toad lives in forest floor litter and is confined
to lower elevations (< 1000 m a.s.l.). It can be found
in both primary and secondary forests, plantations,
and village clearings. B. divergens is mainly active
at night, although it was also observed hunting
during daylight hours. It feeds on a wide variety of
arthropods, favouring ants and termites. The males
call at night, forming calling communities along
the edges of spawning sites.
Tadpoles occur in the shallow areas of stagnant or
slowly flowing water in small streams (m: 1e, f).
They are bottom suspension feeders (INGER 1985,
1986).
Call: A series of short, raspy trills. Sonagram not
published.
Distribution
Bufo divergens is known from Borneo, Sumatra,
and the Natuna Islands. Mt. Kinabalu: This spe-
cies was mentioned for Mt. Kinabalu only once,
i.e. in a check list by SMITH (1931), as “known to
inhabit Mt. Kina Balu above 3 000 feet”, which
refers presumably to the Kiau region. Recently, tad-
poles were found near Poring.
Bufonidae
Bufo juxtasper INGER, 1964
Bufo juxtasper INGER, 1964: 154 – Type locality: Sungei Tawan,
Kalabakan, Tawau District, Sabah
Bufo asper, (not Bufo asper GRAVENHORST, 1829) MOCQUARD
1890: 158; VAN KAMPEN 1923: 82; SMITH 1931: 30
Bufo juxtasper, INGER 1966: 69; MATSUI 1979: 313; MANTHEY
1983: 21; MALKMUS 1985: 13, 1987: 277, 1988b: 9, 1988c:
176, 1992a: 106; INGER & STUEBING 1992: 41-51; MALKMUS
1994b: 220; WONG 1994: 25-37; HOFFMANN 1995a: 9; INGER ,
STUEBING & TAN 1996: 362; INGER & TAN 1996a: 555; LAKIM
et al. 1999: 32; INGER et al. (2000): 16
Diagnosis
A very large, stocky toad, with a broad, blunt head;
ear-openings rather small; a thick supraorbital, but
no parietal crest; toes fully webbed, except for the
4th toe; 1st finger equal in length to 2nd.
Description
Males up to 125 mm, females up to 215 mm; TL/SVL
0.42-0.48; head wider than long; snout pointed;
canthus rostralis rounded, lores vertical; tympanum
smaller than one third of the eye diameter; tips of
fingers and toes swollen; subarticular tubercles
large; supernumerary metacarpal tubercles present;
toe lengths 4>3>5>2=1; toe webbing 1(0), 2(0/0),
3(0/0), 4(1.5/1.5), 5(0); metatarsal tubercles large;
a sharp tarsal ridge; skin above and on the sides
covered with small to large, round warts; parotoids
large, as long as their distance from the tip of the
snout; ventral sides coarsely granular, each gran-
ule tipped with a melanic spinule.
Grey to earth-brown above, lighter on the sides;
venter greyish white with dark mottling and spots;
nuptial pads black; iris yellowish golden with a fine
black network.
Tadpoles: Head-body of tadpoles oval; eyes and
nostrils dorsal, oral disk ventral; labial teeth rows
formula 2/3(1-2). Head-body, and caudal muscle
blackish brown; disk region greyish; venter golden
yellowish; total length up to 20 mm.
Ecology/Ethology
Bufo juxtasper lives in primary forests and dis-
turbed habitats associated with the activities of
man (secondary forests, agricultural regions, vil-
lages). The altitudinal distribution range is mod-
erately wide, and it is usually encountered below
77
 Bufonidae
© A. Nöllert

Fig. 67 Bufo juxtasper from Sg. Kupungit I, Poring.

Fig. 68 A pair of Bufo juxtasper in amplexus,

near Tambunan, Sabah.
© R. Malkmus

78
 Bufonidae

Call: The call consists of a single, resonant,

hoarse chirp. The Dusun name for B. juxtasper
imitates the sound of two calling toads as “bu-
angkut”.
There are two types of calls, a shorter and a
longer one. The former lasts 0.07 to 0.16 s and
is repeated 2 to 2.8 times per second; dominant
frequency 540-730 Hz; harmonics are not evi-
dent. The longer call is a typical trill in which a
© U. Manthey

shorter note (0.05 s; 6 pulses) is followed by a

long one (> 1 s; 130 pulses), dominant frequen-
cies slightly higher than in the shorter call
Fig. 69 Tadpole of Bufo juxtasper from Aceh, Sumatra. (MATSUI 1982a).

Distribution
1000 m a.s.l. This species is active by day and night. Borneo and Sumatra. Mt. Kinabalu: Common in
Toads of this species were found on gravel banks, disturbed habitats along the boundaries of the Park
large boulders, and fallen logs on the banks and in (valley of Sg. Kadamaian near Kiau, Kebeyau;
the middle of the lower Sg. Langanan and Sg. Kada- Poring; lower Sg. Langanan, Sg. Mantukungan;
maian, but also far away from water on the leaf Mamut Copper Mine), but rare in primary forest
litter of the forest floor. When captured, this toad (Sg. Kipungit, Sg. Sasapan; Sayap; lower Sg. Silau-
produces a large quantity of a foul smelling, white Silau, 1500 m a.s.l.).
secretion. Its diet in-
cludes a wide variety
of spiders, worms, and Fig. 70 Oral disc of Bufo juxtasper from Aceh, Sumatra.
insects, particularly
ants and termites. In
the Poring region it
was observed along ant
paths formed by the
large ant, Camponotus
gigas, consuming con-
siderable quantities of
these insects. The
males call from along
stream banks. The spe-
cies name is onomato-
poeic in being reminis-
cent of the call of the
species.
The strings of gelati-
nous, brownish grey
eggs are attached to
aquatic plants and
rocks in spring basins
© U. Manthey

and calm sections of

streams and rivers (m:
1e, f).

79
 Bufonidae
Genus Leptophryne FITZINGER, 1843
The genus Leptophryne comprises two species: L.
cruentata TSCHUDI, 1838, of Java, and L. borbonica
(KUHL & VAN HASSELT, 1827), of the Malay Penin-
sula, Sumatra, Java and Borneo. It is the only ge-
nus of Indo-Malayan bufonids with an arcifero-
firmisternal pectoral girdle.
Leptophryne borbonica
(KUHL & VAN HASSELT, 1827)
Hylaplesia borbonica KUHL & VAN HASSELT in SCHLEGEL, 1827:
294 – Type locality: “East Indies”, restricted to “Java” by
INGER (1966)
Cacophryne borbonica, INGER 1966: 75, 1978: 312; MANTHEY
1983: 21
Leptophryne borbonica, INGER & STUEBING 1992: 41-51; INGER,
STUEBING & TAN 1996: 362; INGER & TAN 1996a: 554
Diagnosis
A small, slender bufonid with very long, slender
limbs; head without bony crests; snout short, pro-
Fig. 71 Leptophryne borbonica (female) from Crocker Range NP, Sabah.
© I. Das
80
jecting slightly over the mouth; tympanum distinct;
fingers and toes long, with small, round disks on
the tips; no webbing on the fingers, toes webbed
over half their lengths.
Description
Males up to 31 mm, females up to 50 mm; TL/SVL
0.56-0.70; head as wide as long; canthus rostralis
sharp, lores vertical, straight; tympanum oval,
one third the eye diameter; finger lengths
3>4>2>1; toe lengths 4>5>3>2>1; subarticular
tubercles large; toe webbing 1(1), 2(1/1-2), 3(2-
3/2-3), 4(3-3.5/3-3.5), 5(2-3); inner metatarsal
tubercle oval, outer one round; nuptial pads
present; skin above and on the sides with numer-
ous warts; parotoids weakly developed; a row of
dorsolateral glands; lower side granular; a row
of conical tubercles on the outer side of the lower
arm and tarsus; males with a median subgular
vocal sac.
Greyish to brown above and on the sides, with
darker and lighter markings, usually forming an X
in the centre of the back; limbs with dark cross-
bars; ventrum yellowish, mottled with brown; throat

© U. Manthey
Fig. 72 Leptophryne borbonica (male) from North
Sumatra, near Berastagi, 1450 m.
greyish to brown; undersides of the legs grey to
yellow with brown spots; iris golden brown.
Tadpoles: Head-body portion of tadpoles oval,
slightly depressed; eyes and nostrils dorsolateral, oral
disk ventral, subterminal; lower lip bordered by pa-
pillae all around; labial teeth rows formula 2(2)/3 or
2(1-2)/3; beaks finely serrated, white, narrowly
Genus Pedostibes GÜNTHER, 1875
The species of the genus Pedostibes are native to
the Malay Peninsula, Sumatra, and Borneo.
This genus contains medium-sized to large, mod-
erately stout toads with long legs. Parotoids present
or absent. Fingers long dilated at tips into truncate
disks; tips of toes round and swollen. Tympanum
present or absent. Males with median subgular vo-
cal sac. The species produce small, pigmented eggs
Bufonidae
after BERRY 1972
Fig. IXL Tadpole of Leptophryne borbonica.
edged with black; head-body and caudal muscle
black; fins blackish grey; total length up to 11 mm.
Ecology/Ethology
Nearly nothing is known on the natural history of
this species other than it is terrestrial and occurs in
primary forest where it favours the proximity of
forest streams, especially seepage areas, from sea
level to 1500 m altitude.
The eggs are not arranged in strings, but deposited
in clumps of 200-300 in calm sections of streams
(m: 1f). The tadpoles are benthic feeders.
Distribution
Malay Peninsula, Sumatra, Java, and Borneo. Mt.
Kinabalu: The only hint towards an occurrence
on Mt. Kinabalu is found in INGER (1966) as “Kota
Belud District, Mount Kina Balu”, which presum-
ably refers to the Sg. Kadamaian region.
in strings, which are deposited in calm side pools
of small forest streams. The only tadpoles known
are those of P. hosii; they are benthic feeders and
similar to those of the genus Bufo.
Pedostibes are arboreal and occur in primary and
old secondary forests at altitudes ranging from
sea level to 1000 m; nearly nothing is known with
regard to the vertical distribution on Mt. Kina-
balu.
Three or four species (see remark) are found on
Borneo, two of them also occur on Mt. Kinabalu.
81
Bufonidae
Key to the Bornean species of Pedostibes (based on INGER 1966)
1 Tarsal ridge present ........................................... 2
• No tarsal ridge ................................................... 3
2 Parotoid oval or circular; lateral and medial bor-
ders convex ............................................. rugosus
• Parotoid triangular; lateral border straight or
concave, medial border convex .................. hosii
3 Parotoid glands present; tympanum visible ........
............................................................... everetti
• No parotoid glands, tympanum invisible ............
......................................................... maculatus
Tab. 3 Distribution of Pedostibes-species on Borneo.
Pedostibes everetti (BOULENGER, 1896)
Nectophryne everetti BOULENGER 1896b: 450 – Type locality:
Mount Kinabalu, Sabah
Nectophryne everetti, VAN KAMPEN 1923: 68; SMITH 1925b: 31,
1931: 30
Pedostibes everetti, BARBOUR 1938: 192; INGER 1966: 90, 1978:
312; MANTHEY & DENZER 1982: 17; MANTHEY 1983: 21;
MALKMUS 1988c: 176; INGER & STUEBING 1992: 41-51; INGER,
STUEBING & T AN 1996: 362; I NGER & T AN 1996a: 555;
MANTHEY & GROSSMANN 1997: 38
Diagnosis
A medium-sized to large, moderately stout
Pedostibes with egg-shaped parotoids; skin of the
upper surfaces very warty; no tarsal ridge; above
green with large brown spots.
Description
Up to 87 mm; head broader than long; no cranial
crests; tympanum one third the eye diameter; fin-
gers webbed on the bases; finger lengths 3>4>2>1;
82
Remark: INGER (1958c) described the species P.
rugosus which, however, is hardly distinct morpho-
logically from P. everetti, stating: “The main differ-
ences being the absence of a tarsal ridge, the smaller
digital disks and the greater separation of parotoid and
eyelid in everetti” (INGER 1966). Further considera-
tion of variation in P. rugosus demonstrated (MANTHEY
& GROSSMANN, 1997), that none of his diagnostic char-
acters were distinct from character states exhibited
by P. everetti.
toe lengths 4>3>5>2>1; toe webbing 1(0), 2(0/0,
3(0/0), 4(2/2), 5(0); two metatarsal tubercles.
Green above and on the sides, with small to large,
irregularly scattered, dark brown to reddish brown
spots; limbs green with brown crossbars; lower
sides cream-coloured, immaculate or mottled with
brown.
Ecology/Ethology
The only report on field observations is found in
MANTHEY & GROSSMANN (1997) reading in transla-
tion: “On Mt. Kinabalu, in the environs of Poring,
we found nearly 200 juvenile toads on the bank of
a partially torrential stream on a clearing in the rain-
forest (650 m a.s.l.) in August. Here, the stream
was blocked by wedged drift wood which had
caused it to form a pool. ... The juveniles measured
10-15 mm and presented themselves openly on the
sparsely vegetated ground with only little leaf lit-
ter.” Some specimens were raised in terraria. While

© U. Manthey
they were terrestrial and diurnal during the initial
4-6 weeks, they then began to climb and shifted
their activity to the hours of dusk and night. After
only one year they had grown to 60-82 mm SVL.
Distribution
This species is only known from “Mount Kinabalu,
Sabah” (BOULENGER 1896, INGER 1966), presumably
from the Kiau region. A more recent record stems
from the Sg. Kipungit II region (650 m a.s.l.)
(MANTHEY 1983).
Bufonidae
© U. Manthey
Above
Fig. 73 Pedostibes everetti, nearly 20 years old from
Sg. Kipungit II, near Poring.
Left
Fig. 74 Pedostibes everetti, juvenile from the same place
as above.
Pedostibes maculatus (MOCQUARD, 1890)
Nectophryne maculata MOCQUARD, 1890: 162 – Type locality:
Kina Balu
Nectophryne maculata, VAN KAMPEN 1923: 71
Pelophryne maculata, BARBOUR 1938: 193
Pedostibes maculatus, INGER 1966: 92, 1978: 312; MANTHEY
1983: 21; INGER & STUEBING 1992: 41-51; INGER, STUEBING &
TAN 1996: 362; INGER & TAN 1996a: 556
Diagnosis
A small, slender Pedostibes with long limbs; fin-
gers long, tips with truncate disks; head without
bony crests; tympanum invisible; parotoid glands
absent; no tarsal ridge.
83
Bufonidae

Description
Males up to 42 mm, females up to 51 mm; TL/SVL
0.54-0.60; snout truncate, projecting in profile; fin-
ger lengths 3>4>2>1; subarticular tubercles mod-
erate large; fingers webbed at the bases; tips of toes
not dilated; toe lengths 4>3>5>2>1; toe webbing
1(1), 2(1/1), 3(1/1.5), 4(3/3), 5(1-1.5); an oval in-
ner and a round outer metatarsal tubercle; numer-
ous round warts with black spines above and on
the sides.
Brown with irregular dark spots above and on the
sides; limbs with dark crossbars; greyish below.

Ecology/Ethology
There is no information on the life history of this
tree toad.

Distribution after MOCQUARD 1890

Distributional data is limited to “Mt. Kinabalu,

Fig. XL Pedostibes maculatus
Sabah”.

The females produce large, non-pigmented, whit-

Genus Pelophryne BARBOUR, 1938
The genus Pelophryne is distributed on the Malay
Peninsula, Sumatra, Borneo, and the Philippines.
The dwarf toads represent the most distinctive ge-
nus of Oriental bufonids: Very small toads (SVL <
31 mm) with a moderately stout body and slender
limbs; hands and feet with fleshy webbings, partly
reduced phalanges; tympanum visible, parotoid
glands absent. Coccyx fused to sacral vertebra and
expanded dorsally; only seven presacral vertebrae.
Males with subgular vocal sac; nuptial pads and
mandibular spines may be present or not.
ish ova in small clutches (usually less than 20 eggs).
Suitable spawning sites are rainwater-filled depres-
sions and phytothelmes (e.g. leaf axils and tree
holes). Larvae small, subterminal, circular oral disk
with reduced labial teeth and weak beaks; spiracle
not visible. They are non-feeding and subsist en-
tirely on yolk.
Pelophryne are terrestrial and can be found in low
vegetation in primary and old secondary forests. The
altitudinal distribution of this genus is wide-rang-
ing, stretching from sea level up to 3100 m a.s.l.
Eight species are known from Borneo, but only one
occurs on Mt. Kinabalu.

Tab. 4 Distribution of Pelophryne-species on Borneo.

84

Preliminary key to the Bornean species of Pelophryne
1 Tips of fingers dilated ....................................... 3
• Tips of fingers not dilated ................................. 2
2 Dorsum black, without extensive light areas ... api
• Dorsum brown, with lighter areas .......... misera
3 Tympanum at least four fifths of the eye diameter
................................................................ macrotis
• Tympanum at most three fifths of the eye diameter
........................................................................... 4
4 5th toe webbed up to the disk .............. guentheri
• 5th toe not webbed up to the disk ...................... 5
5 A yellowish white stripe between eye and tym-
panum .................................................... brevipes
• No such light stripe below the eye ... rhopophilius
Pelophryne misera (MOCQUARD, 1890)
Nectophryne misera MOCQUARD, 1890: 161 – Type locality:
“Nord de Bornéo”, restricted here to “Kadamaian River,
Mt. Kinabalu” (based on data provided by WHITEHEAD 1893)
Nectophryne misera, WHITEHEAD 1893: 177; BOULENGER 1894a:
81; SMITH 1931: 14
Nectophryne guentheri, (not Nectophryne guentheri BOULEN-
GER, 1882) BARTLETT 1894: 204
Pelophryne misera, BARBOUR 1938: 193; INGER 1954a: 250,
1966: 88, 1978: 312; MANTHEY 1983: 21; MALKMUS 1985:
12, 1987: 277, 1988c: 176; INGER & STUEBING 1992: 41-51;
INGER, STUEBING & TAN 1996: 362; MALKMUS 1996c: 280;
INGER & TAN 1996a: 556; INGER & STUEBING 1997: 87
Pelophryne brevipes, (not Hylaplesia brevipes PETERS, 1867)
MALKMUS 1989: 183, 1991a: 27, 1992a: 110, 1994b: 221,
1996a: 129; INGER et al. 2000: 17
Diagnosis
A small montane species with a moderately stout
habitus and slender limbs; snout truncate, vertical
in profile; tips of fingers and toes not wider than
bases of phalanges; fingers with fleshy webbings.
Description
Males up to 21 mm, females up to 23 mm; TL/SVL
0.34-0.41; head as wide as body; canthus rostralis
sharp, concave; lores vertical; tympanum half to two
thirds of eye diameter; finger webbing 1(0), 2(1/1),
3(2/2), 4(1); subarticular and palmar tubercles hid-
den; toe webbing thick, 1(0.5), 2(0.5/0.5), 3(0.5-1/
Bufonidae
Remark: ROUX (1906) placed Pelophryne exigua in
the synonymy of P. guentheri, INGER (1966) referred
it “in the absence of distinguishing characters” to the
synonymy of P. brevipes. In the latest check list of the
frogs of Borneo (INGER & TAN 1996a), P. exigua ap-
pears again as a valid species.
P. signata was likewise synonymised with P. brevipes
by INGER (1966), does consequently not appear in said
check list (INGER & TAN 1996a), but is treated as a
valid species by INGER & STUEBING (1997) without an
explanation.
For the purpose of the present book, P. exigua and P.
signata have been included in the species list with
reservations, but excluded from the key as the latter is
based on distinguishing morphological traits.
0.5-1), 4(2/2-2.5), 5(1); subarticular and metatarsal
tubercles hidden or absent; skin above with irregu-
larly scattered, small, conical warts, forming frag-
ments of dorsolateral rows; sides and venter coarsely
granular; males with or without mandibular spines
and yellow to brown nuptial pads.
Upper side blackish brown, reddish brown to ochre;
dorsum occasionally with 1-3 large, dark spots of
various shapes and an interorbital, trapezoid-shaped
marking; hind limbs with dark crossbars; dark
flanks often sharply set off from lighter dorsum; a
light supralabial spot between eye and tympanum;
lower sides yellowish white, with large, irregularly
Fig. 75 Pelophryne misera from Sg. Silau-Silau.
© A. Nöllert
85
 Bufonidae
© R. Malkmus
Fig. 76 Pelophryne misera from Liwago trail, 1550 m.
scattered brown spots which may fuse to form a
crude and incomplete reticulation; iris golden brown,
with dense black marbling.
Newly metamorphosed toadlets with highly contrast-
ing colour pattern: blackish brown laterally, a white
dorsolateral line between tip of snout and insertion
of hind limb; upper side whitish grey with large
blackish brown markings; upper side of hind limbs
black with narrow white crossbars.
Tadpoles: Head-body of tadpoles ovoid; eyes dor-
solateral, oral disk ventral, subterminal; labial teeth
rows formula 1/0; beaks weak; spiracle not tubu-
lar; body and caudal muscle blackish brown, fins
lighter; total length 13 mm; freshly metamor-
phosed toadlet 5.8 mm.
86
Ecology/Ethology
Pelophryne misera is locally abundant in the
montane forests of Mt. Kinabalu (1450 to 3100 m
a.s.l.). While the toad was found active, foraging
for food, and calling during the day at subalpine
elevations, it could never be discovered outside its
hiding places (beneath leaf litter, in rock crevices,
holes in the ground, dead pitchers of Nepenthes)
before the onset of dusk at altitudes below 2000 m.
Its locomotion is a slow striding gait with the body
raised high. Moving around like this, a foraging
toad will plunge its head deeply into mosses, li-
chens, and leaf litter every now and again. Its prey
(arthropods < 5 mm; ants, spiders, hymenoptera,
wood lice, etc.) is stalked, visually fixed, and then
 Bufonidae
© R. Malkmus

© R. Malkmus
Fig. 77 Eggs of Pelophryne misera from Sg. Liwago. Fig. 78 Tadpole of Pelophryne misera.

Fig. 79 Pelophryne misera, three days old.

“shot down” with the tongue. Males usually call
from the leaves of shrubs, 10-50 cm above the
ground.
Until now, the clutch of only one female from Sg.
Liwago has become known. It was discovered in
mid-August, in the depression of a dead leaf, in
which water had accumulated and consisted of 10
yellowish white eggs with diameters of 2.8 mm on
average. The larvae emerged after 16 days, one of
which metamorphosed under terrarium conditions
after another 44 days.

Call: So far, single notes and note series have been

recorded. Single note (+19°C) repeated at inter-

© R. Malkmus
vals of 4-10 s, duration 90-110 ms, frequency 3.5-
4.5 kHz; the note is pulsed and has a short grace
note. Note series (+19°C) consists of 7-12 pulsed
individual notes (40-50 ms) with
internote intervals of 100-130 ms;
at 7-8 kHz and peaks up to 14 kHz Fig. XLI Sonagram and oscillogram of Pelophryne misera
these calls belong to the highest (Sg. Liwago, 1550 m; 8.00 p.m. +19°C)
tones produced by bufonid anu-
rans.

Distribution
This species is known only from
two localities on Borneo, i.e. Mt.
Murud (Sarawak) and Mt. Kina-
balu. Mt. Kinabalu: Marei Parei
spur (1550 m), slopes of Sg.
Liwago valley (1450-1600 m),
Kamborangoh (2200 m), around
Pakka Cave (3000-3100 m).

87
Megophryidae

Megophryidae
The Oriental megophryids are distributed from Pakistan east to China, the Sunda Islands and the Philip-
pines.
The family is characterised by a arciferous pectoral girdle and a cartilaginous sternum and omosternum.
The presacrals are not fused. The sacrum has substantially expanded diapophyses and is fused with the
coccyx; the epicoracoids overlap at mid-line. The parahyoid is absent, maxillary teeth are present, vomerine
teeth absent (except in the species of the genus Megophrys). The pupil is vertically elliptical, except of
Ophryophryne. Mating takes place in an inguinal amplexus.
All Bornean species have aquatic larvae, most with beaks and labial teeth (except in the genera
Leptobrachella and Megophrys), and depressed, spheroidal body shapes (except in the genus Megophrys).
Megophrys have funnel-mouthed tadpoles.
Bornean megophryids are terrestrial and live in leaf litter layer of primary rain forests, particularly in the
vincinity of brooks. On Mt. Kinabalu they occur up to altitudes of 2200 m (Leptobrachella baluensis,
Leptobrachium gunungense).
On Borneo, the family is represented by 4 genera and 22 species, on Mt. Kinabalu by 4 genera and 13
species.

Key to the Bornean genera of Megophryidae

A B
1 Dermal appendage on upper eyelid and/or snout
(A) .................................................... Megophrys
• No such appendage(s) ....................................... 2
2 Head conspicuously wider than body (B) ...........
.................................................. Leptobrachium
• Head not wider than body ................................ 3
3 Tips of digits swollen, with pointed disks (C);
adults never larger than 25 mm SVL ..................
.................................................. Leptobrachella
• Tips of digits not swollen (D); adults larger than C D
25 mm SVL ....................................... Leptolalax

This genus includes the smallest forms (SVL <

Genus Leptobrachella SMITH, 1925
Representatives of the genus Leptobrachella are
only known from Borneo and the small islands
Pulau Serasan and Pulau Natuna Besar (the latter
is home to the endemic L. natunae GÜNTHER, 1895)
west of Borneo.
23 mm) of megophryid frogs. Habitus moderately
stocky to slender with smooth or shagreened skin
on dorsum and venter and granular flanks. A small,
conical pectoral gland present medial to arm inser-
tion. Limbs slender, fingers and toes ending in
pointed disks; large inner palmar tubercle, long,
low inner metatarsal tubercle; nuptial pads absent.

88

Tab. 5 Distribution of Leptobrachella-species on Borneo.
All species, with the exception of L. palmata, have
rudimentary webbings confined to the bases of the
toes. Maxillary teeth present, vomerine teeth ab-
sent; tympanum distinct.
Larvae have a single filter cavity on either side; they
lack the usual frilled gill filter rows, and also have no
labial teeth. The head-body section is extremely elon-
gated. The larvae of all species may be very similar.
Preliminary key to the Bornean species
of Leptobrachella
(based on INGER & STUEBING 1991)
1 3rd and 5th toes webbed up to the terminal disks .
................................................................ palmata
• Toes only with rudimentary webbing ............... 2
2 Dark lateral stripe between eye and groin continu-
ous ..................................................... serasanae
• With or without small, isolated, dark lateral spots
.......................................................................... 3
3 Belly dark with many small, irregular, light spots; a
dark streak on the supratympanic fold ... baluensis
• Belly light, with or without a faint dark network;
no supratympanic streak ................................... 4
4 An aligned ventrolateral row of elongated, slightly
separated whitish glands between axilla and groin
.............................................................. mjobergi
• Ventrolateral glands round, isolated, not aligned
.......................................................................... 5
5 Adult males 15-17.5 mm SVL; no tarsal gland ..
.................................................................. parva
• Adult males 19-21 mm SVL; tarsal gland present
............................................................ brevicrus
Megophryidae
Adults dwell in leaf litter and appear on the banks
of small, rocky, clear streams during the breeding
season. Their larvae are macrophagous feeders
adapted to a life in well-oxygenated waters dwell-
ing interstices of gravel and rock crevices at the
bottom of riffles.
Six species are known from Borneo, two from Mt.
Kinabalu.
Leptobrachella baluensis SMITH, 1931
Leptobrachella baluensis SMITH, 1931: 12 – Type locality: Kambo-
rangah
Nesobia mjobergi, (not Leptobrachella mjöbergi SMITH, 1925)
INGER 1966: 49 (part), 1978: 312; MANTHEY 1983: 20 (part)
Leptobrachella baluensis, DRING 1983: 96; INGER & STUEBING
1992: 41-51; INGER & TAN 1996a: 552, 1996b: 72; LAKIM et
al. 1999: 33; INGER et al. 2000: 16
Leptobrachella mjobergi, (not Leptobrachella mjöbergi SMITH,
1925) HOFFMANN 1995a: 8
Diagnosis
A highland species with “chirp” call. Flank glands
scattered or fused in an irregular, short, ventrola-
teral series.
Description
A small terrestrial frog; SVL of males 15-19.6 mm,
females up to 23 mm; TL/SVL 0.500-0.527; tips
of fingers and toes slightly dilated, sharply
pointed; finger lengths 3>1>2=4; toe lengths
4>3>5>2>1; subarticular tubercles weak, but
present; dorsum smooth; flanks slightly granular
with small pale glands, the latter elongated in the
posterior portion of the lower flank, forming ven-
trolateral series; males with a median subgular
vocal sac.
89
 Megophryidae

Distribution
There are only a few records from Sabah
and Sarawak. Mt. Kinabalu: Sg. Lumo-
tok Kecil/Sayap (900 m); Kamborangoh
(2200 m).

Leptobrachella parva
DRING, 1983

Leptobrachella parva DRING , 1983: 89 – Type

locality: Gunung Mulu, Fourth Division, Sarawak
Leptobrachella parva, MALKMUS 1994b: 219

Diagnosis
A lowland species with a “buzz” call;
flank glands scattered and not in ventro-
lateral series like in L. baluensis.
© P. Hoffmann

Description
Very similar to L. baluensis; males 15-
Fig. 80 Male of Leptobrachella baluensis from Sg. Lumotok Kecil, 18 mm, a gravid female 17.8 mm; TL/SVL
Sayap (900 m). 0.497-0.548; in contrast to L. baluensis
there are no subarticular tubercles; no dark
tympanic mask or stripe on the flank, and
Brown above with a dark interorbital bar and a flanks not spotted with black, but with small dots
scapular W-shaped marking; a distinct, dark and black areas around white glands.
supratympanic spot or streak; tubercles on flanks
pale, dark outlined; limbs with dark crossbars; Ecology/Ethology
throat and belly greyish, with diffuse dark pigment. Calling males were found in hidden places on peb-
ble beaches and boulders at the edge of streams
Ecology/Ethology and on rocks protruding from the water in dipte-
The few records from Mt. Kinabalu so far all came rocarp forest. L. parva was observed on the leaves
from the levels between the oak-chestnut forest and of low bushes (30-50 cm) along Sg. Langanan.
the mossy forest (900-2200 m) where this species
inhabits the leaf litter layer and rock rubble near Call: (+22.5°C) The call is a buzz lasting 0.15-
streams. The males call in the late afternoon and at 0.16 s, consisting of 24-28 pulses. The pulse rate
night from between rocks or are perched on the increases during the call from an initial 50-67
leaves of shrubs or in low bush vegetation usually pulses per second to 300-330 pulses per second
in the vicinity of streams with strong currents. towards the end. Dominant frequency 11.2-12.8 kHz;
no fundamental frequency; frequency modulation
Call: (+22°C) Call duration 0.14-0.15 s; each call and intensity modulation are not evident (DRING
begins with a soft click, followed by a series of 14 1983a).
pulses (0.06 s) at a constant rate of 275-280 pulses
per second; dominant frequency 12-14 kHz, with a Distribution
strong fundamental below 2 kHz; frequency modu- Only a few records exist from Sabah and Sarawak
lation absent, but amplitude modulation present; from between 150 and 850 m a.s.l. Mt. Kinabalu:
call intervals 0.38-0.40 s (DRING 1983a). Sg. Langanan (850 m).

90

Genus Leptobrachium TSCHUDI, 1838
The species of this genus are widespread in South-
east Asia, ranging from southern China to the Phil-
ippines, the Greater Sunda Islands and Bali.
Its Bornean representatives are moderately large
to large stocky frogs with a head broader than, or
as wide as, the body; snout not projecting, tym-
panum visible. Maxillary teeth present, vomerine
teeth absent. Tips of fingers and toes rounded (ex-
cept in L. nigrops); inner palmar tubercle circu-
lar, not extending along the first metacarpal oval
or round, flat axillary glands and small femoral
glands; skin of body surface with a network of
low ridges delimitating closed cells. No dermal
palpebral projections. Males lack nuptial pads and
have a median subgular vocal sac. Females are
Tab. 6 Distribution of Leptobrachium-species on Borneo.
Preliminary key to the Bornean species
of Leptobrachium
1 Upper part of iris orange ................ hendricksoni
• Upper part of iris blackish ............................... 2
2 Tips of fingers sharply pointed; subarticular
tubercles present .................................... nigrops
• Tips of fingers rounded; no subarticular tubercles
........................................................................... 3
3 Lower side with white and black pattern; weakly
bluish ring around the eye ...................... abbotti
• Lower side with no such pattern; distinct white
ring around the eye .......................... montanum
• Morphologically indistinguishable from L. monta-
num, but with significantly different bioacoustics
....................................................... gunungense
Megophryidae
substantially larger than males, with relatively
shorter tibiae and narrower heads. Their eggs are
pigmented.
The tadpoles of Bornean Leptobrachium are mod-
erately large to large (70-90 mm) with a long, leaf-
shaped tail. Head-body portion robust, oral disk
ventral, subterminal. A continuous fringe of short
papillae surrounds both lips. Black beaks V-
shaped, coarsely serrated; at least four divided
rows of labial teeth on upper lip. Eyes dorsola-
teral, spiracle mid-lateral; anal tube dextral, united
to ventral fin. These larvae are macrophagous
feeders and occur in relatively calm reaches of
streams with weak to moderate currents.
Adults can be found among the forest floor leaf
litter of primary rain forests and on clearings.
Five species are known from Borneo, three of them
from Mt. Kinabalu.
Leptobrachium abbotti (COCHRAN, 1926)
Megophrys abbotti COCHRAN , 1926: 446 – Type locality:
Balikpapan
Leptobrachium hasselti, (not Leptobrachium hasseltii TSCHUDI,
1838) INGER 1966: 29 (part), 1962/63: 46, 1978: 312 (part);
MATSUI 1979: 306 (part); MANTHEY 1983: 20 (part)
Leptobrachium montanum, (not Leptobrachium montanum FI-
SCHER, 1885) INGER & STUEBING 1992: 41-51 (part)
Leptobrachium cf. montanum, (not Leptobrachium montanum
FISCHER, 1885) MALKMUS 1994b: 218 (part)
Leptobrachium abbotti, INGER, STUEBING & TAN 1995: 123,
1996: 362; MALKMUS 1996c: 278; INGER et al. 2000: 16
Diagnosis
A stocky, broad-headed frog with bulging eyes;
short, slender hind limbs; in contrast to L. monta-
num a distinct ventral pattern of black and white;
no white ring around the eye, but a small bluish arc
in the distal portion of the eye.
91
 Megophryidae

4(2.5/3), 5(1.5); inner

metatarsal tubercle low,
oval in shape; dorsum
smooth, often with a net-
work of low ridges; a
supratympanic fold be-
tween eye and axilla; un-
derside weakly rugose
or granular.
Upper side dark brown,
occasionally blackish,
with an indistinct, even
darker pattern between
the eyes and on the back;
upper sides of legs with
dark bars; ventral side of
body white, with large,
dark lead-grey to black
© R. Malkmus

spots between the chest

and the cloaca, which
become more prominent
Fig. 81 Leptobrachium abbotti from Sg. Kipungit I, 550 m. distally, peppered with
white dots; juveniles
with a brick-red supra-
Description tympanic fold, canthus rostralis, and lateral glan-
Males up to 75 mm, females up to 95 mm; TL/SVL dular protuberances.
0.33-0.39; head as wide as long; snout obtusely Tadpoles see “Genus”. Labial teeth rows formula
pointed or truncate, not projecting; canthus rostralis 6-7(2-6 or 2-7)/6(1-5); lateral inframarginal ridges
sharp; tympanum distinct, half the eye diameter; a on lower lip with up to 16 groups of small denticles
large, round inner palmar tubercle; toe lengths on either side; tail slightly convex; tail length
4<3<5<2<1, toes weakly webbed: 1(1), 2(2/1), 3(3/2), 1.5-1.9 times the head-body-length; total length
75-90 mm. Hatchling tadpoles light brown. They
darken gradually towards earth-brown with obscure
Fig. XLII Oral disc of Leptobrachium abbotti. dark spots on head, body and tail. Lower side of
body and caudal muscle with irregular reticulation
of thin black lines.

Ecology/Ethology
On Mt. Kinabalu, L. abbotti presents itself as an
inhabitant of the dipterocarp forests between 500
and 900 m a.s.l. Here, it occurs along streams and
rivers, as well as far away from these, on the leaf
litter layer of the forest floor. It is nocturnal and
feeds on large insects (Orthoptera, Blattaria,
moths), their larvae, and spiders. The males do not
form calling communities. The eggs are cream-col-
oured and black in the entire animal hemisphere.
R. Malkmus
The larvae live in stream microhabitats of the types

92
 Megophryidae

1c, d, e, and f, sometimes in 1a or b. Fig. XLIII Sonagram and oscillograms of Leptobrachium abbotti
They prefer calm reaches and can (Sg. Kipungit I 550 m; 8.00 p.m. +22°C)
be found exposed on rocks and
gravel or hidden among dead
leaves covering the bottom of
stream pools. They feed on dead
leaves or graze on algae which
overgrow rocks. When disturbed
they attempt to reach a shelter with
great speed.

Call: (+22°C) 1-3 loud, distinctly

pulsed notes (duration 190-250 ms,
of which the first note of a call is
the longest; each note consists of
20-30 pulses), emitted at intervals
ranging from a few seconds to
some minutes; dominant fre-
quency 1-2 kHz; pulse repetition
rate 130-170.

Distribution
This species is probably present in
all parts of Borneo. Mt. Kinabalu:
Sg. Kipungit (500-600 m), Sg.
Langanan (700-900 m), Sg. Kada-
maian (below Kiau).

Leptobrachium gunungense Tadpoles very similar to those of L. montanum;

MALKMUS, 1996
Leptobrachium gunungensis MALKMUS , 1996d: 297 – Type
locality: Sg. Carson (= Sg. Tibabar)
Leptobrachium (Leptobrachium) gunungense, DUBOIS & OHLER
1998: 23
labial teeth rows formula 6(2-6)/5(1-4, 1-5) or 6(1-6),
only 4-7 short lateral rows of denticles; total length
(st 37) 63.9 mm; lateral side system often distinctly
visible. In contrast to the larvae found in Sg.
Tibabar, those of Sg. Mesilau have blackish lip
margins.

Diagnosis
Morphologically very similar to L. montanum.
Description
Morphometric ratios almost identical to those in
L. montanum. Together with the latter, L. gunun-
gense forms a pair of species which can hardly be
differentiated morphologically, but which differ
substantially with regard to their bioacoustics and
their molecular genetics (KOSUCH, unpublished).
Ecology/Ethology
Leptobrachium gunungense inhabits the transitory
zone between the upper oak-chestnut forest and the
mossy forest (1750-2200 m a.s.l.) on Mt. Kinabalu.
Here it displays a predilection for the slopes of
creeks where it can sometimes be encountered in
large populations. The males form incessantly call-
ing choruses at the onset of dusk which reach their
greatest intensity between 6.00 and 8.00 p.m. and
are particularly noisy after rain showers. The males

93
 Megophryidae

© H. Sauer
Fig. 83 Tadpole of Leptobrachium gunungense from Sg.
Tibabar, 1800 m.
© R. Malkmus

call out of their hiding places (beneath leaf litter,

from rock cavities and holes in the ground) and are
Fig. 82 Leptobrachium gunungense from Sg. Tibabar, very difficult to locate.
1800 m.
Up to an altitude of 2000 m the tad-
poles of L. gunungense occur in large
Fig. XLIV Sonagram and oscillograms of Leptobrachium gunungense numbers in interstices of the gravel
(Sg. Tibabar, 1800 m; 7.00 p.m. +16°C) sediment and accumulations of leaf
litter in the side pools of streams (m:
1a-f), above 2000 m the population
density decreases rapidly. The wa-
ter temperatures at these altitudes av-
erage only 13-15°C.

Call: (+16°C) The relatively stifled,

nasal call consists of series of 4-16
individual notes (note duration 140-
160 ms; internote interval 240-300
ms; dominant frequency 1-1.4 kHz).
A note is composed of 15-16 pulses
(interpulse interval 3.6-11 ms) with
a distinctly longer inter-pulse inter-
val (16-20 ms) between the first 5
and the subsequent pulses.

Distribution
Leptobrachium gunungense is as yet
only known from the tributaries of
Sg. Kadamaian, Sg. Liwago, and Sg.
Mesilau at altitudes above 1750 m
a.s.l. With L. montanum, which oc-
curs below 1800 m, it shares only a
narrow zone of sympatry.

94
 Megophryidae

Leptobrachium montanum FISCHER, 1885

Leptobrachium montanum FISCHER, 1885: 44 – Type locality:
Pramassan-Alai Mountains, Southeast Borneo
Megalophrys hasselti, (not Leptobrachium hasseltii TSCHUDI,
1838) VAN KAMPEN 1923: 13
Megophrys hasselti, (not Leptobrachium hasseltii TSCHUDI,
1838) INGER 1966: 29 (part), 1978: 312 (part); MATSUI 1979:
306 (part); MANTHEY & DENZER 1982: 13; MANTHEY 1983:
20 (part); MALKMUS 1985: 8, 1987: 276, 1988c: 176
Leptobrachium cf. montanum, INGER & STUEBING 1992: 41-51;
MALKMUS 1994b: 218 (part)
Leptobrachium sp., MALKMUS 1989: 181
Leptobrachium montanum, MALKMUS 1991a: 27, 1992a: 102;
WONG 1994: 29-37; HOFFMANN 1995a: 7; INGER , STUEBING &
TAN 1995: 123, 1996: 362; INGER & TAN 1996b: 75; MALKMUS
1996b: 20-26, 1996c: 278, 1999: 13-19; LAKIM et al. 1999:
33; INGER et al. 2000: 16

Diagnosis

© R. Malkmus
Similar to L. abbotti, but ventral side not patterned
in black and white; a conspicuous white arc in the
upper portion of the eye; larger than 60 mm.

Description
Males up to 62.7 mm, females up to 64.4 mm; TL/SVL
0.33-0.43; habitus and morphological details are
very similar to those of L. abbotti.
Head and upper side of body whitish brown to
ochre, with a medial, highly variable, asymmet-
ric, occasionally light-bordered spotted pattern,
partly flanked by wavy bands; dark spots also in
the tympanic region and on the sides of the head.
If the dorsal side is chocolate-brown, the spotted

© P. Hoffmann
pattern may be hardly recognisable. Upper sides
of limbs with dark crossbars; lower side grey to
blackish, peppered with a multitude of white dots,
lightened to violet in the gular region; axillary
glands whitish. Juveniles show a brick-red supra-
tympanic fold.
Tadpoles similar to larvae of L. abbotti, but without
fine network of lines on lower side and caudal mus-
cle; labial teeth rows formula 6-7 (2-5, 6, 7)/5-6 (1-
4, 5) and 4-8 short inframarginal ridges of denticles
on lower lip; total length up to 73 mm (st 41).
© P. Hoffmann

From top to base: Leptobrachium montanum

Fig. 84 from the Headquarters, 1500 m.
Fig. 85 from the Headquarters, Sg. Silau-Silau, 1550 m.
Fig. 86 from Sayap, 1100 m.

95
 Megophryidae

From top to base: Tadpoles of Leptobrachium montanum

Fig. 87 ventral view.
Fig. 88 from Sg. Silau-Silau, 1550 m.
Fig. 89 from Sg. Liwago.

Although some reports exist on its occasional

diurnality (INGER 1966, MATSUI 1979) it is usually
© H. Sauer

found outside its hiding places (beneath leaf lit-

ter, rocks, dead wood)
only after the onset of
dusk, i.e. between 6.00
p.m. and 6.00 a.m. This
is also the time when its
call can be heard in the
vicinity of streams, but
also far away from these.
Short duets and vocal
trios may arise, but these
© A. Nöllert

never escalate into cho-

ruses like in L. gunun-
gense. L. montanum is
one of the few anuran
species which continue
calling to a considerable
extent even during pro-
longed periods of drought.
When disturbed, it either
attempts to flee with short,
low leaps, or closes its
eyes and crouches to the
ground with its limbs
drawn up to the body
(letisimulation) so that its
phytomimetic properties
render it almost invisible.
It swims by using the
hind limbs only while the
© R. Malkmus

arms are rigidly held for-

Ecology/Ethology
Leptobrachium montanum is one of the most com-
mon terrestrial frogs of Mt. Kinabalu (900-1750 m
a.s.l.) and is encountered on the trails of the
mountain forests almost regularly. It inhabits the
leaf litter layer, even far away from running water.
ward at a right angle. Its
cleansing behaviour in-
cludes numerous ele-
ments of that of Kalophrynus baluensis (see the
respective account).
The ova of gravid females are cream-coloured with
a small area of dark pigment on the animal pole.
The larvae of L. montanum are the most abundant
tadpoles in the forest streams of their distribution

96
 Megophryidae

area. They prefer calm reaches with Fig. XLV Sonagram and oscillograms of Leptobrachium montanum
moderate currents and are active at (Headquarters 1550 m; 7.00 p.m. +20 °C)
day and night. They often hide in
leaf drifts and gravel interstices (m:
1a-f).
Adults feed on grasshoppers, moths,
spiders, roaches, etc.

Call: (+18 to 20°C) 1-3 (4) loud,

very accentuated, pulsed notes (note
duration 125-160 ms; internote in-
terval 110-126 ms) which are emit-
ted at intervals of a few seconds up
to 5 minutes; dominant frequency
500-1100 Hz with harmonics up to
5000 Hz.

Distribution
This species is probably present in
all montane zones of Borneo. Mt.
Kinabalu: Sayap (Sg. Kemantis; Sg.
Minodtuhan); Marei Parei; Bundu
Tuhan; entire Headquarters region up
to 1750 m altitude; East Ridge above
Poring (1000 m).

shape. Small eyes, dorsolateral to dorsal, not vis-

Genus Leptolalax DUBOIS, 1980
This genus is known from southern China to Pe-
ninsular Malaysia and Borneo.
Its Bornean species are small to moderately large,
slender frogs (SVL > 25 mm). Width of head and
body subequal; legs slender and relatively long.
Vomerine teeth absent, tympanum visible. Tips of
digits rounded, not swollen. Webbings between fin-
gers absent, those between toes are limited to the
bases; large inner palmar tubercle not extending
onto first metacarpal; inner metatarsal tubercle low,
no outer one; subarticular tubercles on fingers and
toes replaced by long, low, callous tissue. Males
withmedial subgular vocal sac, no nuptial pads.
Females larger than males, with relatively shorter
tibiae and narrower heads.
The tadpoles of Leptolalax are adapted to a life in
streams, and have an elongated, slightly depressed
ible from below; oral disk ventral and subterminal,
with cup-like expanded lips, at least one lip with
short tooth rows; inframarginal ridges of lower lips
lack small denticles. Heavy beaks, serrated, and
fully pigmented. Spiracle is sinistrally, mid-lateral.
Tail lanceolate, 1.7 to 2.2 times the head-body-
length. Tail with strong muscles and low fins. To-
tal length of larvae 60-65 mm. They inhabit clear
streams in reaches with moderate to swift currents
and occur syntopically with the larvae of Leptobra-
chella in the interstices of gravel beds.
Adults can be found on the floors of primary and
secondary forests where they prefer zones near
streams with dense vegetation. The males call at
night from branches of bushes flanking streams.
Their calls consist of series of pulsating, high-
pitched notes.
Six species are known to inhabit Borneo, four oc-
cur on Mt. Kinabalu.

97
Megophryidae

Key to the Bornean species of Leptolalax

1 Superficial pectoral gland present; uniformly dark 4 Dorsum coarsely shagreened ................... arayai
brown on the lower side of the head ..... maurus • Dorsum smooth ................................................. 5
• No superficial pectoral gland; lower side of the 5 Dark, round markings on the snout ........ hamidi
head not dark brown ......................................... 2
• No dark markings on the snout ................ pictus
2 Ventrum spotted ................................................ 3
• Ventrum not spotted .......................................... 4
3 Forelimb bicoloured, elbow cream-coloured with-
out dark marking, contrasting with the dark lower
arm .......................................................... gracilis
• Forelimb not bicoloured, elbow with the same dark
markings as the lower arm ....................... dringi

Tab. 7 Distribution of Leptolalax-species on Borneo.

Leptolalax arayai MATSUI, 1997 Diagnosis

Leptolalax arayai M ATSUI , 1997: 163 – Type locality:
Headquarters, Kinabalu Park, Sabah
Leptobrachium gracile, (not Leptobrachium gracile GÜNTHER,
1872) BOULENGER 1887: 95; MOCQUARD 1890: 163
Megalophrys gracilis, (not Leptobrachium gracile GÜNTHER,
1872) BOULENGER 1908: 421 (part); VAN KAMPEN 1923: 15 (part)
Megophrys gracilis, (not Leptobrachium gracile GÜNTHER, 1872)
SMITH 1931: 12
Leptobrachium gracilis, (not Leptobrachium gracile GÜNTHER,
1872) INGER 1966: 24 (part), 1978: 312 (part); MATSUI 1979:
305; MANTHEY 1983: 20 (part)
Leptobrachella baluensis, (not Leptobrachella baluensis SMITH,
1931) MALKMUS 1989: 182, 1991a: 27
Leptobrachella cf. baluensis, (not Leptobrachella baluensis
SMITH, 1931) MALKMUS 1992a: 103
Leptolalax gracilis, (not Leptobrachium gracile GÜNTHER, 1872)
INGER & STUEBING 1992: 41-51 (part); MALKMUS & RIEDE 1993:
7; MALKMUS 1994b: 219; WONG 1994: 29-37; INGER, STUEBING
& TAN 1996: 362
Leptolalax sp., MALKMUS 1996c: 279
Leptolalax sp. “a”, MALKMUS 1996b: 20-26; MANTHEY &
GROSSMANN 1997: 75, fig. 41
Leptolalax arayai, MALKMUS & KOSUCH 1999: 161; MALKMUS
1999: 13-19; MALKMUS 2000: 12
A medium-sized form of the genus Leptolalax;
dorsum roughly shagreened; chest and abdomen
without spots; among the Bornean species it is the
one with the largest feet (> 50% SVL).
Description
Males up to 36 mm, females up to 38 mm; TL/SVL
0.60-0.62; head slightly longer than wide; snout
rounded, projecting beyond lower jaw; eye diam-
eter greater than snout length; lores slightly con-
cave; tympanum distinct, half the eye diameter; fin-
ger lengths 3>4=1=2; tibiotarsal articulation of
adpressed limb reaches far beyond the tip of the
snout; toe lengths 4>3>5>2>1; dorsum shagreened,
flanks with round tubercles; venter smooth; a low
supratympanic fold.
Upper sides light to dark brown, often with dark
spots of various sizes; narrow, dark brown cross-
bars on upper sides of hind limbs and lower arms;
flanks light brown, distal third and inner sides of
thighs orange red; tubercles completely or partly

98

© R. Malkmus
© R. Malkmus
© A. Nöllert
From top to base: Leptolalax arayai
Fig. 90 female from Sg. Silau-Silau, 1550 m, and
Fig. 91 male. – Fig. 92 male from Sg. Liwago.
framed with black; lower sides whitish, obscurely
spotted with grey on the belly; anterior portion of
arc of lower jaw blackish with light dots; iris whit-
ish grey to light greyish green, with a fine network
of black veins; one segment (ca. 100-120°) in up-
per portion of iris bright reddish brown.
Tadpoles’ shape as described under “Genus”; both
lips notched in centres, lower one more deeply so;
Megophryidae
R. Malkmus
© M. Schroth
From top to base: Tadpole of Leptolalax arayai
Fig. XLVI oral disc.
Fig. 93 from Sg. Liwago.
both lips covered with large, rounded papillae; labial
teeth reduced to two small rows in centre of upper
lip; lower lip without labial teeth; beaks heavy, ser-
rated, black; tail 1.8-2.0 times head-body-length; to-
tal length (st 28) 61 mm; head and body whitish blue
with a metallic shine above and on sides; undersides
lighter, not pigmented; caudal muscle whitish blue
with small white dots; fins dusky, not transparent.
99
Megophryidae

Ecology/Ethology
Leptolalax arayai is a very charac-
teristic and common faunal element
of the banks of streams crossing
through montane oak-chestnut for-
ests on Mt. Kinabalu occurring at al-
titudes between 1400 and 1750 m
a.s.l., occasionally up to 1900 m. It
finds shelter beneath leaf litter and
bark, in holes in the ground, and
cavities in dead wood. During pro-
longed periods of drought it often
hides in the immediate vicinity of
streams under rocks and wood on
gravel banks, and leaves these only
at the onset of dusk. Advertisement
calls can be heard between 6.00 p.m.
and 5.30 a.m. (occasionally until
6.00 a.m.). The males call from the
leaves and branches of shrubs and
bushes (20 to 200 cm above the
ground) on the banks of streams,
very rarely also from boulders.
The larvae are restricted to the in- Fig. XLVII Sonagram and oscillograms of Leptolalax arayai
terstices of the gravel bottoms of (Sg. Silau-Silau, 1550 m; 7.00 p.m. +20°C)
forest streams (m: 1b, c, e).
L. arayai is an “sit-and-wait”
hunter which feeds primarily on spiders, moths, Leptolalax dringi DUBOIS, 1987
and hairless caterpillars. A female which was star-
tled in its hiding place feigned death with its body Leptolalax dringi DUBOIS, 1987 (1986): 13 – Type locality:
adpressed to the ground for 8 seconds, only to take Mount Mulu, Sarawak
to flight suddenly and escape with 50-80 cm long Leptolalax dringi, INGER, STUEBING & TAN 1995: 119, 1996:
362; LAKIM et al. 1999: 32; INGER et al. 2000: 16
leaps.
Diagnosis
Call: (+18 to 20°C) The 2-15 second long call con- A moderately stocky, large-eyed species with slen-
sist of incessantly repeated, shrill notes. Each note der hind limbs; skin with round tubercles above
(duration 16-29 ms) is composed of a double pulse and on the sides, most concentrated on the snout;
(3.3 and 4 ms, respectively; interpulse interval 8- chest and belly with small black spots.
9 ms) which is repeated at intervals of 100-115 ms;
dominant frequency 5.7-5.9 kHz; frequency modu- Description
lation weak; note repetition rate 9.0-9.3 per sec- Males up to 35 mm, females up to 48 mm; TL/SVL
ond. 0.48-0.56; head width equal to body width behind
the arms; snout obtusely pointed, rounded in pro-
Distribution file, not projecting beyond lower jaw; canthus
Leptolalax arayai is as yet known only from Mt. rostralis rounded, lores concave; eye diameter
Kinabalu where it has been recorded from along greater than distance between eye and snout; tym-
Sg. Liwago and its tributaries in the Headquarters panum distinct, half the eye diameter; finger tips
region. rounded; finger lengths 3>4=2=1; palmar and

100
 Megophryidae

© C.L. Chan
Fig. 94 Leptolalax dringi from Mulu NP, Sarawak.

subarticular tubercles indistinct; toe Fig. XLVIII Sonagram and oscillograms of Leptolalax dringi
lengths 4>3>5>2>1; no webbing; sub- (Gunung Mulu, 1800 m; 1.00 a.m.)
articular tubercles elongated, obscure;
inner metatarsal tubercle oval, no outer
one; dorsal side with round tubercles,
larger on the sides; venter smooth.
Dorsum and head greyish to reddish
brown, usually with black markings; hind
limbs and lower arms brown with dark
crossbars; elbow often lighter; flanks
grey with light tubercles; chest and belly
whitish with dark spots.

Ecology/Ethology
Leptolalax dringi lives in primary and old
secondary forests. Nothing else is known
about its life history.

Call: The call consists of series (dura-

tion 1.8-3.5 s) of 8-10 pulsed notes (du-
ration 35-40 ms; note interval 60 ms; note
repetition rate 11.0-11.5 per second);

101
 Megophryidae
frequency modulation distinct; dominant frequency
7-7.5 kHz with a fundamental below 0.5 kHz.
Distribution
Sarawak and Sabah. Mt. Kinabalu: Records have
become known from Poring (600 m), Sayap and
“Kinabalu”, with the latter referring to a specimen
in the British Museum said to be collected at 1800 m
altitude.
Leptolalax maurus
INGER, LAKIM, BIUN & YAMBUN, 1997
Leptolalax maurus INGER, LAKIM, BIUN & YAMBUN, 1997: 48 –
Type locality: Mesilau Station, 1860 m, Kinabalu Park,
Sabah
Leptolalax maurus, INGER & STUEBING 1997: 59; LAKIM et al.
1999: 33; MALKMUS 2000: 13; INGER et al. 2000: 16
Fig. 95 Leptolalax maurus from western Sg. Mesilau, 1800 m.
© M. Schroth
102
Diagnosis
A small, moderately stocky frog (< 35 mm); entire
upper side of the body dark; belly with a network
of thin lines; a superficial, round pectoral gland on
either side; back and sides covered with homoge-
neous, round tubercles.
Description
Males up to 26.1 mm, females up to 31.8 mm;
TL/SVL 0.50; widths of head and body subequal;
snout obtusely pointed, rounded in profile; eye di-
ameter slightly greater than length of snout; lores
concave; tympanum more than half the eye diam-
eter; tips of fingers and toes rounded; finger lengths
3>4=2=1; toe lengths 4>3>5>2>1; back and flanks
covered with round tubercles, smaller ones on limbs;
ventral surface smooth; pectoral glands visible; a dis-
tinct, angular supratympanic fold is present.
Upper sides uniformly black, peppered with tiny
white dots which concentrate on the flanks; hind
 Megophryidae

Leptolalax pictus MALKMUS, 1992

Leptolalax pictus MALKMUS, 1992b: 3 – Type locality: Sg.
Langanan, Kinabalu, Sabah
Leptolalax gracilis, (not Leptobrachium gracile GÜNTHER, 1872)
MALKMUS 1992a: 104; INGER, S TUEBING & TAN 1995: 122
(part)
Leptolalax pictus, MALKMUS 1992b: 3; MALKMUS & RIEDE 1993:
7; MALKMUS 1994b: 2; HOFFMANN 1995a: 7; INGER, STUEBING
& TAN 1996: 362; INGER & TAN 1996a: 553; MALKMUS 1996b:
20-26; MATSUI 1997: 164; LAKIM et al. 1999: 32; MALKMUS
2000: 14; INGER et al. 2000: 16

Diagnosis
A slender, large species of Leptolalax with a dis-
tinctive dorsal and lateral spotted pattern; skin
smooth; lower side not spotted.
© M. Schroth

Fig. 96 Ventral view of Leptolalax maurus from west-
ern Sg. Mesilau, 1800 m.
Description
Males up to 36 mm, females up to 47 mm; TL/SVL
0.55-0.61; head slightly longer than wide; snout
rounded, projecting beyond lower jaw; lores slightly
concave; tympanum distinct, half the eye diameter;

limbs slightly lighter above, with four black cross- Fig. 97 Dorsal pattern of Leptolalax pictus from Sg.
bars between which are clusters of white dots; fore Kipungit II.
limbs blackish brown, upper side of the upper arm
light brown; lower sides of the legs, and throat, chest,
and belly uniformly blackish brown; belly with an
irregular, fine, white network; iris reddish brown.

Ecology/Ethology
Leptolalax maurus has as yet been found only at a
few sites in oak-chestnut forests (1860-1900 m
a.s.l.) and always among leaf litter. During an ex-
tended dry period (January through March 1998),
a male was discovered around noon (1.00 p.m.)
under a rock flake lying across a spring stream
(western Sg. Mesilau).
The eggs of L. maurus have the greatest diameter
of all known species of Leptolalax. Increased egg
sizes are, however, also typical for the high-
montane species of other genera (Philautus,
Pelophryne, Ansonia).

Distribution
© P. Hoffmann

This species has as yet been found only in envi-

rons of the type locality, the region of eastern and
western Sg. Mesilau.

103
 Megophryidae
© R. Malkmus
Fig. 98 Female of Leptolalax pictus from Sg. Kipungit II.
finger lengths 3>4=1=2; hind limbs long, tibiotarsal
articulation of adpressed limb reaching beyond tip of
snout; toe lengths 4>3>5>2>1; dorsum and venter
smooth; small tubercles on flanks and eyelids.
Fig. 99 Calling male of Leptolalax pictus from Sg.
Langanan.
© H. Sauer
104
Dorsal side light brown with dark brown, irregu-
larly scattered spots of highly variable size and
shape which themselves are bordered by fine, white,
partly interrupted outlines; an equally coloured,
broad orbital band; flanks with numerous small to
medium-sized, dark brown, white-bordered spots;
4-5 large and numerous small brown spots on the
sides of the head between tympanum and tip of
snout; upper side of lower arm and thigh with dark
crossbars; ventral side of body white, with a dark
tinge distally; iris greyish golden, bilaterally with
segments of 10-20% black and a reddish brown seg-
ment of about 120° in upper half; entire iris overlain
with black reticulation that appears fragmented.
Ecology/Ethology
Leptolalax pictus is a common appearance along
streams in the upper dipterocarp forest and oak-
chestnut forest (500-1100 m a.s.l.) on Mt. Kina-
balu. It activity patterns are identical with those of
 Megophryidae

L. arayai. The males call from shrubs and leaves Fig. L Sonagram and oscillograms of Leptolalax pictus
and twigs of bushes in the vicinity of streams, oc- (Sg. Kipungit, I 600 m; 7.00 p.m. +20°C)
casionally from directly above the water.

Call: (+19 to 22°C) The call is shorter (1.9-4.9 s)

than in L. arayai, use a higher frequency band (6.8-
7.2 kHz), and are more often paused (duration 4-5 s).
Each note (duration 30-35 ms; internote interval
45 ms) is marked by an initial frequency drop from
7.2 to 6.8 kHz. The note repetition rate is 11-13 per
second.

Distribution
Leptolalax pictus has as yet become known only from
North Borneo. Mt. Kinabalu: Poring: Sg. Kipungit,
Sg. Mamut, Sg. Langanan; Sayap: Sg. Kemantis.

Genus Megophrys
KUHL & VAN HASSELT, 1822
Fig. IL Distribution of
The genus Megophrys is native to continental Asia,
Leptolalax pictus in the
Poring area. Sumatra, Borneo, Java, and parts of the Philippines.
The Bornean species are medium-sized to large frogs
with relatively short, slender hind limbs and large,
wide heads. Most species have characteristic dermal
palpebral projections (particularly above the eyes and
on the snout) and a sharply curved ridge of skin be-
tween the eye and the arm. Tympanum is visible, but
often indistinct. Adults possess a conical axillary gland
at the insertion of the arm. Tips of fingers and toes
rounded, outer palmar tubercel and subarticular tu-
bercles absent or indistinct; inner palmar tubercle oval,
extending along first metacarpal. Webbing reduced
to basal rudiments. Males possess vocal sacs and nup-
tial pads. The larvae are slender and have funnel-
mouths; beaks soft, serrated, lacking labial teeth, but
with many short, low ridges; oral disk terminal, lips
expanded into a horizontal, dorsally-oriented funnel;
spiracle sinistral; anal tube medial, fused to the ven-
tral fin or not. The tadpoles dwell streams in sections

105
Megophryidae

with moderate currents. They are adapted to feeding species present on Mt. Kinabalu form calling cho-
on plankton in the surface film. ruses.
Adults are terrestrial and can be found in the layer of Five species are known from Borneo, four from Mt.
leaf litter in primary forests. The males of all three Kinabalu.

Tab. 8 Distribution of Megophrys-species on Borneo.

Key to the Bornean species of Megophrys
1 Head distinctly narrower than body ......... dringi
• Head a little narrower or as wide as body ........ 2
2 Two continuous dermal lateral or dorsolateral folds;
dermal rostral appendage present ............ nasuta
• Dermal folds not continuous; no rostral appendage
............................................................................ 3
3 Large frog, SVL > 90 mm; angle of jaw and end
of supratympanic fold with large projections .....
.......................................................... kobayashii
• Never larger than 70 mm; no such projections or
only small ones ................................................. 4
4 Vomerine teeth present ....................... baluensis
• Vomerine teeth absent; no dorsolateral folds ......
........................................................ edwardinae
Diagnosis
A moderately large, stout terrestrial frog. It is dis-
tinctly smaller (always < 70 mm) than M. koba-
yashii and M. nasuta. Snout very short and with-
out rostral appendage; dorsolateral, or lateral, glan-
dular lumps grouped in rows, but not forming con-
tinuous folds.
Description
Males up to 45 mm, females up to 70 mm; TL/SVL
in females 0.36-0.38, in males 0.40-0.46; head
Fig. 100 Semiadult Megophrys baluensis from Sg.
Tibabar, 1700 m.

Megophrys baluensis (BOULENGER, 1899)

Leptobrachium baluense B OULENGER , 1899: 453 – Type
locality: Mount Kina Balu
Leptobrachium baluense, HANITSCH 1900a: 74
Megalophrys baluensis, BOULENGER 1908: 429; VAN KAMPEN
1923: 16
Megophrys baluensis, SMITH 1931: 30; INGER 1954a: 250, 1966:
46, 1978: 312; MATSUI 1979: 310; MANTHEY 1983: 20; MALKMUS
1985: 9, 1987: 277, 1988c: 176, 1989: 181; INGER 1989: 229;
INGER & STUEBING 1989: 41; MALKMUS 1991: 27; INGER &
STUEBING 1992: 41-51; MALKMUS 1994b: 220, 1994c: 86, 1995c:
© R. Malkmus

8, 1996b: 20-26, 1996c: 280; INGER, STUEBING & TAN 1996: 362;
INGER & TAN 1996a: 553; MALKMUS & RIEDE 1996c: 152; LAKIM
et al. (1999): 32; MALKMUS 1999: 13-19; INGER et al. 2000: 16
Megophrys cf. baluensis, MALKMUS 1992a: 104

106

Fig. 101 Male of Megophrys baluensis from the Headquarters.
wider than long; very short snout, vertical in pro-
file, without dermal appendages; short, pointed
dermal appendages above the eyes, followed by
conical tubercles; tips of phalanges rounded; no
webbings between fingers; occasionally rudimen-
tary webbings between toes; finger lengths
3>2=4>1; inner metatarsal tubercles low, oval, no
outer one; subarticular tubercle indistinct; toe
lengths 4>3>4>2>1; tympanum usually clearly
visible, smaller than half the eye diameter; upper
surface covered with irregularly scattered, usually
elongate-oval glandular rises which form rows,
and partly ridges, dorsolaterally and laterally; a
distinct, angled supratympanic fold between eye
and axilla; males with nuptial pads on 1st and 2nd
fingers.
Megophryidae
© A. Nöllert
Coloration and pattern highly variable and perfectly
adapted to pattern and coloration of leaf litter; up-
per sides dark brown, reddish brown to ochre,
lighter laterally and on limbs; lower side yellow-
ish brown to dark flesh-red, with dark brown cloud-
ing; throat with dark or light median stripe; a trian-
gular, blackish brown marking between occiput and
mid-dorsum; between eye and margin of upper jaw
alternating yellowish white and dark brown verti-
cal bars; iris golden brown with black reticulation.
A 3 cm long semi-adult specimen was light loam-
yellow with white ventrum, clouded with brick-red
towards chest.
Tadpoles similar to larvae of M. nasuta. Nostrils
elliptical, rim low, with very flat mid-dorsal pro-
jection; total length (st 31) 36.7 mm; tail length
107
Megophryidae
R. Malkmus
Fig. L Oral funnel of Megophrys baluensis.
tail length 2.09-2.14 times head-body-length; tail
tapering to pointed tip; body above and on sides
dark brown with large loam-coloured spots; belly
greyish brown, without pattern; caudal muscle
brown with light spots, particularly on upper half;
both beaks unpigmented with blackish band in the
centre; fins dusky, with isolated dark flecks.
Ecology/Ethology
On Mt. Kinabalu, M. baluensis occurs in the vicin-
ity of streams cutting through oak-chestnut and
Fig. LI Sonagram and oscillograms of Megophrys baluensis
(Sg. Lanaga, 1500 m; 7.00 p.m. +19°C)
108
lower mossy forests (1280-1950 m a.s.l.). It prefers
the banks of small running waters, particularly fields
of rock rubble which are densely overgrown by
shrubs (Begonia, Gunnera, Impatiens). Here, the
frogs live in very stationary calling communities.
Five of those colonies could be traced always in the
same spots on tributaries of Sg. Liwago between
1984 and 1998. Choruses can be heard between dawn
and dusk with maximum intensities between 5.30
and 6.00 a.m. and 6.00 and 7.00 p.m., respectively.
These are started by one male beginning to call, trig-
gering others to respond, and so building up to a
chorus which then lasts for 30-100 seconds. The
frogs call from rock crevices, cavities among roots,
and holes in the ground. They leave their shelters at
night and forage on the forest floor, occasionally
climbing on low bushes up to about 30 cm above
the ground. Their tadpoles dwell mountain brooks
with weak current (m: 1d, e).
M. baluensis shows ritualised cleansing motions,
otherwise known from species of the genera Lepto-
brachium and Kalophrynus. Captured specimens
show a pulsating trembling shortly before they at-
tempt to flee.
Call: (+17 to 19°C) The calls are emitted
in series of 3-8 loud, metallic, sharp, and
hard strikes at intervals of 550-700 ms.
They consist, especially in their second
third, of distinctly differentiated pulses
and last 90-100 ms; carrier frequency
2.8 kHz.
Distribution
Until recently M. baluensis used to be
listed for the fauna of Sarawak, Sabah
(INGER 1966, INGER et al. 1985, INGER
& STUEBING 1992) and Brunei (DAS
1995).The latest check list of the frogs
of Borneo (INGER & TAN 1996a), how-
ever, lists this species only for Sabah,
and INGER & STUEBING (1997) consider
it to be endemic for Mount Kinabalu,
stating: “M. baluensis is so far known
only from Mount Kinabalu”. Mt. Kina-
balu: Tributaries of Sg. Liwago (up to
1950 m on Sg. Tibabar); Bundu Tuhan;
eastern Sg. Mesilau (1800 m).

Fig. LII Distribution of Megophrys baluensis and M. kobayashii in the area of the Headquarters.
Megophrys edwardinae INGER, 1989
Megophrys edwardinae INGER, 1989: 229 – Type locality:
Nanga Tekalit, Kapit District, Sarawak
Megophrys edwardinae, INGER, S TUEBING & TAN 1996: 362;
INGER et al. 2000: 16
Diagnosis
A medium-sized Megophrys; dorsum with scat-
tered, round and elongate tubercles; a “horn” on
the rough-surfaced upper eyelid; no continuous dor-
solateral folds.
Description
Males up to 42 mm, females up to 82 mm; TL/SVL
0.43-0.50; head wider than long; snout obtusely
pointed, rounded in profile; canthus rostralis sharp,
curved; lores oblique, concave; eye diameter equal
to eye-snout distance; tympanum obscure; vomer-
ine teeth absent; fingertips rounded; palmar tuber-
cles weak; finger lengths 3>1>2=4; tips of toes
rounded; rudimentary webbings on bases of toes;
toe lengths 4>3>5>2>1; subarticular tubercles ob-
scure; inner metatarsal tubercle low, no outer one;
Megophryidae
skin on dorsum and head with many irregular tu-
bercles and short ridges; a large conical tubercle at
rictus; upper eyelid with tubercles and ridges, cen-
tral margin of eyelid with triangular projection;
sides of the head with ridges radiating from coni-
cal tubercles; a strong supratympanic fold present;
throat rugose; teat-like projection on chest near
axilla; belly smooth.
Brown above and on sides; an orange patch on shoul-
der; two dark bars between eye and upper lip; sides
light brown with irregular dark markings; ventrum
pale brown with dark spots; lower lip barred; dorsal
surfaces of limbs with dark crossbars.
Ecology/Ethology
This species lives on the leaf-littered floor of pri-
mary forests. Nearly nothing is known about its
life history. It was found in lowland and hilly re-
gions at altitudes between 200 and 600 m a.s.l.
Distribution
Megophrys edwardinae is known from Sarawak and
Sabah. Mt. Kinabalu: Only known from the Po-
ring region.
109
 Megophryidae
Megophrys kobayashii
MALKMUS & MATSUI, 1997
Megophrys kobayashii MALKMUS & MATSUI, 1997: 31 – Type
locality: Headquarters, Mount Kinabalu, Sabah
Megophrys monticola, (not Megophrys montana KUHL & VAN
HASSELT, 1822) SMITH 1931: 12
Megophrys monticola nasuta, (not Ceratophryne nasuta SCHLE-
GEL, 1858) INGER 1966: 41 (part), 1978: 312 (part); MATSUI
1979: 310; MANTHEY 1983: 20 (part)
Megophrys nasuta, (not Ceratophryne nasuta SCHLEGEL, 1858)
INGER & STUEBING 1992: 41-51 (part); INGER, STUEBING & TAN
1996: 362 (part)
Megophrys sp., MALKMUS 1995c: 7, 1996b: 20-26, 1996c: 279
Megophrys kobayashii, MALKMUS 1999: 13-19
Diagnosis
A large form of Megophrys; in contrast to M. nasuta
with no long continuous dorsolateral folds; snout
rounded without rostral appendage. M. kobayashii
is most similar to M. ligayae from the Philippines.
Fig. 102 Megophrys kobayashii from Sg. Silau-Silau, 1500 m.
© R. Malkmus
110
Description
Females up to 109 mm; TL/SVL 0.45-0.49; head
and body stout, head wider than long; snout rounded
in profile, projecting beyond lower jaw; no rostral
appendage; snout somewhat shorter in length than
eye diameter; upper eyelid drawn out into a triangu-
lar projection; canthus rostralis sharp, lores concave;
tympanum visible; supratympanic fold distinct;
vomerine teeth present; tips of fingers rounded; fin-
ger lengths 3>1>2=4; tips of toes blunt; toe lengths
4>3>5>2>1; inner metatarsal tubercle ellipsoid; ti-
biotarsal articulation reaching up to the eye when
leg is extended; very short pairs of dorsolateral lon-
gitudinal folds; skin of head fused to skull; skin be-
hind head calcified; eyelid with conical tubercles;
lateral and ventral skin rugose or coarsely granular;
sides with irregularly scattered, large, round and
elongated tubercles; angle of jaws and end of
supratympanic fold with irregular projections; arms
and hind limbs with tubercles and short folds.

© A. Nöllert
Fig. 103 Megophrys kobayashii from Silau-Silau trail,
near the Headquarters.
Dorsum light brown; two conical, black tubercles
on head; large dark blotch below canthus rostralis
and behind eye; lower lip dark brown, with narrow
white bars; very large brown, white-bordered mark-
ing between posterior lower lip and upper arm;
lower arms with large brown spots; hind limbs light
brown with darker fields around ridges; large dark
Fig. LIII Tadpole of Megophrys kobayashii.
R. Malkmus
Megophryidae
© M. Schroth
Fig. 104 Juvenile of Megophrys kobayashii from Sg.
Silau-Silau/Sg. Liwago, 1450 m.
spot on knee; lateral tubercles and ridges light
brown with irregular dark spots; throat brown,
medially with darker spots; abdomen light brown
with two parallel rows of large dark spots.
Tadpoles similar to larvae of M. baluensis and M.
nasuta; rim of nostril mid-dorsally with pointed
projection; total length (st 37) 41.8 mm; tail length
2.1 times head-body-length; tail tip rounded; body
and caudal muscle darker in coloration than in M.
baluensis and M. nasuta; with fewer lighter flecks;
a light vertical bar on lower half of tail muscle near
its root; fins dusky, marbled with brown; centre of
upper lip, margin of oral disk, the region lateral to
beaks, and lateral tips of funnel black.
Ecology/Ethology
M. kobayashii dwells the layer of leaf litter on the
floor of lower montane rain forests (1450-1675 m
a.s.l.), preferring the neighbourhood of small
streams, and hiding in holes in the ground, cavities
among and in roots and among rocks. Its calling
activity is limited to short periods of time in the
early morning (6.00 to 6.30 a.m.) and in the late
afternoon (6.00 to 6.30 p.m.), but calls can also be
heard during rain showers (between 3.00 and 6.00
p.m.) when the daylight fades. It is usually one
specimen which begins to call, triggering others to
respond, and a full chorus is in place within 3-5
seconds. This species lives in permanent colonies.
Larvae were found in small mountain brooks with
weak current (m: 1d,e). M. kobayashii appears to
be rare.
111
Megophryidae

Call: (+21°C) a very powerful

voice; call sequences last be-
tween 3 and 5 seconds and con-
sist of 8-13 notes with distinct
pulse modulation; note duration
varies from 135 to 188 ms (mean
163 ms); internote interval 134-
182 ms (mean 150 ms); carrier
frequency 1.3-1.9 kHz.

Distribution
M. kobayashii is known only
from the Headquarters region on
Mt. Kinabalu, i.e. Sg. Silau-
Silau, Sg. Lanaga; Lumu Lumu Fig. LIV Sonagram and oscillogram of Megophrys kobayashi
(see also map on page 109). (Headquarters, 1550 m; 6.30 p.m. +21°C)

Megophrys nasuta (SCHLEGEL, 1858)
Ceratophryne nasuta SCHLEGEL, 1858: 75 – Type locality: Su-
matra
Megalophrys nasuta, MOCQUARD 1890: 163; HANITSCH 1900a:
74; VAN KAMPEN 1923: 10
Megophrys monticola nasuta, INGER 1954a: 250, 1956: 393,
1962/63: 46, 1966: 41 (part). 1978: 310 (part); MANTHEY
1983: 20 (part); MALKMUS 1985: 9, 1987: 277, 1988c: 176
Megophrys nasuta, SMITH 1931: 9; INGER 1985: 13; INGER &
STUEBING 1992: 41-51 (part); MALKMUS 1992a: 104; WONG
1994: 29-37; HOFFMANN 1995a: 8; MALKMUS 1995c: 6, 1996c:
279; INGER , STUEBING & TAN 1996: 362 (part); LAKIM et al.
1999: 33; INGER et al. 2000: 16
Diagnosis
A large Megophrys; dermal rostral appendage
present, and upper eyelid drawn out into a triangu-
lar projection; two pairs of parallel, longitudinal,
dorsolateral folds continuous between head and
groin.
granular; 2 pairs of narrow dorsolateral folds; trian-
gular dermal appendages on eyelids and rostrum are
smaller in specimens from Mt. Kinabalu than in those
from other parts of Borneo (INGER 1966).
Specimens from Mt. Kinabalu are light to dark red-
dish brown above, sometimes with a dark-clouded
central portion; a blackish brown, wide, V-shaped
marking between eyes and central occiput; black
bar behind and below eye; dark crossbars on upper
side of limbs; black spot on knee; throat brownish,
grey to reddish, partly with small white spots; belly
marbled light brown on a dirty whitish ground,
lighter distally; iris blackish brown.
The structure of the skin, its body shape, and the
colour pattern of this frog are an example for
Fig. 105 Tadpole of Megophrys nasuta from Sg. Keman-
tis, Sayap.

Description
Males up to 105 mm, females up to 125 mm;
TL/SVL 0.34-0.46. A stocky frog with short, slen-
der hind limbs; head wider than long; tympanum
present, but usually hidden by skin; vomerine teeth
present or absent; shapes and relative lengths of fin-
© M. Schroth

gers and toes identical to those of M. kobayashii; dor-

sum smooth with 1-4 large, conical tubercles of black
coloration; sides and ventrum rugose to coarsely

112
 Megophryidae

© M. Schroth
Fig. 106 Megophrys nasuta from Sayap.

phytomimesis (dead leaves on the forest floor) de- rubble. The frogs spend the day hidden in cavi-
veloped to perfection. ties among the rocks, beneath dead wood, and in
Tadpole: Head-body of tadpoles elongate, almost sub-terrestrial tunnels, and emerge with the onset
circular in cross section; eyes laterally, oral disk of dusk in order to forage. Their huge mouths en-
terminal; lips expanded to form a horizontal fun-
nel; open funnel as wide as body; lips without la-
bial teeth, but with many short, low ridges; beaks Fig. 107 Megophrys nasuta from Poring region.
thin, serrated; total length 42.2 mm (st 41); tail
length 1.9-2.4 times the head-body-length. Body
brown dorsally and laterally; belly with or with-
out dark spots; caudal muscle brown; dorsal fin
dusky; ventral fin without pigmentation in its
proximal two thirds, dusky near the tip.

Ecology/Ethology
On Mt. Kinabalu, M. nasuta is an inhabitant of
the layer of leaf litter covering the floors of the
submontane dipterocarp and lower oak-chestnut
forest (500-1300 m a.s.l.). It is not only encoun-
© T. Eltz

tered in the vicinity of water bodies, but also far

away from these, particularly on slopes of rock

113
 Megophryidae

able them to devour snails with

shell diameters of up to 50 mm,
large Orthoptera, moths, cater-
pillars, beetles, and even scorpi-
ons of up to 10 cm in length. Oc-
casionally, a diurnal specimen
may be encountered. The males
call from the vicinity of streams,
particularly during the periods of
dawn and dusk. They form very
noisy choruses which, however,
rarely last longer than 30 min-
utes. The white eggs (2 mm in
diameter) are deposited on the
underside of rocks and fallen
logs in those sections of streams
where the current is moderate
to weak (m: 1b-i). The larvae
prefer calmer sections of forest
© U. Manthey

streams, particularly accumula-

tions of leaf litter along the
edges of basins and on the bot-
Fig. 108 Megophrys nasuta one day old from Pulau Pinang, West Malay- tom (m: 1d, e, f). They “suck
sia. up” floating particles from the
Fig. LV Sonagram and oscillograms of Megophrys nasuta water surface by expanding
(Poring region; 6.00 p.m. +20°C) their funnel-mouth. The latter
does not serve as an adhesive
organ (BOSCHMA 1922, SCHMIDT
1976).

Call: A resonant “honk” note.

The call consist of 9-34 of such
honks. The very loud, initial
notes of a call are often drawn-
out and resemble the song of ci-
cadas. The notes last 160-170 ms;
internote interval 150-160 ms;
dominant frequency 2-2.5 kHz.

Distribution
Megophrys nasuta is native to
the Malay Peninsula, Sumatra,
and Borneo. Mt. Kinabalu:
Koung, Kiau; Sayap: Sg. Wariu,
Sg. Kemantis, Bundu Tuhan;
Poring: Sg. Mantukungan, Sg.
Mamut, Sg. Kipungit to Sg.
Langanan (500-950 m).

114
 Microhylidae

Microhylidae

This family is rich in species and nearly cosmo-
politan in tropical and subtropical latitudes.
The Bornean species are small to medium-sized frogs
with a roundish shape and a small head. They are
characterised by a firmisternal pectoral girdle. The
omosternum is absent, a cartilaginous sternum
present. The vertebral column is displasiocoelous,
the sacrum is furnished with dilated diapophyses.
Maxillae and premaxillae are edentate (except in
Calluella). The tips of the fingers and toes may or
may not be expanded, and there are no intercalary
phalanges. The toes are webbed over half their
lengths, or only on the bases (except in Microhyla
berdmorei). A feebly raised skin fold is found be-
tween the snout and the groin. Amplexus is axillary.
The Bornean species produce small, pigmented,
aquatic eggs from which free-swimming tadpoles
emerge. These possess small terminal mouths with-
out cornified beaks and denticles. The tadpoles of
Kalophrynus are nidicolous. They are character-
ised by a median spiracle, laterally positioned eyes,
and the nares do not appear until shortly before
metamorphosis.
Bornean microhylids include both terrestrial
(Calluella, Chaperina, Gastrophrynoides, Kalo-
phrynus, Kaloula, Microhyla) and arboreal forms
(Metaphrynella). They dwell primary forests and
secondary growth; some species occur in disturbed
habitats associated with the activities of man. On
Mt. Kinabalu, microhylids have a vertical distri-
bution ranging up to 1800 m.
Bornean microhylids comprises 20 species repre-
senting 7 genera. Mt. Kinabalu is home to 5 gen-
era with 8 species.

Key to the Bornean genera of Microhylidae (after MANTHEY & GROSSMANN 1997)

1 Ventral side with dark reticulated pattern (A) .... A B

........................................................... Chaperina
• Ventral side with different colour pattern ........ 2
2 Finger tips spatulate (B) ................................... 3
• Finger tips with different shape ........................ 4
3 Lower sides of outer fingers with double, enlarged
basal tubercle (C) ....................... Metaphrynella
• Lower sides of fingers without enlarged basal D
C
tubercle .................................................. Kaloula
4 Tympanum invisible .......................................... 5
• Tympanum distinctly visible (D) ... Kalophrynus
5 Maxillary and vomerine teeth absent; inner meta-
tarsal tubercle low ............................................. 6
• Maxillary and vomerine teeth present; inner meta-
tarsal tubercle spatulate (E) ................ Calluella
6 Snout short (< two times the eye diameter); toe
E
tips not T-shaped ............................... Microhyla F
• Snout long (> two time the eye diameter); toe tips
T-shaped (F) .......................... Gastrophrynoides

115
 Microhylidae
Genus Chaperina MOCQUARD, 1892
Chaperina is a monotypical genus. Its only spe-
cies, Chaperina fusca, occurs all over the island
(Sabah, Sarawak, Brunei, and Kalimantan), in the
southern part of the Philippines, and on the Malay
Peninsula.
Chaperina fusca MOCQUARD, 1892
Chaperina fusca MOCQUARD, 1892: 35 – Type locality: Sintang,
Borneo
Chaperina fusca, SMITH 1931: 21; INGER 1956: 398, 1966: 140,
1978: 312; MATSUI 1979: 322; MANTHEY 1983: 21; INGER &
STUEBING 1992: 41-51; ROZKOŠNÝ & KOVAC 1994: 865;
HOFFMANN 1995a: 10, 1995c: 27; INGER, STUEBING & TAN 1996:
362; LAKIM et al. 1999: 32; INGER et al. 2000: 16
Diagnosis
A tiny, long-limbed frog (< 30 mm) with a very
characteristic pattern on the ventral side, i.e. yel-
low spots separated by a dark brown network. Cu-
riously, the yellow colour will rub off onto human
fingers. There are small, flexible, spine-like der-
mal projections on the heels and elbows.
Description
Males up to 21 mm, females up to 26 mm; TL/SVL
0.49-0.56; habitus slender to moderately stocky;
snout rounded, projecting, longer than eye diam-
Fig. 109 Chaperina fusca from Sg. Wariu, Sayap.
© M. Schroth
116
© P. Hoffmann
Fig. 110 Chaperina fusca from Poring region, 650 m.
eter; canthus not distinct, lores almost vertical; tym-
panum obscure; pupil horizontal; tips of outer three
fingers dilated into round disks; tip of 1st finger not
expanded; finger lengths 3>2=4>1; subarticular
tubercles small; tips of toes expanded into disks;
webbing rudimentary, limited to bases of toes; in-
ner metatarsal tubercle oval; no outer metatarsal
tubercle; skin above smooth or with low tubercles,
sides with low, round tubercles; ventral surface
smooth; supratympanic fold between eye and ax-
illa; males with median subgular vocal sacs, nup-
tial pads absent.
Dorsum and sides black, with or without small
white, greenish, or bluish dots; limbs brown with
dark bars; below a dark brown network enclosing
large yellow cells; iris dark brown.
Head-body section of tadpoles oval, slightly flat-
tened above, rounded below; snout truncate; tail
tapering abruptly near end to a rounded tip; mouth
terminal; lips not expanded; no labial teeth; no
keratinized beaks or papillae; head-body-length (st
40) 21 mm; head-body portion black above, lighter
below; caudal muscles grey with black line at dor-
sal edge; caudal fins unpigmented; above iris a
small, halfmoon-shaped, bright metallic golden
spot. Freshly metamorphosed juveniles are black
below, peppered with fine white dots.
 Microhylidae

1 2

© P. Hoffmann
3 4

Fig. 112 Eggs of Chaperina fusca from Poring region,

650 m.

the walls of small water bodies or to submerged

5 pool next to Sg. Lidian (surface area 10 × 20 cm,
6
Fig. 111 Ventral views of Chaperina fusca from Poring
(1-5), photos: P. Hoffmann, and from Sg. Kemantis,
Sayap (6), photo: M. Schroth.
leaf litter (m: 1g, 2b, d, e, g). We found adults and
larvae at various stages of development in a rock
depth 5-10 cm, filled with rotting mud and bam-
boo leaf litter), and another reproduction commu-
nity in a phytothelme in a buttress tank at Sayap
(surface area 20 × 30 cm, depth 25 cm, 1 m above
the ground). Here, we observed nightly territorial
encounters between two males which lasted a cou-
ple of minutes. The opponents would raise against
one another, chirp, and try to push each other away.
Matings and oviposition take place at night.
Fig. 113 Tadpole of Chaperina fusca from Pulau Tioman,
West Malaysia.

Ecology/Ethology
On Mt. Kinabalu, Chaperina fusca is an inhabitant
of the primary dipterocarp forest (500-1250 m). It
lives in the layer of leaf litter, occasionally climb-
ing low vegetation. It is generally diurnal. The
males often form choruses in and around small
ephemeral water bodies with maximum depths of
© U. Manthey

less than 25 cm rich in humic acids and infusoria

(the tadpoles feed primarily on infusoria). A clutch
comprises 6-10 grey eggs which are attached to

117
Microhylidae

ROZKOŠNÝ & KOVAC (1994) found

tadpoles in bamboo internodes
(Gigantochloa levis) near Poring.
The internodes lying on the sur-
face and measured 10 cm in di-
ameter, were 30-40 cm long, and
held rain water and fallen bam-
boo leaves. Other aquatic inhab-
itants were larvae of Coleoptera,
Diptera, and Odonata.

Call: (+22°C) Two squeaking,

nasal notes which are particularly
distinct in their pulsing in their
first and last thirds (note duration
400-550 ms; internote interval
400 ms), followed by an inter-
val of 3-4 s and three short (100-
150 ms) postnotes; dominant
frequency 2.5-3.5 kHz; frequen-
cy modulation and pulse modu-
lation present. Furthermore, long
drawn-out sounds (up to 4 s in
duration) with distinctly modu-
lated pulsing were noted (fre-
quency 2-3 kHz).

Distribution
Chaperina fusca is known from
the Peninsular Malaysia, Borneo,
and the southern parts of the Phil-
ippines. Mt. Kinabalu: Sayap:
Gua Melayu, between Sg. Wariu
and Sg. Kemantis (1050 m);
Kiau; Kenokok (1000 m); Sg.
Lidian (1250 m); Sg. Kipungit
(500-600 m); East Ridge above
Poring (1000 m). It is very likely
that the species occurs also at
greater altitudes on Mt. Kinabalu
since it was recorded from 1800 m
on Mt. Murud, Sarawak (SMITH
1925a).

Fig. LVI Sonagrams and oscillo-

grams of Chaperina fusca (Sayap re-
gion, 1000 m; 8.00 p.m. + 20-22°C)

118

Genus Kalophrynus TSCHUDI, 1838
Species of the genus Kalophrynus occur in parts of
Southeast Asia, on Sumatra, Java, Borneo, and the
Philippines.
The Bornean species are small to medium-sized frogs
with pointed snouts, short hind limbs, and relatively
narrow heads. Tympanum present, but often indis-
tinct; skin usually granular above and below. Tips
of fingers and toes not dilated, 4th finger and 5th toe
very short; webbings limited to less than half length
of toes (except in K. pleurostigma). Maxillary and
vomerine teeth absent. Males with median subgular
vocal sacs and nuptial pads.
Tab. 9 Distribution of Kalophrynus-species on Borneo.
Key to the Bornean species of Kalophrynus (partly after INGER 1966)
1 Fleshy webbings present on palm ....... nubicola
• No fleshy webbing on palm .............................. 2
2 5th toe projecting as far as or farther than 3rd toe
............................................................ punctatus
• 5th toe not projecting as far as 3rd ...................... 3
3 Portion of 4th finger projecting from palm shorter
than terminal phalanx of 3rd finger ................... 4
• Portion of 4th finger projecting from palm longer
than terminal phalanx of 3rd finger ......................
....................................................... pleurostigma
4 Only 1 tubercle under the 4th finger between the
palmar tubercle and tip of the finger ...................
........................................................ subterrestris
• 2 tubercles under the 4th finger ......................... 5
Microhylidae
These frogs deposit their eggs in aquatic micro-
habitats such as tree holes and bamboo internodes.
Head-body of larvae oval, flattened above; snout
broadly rounded, mouth terminal. Lips not ex-
panded; eyes laterally, visible from below. Tail
lanceolate. Spiracle mid-ventral, without free
flap; anal tube median, not open; gut with only
one or two coils. The larvae do not feed (no
beaks, no labial teeth or papillae) and complete
their development with the nutrition provided by
their yolk.
Adults are terrestrial, dwelling the leaf litter on the
floors of primary forests. Seven species are known
from Borneo, two of which appear on Mt. Kina-
balu.
5 A light stripe or line from canthus to groin ..... 6
• No light stripe or line on snout or side ...............
......................................................... intermedius
6 Bluish or whitish inguinal spots in a large dark
area; adult females < 35 mm; lowland species ...
....................................................... heterochirus
• Inguinal spots present or absent; adult females >
40 mm; montane species .................... baluensis
119
 Microhylidae
© H. Sauer
Fig. 114 Kalophrynus baluensis from Sg. Silau-Silau, 1500 m.
Kalophrynus baluensis KIEW, 1984
Kalophrynus baluensis KIEW, 1984: 151 – Type locality:
Kamborangoh at 7,200 feet on Mount Kinabalu
Kalophrynus pleurostigma, (not Kalophrynus pleurostigma
TSCHUDI, 1838) SMITH 1931: 21 (part); INGER 1954a: 250
Kalophrynus pleurostigma pleurostigma, (not Kalophrynus
pleurostigma TSCHUDI, 1838) MATSUI 1979: 320
Kalophrynus baluensis, INGER & STUEBING 1989: 59; MALKMUS
1991a: 28, 1992a: 111; INGER & STUEBING 1992: 41-51; MALKMUS
1994a: 9, 1994b: 224, 1996b: 20, 1996c: 282; INGER, STUEBING
& TAN 1996: 362; INGER & TAN 1996a: 557; MALKMUS & RIEDE
1996c: 152; INGER & STUEBING 1997: 96; INGER et al. 2000: 16
Diagnosis
A medium-sized brown frog; habitus stout; snout
pointed, shorter than the eye diameter; yellow
patch(es) bordered with dark brown present or ab-
sent in the inguinal region; 5th toe with 1 subarti-
cular tubercle.
120
Description
A moderately large, stout microhylid; males up to
36, females up to 47 mm; snout short, pointed; hind
limbs short (TL/SVL 0.39-0.45); head wider than
long, canthus rostralis distinct; tympanum 1/2 to
3/4 of eye diameter; supratympanic fold not rec-
ognisable; subarticular and metatarsal tubercles
weakly developed; tips of fingers and toes round,
not dilated; finger lengths 3>2>1>4; toe lengths
4>3>5>2>1; rudimentary webbings on bases of
toes; skin finely tubercular, particularly on upper
side of body and limbs.
Upper side medium to light brown, with individu-
ally highly variable, asymmetric, medially oriented
spotted pattern ; hind limbs with dark crossbars; a
light dorsolateral stripe between rostrum and fe-
mur, paralleled below by a dark brown one; sides
light brown, with partly densely arranged dark
brown dots; a dark reticulated pattern in the inguinal
 Microhylidae

region, partly with 1-2 large, yel-

low ocelli; lower side whitish
grey with small brown dots which
become more numerous towards
the throat; posterior half of belly
sometimes with dark-bordered
white spots; iris pale golden with
black dots.
The colour pattern and body
shape show distinct phytomime-
tic properties.
Tadpoles probably similar to
those of K. pleurostigma.

Ecology/Ethology
Kalophrynus baluensis inhabits
the leaf litter layer of the oak-
chestnut forests between 1400 and
1800 m a.s.l. Along larger streams
with a high noise level, the spe-
cies keeps a distance of at least
20-30 metres from the water. It has
also formed a colony in the midst
of the chalet settlement of the
Headquarters and is locally com-
mon. The males call between 6.00
p.m. and 6.00 a.m., but can occa-
sionally also be heard during the

© R. Malkmus
day. The maximum calling activ-
ity is reached between 6.00 and
9.00 p.m. after rain showers. The
frogs remain mute during pro- Fig. 115 Kalophrynus baluensis from the Headquarters.
longed dry periods. The call is
emitted from holes in the ground,
rock crevices, and cavities in roots and dead wood crickets, hairless caterpillars, and particularly spi-
and is very difficult to pinpoint. Depending on the ders) in slow motion, fixes it visually, and then
state of excitement and the prevailing weather con- seizes it with the tongue in a sudden forward mo-
ditions, the calls are emitted at intervals of 3-10 s tion.
and often answered by other males leading to the Defence behaviour includes fleeing via a series of
formation of a chorus. These do, however, not rep- quick, 20-30 cm long leaps. When touched it ex-
resent insular, geographically defined calling com- pels large amounts of a white, very sticky secre-
munities which (like in Megophrys baluensis) are tion from its dorsal glands which is probably ef-
isolated from one another acoustically, but rather fective enough to deter even anurophagous snakes.
variable concentrations of groups of individuals The frogs uses its hands to rub its chest, belly, and
within a widespread population. Nothing is known flanks, while the hind limbs take care of the distal
about its reproductive biology. portions of the body. It will then make a hand-
Kalophrynus baluensis forages for food at night, stand raising the posterior body off the ground.
whereby it stalks its prey (ants, termites, small Without losing balance, the hind limbs rub over

121
Microhylidae

the posterior two thirds of the body

in circular motions. It is to be pre-
sumed that this serves to distribute
glandular secretions over the skin.

Call: (+20°C) The calls are emitted

as euphonic, very weakly pulsed sin-
gle notes at intervals of 6-10 s. Fre-
quency 1.8 to 5 kHz; carrier fre-
quency 2.6 kHz; harmonics at 7 kHz;
duration 125 ms on average; fre-
quency modulation present.

Distribution
This species is so far known only
from Mt. Kinabalu where records are
as yet further limited to the Head-
quarters and Mesilau region. As the
very characteristic voice of this
microhylid has never been recorded
from above 1800 m altitude, it ap-
pears likely that the altitudinal infor-
mation relating to the type locality Fig. LVII Sonagram and oscillograms of Kalophrynus baluensis
(2200 m) is erroneous (SMITH 1931). (Headquarters, 1550 m; 7.00 p.m. +20°C)

Fig. LVIII Distribution of Kalophrynus baluensis in the area of the Headquarters.

122
 Microhylidae

Kalophrynus pleurostigma Tadpoles endotrophic; head-body portion oval,

snout broadly rounded; mouth terminal; lips not
TSCHUDI, 1838 expanded; spiracle situated mid-way between eye
Kalophrynus pleurostigma TSCHUDI, 1838: 86 – Type locality: and end of body, without free flap; tail lanceo-
Sumatra late, tip bluntly rounded, 1.6-1.9 times the head-
Kalophrynus pleurostigma pleurostigma, INGER, 1966: 132 body-length; total length of mature larvae up to
Kalophrynus pleurostigma, SMITH 1931: 21; INGER 1956: 396, 14 mm. Head-body grey to olive above, heavily
1978: 312; MANTHEY 1983: 21; MALKMUS 1985: 8, 1987: 279,
1988c: 176, 1988d: 9; INGER & STUEBING 1992: 41-51; INGER,
dusted below; caudal muscle brown; fins dusky
STUEBING & TAN 1996: 362; MANTHEY & GROSSMANN 1997: or not.
54; LAKIM et al. 1999: 32; INGER et al. 2000: 16
Ecology/Ethology
Diagnosis K. pleurostigma inhabits the leaf litter layer of dip-
A medium-sized brown frog; habitus stout; head terocarp primary forests between altitudes of 500
narrow; snout pointed, its length equal to eye di- and 800 m on Mt. Kinabalu. Males call at night, to
ameter ; a black ocellus at each groin; similar to K. a considerably lesser extent also during the day,
baluensis, but larger. from holes in the ground, below leaf litter, or the
edges of spawning sites. This species chooses shal-
Description low temporary pools on the forest floor for breed-
Males up to 50 mm, females up to 57 mm; hind limbs ing, such as holes in logs and road ruts filled with
short, TL/SVL 0.36-0.46; head wider than long; nos- rainwater (m: 2b, d, e, h). The eggs are spherical,
tril closer to tip of snout than to eye; loreal region unpigmented, and deposited in clusters, 2.5-3 mm
almost vertical; tympanum usually distinct, 0.5-0.75 in diameter. The development from gastrulation to
times eye diameter; finger tips round, not expanded; metamorphosis takes 16 days at a water tempera-
finger lengths 3>2>1>4; subarticular tubercles dis- ture of 24-29°C. Newly metamorphosed froglets are
tinct; tips of toes like those of fin-
gers; toe lengths 4>3>5>2>1; de-
gree of toe webbing very variable, Fig. 116 Kalophrynus pleurostigma from Poring region.
usually webbed over 1/3 to 3/4
of their lengths; dorsal skin
granular (with keratinized
spines in male specimens); belly
coarsely granular; supratympanic
fold present, but often obscure;
vocal sac and nuptial pads
present.
Reddish brown to loam-coloured
or greyish brown above, usually
with large dark blotches; dark
crossbars on upper sides of hind
limbs; flanks dark, sharply set off
from dorsal coloration by a light
line running from tip of snout to
groin; a black spot in front of the
insertion of the hind limb; lower
side yellowish white; throat usu-
© R. Malkmus

ally dark brown, peppered with

white dots; belly with dark
spots; iris bright reddish golden.

123
Microhylidae
excellent jumpers performing leaps of up to 60 cm
(LIM & NG 1991).
The defence, comfort, and hunting behaviours of
the adults are very similar to those shown by K.
baluensis. Their diet consists of spiders, small cat-
erpillars and beetles, but mainly of ants and ter-
mites. The stomach of a single individual contained
up to 150 ants. When foraging the frogs may also
ascend into the lowest strata of shrubs up to 20 cm
above the ground.
Call: The call consists of a long series of unpulsed
notes. The calls emitted by a frog from Sepilok
(eastern Sabah) were irregular both in length
Genus Kaloula GRAY, 1831
Species of the genus Kaloula are present from
Korea and northern China to the Lesser Sunda Is-
lands, the Philippines, and Sri Lanka.
The Bornean species are medium-sized, semi-
fossorial, stocky frogs with short thick hind limbs;
head wider than long, snout broadly rounded, ver-
tical in profile; canthus rostralis indistinct; tympa-
num concealed by skin; tips of fingers not ex-
panded, webbing rudimentary; large inner meta-
tarsal tubercle present. Males possess a median
subgular vocal sac and coarsely granular skin on
the throat.
The frogs deposit their eggs in temporary pools.
Tadpoles with oval body, flattened above; snout
broad and truncate; eyes lateral, 6-7 times as widely
separated from each other as the nostrils; mouth
terminal, no beaks or labial teeth. The tadpoles of
Kaloula differ from all other Bornean microhylid
larvae by having the opening of the spiracle at the
end of the body above the anal tube. Anal tube
medial, projecting over the lower caudal fin. Tail
Tab. 10 Distribution of Kaloula-species on Borneo.
124
(27-285 ms) and interval (138-348 ms), with a
dominant frequency of 420-625 Hz at +27.5°C. Fre-
quency modulation present.
Distribution
Kalophrynus pleurostigma is found on the Malay
Peninsula, Sumatra, Borneo, and the Philippines. The
taxonomic status of the population from Java remains
undetermined (MATSUI, CHANARD & NABHITABHATA
1996). Mt. Kinabalu: No reliable records exist from
localities above 800 m altitude (Poring - Sg. Langa-
nan), with those from Bundu Tuhan, Lumu Lumu,
Kamborangoh (SMITH 1931, INGER 1966, MATSUI
1979) being most likely referable to K. baluensis.
pointed, almost twice as long as head-body length.
Larvae are suspension feeders.
Adults are terrestrial, but may ascend bushes and
trees during periods of heavy rainfall. They also
occur in human settlements, agriculturally used
land, and secondary scrub; K. baleata is also found
in primary forests. They are nocturnal and find
suitable shelters for the day in holes in the ground,
beneath piles of rocks, and fallen logs.
The genus comprises 11 species, 2 of which are
widely distributed on Borneo. Their occurrence on
Mt. Kinabalu is limited to cultivated lands in the
marginal zones.
Key to the Bornean species of Kaloula
1 Webbing reduced to basal fragments; a wide, pale
brown stripe between eye and groin and from eye
to arm ..................................................... pulchra
• Webbing reaches the middle subarticular tubercle
on inner edge of 4th toe; pattern different from
above ...................................................... baleata
 Microhylidae

Kaloula baleata (MÜLLER, 1836)

Bombinator baleatus MÜLLER, 1836: 96 – Type locality:
Krawang, Java

Diagnosis
A stocky frog with short, thick limbs and a short,
broadly rounded snout; lores not concave; tips of
fingers dilated into truncate disks; the webbing of
the toes reaches the middle articular tubercle on
the inner edge of the 4th toe.

© P. Hoffmann
Description
Males up to 61 mm, females up to 67 mm; TL/SVL
0.29-0.36; otherwise see Diagnosis. Finger lengths
3>4>2>1; toe lengths 4>3>5>2>1; webbing 1(1.5),
2(1.5/1.5), 3(2/2), 4(3/3), 5(2); inner metatarsal tu-
bercle oval, compressed, elevated; outer one round;
skin granular above, or with low, rounded tubercles;
smooth or rugose below; throat of males granular;
often with a fold between eye and insertion of fore-
limb. Dorsum dark or pale brown with a large dark
spot in the centre; sides brown with dark spots; of-

© U. Manthey
ten with white spots in the groin region; pale brown
below, with darker spotting, lighter towards the belly;
iris golden brown.
Tadpoles very similar to those of K. pulchra. From top to base: Male of Kaloula baleata
Fig. 117 from Poring, 600 m.
Ecology/Ethology Fig. 118 from Taman Negara NP, West Malaysia.
K. baleata is a commensal species which finds ref-
uges for the day under rocks, wood, and in the soil,
but which has also been found in tree holes up to Distribution
2.5 m above the ground. Following heavy rainfalls, Peninsular Malaysia, the Greater Sunda Islands,
the males form noisy choruses along ponds, road Palawan, and the Lesser Sunda Islands to Flores.
ditches, and the margins of rice paddies. They in- Mt. Kinabalu: Rainwater puddles on the road side
flate their bodies substantially during calling. The at Poring (600 m a.s.l.), edges of primary forest.
eggs are small and are deposited in clusters in
puddles, road ditches, and small ponds (m: 2b).
The food of K. balaeta comprises small insects,
particularly ants.

Call: (+23°C) A series of loud groaning honks

in slow sequence. The notes are pulsed with a
duration of 280-300 ms, with a distinct decrease
in intensity; dominant frequency 1-3 kHz.

Fig. LIX Sonagram and oscillogram of Kaloula

baleata

125
 Microhylidae

Kaloula pulchra GRAY, 1831

Kaloula pulchra GRAY, 1831: 38 – Type locality: China

Kaloula pulchra, MATSUI 1979: 319; INGER , STUEBING & TAN
1996: 362

Diagnosis
A very stocky, rotund frog with short, thick hind
limbs and a short, rounded snout; lores weakly con-
cave; fingers long, their tips expanded into trun-
cate disks; toe webbing not extending beyond the
level of the proximal subarticular tubercle of the

© U. Manthey
4th toe; a pair of prominent flanges on the sole of
each foot between the heel and the base of the toes.

Description Fig. 120 Ventral view of Kaloula pulchra from Laos.

Males up to 70 mm, females up to 75 mm; TL/SVL
0.25-0.35; habitus similar to that of K. baleata; fin-
ger lengths 3>4>2>1; toe lengths 4>3>5>2>1; web-
bing rudimentary; inner metatarsal tubercle strongly
raised, outer one transversely oval; skin thick, finely
pebbled or tubercular above; below smooth or faintly
granular; lower back with widely scattered bumps;
indistinct fold from eye to axilla.
Dorsum brown with scattered dark spots; a wide ir-
regular, whitish to pale brown stripe from eye to groin
and from eye to insertion of arm; throat brown (to
black in males); chest, belly, and undersides of legs
yellowish with brown spots, sometimes forming a
network.

Fig. 119 Kaloula pulchra from Laos.

© U. Manthey

126

© U. Manthey
Fig. 121 Tadpole of Kaloula pulchra from Laos.
Tadpoles olive brown to black above, sometimes
with golden dots, and lighter beneath spotted with
white; fins transparent, with small dark patches;
maximum head-body-length 14 mm, tail 26 mm.
For further description see under genus.
Ecology/Ethology
The bullfrog is a distinct commensal commonly
encountered within human settlements and
anthropogenously disturbed habitats, in particular
in land used for agriculture. It survives periods of
drought in underground hiding places which it
excavates itself and which also serve as shelters
during the day. The males form choruses after
rainfalls which may often include very many indi-
viduals (> 100) along temporary stagnant water
bodies. Here, the extremely inflated frogs can of-
ten be seen floating on the water surface. Females
which are attracted by the calling are immediately
seized on arrival in an axillary amplexus. They dip
their heads into the water at intervals, simultane-
ously raising the rear and releasing spawn. Any
temporary pool may be utilised for breeding pur-
poses by this species (m: 2b, c). The small eggs are
laid in floating masses and develop at great speed.
The frogs feed on small insects with a predilection
for ants.
Call: A loud groaning honk consisting of a single,
partially pulsed note (0.56-0.60 s). Each note is
composed of 18-21 pulses at a rate of 32-35 pulses
Microhylidae
per second; dominant frequency 2500 Hz; no fre-
quency or intensity modulations (HEYER 1971).
Distribution
The species is present throughout Southeast Asia. It
has probably been introduced to the Sunda Islands
and now occurs eastwards up to the Lesser Sunda
Islands and Sulawesi. It is commonly presumed that
it has reached Borneo only more recently. Mt. Kina-
balu: Only a single female specimen has become
known from the Headquarters region, 1665 m
(MATSUI 1979). This is likely to have been a translo-
cated specimen as this species is not normally a
montane one and has never since been found on Mt.
Kinabalu again.
Genus Metaphrynella PARKER, 1934
The genus Metaphrynella comprises only two spe-
cies: the montane M. pollicaris (BOULENGER, 1890)
of the Malay Peninsula and the Bornean endemic
M. sundana (PETERS, 1867) known from Sabah,
Sarawak, Brunei, and Kalimantan.
Metaphrynella sundana (PETERS, 1867)
Calohyla sundana PETERS, 1867: 35 – Type locality: Pontianak
(Borneo)
Metaphrynella sundana, MALKMUS 1988c: 176, 1988d: 7, 1989:
185, 1992a: 110; INGER & STUEBING 1992: 41-51; MALKMUS
1994b: 224, 1995b: 35; HOFFMANN 1995a: 10, 2000: 37; MALK-
MUS 1996b: 20-26, 1996c: 282; INGER, STUEBING & TAN 1996:
362; HOFFMANN 2000:37; LAKIM et al. 1999: 32; INGER et al.
2000: 17
Diagnosis
A tiny, stocky, grey to black, arboreal microhylid
with a pointed snout and moderately short limbs.
The underside of each finger with enormously en-
larged tubercles at the base which form accessory
adhesive organs. Tips of the fingers dilated, tips of
the toes not expanded.
Description
Males up to 23 mm, females up to 25 mm; TL/SVL
0.42-0.47; head as wide as long; snout projecting,
127
 Microhylidae

Fig. 122 Metaphrynella sundana

from Poring.
© M. Maronde

Fig. 123 Metaphrynella sundana

from Sg. Langanan.
© M. Schroth

128
 Microhylidae
© P. Hoffmann

Fig. 124 Hand of Metaphrynella sundana from Poring,

about 600 m.

© P. Hoffmann
a little longer than eye diameter; lores almost ver-
tical; tympanum visible, but somewhat concealed,
3/5 of eye diameter; fingers with rudimentary, Fig. 125 Tree hole with foam and eggs of Metaphrynella
fleshy webbings; finger lengths 3>4>2>1; toe sundana near Poring, 550 m.
lengths 4>3>5>2>1; toes webbed for 1/2 to 3/4; a
weak, elongated inner, but no outer metatarsal tu-
bercle; tibio-tarsal articulation reaching tympanic continue throughout the night until about 5.30 a.m.
region; skin above and below smooth or feebly The largest calling community was discovered in a
granular, sides with larger tubercles; a curved bamboo forest bordering the Sg. Langanan. The
supratympanic fold between eye and axilla; males frequency and timbre of a call is often substantially
with a subgular vocal sac, but no nuptial pads. altered by the size and shape of the treehole. Call-
Upper sides grey to brown, with irregularly scat- ing communities are always farther than 100 m
tered black spots which are peppered with small
white dots; hind limbs with or without black
crosslines; elbows often yellowish brown; lower Fig. 126 Eggs of Metaphrynella sundana from Poring.
sides dark grey to blackish brown with lighter
clouding, occasionally yellowish; iris reddish
golden with fine black dotting.

Ecology/Ethology
This microhylid can be found in tree holes (often
in bamboo internodes) filled with rainwater in
dipterocarp primary forests, on Mt. Kinabalu at
500-850 m a.s.l. The males call at night from
treeholes 0.3 to 10 m above the ground with the
openings of these cavities not exceeding 10-12 cm
© P. Hoffmann

in diameter. Calling activity was noted at Poring to

begin as early as 4.00 p.m. in some instances, but
normally commences from 5.00 p.m. These calls

129
Microhylidae

away from mountain streams if the latter create a Fig. LX Sonagram and oscillograms of Metaphrynella
high background noise. A male was observed emit- sundana (Bat Caves/Poring, 580 m; 9.00 p.m. +23°C)
ting advertising calls during amplexus.
In February in Poring region a female was observed
depositing 1300 eggs (egg diameter 1 mm) in a jelly
string (5-6 mm diameter) in a water filled tree hole,
1.8 m above ground. The water surface was cov-
ered with a light whitish foam (HOFFMANN 2000).
A clutch covered in foam was found in a bamboo
internode (m: 2g) at Poring.

Call: (+23 to 25°C) The call consists of a single,

unpulsed piping note (80-150 ms) emitted at in-
tervals of mostly > 3 s; dominant frequency 800-
1500 Hz; frequency modulation present, with peaks
up to 21000 Hz.

Distribution
Metaphrynella sundana is endemic to Borneo. Mt.
Kinabalu: As yet, this species has only been re-
corded from the region between Poring and Sg.
Langanan.

sess a median subgular vocal sac.

Genus Microhyla TSCHUDI, 1838
The genus Microhyla is distributed from China to
India and Sri Lanka, in Southeast Asia eastwards
up to Bali.
All five of the Bornean species are small (except
for M. berdmorei under 30 mm SVL), and have
conical body shapes with a rounded snout and rela-
tively long, strong hind limbs. They all share a
distinct tendency for a reduction of the length of
the first finger. Maxillary and vomerine teeth ab-
sent, tympanum usually not visible. Males pos-
Eggs are deposited in very small pools. Tadpoles
without beaks, eyes lateral, mouth terminal, spiracle
medial, ventral, with movable flaps, or with an
opening at the end of the tube near the anal tube.
The shapes of the tail and the lower lip are often
indicative of the species.
Adults of the Bornean species live in the leaf litter
layer on the floor of primary and old secondary for-
ests below 700 m altitude.
Five species are known from Borneo, two from Mt.
Kinabalu.

Tab. 11 Distribution of Microhyla-species on Borneo.

130
 Microhylidae

toe webbing 1(1), 2(1/1), 3(2/2), 4(3/3), 5(1.5-2); an

Key to the Bornean species of
Microhyla
1 No disks and dorsal grooves at bases of toes ......
........................................................... maculifera
• Disks and dorsal grooves at bases of toes present
........................................................................... 2
2 3rd and 5th toes webbed up to disks ................... 3
• 3rd and 5th toes not webbed up to disks ................
........................................................... borneensis
3 1st finger 1/3 of length of 3rd; toes fully webbed .
............................................................ berdmorei
• 1st finger reduced to a rudiment ........................ 4
4 Belly pigmented, but with an irregularly shaped
area free of melanophores; dark cross bars on hind
limbs bordered with light ................... petrigena
• Belly without melanophores; dark cross bars on
hind limbs without light bordering ..... perparva
oval inner metatarsal tubercle and a conical outer one;
tibio-tarsal articulation reaching a little beyond tip of
snout; skin above and below smooth, with scattered
low tubercles, with tubercles and low ridges on sides;
weak supratympanic fold; males with subgular vocal
sacs; nuptial pads wanting.
Dorsum light brown, with dark brown to purple-
coloured markings between eyes; large light-edged
mark on middle of back; lips spotted with black;
broken black stripe on side; hind limbs dorsally
with crossbars; dark spot on knee; lower surfaces
yellowish, mottled with brown on throat and chest.
Head-body portion of tadpoles almost triangular,
with very wide snout, flat above, sloping upward
from snout to root of tail which is then prominently
raised; tail tapering gradually from root into a long,
narrow filament; eyes lateral, nostrils dorsal, mouth
terminal; lips not expanded; tail length 2.5-2.8
times head-body-length. Head-body grey, lighter
ventrally; fins dusted with melanophores. Total
length 22.4 mm (st 32-40).

Microhyla borneensis PARKER, 1926 Ecology/Ethology

Microhyla borneensis is an inhabitant of the low-
Microhyla borneense PARKER 1928: 473 – Type locality: “Kidi land primary forests dwelling the leaf litter layer
District, Sarawak” (= Bidi District?)
on the forest floor and low vegetation (< 30 cm). It
Microhyla borneensis, PARKER 1934: 128; MATSUI 1979: 323;
INGER & STUEBING 1992: 41-51; INGER, STUEBING & TAN 1996: is both diurnal and nocturnal. This species breeds
362; INGER et al. 2000: 17 in small accumulations of rainwater, sometimes in
wallows used by wild boars (m: 2c). The tadpoles
Diagnosis
A tiny brown frog with a moderately
Fig. 127 Microhyla borneensis
stout body and long limbs; dorsal sur-
faces of disks of fingers and toes with
longitudinal grooves; 3rd and 5th toe not
webbed up to disks; 1st finger less than
1/3 length of 3rd; tympanum invisible.

Description
Males up to 18 mm, females up to 23 mm;
TL/SVL 0.62-0.70; head broader than
long; snout slightly pointed, projecting
beyond mouth, longer than eye in diam-
eter; canthus rostralis rounded, lores
© R.B. Stuebing

weakly concave; tips of outer 3 fingers

slightly dilated into disks; finger lengths
3>4>2>1; tips of toes more expanded than
those of fingers; toe lengths: 4>3>5>2>1;

131
 Microhylidae
are mid-water suspension feeders. Nearly nothing
is known about its natural history.
Distribution
Microhyla borneensis is known from the Malay
Peninsula and Borneo. Mt. Kinabalu: Only one
specimen has as yet been recorded, a female from
Poring/Hot Springs (555 m).
Microhyla petrigena
INGER & FROGNER, 1979
Microhyla petrigena INGER & FROGNER, 1979: 318 – Type
locality: Nanga Tekalit, Kapit District, Sarawak
Microhyla petrigena, INGER, STUEBING & TAN 1996: 362; INGER
et al. 2000: 17
Diagnosis
A very small, moderately stout, brown frog with
long hind limbs; 1st finger reduced to a rudiment;
all toes but the 4th fully webbed.
Description
Males up to 16 mm, females up to 18 mm; TL/SVL
0.60-0.70; head obtusely pointed; tympanum invis-
ible; fingers with slightly expanded tips; 1st finger
Fig. 128 Microhyla petrigena
© R.B. Stuebing
132
rudimentary; finger lengths 3>4>2(>1); 2 low meta-
carpal tubercles; tips of toes more expanded than
those of fingers, with longitudinal groove on dor-
sal side; toe lengths 4>3>5>2>1; toe webbing 1(0),
2(0/0), 3(0/0), 4(0.5/0.5), 5(0); low inner metatar-
sal tubercle, no outer one; males with subgular
vocal sac.
“Top of head dark grey; a constricted dark grey fig-
ure on back beginning at level of shoulders and
extending to groin then curving forward in a broad
band low on side and reaching just behind eye;
except for these dark areas, back and sides grey
white; dorsal surface of limbs russet with broad
darker bands bordered by light grey to ivory, dark-
edged stripes; throat, chest and belly dark greyish;
belly with wide irregular, meandering white area”
(INGER & FROGNER 1979).
Head-body portion of tadpoles oval in shape, flat-
tened above, snout broadly rounded; eyes lateral,
visible from below; nostril dorsal; mouth termi-
nal; anal tube in ventral fin; tail abruptly tapering
subterminally to a short tip; tail length 1.5-2.0
times head-body-length. Head-body black above
and on sides, whitish below; caudal muscle black;
fins without pigment except for a vertical black
band near end of tail. Total length up to 17 mm
(st 37).
Ecology/Ethology
Microhyla petrigena is a low-
land species, dwelling the
floor litter in primary forests
where it feeds on tiny inver-
tebrates, mainly ants. Micro-
hyla petrigena breeds in wa-
ter-filled rock potholes (diam-
eter and depth < 30 cm) along
small forest streams (m: 1g).
The tadpoles are mid-water
suspension feeders filtering
tiny organisms. Nearly noth-
ing is known about its life his-
tory.
Distribution
This species is endemic to
Borneo. Mt. Kinabalu: Only
known from the Poring area.

Ranidae
Ranid frogs are almost cosmopolitan in distribu-
tion and absent only from southern South America,
the West Indies, most of the Australian continent,
and many Oceanic islands.
The Bornean representatives of this family exhibit
all anatomical features typical for ranid frogs. They
have a firmisternal pectoral girdle, a sacrum with
cylindrical diapophyses, and the omosternum and
sternum are ossified. Bony postzonal sternal ele-
ments are present, and so are eight holochordal,
procoelous presacral vertebrae and palatines. A
parahyoid is absent, and the cricoid ring is com-
plete. Maxillae and premaxillae are dentate. The
fingers are generally free of webbings, while
webbings between the toes are well-developed (ex-
cept in the genus Taylorana DUBOIS, 1987). The
pupil is horizontal. Mating takes place in an axil-
lary amplexus.
Ranids are extremely variable with regard to size
and habitus. They have conquered a multitude of
ecological niches. The majority are ground dwell-
ers, but many can climb as well. Many species are
closely associated with water and never wander far
from it. Others are, in contrast, typical inhabitants
of the forest floor, e.g. Limnonectes finchi and Rana
luctuosa, which may be encountered far away from
any water body. Various species, e.g. Rana signata
Key to the Bornean genera of Ranidae (based on MANTHEY & GROSSMANN 1997)
1 Skin membrane between the two outer metatarsals
present (A) ......................................................... 2
• No such membrane; tips of fingers and toes
spatular (B) .......................................... Ingerana
2 Tips of fingers with marginal folds on the disks
(C) and usually dilated (D) ............................... 6
• Tips of fingers without marginal folds on the disks
and not or only weakly dilated ......................... 3
3 Vomerine teeth present (E) ............................... 4
• Vomerine teeth absent ..................... Occidozyga
Ranidae
and Meristogenys orphnocnemis, are known to
leave the water bodies where they have completed
their metamorphosis; they spend their early lives
in the forest, only to return to aquatic habitats on
reaching maturity.
Spawning usually takes place in the water, although
there are exceptions in the form of species like
Limnonectes finchi and L. palavanensis which de-
posit their eggs on land and later transport the tad-
poles to a water body. The larval development in-
volves a stage of free-swimming tadpoles (except
in Taylorana). All tadpoles have beaks, denticles,
and a sinistral spiracle. Most ranid larvae can be
found in ponds, but some have adapted to streams
(e.g. Meristogenys, Huia).
Bornean Ranidae are classified as belonging to the
genera Fejervarya, Hoplobatrachus, Huia, Ingera-
na, Limnonectes, Meristogenys, Occidozyga, Rana,
and Staurois, and comprise altogether 38 species
at present.
Representatives of this family are numerous up to
an altitude of about 1500 m on Mt. Kinabalu, but
only a few species are present above this elevation
(e.g. Limnonectes kuhlii, Meristogenys kinabalu-
ensis, Staurois tuberilinguis). The 22 species repre-
senting 8 genera which occur on Mt. Kinabalu ac-
count for some 60% of the ranid species of Borneo.
A B
C D E
133
Ranidae

F G
4 Dark ventro-lateral line between axilla and groin
present (F) ......................................... Fejervarya
• No dark ventro-lateral line ............................... 5
5 Dorsum with elongate skin folds, tips of fingers
pointed (G) ............................... Hoplobatrachus
• Dorsum without elongate skin folds, if present tips
of fingers rounded and blunt (H); males with
fanged like process (I) on the lower jaw (except I
L. finchi, L. palavanensis) ............ Limnonectes H
6 Digital bones T-shaped distally (J) ................... 7
• Digital bones not dilated distally .............. Rana
7 Habitus very slender; transverse folds beneath the
phalangeal tips (K) ................................ Staurois J K
• Habitus not so slender; no such transverse fold .
........................................................................... 8
8 Tympanum sunken ..................................... Huia
• Tympanum not sunken ................. Meristogenys

differences in the mouthparts of their tadpoles (FEI

Genus Fejervarya BOLKAY, 1915
A genus of relativ small to moderatly large ro-
bust bodied frogs. The skin texture is relatively
smooth, but covered with longitudinal oriented
skin glands in the form of folds. The tips of toe
and fingers are featureless, without any enlarge-
ments or discs.
Recently, Fejervarya was elevated from a subge-
nus of Limnonectes to genus rank by DUBOIS &
OHLER (2000). They summed up some phenetic
differences that exist between Fejervarya and
Limnonectes, such as the shape of the digits of
adults (OHLER & DUBOIS 1999), their types of male
secondary characters (BOULENGER 1920) or the
et al. 1991). Additionally, DUBOIS & OHLER (2000)
pointed out the existence of a dark ventro-lateral
line from armpit to groin, a unique common de-
rived character only to be found in Fejervarya.
More significantly even, the analyses based on
mtDNA sequencing of VENCES (1999), MARMARYOU
et al. (2000), BOSSUYT & MILINKOVITCH (2000) and
EMERSON et al. (2000) suggest that Fejervarya is
not the sister-group of Limnonectes (see also ge-
nus Limnonectes).
Unfortunately, no up to date complete diagnosis of
the genus Fejervarya was published so far.
The genus Fejervarya comprises 14 species (DUBOIS
& OHLER 2000), two of which are present on Bor-
neo. Mt. Kinabalu is home to one of these species.

Tab. 12 Distribution of Fejervarya-species on Borneo.

134
 Ranidae

Fejervarya limnocharis
(GRAVENHORST, 1829)

Rana limnocharis GRAVENHORST, 1829: 42 – Type locality: Java

Rana limnocharis limnocharis, MATSUI et al. 1985: 153
Rana limnocharis, MALKMUS 1994b: 244, 1996c: 285; HOFF-
MANN 1995b: 11
Rana (Limnonectes) limnocharis, HOFFMANN 1998b: 89

Diagnosis
A small to moderately large terrestrial frog with a
very granular back. In contrast to F. cancrivora the
webbing of toes is deeply excised, leaving at least
one phalanx of each toe free.

© P. Hoffmann
Description
Males 32-50 mm, females 48-60 mm; habitus slen-
der to somewhat stout; head longer than wide; snout
round or slightly pointed; vomerine teeth present;
tympanum visible, about 60% of the diameter;
supratympanic fold present; 1st finger longer than
the 2nd; finger tips not dilated; marginal disk folds
absent; fingers without webbings; toe tips not di-
lated; webbing formula 1(1), 2(1/1), 3(1/1), 4(3/
3), 5(1-2); inner metatarsal tubercle oval, outer one
round and very small; dorsal skin with irregular
longitudinal folds.
Males during the mating period with a grey, M-shaped
gular pattern; subgular vocal sac entire; 1st finger

© P. Hoffmann
with whitish to light grey nuptial pads.
Dorsal coloration grey to brown with irregular dark
spots; ventral side white. Specimens with a white
dorsal stripe are found syntopically with those
where such is absent.
Body shape of tadpoles oval; fringe of fins taper-
ing to a point; upper side of body olive green with
black speckles; lower side white; tail tip striped
with black or entirely black; head-body-length
13-15 mm; total length up to 45 mm; labial teeth
rows formula 2(2)/3.

Ecology/Ethology
© M. Schroth

Fejervarya limnocharis is a nocturnal ground

dweller, usually found in the environs of water
bodies (ponds, canals, etc.). It is an explicit
commensal which avoids primary forests. Follow- From top to base
ing heavy nightly downpours, the males form large Fig. 129-131 Different dorsal patterns of Fejervarya
calling communities on the banks of stagnant wa- limnocharis from Poring.

135
Ranidae

ter bodies in the morning hours.

The frogs are often encountered
sitting in the waste water ditches
of human settlements. This spe-
cies shows a predilection for
lower altitudes. It feeds on a va-
riety of insects, millipedes, and
occasionally snails.
The larvae favour stagnant wa-
ter bodies (m: 2b).

Call: A series consists of rapidly

emitted, raspy chirps which are
repeated at irregular intervals.
Note duration 8-120 ms; inter-
note interval 32-440 ms; domi-
nant frequency 1-3 kHz; harmon-
ics at 5 kHz.

Distribution
This species is widely distributed
from Pakistan, India and Sri
Lanka to Japan, to Borneo and
Flores. Mt. Kinabalu: Sayap,
Poring, MATSUI (1985) reported
“one adult female between
Layang Layang and Kambo-
rangoh, 2000 m”, which probably Fig. LXI Sonagram and oscillograms of Fejervarya limnocharis
refers to a translocated specimen. (Peninsular Malaysia, 40 m; 8.00 p.m.)

The distribution of the genus Huia covers the en-

Genus Huia YANG, 1991
Taxonomic remark:
The species originally included in the composite
genus Amolops were rearranged by YANG (1991)
and are now allocated to the genera Amolops,
Huia, and Meristogenys. The respective revision
was based mainly on larval traits such as the
number of continuous lower rows of denticles in
the tadpoles of Amolops being three, whereas they
count at least four, usually more, in Huia and
Meristogenys. The upper beak is entire and M-
shaped in the tadpoles of Huia, and divided in
those of Meristogenys.
tire Sunda region and Yunnan (China).
It includes more or less slender, long-legged frogs
which are capable of very long leaps and perfectly
adapted to a life in mountain stream habitats. The 1st
finger is of equal length, or slightly longer, than the
2nd. The tips of the fingers and toes are dilated, the
toes are connected by almost fully webbings. The
length of the tibia is > 0.7 times the SVL. Like those
of Meristogenys, the tadpoles of Huia are adapted
to rapidly flowing streams by a large ventral sucker
disk (gasteromyzophorous).
The genus is represented on Borneo by only one
species which occurs nearly all over the island
(Sabah, Sarawak, and Kalimantan).

136

Huia cavitympanum (BOULENGER, 1893)
Rana cavitympanum BOULENGER, 1893a: 525 – Type locality:
Kina Balu
Rana cavitympanum, BARTLETT 1894: 203; HANITSCH 1900a:
73; BOULENGER 1920: 193; VAN KAMPEN 1923: 211; SMITH
1931: 30; INGER 1962/63: 45
Amolops cavitympanum, INGER 1966: 260, 1978: 313; MATSUI
1979: 337; MANTHEY 1983: 22; INGER & STUEBING 1992: 42;
MALKMUS 1994b: 224
Huia cavitympanum, YANG 1991: 29; MALKMUS 1992a: 118;
HOFFMANN 1995b: 9; INGER, STUEBING & TAN 1996: 363; INGER
& TAN 1996a: 559; LAKIM et al.1999: 32; INGER et al. 2000:
16
Diagnosis
A moderately large terrestrial frog with a relatively
contrasting colour pattern. Its sunken tympanum
makes this species unmistakable.
Fig. 132 Huia cavitympanum from Sayap, Sg. Kemantis 1100 m.
Ranidae
Description
Males 42-52 mm, females 68-80 mm; TL/SVL
0.73-0.76; habitus slightly stout; head longer than
wide; vomerine teeth present; nostrils mid-way
between eyes and tip of snout, or slightly closer to
the latter; canthus rostralis slightly rounded; tym-
panum visible, in a depression on the side of head;
distinct, strongly angled supratympanic fold be-
tween eye and axilla; 1st finger longer than 2nd; fin-
ger tips dilated, outer ones twice as wide as others;
diameter of tip of 3rd finger equal to that of tympa-
num; marginal disk folds present; webbing limited
to basal portion of fingers or absent; weak super-
numerary metacarpal tubercle present; tips of toes
dilated and rounded, slightly smaller than finger
tips; toes fully webbed; small, oval inner metatar-
sal tubercle present; outer metatarsal tubercle small
or absent; dorsolateral folds present; dorsal skin
© P. Hoffmann
137
Ranidae
smooth, lateral with numerous oval glands, smooth
on throat and chest; ventral skin slightly granular.
Males with yellowish nuptial pads on 1st metacar-
pal, and a pair of subgular vocal sacs in gular pock-
ets.
Coloration above chocolate-brown; light dorsola-
teral stripe from behind the eye; sides and upper
sides of limbs yellowish brown with dark brown
spots or cross bars; venter white to yellowish or-
ange; iris light golden above, cinnamon-coloured
below; lips with dark spotting.
Body of tadpoles oval, flattened; head-body-length
up to 25 mm, total length up to 70 mm; eyes and
nostrils positioned dorsally; distance between nos-
trils larger than distance between nostril and eye;
distance between nostril and tip of snout almost
twice the distance between the nostrils; lips form a
large oral disk and lack papillae; labial teeth rows
formula 8(3-9)/5(1) to 12(4-12)/6(1); both beaks
entire; ventral sucker disk covering almost the en-
tire belly; caudal fin limited to posterior two thirds
of tail; coloration dorsally and laterally very dark,
Genus Ingerana DUBOIS, 1986
The genus Ingerana comprises several diurnal and
nocturnal, relatively small species which are rec-
ognized by their finger and toes disks being spatu-
late in shape. The toes are not fully webbed to the
disks. The larvae of these frogs are unknown.
On Borneo, a single species is represented, which
is known from Sarawak, Brunei, Sabah, and Mt.
Kinabalu.
Ingerana baluensis (BOULENGER, 1896)
Cornufer baluensis BOULENGER 1896b: 449 – Type locality:
Mount Kina Balu, North Borneo
Cornufer baluensis, VAN KAMPEN 1923: 241; SMITH 1931: 30
Micrixalus baluensis, INGER 1966: 255, 1978: 313; MANTHEY
1983: 22; INGER & STUEBING 1992: 42
Ingerana baluensis, MALKMUS 1994b: 244; INGER , STUEBING &
TAN 1996: 363; INGER & TAN 1996a: 559; INGER et al. 2000:
16
138
marbled lighter; cream-coloured ventrally. Tadpoles
at an advanced stage of development show a light
line on the tibia.
Ecology/Ethology
Huia cavitympanum is a nocturnal terrestrial frog
inhabiting the banks of fast-flowing streams in pri-
mary forests (250 to 900 m a.s.l.). Here, it is com-
monly found perched on rocks or hiding beneath
them. Adults wander widely through the forests
(INGER & STUEBING 1997). The males call individu-
ally. Tadpoles live in the strongest currents of me-
dium-sized streams, clinging to bedrock or boul-
ders (m: 1a, b).
Call: The call consists of a weak, high-pitched chirp.
Distribution
Endemic to Borneo with records from Sabah,
Sarawak, and Kalimantan. Mt. Kinabalu: Sg.
Wariu, Sg. Kemantis, Sg. Kipungit, Sg. Langanan,
Sg. Mamut.
Diagnosis
A small terrestrial frog with conspicuous spatulate
finger and toe tips.
Description
SVL males 20-25 mm, females 24-30 mm; TL/SVL
0.48-0.56; head wider than long; snout rounded;
vomerine teeth present; canthus rostralis rounded;
diameter of tympanum 40-50% of eye diameter;
supratympanic fold present; 1st finger slightly
longer than 2nd; finger tips spatulate, narrower than
tympanum; marginal disk folds present; webbings
between fingers limited to the bases; no supernu-
merary metacarpal tubercles; tips of toes spatu-
late, wider than the dilations of the fingers;
webbings between the toes covering about their
entire lengths; inner metatarsal tubercle oval, no
outer one; dorsolateral folds absent; skin granu-
lar; males with subgular vocal sacs, but no nup-
tial pads.
Dorsal coloration light reddish brown with obscure
dark markings; light dorsolateral stripe; limbs pale

Fig. 133 Ingerana baluensis from Sg. Kadamaian, 900 m.
brown with dark cross bars; venter cream-coloured;
throat, more rarely also the chest, spotted or mar-
bled with brown; lips striped; iris golden brown
with fine black reticulation.
Ecology/Ethology
Ingerana baluensis prefers moderate altitudes of
montane forests where it is primarily found along
mountain streams. Here, the frogs can be discov-
ered in rock crevices with spring water seepage on
gravel banks, such as the 6-8 m wide Sg. Kada-
maian. Their eggs are large (2.2-2.5 mm in diam-
eter) and unpigmented. The frogs are, however,
occasionally also found far away from water. They
are excellent climbers on the vertical rock faces
and capable of long leaps.
Distribution
Endemic to Borneo, the frog has been recorded
from Sabah, Sarawak, and Brunei. Mt. Kinabalu:
Poring, Sayap, Sg. Kadamaian above Kiau.
Ranidae
© M. Schroth
Genus Limnonectes FITZINGER, 1843
The genus Limnonectes comprise mainly of noc-
turnal, moderately to very large terrestrial frogs that
were previously assigned to the genus Rana.
The 2nd finger is never longer than the 1st. In con-
trast to the genus Rana, the finger tips are only
slightly or not dilated. Limnonectes is further dis-
tinguished from Rana by the absence of marginal
disk folds on the finger tips (except in the subgenus
Elachyglossa ANDERSSON, 1916) and the presence
of well-developed odontonoid processes at the
lower jaw of male specimens. They mostly lack
advertisement calls and often also lack dorsolateral
folds which are often distinctly developed in Rana.
These frogs do not usually wander far from water,
but there are exceptions, such as L. finchi and L.
palavanensis.
The genus Limnonectes is present on Borneo with
12 species and thus forms one of the most diverse
139
Ranidae
genera of Bornean anurans. Mt. Kinabalu is home
to four of these species.
Taxonomic remark: The monophyly of the genus
Limnonectes, as previously proposed by DUBOIS
(1986 “1987”, 1992), was often questioned. Based
on morphological and molecular data, EMERSON et
al. (2000) showed that at least L. cancrivora and L.
Key to the Bornean species of Limnonectes
1 Upper eyelid smooth ....................................... 2
• Upper eyelid with spiked tubercles ...................
......................................................... malesianus
2 Tympanum obscured by skin ........................... 3
• Tympanum clearly visible or only partially ob-
scured ............................................................... 5
3 Toes not fully webbed ..................................... 4
• Toes fully webbed .................................... kuhlii
4 Dorsolateral fold absent; lemon-yellow below ...
............................................................... laticeps
• Interrupted dorsolateral fold present; white be-
low ...................................................... rhacodus
5 Tympanum fully visible ................................... 6
• Upper posterior third of tympanum obscured by
skin ..................................................... asperatus
6 Lower jaw with two tooth-like bony projections
near the symphysis .......................................... 7
• Lower jaw without projections ..................... 11
Tab. 13 Distribution of Limnonectes-species on Borneo.
140
limnocharis do not represent the actual sister group
of the remaining taxa from the genus Limnonectes
in their analyses. Therefore it seems to be likely to
place these two species in the genus Fejervarya, as
previously proposed by DUBOIS & OHLER (2000)
and actually accepted by INGER et al. (2000) for L.
limnocharis (see also genus Fejervarya section).
7 Dorsolateral fold absent .................................. 8
• Interrupted dorsolateral fold present ..... ibanorus
8 Toes fully webbed ............................................. 9
• Toes not fully webbed ................................... 10
9 Canthus rostralis distinct ................... leporinus
• Canthus rostralis indistinct ...................... ingeri
10 SVL less than 55 mm, males with vocal sac .....
........................................................ kenepaiensis
• SVL greater than 55 mm, males without vocal
sac ............................................... paramacrodon
11 Dorsal skin smooth, except for reverse V-shaped
marking ........................................ palavanensis
• Dorsal skin not smooth ............................ finchi

Fig. 134 A male of Limnonectes finchi from Poring is carrying tadpoles.
Limnonectes finchi (INGER, 1966)
Rana microdisca finchi INGER, 1966: 222 – Type locality:
Kalabakan, Tawau District, Sabah
Rana finchi, INGER & STUEBING 1992: 42; INGER, STUEBING &
TAN 1996: 363; MALKMUS 1996c: 293; LAKIM et al. 1999: 32
Limnonectes finchi, INGER et al. 2000: 16
Diagnosis
A small ground-dweller with a conspicuously
granular back, weakly dilated toe tips, and not fully
webbed toes.
Description
Males 32-39 mm, females 39-45 mm; TL/SVL
0.53-0.61; habitus rather slender; head slightly
longer than wide; vomerine teeth present; tympa-
num visible, its diameter ca. 2/3 of that of eye; 1st
and 2nd finger equal in length; finger tips rounded;
toe tips relatively weakly dilated into disks; web-
bing formula 1(0), 2(0/0), 3(0/0), 4(2-3/2-3), 5(0);
Ranidae
© P. Hoffmann
outer metatarsal tubercle absent; dorsolateral folds
interrupted; dorsal skin with numerous, usually lon-
gitudinal tubercles; a reverse V-shaped dermal el-
evation on back behind the shoulder girdle.
Dorsal coloration grey to greyish brown with dark
spots; venter white to yellowish; throat speckled;
lips striped; light interorbital band.
Head-body-length of tadpoles up to 8 mm, total
length up to 25 mm; tails of tadpoles with dark
crossbars; labial teeth rows formula 2(2)/3(1).
Ecology/Ethology
Limnonectes finchi is a nocturnal ground-dwell-
ing frog of primary and secondary rain forests.
The males occupy calling sites beneath leaf litter
on the ground. Most of the time the frogs live far
away from any water bodies which they access
only for the purpose of releasing tadpoles. The
about 50 eggs are 2 mm in diameter, with a dark
brown and a light brown hemisphere. They are
deposited beneath dead leaves on the forest floor
141
Ranidae
within a clutch. The males guard the clutch and
transport the larvae to water bodies once they have
reached stage 24 or 25 (following GOSNER 1960).
Up to 30 tadpoles of 8-9 mm in length may be
transported in one batch. A male was discovered
near Poring carrying seven tadpoles to an ephem-
eral pond next to the road along the boundary of
Kinabalu Park. This pond is fed only by a small
temporary trickle of water, and its water was
murky and practically stagnant, its banks were
muddy and overgrown to the waterline with low
bushes and grasses. Neither the males nor the tad-
poles show any morphological adaptations for this
type of larval translocation. Free-swimming tad-
poles were also found in rainwater puddles on the
forest floor (m: 2b). Adults feed on small insects
and spiders.
This species’ altitudinal distribution ranges from
200 to 1150 m, but the population densities are
higher at low elevations.
Call: The call consists of a short trill.
Distribution
Endemic to Sabah. Mt. Kinabalu: Poring.
Limnonectes kuhlii (TSCHUDI, 1838)
Rana kuhlii TSCHUDI, 1838: 40 – Type locality: Java
Rana paradoxa MOCQUARD, 1890: 148 – Type locality: Kina
Balu (syn. after BOULENGER 1891)
Rana kuhlii, BOULENGER 1920: 62; VAN KAMPEN 1923: 178; SMITH
1931: 30; INGER 1954a: 250, 1956: 403, 1962/63: 46, 1966:
196, 1978: 313; MATSUI 1979: 325; INGER 1985: 48; INGER &
STUEBING 1992: 42; WONG 1994: 32; HOFFMANN 1995b: 11;
INGER & STUEBING 1997: 140; LAKIM et al. 1999: 33
Limnonectes kuhlii, MALKMUS 1994b: 225, 1996c: 285; INGER,
STUEBING & TAN 1996: 363; INGER & TAN 1996b: 75; INGER
et al. 2000: 16
Dicroglossus kuhli, MANTHEY & DENZER 1982: 13; MANTHEY
1983: 22; MALKMUS 1985: 8, 1987: 280, 1988c: 176, 1988d:
7, 1989: 186, 1991a: 30
Rana (Limnonectes) kuhlii, MALKMUS 1996c: 285
Taxonomic remark: Limnonectes kuhlii is consid-
ered a species complex. Morphological and mo-
lecular evidence by KOSUCH & OHLER (unpublished)
indicates that L. kuhlii from Mt. Kinabalu is not
conspecific with neither topotypical (Javanese)
142
specimens nor those from Kalimantan, Sumatra and
the Asian mainland. A second indication for the
existence of a species complex are the results of a
molecular study of EMERSON et al. (2000), refer-
ring to L. kuhlii from Sabah, Brunei, Java and Tai-
wan. Further analyses are required to demonstrate
if those frogs from Mt. Kinabalu represent a new
species or refer to the currently synonymized taxa
Limnonectes conspicillatus (GÜNTHER, 1872, type
locality Matang, Sarawak) or L. paradoxa (MOC-
QUARD, 1890, type locality Kina Balu, Sabah). Com-
parative studies of and with these two taxa still have
to be undertaken.
Diagnosis
A moderately large to large ground-dweller, rarely
observed outside water. It has a stout build, is dor-
soventrally compressed, and has eyes which are
directed upward in a conspicuous manner.
Description
Males 44-74 mm, females 50-67 mm; habitus
stout; limbs short and muscular; head wide, snout
slightly rounded; eye directed upward; lower jaw
with two tooth-like bony protrusions near the
symphysis; males with vomerine teeth; tympa-
num invisible; supratympanic fold distinct,
slightly arched; 1st finger longer than 2 nd; finger
tips rounded; no marginal disk folds; 2nd and 3 rd
finger with dermal fringes; subarticular tubercle
oval, but not elongated; no supernumerary meta-
carpal tubercles; toe tips disk-like, weakly di-
lated; toes fully webbed; inner metatarsal tuber-
cle oval, shorter than 1 st toe; outer metatarsal
tubercle absent; dorsolateral folds absent; skin
above coarse, almost smooth below. Males with-
out vocal sacs, larger than females, their bony
protrusions in the lower jaw also larger, and the
head wider; no nuptial pads, but a differently
coloured marking of their respective positions
on the 1st finger.
Coloration variable and differing regionally; dor-
sum yellowish, reddish, brownish, or greenish,
usually with dark spots, and always with a dark
interorbital band; venter cream-coloured, often
with small grey spots; specimens with and others
without a white dorsal stripe are found synto-
pically.

© P. Hoffmann
Fig. 135 Limnonectes kuhlii from the Headquarters,
1550 m.
Body of tadpoles oval, slightly flattened; brown
with irregular dark spots; tail tapering to a point,
roughly spotted in its first half, in the second half
with four stripes which are not sharply delineated;
head-body-length up to 14 mm, total length up to
38 mm; labial teeth rows formula 2(2)/3(1).
Ecology/Ethology
Limnonectes kuhlii is basically a nocturnal
ground dweller, but not exclusively so. It may
also be found active on day climbing up to 50 cm
high in vegetation. The frogs are almost always
encountered in or near water, in primary forests
as well as in the surrounding of human settle-
Fig. LXII Oral disc of Limnonectes kuhlii (Sg. Kemantis,
1000 m).
R. Malkmus
Ranidae
© R. Malkmus
Fig. 136 Male of Limnonectes kuhlii from the Head-
quarters, 1550 m.
ments. They show a predilection for the banks
of water bodies with moderate to weak currents,
where they often reside on the lee-side of boul-
ders or fallen logs, but also in ditches and ca-
nals. A morphological peculiarity is a hardened
spinous process at the end of the terminal
phalange of at least the 3rd toe which is used in
defense. By lateral movements it will pierce the
skin of an attacker, and this also applies to the
hand of a collector (comp. MALKMUS 1996, MAL-
KMUS , B RÜHL & E LTZ 1999).
The eggs are deposited in the water and attached to
vegetation, e.g. to overhanging tufts of grass. They
hatch after about 10 days. Tadpoles are primarily
Fig. 137 Tadpole of Limnonectes kuhlii from Sg.
Kemantis, 1000 m.
© M. Schroth
143
Ranidae

found in calm sections of streams near the banks,
in residual ponds, and seepages (m: 1e, f, h, i;
2a).
The altitudinal distribution ranges from sealevel to
1750 m a.s.l., but the species becomes very rare
above 1600 m.
Call: Specimens at low elevations do not call.
“Males living at higher elevations at Mt. Kinabalu
emit a soft tweet” (INGER, STUEBING & TAN 1997).
Specimens observed near the Headquarters often
emit dog-like barking and growling sounds.
Distribution
(see the introductory taxonomic comments) India,
southern China, Southeast Asia, Greater Sunda Is-
lands. Mt. Kinabalu: Sg. Kemantis, Sg. Kinateki,
Sg. Kadamaian, Kenokok, Marei Parei, Kiau,
Tenompok, Bundu Tuhan, Sg. Silau-Silau, Sg.
Liwago, Kamborangoh (up to 1750 m), Pinokok,
Poring, Sg. Mantukungan, Sg. Kipungit, Sg.
Langanan, Sg. Mamut, the record “Kamborangoh,
alt. 7.200 feet” (SMITH 1931) is doubtful.
Limnonectes leporinus
(ANDERSSON, 1923)
Rana macrodon var leporina ANDERSSON, 1923: 121, 124 – Type
locality: Tumbang Maruwei and North-eastern Borneo,
Bulungan
Rana macrodon, SMITH 1931: 30; SMEDLEY 1931: 61
Rana blythi (not Rana blythii BOULENGER, 1920), INGER 1966:
162 (part.)
Rana blythi (not Rana blythii BOULENGER, 1920), INGER &
STUEBING 1992: 42; HOFFMANN 1995b: 10; INGER , STUEBING
& TAN 1996: 363
Limnonectes blythi (not Rana blythii B OULENGER, 1920),
MALKMUS 1994b: 226
Limnonectes leporina INGER & TAN 1996a: 560; INGER et al.
2000: 16
Rana leporina LAKIM et al. 1999: 33
biochemical (KOSUCH 1994) and molecular studies
(EMERSON et al. 2000).
Diagnosis
A large, robust terrestrial frog which can easily be
confused with Limnonectes ingeri. L. leporinus
exhibits a dark stripe below the canthus rostralis,
while L. ingeri does not. L. leporinus furthermore
has a narrower head, longer hind limbs, and a more
sharply defined canthus rostralis than L. ingeri.
Description
Males up to 125 mm, females up to 175 mm; ma-
turity on reaching 85-90 mm; TL/SVL 0.52-0.62;
body massive; head longer than wide, snout
pointed; vomerine teeth present; canthus rostralis
sharply defined; tympanum visible; supratympanic
fold present; lower jaw with two tooth-like, bony
protrusions near the symphysis; 1st finger longer
than the 2nd; finger tips rounded; marginal disk folds
absent; toe tips dilated; toes fully webbed; inner
metatarsal tubercle oval, no outer one; dorsolateral
folds absent; skin smooth. Males without vocal
sacs; with a larger head than females and larger
mandibular protrusions.
Light brown to reddish dorsally, juveniles with dark
pattern; limbs with dark crossbars; whitish ventrally;
throat finely dotted; dark interorbital band present;
lips banded with black; a dark stripe below the can-
thus rostralis and from eye to tympanum.
Body shape of tadpoles oval, about 0.75 times as
wide as long, slightly flattened; eyes and nostrils
Fig. 138 Limnonectes leporinus from Sg. Sasapang.

Taxonomic remark: INGER & TAN (1996a) exam-

ined the type of leporinus ANDERSSON, 1923 and
found it to be identical to all Bornean frogs pre-
viously named blythii BOULENGER, 1920, but not
© R. Malkmus

to be conspecific with blythii from the Malay Pen-

insula. These morphological results have been
confirmed by high genetic differences found in

144
 Ranidae

positioned dorsally, mouth ventrally and subter- Kipungit, Sg. Mamut, Sg. Mantukungan, Sg.
minally; upper lips short; anal tube dextral; tail Sasapan.
muscular; Coloration “graybrown with dark
markings, which form an almost zigzag row of
dark spots on the tail” (INGER & STUEBING 1997). Limnonectes palavanensis
Head-body-length before metamorphosis ca.
36 mm, SVL of freshly metamorphosed froglets (BOULENGER, 1894)
9-11 mm.
Rana palavanensis BOULENGER 1894a: 85 – Type locality:
Palawan, Philippines
Ecology/Ethology
Rana palavanensis, BOULENGER 1920: 59; VAN KAMPEN 1923:
Limnonectes leporinus is a nocturnal ground- 182; SMITH 1931: 30; INGER & STUEBING 1992: 42; INGER,
dweller inhabiting the banks of streams and rivers. STUEBING & TAN 1996: 363; HOFFMANN 1994: 223, 1995b:
Resting places are normally not in the water. The 11; INGER & TAN 1996a: 563; LAKIM et al. 1999: 32
frogs can be found at night sitting among leaf litter Rana microdisca palavanensis, INGER 1954a: 250, 1966: 222,
1978: 313; MATSUI 1979: 333
near their water body, on banking rocks, or on fallen Hylarana microdisca palavanensis, MANTHEY 1983: 22
logs. They spend the day hidden in moist shelters, Limnonectes palavanensis, MALKMUS 1994b: 244; INGER et. al.
e.g. beneath rocks on the banks. They prefer pri- 2000: 16
mary rain forests, but also accept biotopes in sec-
ondary forests. This species is not present in culti- Diagnosis
vated land where it is replaced by L. ingeri. A small terrestrial frog with a characteristic dorsal
The males excavate spawning craters with diam- pattern in the form of a reverse V.
eters of 30 to more than 50 cm in the gravel sedi-
ment of shallow sections of rain forest rivers in Description
which the females deposit their eggs at night. The Males 27-30 mm, females 35-40 mm; habitus rather
eggs are uniformly yellowish grey (vs. light with a slender; vomerine teeth present; diameter of the
dark hemisphere in L. ingeri). The males have not tympanum ca. 2/3 of that of the eye; 1st and 2nd
really an advertisement call, but both sexes emit finger of equal lengths; finger tips rounded; mar-
equal-sounding, deep, soft sounds during the mat- ginal disk folds absent; toe tips disk-like and
ing season whereby, according to EMERSON & INGER slightly dilated; outer metatarsal tubercle absent;
(1992), the sounds emitted by the males must
be regarded as responses to female calls.
Tadpoles may be found in a variety of habitats Fig. 139 Limnonectes palavanensis from Sayap, Sg. Wariu,
although there appears to be a predilection for ca. 850 m.
leaf litter sediment whose layered structure pro-
vides ideal situations (m: 1d, e). Strong currents
are clearly avoided. The larvae are generalized
suspension feeders with stomach analyses re-
vealing mainly algae (blue algae, but also
euglenoids and diatoms), plant matter, and fun-
gal hyphens. Adult specimens prefer larger prey
items including crabs and even other frogs.
Its altitudinal distribution extends from 65 to
800 m, but the frogs are most numerous be-
low 300 m a.s.l.
© P. Hoffmann

Distribution
Endemic to Borneo with records from Sabah,
Brunei, and Kalimantan. Mt. Kinabalu: Sg.

145
 Ranidae
© P. Hoffmann
Fig. 140 Limnonectes palavanensis from Sayap, Sg. Wariu, ca. 850 m.
toe webbing 1(0), 2(0/0), 3(0/0), 4(2-3/2-3), 5(0);
dorsolateral folds present; skin above smooth ex-
cept in the area of the reverse V-marking.
Dorsal coloration medium brown to reddish
brown with continuous, light dorsal stripes; dorsal,
reverse V-marking dark; flanks light grey to grey-
ish brown, with a tinge of orange; lower side of
head greyish white; chest, belly, and lower sides
of legs usually lemon-yellow; dark spots in pos-
terior dorsolateral region; coloration of the limbs
corresponding to that of the flanks, with dark
brown crossbars; light and dark interorbital bands;
lips striped with black, usually spotted with yel-
low or white.
Tadpoles small; tail tapering abruptly near the end
to a slim tip; coloration light or dark brown with
darker dots; tail speckled with black; head-body-
length up to 8 mm; total length up to 30 mm; labial
teeth rows formula: 2(2)/3(1).
146
Ecology/Ethology
Limnonectes palavanensis is a nocturnal ground
dweller of the primary rain forests, primarily in
mountainous country. The frogs are often encoun-
tered in the leaf litter layer or in low vegetation (up
to 20 cm high) far away from the nearest water
body. The males occupy advertisement sites be-
neath leaf litter on the forest floor. Adults prey upon
ants, termites, and other small terrestrial inverte-
brates.
Oviposition takes place on land beneath leaf litter.
The clutches are comparatively small with a maxi-
mum of 35-50 eggs and the eggs being rather large
in comparison to both the body size of the females
and other ranid frogs. Eggs up to 2.6 mm in diam-
eter, a light and a dark hemisphere whereby the
light one is lighter than in the eggs of L. finchi.
Parental care and transport of larvae like in L. finchi.
Free-swimming tadpoles was observed in small,
 Ranidae

shallow pools of temporary streams

and other shallow accumulations of
water (m: 1f, i).
The altitudinal distribution ranges
from sea level to 1300 m.

Call: Series (duration 5-6 s) of 20

or more slightly pulsed notes; note
duration 50-70 ms; internote inter-
val 140-250 ms; frequency modu-
lation in each note rising from 2.4
to 3.3 kHz with harmonics at 9-
10 kHz.

Distribution
Palawan (Philippines) and Borneo.
Mt. Kinabalu: Kiau, Sayap, Bundu
Tuhan.

Fig. LXIII Sonagram and oscillo-

grams of Limnonectes palavanensis
(Gunung Mulu, 150 m; 8.00 p.m.)

and dorsolateral folds are all present. The skin is

Genus Meristogenys YANG, 1991
The distribution of the genus Meristogenys is lim-
ited to Borneo.
It was not before 1991 that YANG split the compos-
ite genus Amolops into Amolops, Huia, and Meris-
togenys, mainly on the basis of larval differences
(see also Genus Huia).
The genus Meristogenys comprises small (espe-
cially small males in many species) to large, mod-
erately slender, and long-limbed frogs. The head is
longer than wide with a rounded snout. The eyes
are large, and the tympanum is visible. The tibia
length corresponds with more than 0.7 times the
SVL. The 1st finger is generally longer than the 2nd;
the tips of the fingers and toes are dilated, with the
diameters of the finger tips being smaller than the
diameter of the tympanum. The toes are fully
webbed, and marginal disk folds, vomerine teeth,
finely granular to smooth.
The reproductive biology could be studied in de-
tail in M. cf. orphnocnemis (MALKMUS 1998). Pairs
migrate in amplexus to a mountain brook where
the eggs are deposited under water on smooth rocks
exposed to the current. They arrange themselves
spontaneously after spawning to form a single-lay-
ered egg disk in which the eggs sit side by side
connected to one another by the jelly mass. After
hatching the larvae form a school which remains
stationary in an unusual density next to the now
empty egg disk before they eventually disperse af-
ter 3-5 days. It is still unknown whether the repro-
ductive biology of Meristogenys is different from
that of Amolops or Huia.
Like in the genus Huia, the tadpoles possess a
large, circular, ventral sucker as an adaptation to
their life in streams with strong currents. The head
is substantially broadened and flattened below.

147
 Ranidae
© R. Malkmus
© R. Malkmus
From top to base: Meristogenys cf. orphnocnemis
Fig. 141 A pair from Sg. Liwago after egg deposition.
Fig. 142 Tadpoles form a school after hatching.
Tab. 14 Distribution of Meristogenys-species on Borneo.
148
© M. Schroth
Fig. 143 Tadpole of Meristogenys cf. kinabaluensis.
They exhibit an M-shaped, keratinized upper beak
which is costate distally. The V-shaped lower beak
may sometimes also be divided. The muscular tail
is 1-1.5 times as long as the body. The tadpoles
possess small venom glands on the body and tail
whose arrangements are indicative of the species.
The larvae of the species Huia cavitympanum dif-
fer from those of the species of Meristogenys by
having a larger number of rows of denticles on the
anterior labium and by the structure of the lower
beak.
In contrast to most of the other anuran genera the
tadpoles of the species of Meristogenys are more
easily distinguished than the adults whose body
shapes and colour patterns are very similar. As the
morphological points of difference for the adults
have often been used in the literature in inaccurate
or contradictory manners, we shall refrain from
adding an identification key to the genus here.
Of the 8 species in this genus, 4 occur on Mt.
Kinabalu.

Meristogenys amoropalamus
(MATSUI, 1986)
Amolops amoropalamus MATSUI, 1986: 623 – Type locality:
Bank of Sg. Pa Riman, 1300 m alt., Gunung Tapai Sai, Krayan
Country, Sarawak border of East Kalimantan
Amolops sp. A, MATSUI 1979: 340
Amolops amoropalamus, MALKMUS 1989: 190, 1994b: 227,
1996c: 283
Meristogenys amoropalamus, YANG 1991: 33; MALKMUS 1992a:
117, 1994c: 89, 1998: 22; INGER , STUEBING & TAN 1996: 363;
LAKIM et al. 1999: 33; INGER et al. 2000: 16
Diagnosis
A moderately large terrestrial frog with a conspicu-
ous iris which is divided in brown and golden sec-
tions; distinguished from M. kinabaluensis by the
outer metatarsal tubercles, from M. orphnocnemis
by the not fully webbed toes, and by the smaller
tympanum from M. whiteheadi (3/5 or more of eye
diameter).
Description
Males 30-40 mm, females > 60 mm; TL/SVL 0.72;
habitus moderately slender; head longer than wide;
Fig. 144 Meristogenys amoropalamus from Sg. Liwago, 1500 m.
Ranidae
snout slightly pointed; vomerine teeth small; nos-
trils closer to tip of snout than to eyes; canthus
rostralis distinct; tympanum about 80% of eye di-
ameter; supratympanic fold present; 1st finger lit-
tle longer than 2nd; dilation of finger tips about 1/3
of tympanum diameter; marginal disk folds present;
subarticular tubercle oval, with weak, supernumer-
ary tubercles on outer two fingers; toe tips dilated;
toe webbing 1(0), 2(0/0), 3(0/0), 4(2/2), 5(0); in-
ner and outer metatarsal tubercles present, the in-
ner one being long and narrow; dorsolateral folds
present; dorsal and lateral skin slightly granular,
smooth ventrally; males with small nuptial pads on
1st finger; a pair of vocal sacs in gular pockets.
Dorsal coloration light brown with dark dotted pat-
tern; ventral side whitish with dark dots, particu-
larly on hind limbs which also sport dark cross-
bars; tibia dusky ventrally; upper lip with light
spots, lower lip with dark bands; dark band from
eye to behind the tympanum; iris golden in upper
part, reddish brown in lower.
Habitus of tadpoles oval and flattened, with large
ventral sucker; eyes situated dorsolaterally; nostrils
© M. Schroth
149
 Ranidae
much closer to eyes than to tip of snout; margin
of upper lip smooth, lower lip with papillae; la-
bial teeth rows formula 7(4-7)/6(1) at stage 28
with a tendency to have increased to 8(5-8)/10
(1) by stage 38; upper beak divided; dorsal root
of the caudal fin at end of body, ventrally at the
end of first third of tail; groups of glands scat-
tered over the body with each of the granular
openings visible; more glands on bases of both
fins; dorsal coloration dark grey, light grey
ventrally.
Ecology/Ethology
Meristogenys amoropalamus is a nocturnal ground
dweller of the primary forests closely associated
to rapidly flowing streams (m: 1a, b, c, e). Its
altitudinal distribution ranges from 1300 to 1800 m
a.s.l.
Distribution
Endemic to Borneo with records from Sabah,
Sarawak, and Kalimantan. Mt. Kinabalu: Sayap,
Sg. Penataran, Sg. Silau-Silau, Sg. Liwago (1500 m),
Sg. Mesilau (Mesilau Cave, 1800 m).
Fig. 145 Meristogenys kinabaluensis from Sayap, Sg. Kemantis, ca. 1050 m.
© P. Hoffmann
150
Meristogenys kinabaluensis
(INGER, 1966)
Amolops kinabaluensis INGER, 1966: 266 – Type locality: Kiau,
Mount Kina Balu, Sabah
Rana whiteheadi, (not Rana whiteheadi BOULENGER , 1887)
MOCQUARD 1890a: 146 (part); VAN KAMPEN 1923: 210; SMITH
1931: 30; WHITEHEAD 1893: 121
Amolops kinabaluensis, INGER 1978: 313, 1985: 65; MATSUI
1979: 339; MANTHEY 1983: 22; MALKMUS 1987: 282, 1989:
189, 1991a: 28, 1994b: 227, 1996c: 283; INGER & STUEBING
1992: 42; WONG 1994: 32; INGER, STUEBING & TAN 1996: 313
Meristogenys kinabaluensis, YANG 1991: 35; MALKMUS 1992a:
117, 1998: 22; HOFFMANN 1995b: 9; INGER & TAN 1996a: 560,
1996b: 76; LAKIM et al. 1999: 33; INGER et al. 2000:16
Diagnosis
The largest species of Meristogenys. It differs from
all other species of the genus by the absence of
outer metatarsal tubercles.
Description
Males 58-68 mm, females 75-93 mm; TL/SVL
0.58-0.65 in males, 0.49-0.51 in females; head
 Ranidae

wide; body and limbs slender; vomerine

teeth present; nostrils closer to tip of snout
than to eyes; interorbital distance smaller
than diameter of upper eyelid; weak
supratympanic fold between eye and axilla;
1st finger longer than 2nd; finger tips dilated,
about twice width of finger; marginal disk
folds present; no webbings between fingers;
subarticular tubercle oval, with supernumer-
ary tubercles on outer three fingers; toe tips
dilated; toes fully webbed; inner metatarsal
tubercle oval and about 1/3 of length of 1st

© M. Schroth
toe; outer metatarsal tubercles absent; dor-
sal skin smooth, with narrow, interrupted
dorsolateral folds; ventral skin smooth save
for posterior abdomen. Males have statisti- Fig. 146 Meristogenys kinabaluensis from Sg. Liwago, ca. 1450 m.
cally larger tympanae and a pair of vocal sacs
in gular pockets.
Females dark brown dorsally, with small, darker branches along the banks of fast-flowing streams
spots; males more contrasting, yellowish green with (m: 1a, b, c, e). Eggs are 2-2.5 mm in diameter;
an irregular, dark pattern; limbs with dark bars; with a pigmented hemisphere.
ventral coloration yellowish white with a share of The altitudinal distribution ranges from 650 to
brown, particularly on throat; lower sides of legs 2000 m a.s.l.
pale brown.
Habitus of tadpoles oval and flattened, with a large Call: The call consists of a high-pitched squeak,
ventral sucker; head-body-length up to 25 mm, to- often emitted at long intervals, i.e. several minutes.
tal length up to 67 mm; labial teeth rows formula
6(4-6)/6(1) to 7(4-7)/7(1); upper beak divided, Distribution
lower one entire. Body mostly brown or black on Endemic to Borneo with records from western Sabah
upper sides; fins dusky, with a fine veining; groups and northern Sarawak. Mt. Kinabalu: Sayap, Sg.
of glands on upper surface of body, but not on tail. Panataran (4800 ft.), Marei Parei, Sg. Kadamaian,
Sg. Kinateki, Kiau, Sg. Tibabar (1800 m), Sg. Silau-
Ecology/Ethology Silau, Sg. Liwago (1400-1500 m), Sg. Mesilau
Meristogenys kinabaluensis is nocturnal. The frogs (1800 m), Sg. Kipungit, Sg. Langanan, Sg. Mantu-
are mostly found perched on boulders or thick kungan.

Fig. LXIV Tadpole of Meristogenys kinabaluensis from Sg. Liwago showing position of groups of glands.

R. Malkmus

151
Ranidae
Meristogenys orphnocnemis
(MATSUI, 1986)
Amolops orphnocnemis MATSUI, 1986: 625 – Type locality:
Kampong Bundu Tuhan, near Kinabalu, 990 m alt., Sabah
Amolops jerboa (not Rana jerboa GÜNTHER , 1872), MATSUI
1979: 338 (part)
Amolops from Kinabalu “small”, INGER & GRITIS 1983: 6
Amolops orphnocnemis, INGER & STUEBING 1992: 42; MALKMUS
1994b: 244, 1996c: 283
Meristogenys orphnocnemis, YANG 1991: 36; WONG 1994: 32;
HOFFMANN 1995b: 10; INGER, STUEBING & TAN 1996: 363;
INGER & TAN 1996a: 560; MALKMUS 1999: 15; LAKIM et al.
1999: 33; INGER et al. 2000: 16
Meristogenys jerboa (not Rana jerboa G ÜNTHER , 1872),
MALKMUS 1992a: 114
Diagnosis
Distinguished from M. whitehead and M. amoropa-
lamus by the fully webbed toes, from M. kinabalu-
ensis by the presence of an outer metatarsal tu-
bercle.
Description
Males up to 43 mm, females larger than 60 mm;
TL/SVL 0.71; habitus slender; head longer than
wide; snout slightly pointed; vomerine teeth
present; canthus rostralis distinct; nostrils closer
to tip of snout than to eyes; 1 st finger slightly
longer than 2 nd; marginal disk folds present;
subarticular tubercle oval; no supernumerary tu-
bercles, but with light spots indicating their posi-
tions on outer two fingers; toe tips dilated; toes
fully webbed; inner metatarsal tubercle oval, outer
one round; skin coarse, with distinct dorsolateral
folds.
Males with nuptial pads on 1st finger and a pair of
vocal sacs in gular pockets; tympanum diameter
greater (0.62 times eye diameter) than in females
(0.42 times).
Dorsal coloration chocolate-brown, lighter later-
ally; ventral side whitish to cream-coloured; limbs
greyish brown above, with darker crossbars.
Habitus of tadpoles oval and compressed dorsov-
entrally, with large ventral sucker; labial teeth rows
formula 5(4-5)6(1) at stage 38 to 5(4-5)/9(1) at
stage 41; upper and lower beaks divided. The oral
disk transforms into a mouth during metamorpho-
sis only after the resorption of the larval tail. This
152
must be viewed as an aid for the larva to withstand
the strong current of its habitat.
Ecology/Ethology
Meristogenys orphnocnemis is both diurnal and
nocturnal with adults inhabiting the banks of fast-
flowing forest streams. The males can often be
seen perched in loose groups at about knee-height
on the vegetation-covered banks next to cascades.
The heavier females are mostly found on the
ground. Juveniles may also be encountered far
away from water on the forest floor or in low veg-
etation. The males call individually or in loose
groups while perched on slightly raised advertise-
ment sites, particularly on the banks of rapids or
small waterfalls.
M. orphnocnemis is the only species of Meristo-
genys whose reproductive biology is known (see
also description of the genus above). Before and
during spawning the sexual dimorphism becomes
evident with the large females carrying the much
smaller males (53-55% of the females’ SVL) in am-
plexus to the spawning sites. Once they have ar-
rived, the female spends some time in carefully se-
lecting a suitable spot by probing the surface with
her hind limbs and ventrum in back-and-forth mo-
tions. The adaptation to spawning in strong cur-
rents (m: 1a, b, c, e) must be viewed as providing
the best possible protection for the larvae from po-
tential predators (such as the larvae of dragonflies
and beetles) and competitors for food, as these pre-
fer the calmer sections of the streams.
The about 800 eggs of a female’s clutch are tightly
arranged in an egg disk and held in place by jelly.
They are attached to a smooth rock surface exposed
to a strong current. The jelly is transparent when
produced, but becomes milky within 1-1.5 hours
after spawning, after the animal poles of the eggs
have turned upwards, and then changes into a bright
whitish blue. The tadpoles hatch after about 24
days, but stay in a dense school for another four
days, slowly dispersing thereafter. When disturbed,
communal larvae immediately sought the contact
with their siblings, while those older than four days
reacted with taking to flight.
The known altitudinal distribution ranges from
190 to 1500 m, after LAKIM et al. (1999) up to
1800 m.
 Ranidae

Call: (+22°C) The call resembles a

muffled crack of a whip. It is emit-
ted sporadically at intervals between
10 s to several minutes; call duration
40 ms; the first half of the call weakly
pulsed, the second distinctly pulsed;
pulse modulation present; frequency
decreasing from 9.5 to 6.5 kHz; har-
monics between 9.5 and 11.2 kHz.

Distribution
Endemic to Borneo with records
from Sabah and Kalimantan. Mt.
Kinabalu: Sayap, Marei Parei, Sg.
Kinateki, Kiau, Sg. Kadamaian, Sg.
Liwago, Mesilau, Bundu Tuhan,
Pinokok, Sg. Mantukungan, Sg.
Mamut, Sg. Kipungit, Sg. Langanan,
Sg. Tananansad, Sg. Sasapan, Sg.
Manggis.

Fig. LXV Sonagram and oscillograms

of Meristogenys orphnocnemis
(Sg. Kipungit, 550 m; 8.00 p.m. +22°C)

amoropalamus by having a smaller tympanum and

Meristogenys whiteheadi being much larger, and from M. kinabaluensis in
(BOULENGER, 1887) having a larger tympanum.

Rana Whiteheadi BOULENGER, 1887b: 96 – Type locality: Mount Description

Kina Baloo, North Borneo
Rana whiteheadi, (part) MOCQUARD 1890a: 146; BARTLETT 1894:
203; HANITSCH 1900a: 73; BOULENGER 1920: 195; VAN KAMPEN
1923: 210; SMITH 1931: 17
Amolops jerboa (not Rana jerboa GÜNTHER, 1872), (part) INGER
1966: 264, 1978: 313
Amolops cf. whiteheadi, MALKMUS 1994b: 227, 1996c: 283
Meristogenys whiteheadi, YANG 1991: 40; INGER, STUEBING &
TAN 1996: 363; INGER & TAN 1996a: 561; LAKIM et al. 1999:
33; INGER et al. 2000: 17
Amolops whiteheadi, INGER & GRITIS 1983: 5
Diagnosis
Three species of this genus, i.e. M. amoropalamus,
M. kinabaluensis, and M. whiteheadi, share the trait
of the 4th toe being webbed up to the distal subar-
ticular tubercle. M. whiteheadi differs from M.
Males up to 64 mm, females up to 93 mm; TL/SVL
0.69-0.70; head about as long as wide; snout round
to slightly pointed; vomerine teeth present; nos-
trils closer to tip of snout than to eyes; canthus
rostralis distinct; tympanum 40-60% of eye diam-
eter; supratympanic fold present; 1st finger as long
as or longer than 2nd; finger tips dilated; marginal
disk folds present; subarticular tubercle very dis-
tinct; toe tips dilated; toes fully webbed except for
4th; inner metatarsal tubercle oval, outer one very
small or even absent; dorsolateral folds present;
skin granular, smoother ventrally, particularly to-
wards throat.
Males with large nuptial pads on 1st finger and a
pair of vocal sacs in gular pockets.

153
 Ranidae
© A. Nöllert
Fig. 147 Meristogenys whiteheadi from Crocker Range NP, Sabah.
Dark greyish brown to dark reddish brown dorsally,
occasionally with darker spots, sometimes with
small light dots; ventral side whitish to cream-col-
oured; a dark streak below canthus rostralis and
supratympanic fold; hind limbs with more or less
indistinct crossbars.
Habitus of tadpoles similar to the preceding spe-
cies; with a large ventral sucker; total length 65-
77 mm; labial teeth rows formula 5(3-5)/5(1);
upper and lower beaks divided.
Ecology/Ethology
Meristogenys whiteheadi is a nocturnal dweller of
the banks of fast-flowing streams (m: 1a, b, c, e)
between 190 and 1500 m altitude, after LAKIM et al.
(1999) up to 1800 m. It is the most common species
on Sg. Liwago (1500 m).
Distribution
Endemic to Borneo with records from Sabah and
Sarawak. Mt. Kinabalu: Kiau, Kenokok, Lobang,
Sg. Liwago, Mesilau, Pinokok, Sg. Kipungit, Sg.
Langanan.
154
Genus Occidozyga
KUHL & VAN HASSELT, 1822
The genus Occidozyga is distributed from Bangla-
desh, Myanmar and Thailand through the Sunda
Islands to the Philippines.
It comprises small, dorsally flattened, and quite
plump frogs with short, muscular hind limbs, which
are very well adapted to their environments with
regard to their colour patterns. The tympanum is at
least partly concealed by skin, the snout is flattened,
and the eyes bulge substantially. The tips of the
toes are slightly dilated to form narrow, roundish
disks. While maxillar teeth are present, vomerine
teeth are absent.
The two Bornean representatives of this genus live
in or on small puddles, seepage springs, or in
swampy areas. Their tadpoles can be found in ac-
cumulations of water of all sorts. The larvae are
quite remarkable as to their body shapes and be-
havioural patterns. They have very slender bod-

ies and tails with the latter being furnished only
with narrow fins. The oral disk is situated termi-
nally, the upper lip is very narrow, the lower horse-
shoe-shaped. There are no denticles, but the up-
per border of the disk has a knob-like structure.
The beaks are heavy and finely serrated. The
spiracle is sinistral. The tadpoles of both Bornean
species are carnivorous feeding on small insects
and their larvae.
The genus Occidozyga is represented on Borneo
by two species which also occur on Mt. Kinabalu.
Tab. 15 Distribution of Occidozyga-species on Borneo.
Occidozyga baluensis (BOULENGER, 1896)
Oreobatrachus baluensis BOULENGER , 1896a: 401 – Type
locality: Mount Kina Balu, North Borneo
Oreobatrachus baluensis, HANITSCH 1900a: 74; VAN KAMPEN
1923: 229
Phrynoglossus baluensis, SMITH 1931: 15
Ooeidozyga baluensis, INGER 1954a: 250, 1956: 401, 1978: 313,
1996: 238; MATSUI 1979: 336; MANTHEY 1983: 23
Occidozyga baluensis, INGER & STUEBING 1992: 42; MALKMUS
1994b: 244; INGER, STUEBING & TAN 1996: 363; INGER & TAN
1996a: 561; INGER et al. 2000: 17
Diagnosis
A small terrestrial frog of stout, compressed habi-
tus, short, muscular hind limbs, and not fully
webbed toes. A faint, U-shaped ridge, open below,
in the middle of the back.
Description
Males up to 26 mm, females up to 35 mm; TL/SVL
0.46-0.57; snout rounded; vomerine teeth absent;
nostrils situated laterally; tympanum partly covered
by skin, ca. 0.5-0.6 times the diameter; supratym-
panic fold present; 1st finger as long as or shorter
than 2nd; finger tips not expanded; ; toe tips dilated;
webbing on 4th toe does not reach toe tip; weakly
developed subarticular tubercles; inner metatarsal
Ranidae
Key to the Bornean species of
Occidozyga
1 Ventral side whitish, faintly dotted or immaculate;
webbing formula 4(0/0) ............................ laevis
• Ventral side spotted; webbing formula 4(2-3/2-3)
............................................................. baluensis
tubercle oval, outer one small or even absent; dor-
solateral folds present; dorsal skin with irregular
glandular ridges, usually a dorsolateral one behind
eyes and an inverse U-shaped one between shoul-
ders; numerous small, round warts on sides; males
with a pair of vocal sacs; yellowish nuptial pads
cover the dorsal and medial surface of 1st finger
between base and terminal phalange.
Dorsal coloration between head and glandular ridge
reddish brown between shoulders, remainder of
back light brown; ventral sides light brown with a
brown or black banded pattern. Specimens with
brown as well as those with entirely black backs
are known from Sayap.
Head-body-length of tadpoles up to 8 mm, total
length up to 26 mm; light brown, with a black streak
between lower anterior margin of eyes and snout,
and a dark, vertical bar below eye; caudal muscle
and low upper fin with small black dots (see also
description of genus).
Ecology/Ethology
This frog is a diurnal and nocturnal ground dweller,
occurring in montane regions as well as in the low-
lands in wet situations among leaf litter, roots, and
on gravel ground. Besides on poorly drained
patches in the forests, it is commonly encountered
155
 Ranidae

Fig. 148 and 149 (below right)

Occidozyga baluensis from a small tributary
to Sg. Kemantis, Sayap, ca. 1100 m.
© M. Schroth

© M. Schroth
© P. Hoffmann

Fig. 150 Occidozyga baluensis from a small tributary to Sg. Kemantis, Sayap, ca. 1100 m.

156
 Ranidae

on the banks of small running waters

and seepage springs (m: 2a). Its altitu-
dinal distribution ranges from 65 to
1370 m.

Call: (+23°C) The note series consists of

hoarse, rasping, distinctly rhythmic and
strongly pulsed elements of 50-300 ms
in duration; internote interval 150-
900 ms; dominant frequency 1-3.3 kHz.

Distribution
Endemic to Borneo with records from
Brunei, Sabah, Sarawak, and Kaliman-
tan. Mt. Kinabalu: Poring, Kiau, Fig. LXVI Sonagram and oscillogram of Occidozyga baluensis
Sayap, Bundu Tuhan, Pinokok. (Gunung Mulu, 150 m; 8.00 p.m. +23°C)

Occidozyga laevis (GÜNTHER, 1859)

Oxyglossus laevis GÜNTHER, 1859 (1858b): 7 – Type locality:

Philippines
Occidozyga laevis, MALKMUS 1994b: 244; LAKIM et al. 1999:
33; INGER et al. 2000: 17

Diagnosis
A small terrestrial frog of stout, compressed habi-
tus with fully webbed toes. This species closely

© U. Manthey
resembles juvenile Limnonectes kuhlii, except that
in O. laevis the inter-ocular distance is equal to or
smaller than the eyelid. Furthermore, O. laevis has
only one tooth-like protrusion on the tip of the lower Fig. 151 Occidozyga laevis from Taman Negara, Penin-
jaw as compared to two in the case of L. kuhlii. sular Malaysia.

Description
Males 20-31 mm, females 25-48 mm; TL/SVL
0.41-0.51; head flattened; vomerine teeth absent;
nostrils situated dorsally to dorsolaterally; can-
thus rostralis strongly rounded or indistinct; tym-
panum concealed by skin and therefore invisible;
supratympanic fold weak, between eye and ax-
illa; length of 1st finger equal to or longer than 2nd
finger tips not dilated; no marginal disk folds; toe
tips dilated, tip of 5th toe only twice as thick as all
others; toes fully webbed; inner metatarsal tuber-
cle shorter than half 1st toe; outer metatarsal tu-
bercle minute or (usually) absent; no dorsolateral
folds; dorsal skin smooth in anterior part, with
some small white rises posteriorly; similar rises
on dorsal and lateral sides of hind limbs, in males
also on throat; skin on flanks with round to oval
warts.
Dorsal coloration grey or brown with a darker pat-
tern. A morph with a white vertebral stripe exists.
Ventral side cream-coloured, immaculate or with
some small dots; ventral side of hind limbs usually
with small dots.
Tadpoles small, very slender; with a long tail and
very low caudal fin; body and margins of the fins
with dark spots; head-body-length up to 10 mm,
total length up to 25 mm.

157
 Ranidae
© U. Manthey
Fig. 152 Ventral pattern of Occidozyga laevis from Taman
Negara, Peninsular Malaysia.
Genus Rana LINNAEUS, 1758
The almost cosmopolitan genus Rana ranks among
the most diverse within the order anura. The frogs
of this genus are heterogeneous morphologically as
well as ecologically. They are in fact so heterogene-
ous that it is impossible to formulate a generalised
description in the same manner as for the other gen-
era. The tips of the fingers and toes are usually dis-
tinctly dilated, marginal disk folds are present. Dor-
solateral folds are often present and distinct.
On Borneo, this genus is represented mainly by
nocturnal terrestrial frogs (although Rana hosii is
Tab. 16 Distribution of Rana-species on Borneo.
158
Ecology/Ethology
Occidozyga laevis is a nocturnal species. It lives
solitarily in slow-flowing water bodies (m: 1f), rice
paddies, and small puddles (m: 2b), and are rarely
encountered outside of water. They feed on insects
and freshwater crustaceans. Altitudinal distribution:
0-500 m.
Call: The call consists of an irregularly spaced se-
ries of single harmonic notes, lasting 0.03-0.08 s,
ended by a lower series of 8-9 short pulses per sec-
ond.
Distribution
From Thailand to the Philippines and Flores, al-
though it is at present not quite certain if it is the
same species throughout. Mt. Kinabalu: Poring.
often found climbing). Most species live con-
stantly, or at least after having reached maturity,
in the vicinity of water. The habitats are quite di-
verse with some species living on the banks of
moderately large to large, others in or on narrow
streams, others again in swamps or in flooded rice
paddies. One species, R. luctuosa, is in contrast a
strict dweller of forest floors which can also be
encountered far away from water. Like the adults,
the tadpoles of the individual species develop in
a variety of types of water bodies and are adapted
accordingly.
The genus Rana is represented on Borneo by 9 spe-
cies, 7 of which also occur on Mt. Kinabalu.
 Ranidae

Key to the Bornean species of Rana

1 Dorsal coloration conspicuously black and yel- 6 Supernumerary metacarpal tubercles on every
low or black and orange ......... signata/picturata metacarpal .......................................................... 7
• Dorsal coloration different from the afore ....... 2 • Supernumerary metacarpal tubercles only on
2 1st finger shorter than 2nd ................................... 3 metacarpals 2-4 ............................ nicobariensis
• 1st finger longer than 2nd .................................... 4 7 Males smaller than 50 mm, females usually
smaller than 65 mm; tympanum black ................
3 Outer metatarsal tubercle absent ............... hosii
............................................................. baramica
• Outer metatarsal tubercle present, round ............
.......................................................... chalconota • Males larger than 50 mm, females usually larger
than 65 mm; tympanum brown ....... glandulosa
4 Web not reaching the disk of the 1st toe ............ 5
• Web reaching the disk of the 1st toe ..... erythraea
5 Supernumerary metacarpal tubercles present .... 6
• Supernumerary metacarpal tubercles absent .......
.............................................................. luctuosa

Rana chalconota (SCHLEGEL, 1837)

Hyla chalconota SCHLEGEL, 1837: 23 (part) – Type locality:

Java
Rana chalconota, SMITH 1931: 17; INGER 1956: 408, 1978: 312;
INGER & STUEBING 1992: 42; MALKMUS 1994b: 244, 1996c:
284; INGER , STUEBING & TAN 1996: 363; LAKIM et al. 1999:
33; Inger et al. 2000: 17
Rana chalconota raniceps, INGER 1966: 177
Hylarana (Rana) chalconota, MANTHEY & DENZER 1982: 19
Hylarana chalconota, MANTHEY 1983: 22

Diagnosis

Rana chalconota differs from R. nicobariensis in
the ratio of finger lengths and foot webbing: In R.
chalconota the 1st finger shorter than the 2nd and
the 1st three toes and the 5th are broadly webbed to
the disks, whereas in R. nicobariensis the 1st finger
is longer than the 2nd; and the 1st three toes and the
5th are not broadly webbed to the disks.
Description
Males 32-44 mm, females 46-60 mm; TL/SVL
0.51-0.60; habitus slender; head longer than wide;
vomerine teeth present; tympanum distinct, 2/3 to
3/4 of the diameter; supratympanic fold absent; 1st
finger shorter than 2nd, 1st finger tip weakly, all oth-
ers substantially dilated; marginal disk folds
present; toe tips dilated, narrower than on the outer
© P. Hoffmann
Fig. 153 Male of Rana chalconota from Poring, 550 m.
fingers; webbing formula 1(0), 2(0/0), 3(0/0), 4(1-
2/1-2), 5(0); inner metatarsal tubercle oval, outer
one round; dorsolateral folds present; dorsal skin
finely granular; ventral skin smooth on chest and
throat, granular on posterior portion of belly.
Males with yellowish nuptial pads on 1st finger and
a pair of subgular vocal sacs; tympani larger in
males than in females.
Dorsal coloration yellowish green, normally with
brown dots; cream-coloured ventrally; legs with

159
Ranidae

dark bars, sometimes without; upper lip cream-yel-
low; lower sides of legs usually reddish.
Head-body-length of tadpoles up to 18 mm, total
length up to 50 mm; body oval, slightly flattened,
width about 2/3 of the length; mouth positioned
ventrally, subterminally; labial teeth rows formula
4(2-4)/3 to 5(2-3)/3(1); dorsal fin slightly deeper
than ventral fin. This tadpole is one of the very few
distinctly coloured ones on the entire island of
Borneo. Body light yellow to straw-coloured, with
black streaks or dots below, in front of, and behind
the eye; lower side with many, small, white glands,
arranged in two longitudinal sections on either side
of the belly.
Ecology/Ethology
Rana chalconota is a nocturnal inhabitant of the
bank regions of rivulets cutting through primary
and secondary forests, in swampy areas on the edge
of cleared areas. The frogs are usually perched be-
neath or on dead wood or among banking vegeta-
tion up to 2 metres above the ground, but usually
lower. The species is absent in deforested and other
sun-exposed areas and thus differs distinctly from
R. nicobariensis in its choice of habitat. Calling
males and tadpoles were found between April and
September.
The brown-pigmented eggs are deposited in groups
of 1000-3200 directly into the water. The larvae
can be observed swimming slowly above the
substrate, predominantly leaf litter, in calm water
and forest ponds (m: 1g, i). They are absent from
water bodies with currents. Stomach analyses re-
vealed algae and protists at equal shares, but also
plant particles and fungal hyphens. Adults feed on
a variety of insects and spiders.
Its vertical distribution ranges from 65 to 915 m,
with a predilection for elevations below 300 m.
Rana erythraea (SCHLEGEL, 1837)
Hyla erythraea SCHLEGEL, 1837: 27 – Type locality: Java
Rana erythraea, INGER 1978: 325; HOFFMANN 1995b: 10
Diagnosis
A medium-sized to large, terrestrial frog with the
typical appearance of a “water frog” and wide, light,
dorsolateral stripes.
Description
Males 32-45 mm, females 48-75 mm; TL/SVL
0.50-0.57; head longer than wide; snout pointed;
vomerine teeth present; tympanum visible, ca. 0.75
times the eye diameter; supratympanic fold weak;
1st finger as long as or longer than 2nd finger tips
dilated, the largest one about 0.5 times the tympa-
num diameter; marginal disk folds present; outer fin-
ger with dermal fringe; toe tips dilated, but not as
widely as on fingers; webbing formula 1(0), 2(0/0),
3(0/0), 4(0/1-2), 5(0); smaller inner metatarsal tu-
bercle oval, outer one round; dorsolateral folds dis-
tinct; ventral and dorsal skin smooth.
Males with larger tympani (0.089-0.118 times the
SVL in males, 0.073-0.089 in females), and a
coarser skin; weakly developed humeral glands
present and nuptial pads on first fingers.
Light green dorsally, light brown laterally; with
cream-white or yellow dorsolateral stripes; a dark
stripe from canthus rostralis along dorsolateral folds
Fig. 154 Rana erythraea from Poring, 550 m.

Call: The call consists of a staccato series of click-

ing notes. Males form large calling communities
at the side pools of medium-sized streams and the
edges of ponds.

Distribution
© P. Hoffmann

Malay Peninsula, Sumatra, Java, and Borneo. Mt.

Kinabalu: Poring, Sg. Kipungit, Sg. Langanan,
Sayap, Kiau.

160
 Ranidae

up to anus; ventral side and up-

per lip white; legs olive green
with black crossbars or rows of
dots.
Head-body-length of tadpoles
up to 13 mm, total length up to
58 mm; body about twice as long
as wide; eyes situated dorsally,
oral disk subterminally, ven-
trally; lower lip and sides of up-
per lip with papillae; labial teeth
rows formula 1/1 to 1(1)/2(1);
anal tube dextral; tail convex,
pointed, and about twice as long
as body; coloration green or
brown dorsally, with dark speck-

© M. Kunkel
les; a dark streak from side of
head through eye up to tail; cau-
dal fin spotted; later develop-
mental stages with light lateral Fig. 155 Rana erythraea from North Sumatra.
and dorsal stripes.

Ecology/Ethology Rana hosii BOULENGER, 1891

A shy, nocturnal, terrestrial commensal which also
climbs in low vegetation. It actually avoids primary
forests and owes its large distribution range to its
ability to adapt to situations in and around human
settlements. It is usually encountered in the vicin-
ity of water, very often in paddy fields. The males
are often perched on grass or reeds, while the fe-
males are more often seen on the banks. The frogs
reproduce throughout the year. The tadpoles are
found only in stagnant water bodies (m: 2b).
The diet of R. erythraea includes small terrestrial
invertebrates like small millipedes, crickets, and
ants. This species prefers low elevations.
Call: Although often found sitting close to each
other, the males do not form calling communities.
The call is similar to that of R. nicobariensis, but
the call duration is longer, the note length is shorter,
and the number of notes per call and the note rep-
etition rate is greater in R. erythraea.
Distribution
From eastern India to Indonesia (Sumatra, Java,
Sulawesi), Borneo and the Philippine Islands. Mt.
Kinabalu: Poring.
Rana hosii BOULENGER, 1891c: 290 – Type locality: Mt. Dulit,
Sarawak
Rana hosei, INGER 1966: 187, 1962/63: 45, 1978: 312; INGER &
STUEBING 1992: 42; MALKMUS 1994b: 226, 1996c: 284;
HOFFMANN 1995b: 10
Hylarana hosei, MANTHEY 1983: 22; MALKMUS 1987: 281,
1988c: 176
Rana (Hylarana) hosei, MALKMUS 1992a: 113
Rana hosii, INGER, STUEBING & TAN 1996: 363; HOFFMANN
1998b: 89; LAKIM et al. 1999: 33; INGER et al. 2000: 17
Diagnosis
A moderately large to large inhabitant of the ground
and low vegetation levels with a slender habitus
and brightly green coloured back.
Description
Males 50-68 mm, females 85-100 mm; TL/SVL
0.57-0.66; vomerine teeth present; tympanum dis-
tinct, 0.75 times eye diameter in males, 0.4 times
in females; supratympanic fold absent; 1st finger
as long as or shorter than 2nd; finger tips substan-
tially dilated, outer ones for more than twice the
width of finger; tip of 3rd finger equal to or wider
than tympanum diameter of females; marginal disk

161
 Ranidae
© A. Nöllert
Fig. 156 Female of Rana hosii from Sg. Kipungit I, Poring, 650 m.
folds present; fingers with narrow dermal fringes;
toe tips dilated, but not as much as finger tips; toes
fully webbed; inner metatarsal tubercle oval, no
outer one; dorsolateral folds weak.
Males with a pair of subgular vocal sacs, separated
from one another by a thin septum; nuptial pads
present; tympani larger than in females (tympanum/
SVL 0.069-0.106 in males, 0.048-0.059 in females).
Fig. 157 Tadpole of Rana hosii from West Malaysia.
© U. Manthey
162
Usually dorsally bright green, interspersed with
darker spots in juveniles, finely pebbled on the
back; dark brown laterally, whitish to grey ven-
trally; dorsal sides of legs brown with dark cross-
bars.
Body of tadpoles oval, slightly flattened; head-
body-length up to 11 mm, total length up to 38 mm;
oral field ventral, subterminal; upper jaw white with
Fig. LXVII Oral disc of R. hosii from West Malaysia.
R. Malkmus

narrow, blackish brown anterior margin; lower jaw
uniformly white; labial teeth rows formula 4(2-4)/
4(1) to 6(3-6)/4(1); dorsal coloration dark grey,
unpigmented ventrally; a ventrolateral skin fold
over the entire body. Together with the strongly flat-
tened ventral side this ventrolateral skin fold has
the effect of a sucker when pressed to the smooth
surface of a rock.
Ecology/Ethology
Rana hosii prefers the banks of clear, fast-flowing
streams and rivers in primary and little-disturbed
rain forests at low to medium altitudes. During the
day, the males hide among rocks on the banks while
the females spend the days 1-2 m up in the vegeta-
tion. Already SMITH (1930) remarked: “The habits
of this species are those of a tree frog”, a hint for
which are also the extremely widened finger tips.
The species possesses a relatively potent skin poi-
son (vernacular name “Poisonous rock frog”) which
is said to be strong enough to kill a cat.
White to cream-coloured eggs (ca. 2.2 mm in di-
ameter) without a halfmoon are deposited in lumps
of ca. 500-2000 directly into the water (m: 1b, c).
The altitudinal distribution stretches from near sea
level to 1500 m.
Call: The males call day and night in loose groups
from raised sites and are often found in the imme-
diate environs of water, in vegetation or climbing
on rock cliffs.
Fig. 158 Eggs of Rana hosii from West Malaysia.
© U. Manthey
Ranidae
The call consists of either one or two notes, with
the call of one note lasting 70-80 ms, the one of
two 160-170 ms. The frequency range varies and
may even vary with the same individual. Domi-
nant frequency 3.3-4.5 kHz, decreasing towards the
end of the call (+22°C).
Distribution
Malay Peninsula, Sumatra, Borneo, and Java. Mt.
Kinabalu: Sayap, Sg. Kipungit, Sg. Mamut, Sg.
Langanan (750 m), Sg. Manggis. INGER, STUEBING
& TAN (1996) listed the species for Sg. Luidan and
Sg. Silau-Silau (1300-1500 m), records that were
contradicted by INGER & STUEBING (1997) who gave
its vertical range as “near sea level to about 750 m”.
Rana luctuosa (PETERS, 1871)
Limnodytes luctuosus PETERS 1871: 579 – Type locality: Sarawak
Rana decorata MOCQUARD 1890a: 145, Pl. 10: Fig. 1-1b – Type
locality: Kina Balu (syn. after BOULENGER 1891b)
Rana luctuosa, HANITSCH 1900a: 73; VAN KAMPEN 1923: 196;
SMITH 1931: 16; INGER 1954a: 250, 1966: 206, 1978: 312,
1985: 56; MATSUI 1979: 333; INGER & STUEBING 1992: 42;
MALKMUS 1994b: 244; INGER, STUEBING & TAN 1996: 363;
INGER & TAN 1996a: 563
Hylarana luctuosa, MANTHEY 1983: 22
Diagnosis
A medium-sized to large ground dweller with a
moderately stout habitus. Its conspicuous, reddish
brown dorsal coloration with the light dorsolateral
stripes is an unmistakable trait.
Description
SVL 53-60 mm; TL/SVL 0.51-0.58; habitus moder-
ately robust; head longer than wide; snout slightly
pointed; vomerine teeth present; nostrils closer to
tip of snout than to eyes; diameter of tympanum
about equal to that of eye; tympanic fold absent;
1st finger longer than 2nd finger tips only slightly
dilated; marginal disk folds present; no supernu-
merary metacarpal tubercles; webbing formula
1(2), 5(2), other toes with skin fringes to disc; in-
ner metatarsal tubercle oval, weakly developed; a
smaller, round outer metatarsal tubercle is recog-
nizable only in about every second specimen; skin
smooth, without dorsolateral folds.
163
 Ranidae
Males without vocal sacs and without nuptial pads,
but with humeral glands of ca. 4 mm in length.
Dorsal coloration reddish brown with sharply de-
fined, wide dorsolateral stripes running through the
eyes up to tip of snout, separating brown dorsal
coloration from black on sides of head and body;
flanks and lower side below black area dark grey,
lighter towards mid-venter; limbs grey with con-
© M. Matsui
© M. Matsui
164
spicuous light bars; eyes dark, top third lighter with
a white band on upper side.
Head-body-length of tadpoles up to 26 mm, total
length up to 65 mm; spiracle closer to eye than to
base of tail; labial teeth rows formula 4(2-4)/4(1)
to 6(2-6)/4(1); coloration of larvae greyish brown
with irregular dark spots on body and tail.
Ecology/Ethology
Rana luctuosa is a nocturnal, solitary inhabitant of
the leaf litter layer which congegrates at stagnant
water bodies only during the mating season. The
reduced webbing is typical for semi-terrestrial frogs.
The tadpoles live primarily in muddy ponds includ-
ing rhinocerus wallows (m: 2b, c).
Altitudinal distribution 150-1400 m.
Fig. 159 (left) Ventral pattern of Rana luctuosa from
Bundu Tuhan.
Fig. 160 (below) Same specimen as left.

Call: The males form calling communities at rain-
water pools with each male calling from a small
hole it has excavated adjacent to the pool. The call
resembles the miaow of a cat.
Distribution
Malay Peninsula, Sarawak, West Sabah. Mt. Kina-
balu: Kiau, Bundu Tuhan.
Rana nicobariensis (STOLICZKA, 1870)
Hylorana nicobariensis STOLICZKA, 1870: 150 – Type locality:
Nicobar Islands
Rana nicobariensis, BOULENGER 1885b: 389; SMITH 1931: 9;
INGER 1966: 223, 1978: 313; MALKMUS 1987: 281, 1994b:
244; HOFFMANN 1995b: 11, 1998b: 89; INGER, STUEBING &
TAN 1996: 363
Rana nicobariensis nicobariensis, INGER 1956: 406
Hylarana nicobariensis, MANTHEY 1983: 22
Diagnosis
A medium-sized terrestrial frog of slender habitus.
The webbing is only weakly developed in R.
nicobariensis, distinguishing it from R. chalconota.
The 1st finger is longer than the 2nd in R. nicoba-
riensis and shorter in R. chalconota.
Description
Males up to 47 mm, females up to 53 mm; TL/SVL
0.50-0.60; head longer than wide; snout pointed;
vomerine teeth present; nostrils much closer to tip of
snout than to eyes; eye diameter slightly greater than
distance between nostril and eye; tympanum visible,
about 75-80% of the diameter; supratympanic fold
Fig. 161 Rana nicobariensis from Sabah.
© J.C. Murphy
Ranidae
© J.C. Murphy
Fig. 162 Rana nicobariensis from Sabah.
absent; 1st finger as long as or slightly longer than 2nd;
finger tips dilated; diameter of finger disks smaller
than half tympanum diameter; marginal disk folds
present; toe tips more expanded than finger tips; web-
bing formula 1(1), 2(1/0), 3(1-2/0), 4(2-3/2-3), 5(0);
inner metatarsal tubercle small and oval, outer one
round; narrow dorsolateral folds present; dorsal and
lateral skin slightly granular, smooth ventrally.
Males with slightly longer heads and larger tympani
than females, and with pairs of subgular vocal sacs,
yellowish nuptial pads, and large humeral glands.
Colour pale brown with black spots dorsally, darker
brown laterally and particularly so on sides of head;
upper lip white; lower side dirty white with a faint
grey pattern; hind limbs with dark crossbars.
Body shape of tadpoles oval; light brown with dark
brown pattern; tail with dark reticulated pattern; head-
body-length up to 15 mm, total length up to 47 mm;
lower lip with conspicuous, long, finger-like projec-
tions; labial teeth rows formula 1/2(1) to 1/3(1).
Ecology/Ethology
The frogs are widely distributed, terrestrial com-
mensals which are limited on Mt. Kinabalu to the
margins of the national park, but are more commonly
encountered in the surrounding villages. They are
often found sitting in drainages and other artificial
water bodies and flooded meadows. They are both
diurnal and nocturnal. Males are very common lo-
cally, but do not appear to form calling communi-
ties. The reproductive season lasts throughout the
165
Ranidae

year. Their diet comprises insects of appropri-

ate sizes.
The larvae develop in the shallow water of
ditches and temporary pools (m: 2b).
Altitudinal distribution 0-925 m a.s.l.

Call: (+21°C) The characteristic, cackling calls

consist of rapid sequences of 6-10 loud, sharply
pulsed notes of 50-60 ms in duration with a
note repetition rate of 7 per second, an internote
interval of 70-80 ms, and a frequency between
1 and 4 kHz; harmonics are not evident.

Distribution
From the Nicobars through the Malay Penin-
sula to Bali, Borneo, and Palawan . Mt. Kina-
balu: Poring, Kiau.

Fig. LXVIII Sonagram and oscillograms of Rana

nicobariensis
(Poring Hot Springs, 500 m; 10.00 p.m. +22°C)

Taxonomic remarks: Rana signata and Rana

Rana signata (GÜNTHER, 1872)
Polypedates signatus GÜNTHER, 1872: 600 – Type locality:
Matang, Sarawak
Rana obsoleta MOCQUARD 1890a: 147 – Type locality: Kina
Balu (syn. after BOULENGER 1920)
Rana signata, VAN KAMPEN 1923: 226 (part); INGER & STUEBING
1992: 42; MALKMUS 1994b: 244; WONG 1994: 32; HOFFMANN
1995b: 12; INGER, STUEBING & TAN 1996: 363; INGER & TAN
1996a: 564; MANTHEY & GROSSMANN 1997: 116
Rana signata signata, (part) INGER 1956: 407, 1966: 233, 1978: 313
Hylarana signata, (part) MANTHEY 1983: 22; MALKMUS 1988c:
176, 1989: 187, 1996c: 284
Rana (Hylarana) signata, MALKMUS 1992a: 113
Rana picturata BOULENGER, 1920
Rana picturata BOULENGER, 1920: 179 – Type locality: Bidi
Caves, Sarawak, Barabas, S.-E. Borneo, and Kina Balu
Rana signata signata (not Rana signata (GÜNTHER, 1872)),
(part) INGER 1966: 223
Rana (Hylarana) signata (not Rana signata (GÜNTHER, 1872)),
(part) VAN KAMPEN 1923: 226
Rana signata (not Rana signata (GÜNTHER, 1872)), MANTHEY
& GROSSMANN 1997: fig. 76
picturata were for a long time considered conspe-
cific, until INGER & TAN (1996a: 564) separated
them on the basis of their colour patterns. In R.
signata, the orange to red stripe runs without inter-
ruption from the eyelid over the back, whereas if
such stripe is present in R. picturata, it is always
interrupted. The back itself is marked with many
(R. picturata) or a few to no (R. signata) spots in a
colour matching the dorsolateral stripes. Another
indication of the presence of two rather than one
species is the inconsistency in the descriptions of
the vocalizations of “R. signata” by GRANDISON
(1972), INGER & STUEBING (1992), SANCHEZ-HERRAIZ
et al. (1995), and MANTHEY & GROSSMANN (1997).
Since, however, no morphological or ecological
differences have been confirmed up to now other
than those of the colour pattern, both forms are
treated here jointly due to their great similarity.
Diagnosis
A small to medium-sized ground dweller with a mod-
erately slender build. Due to the highly contrasting,

166

Fig. 163 Rana picturata from Sg. Kipungit I, Poring.
yellow to orange red and black colour pattern, R.
signata and R. picturata cannot be confused with
any other Bornean frog.
Description
Males 33-47 mm, females 48-69 mm; TL/SVL
0.48-0.57; habitus moderately stout; head slightly
pointed; canthus rostralis distinct; vomerine teeth
present; nostrils closer to tip of snout than to eyes;
supratympanic fold often absent; 1st finger longer
than 2nd; finger and toe tips slightly expanded;
marginal disk folds present; supernumerary meta-
carpal tubercles present; toe tips dilated into disk-
like structures; toes webbed for about half their
lengths; oval inner and large round outer metatar-
sal tubercles present; dorsolateral folds often ab-
sent; dorsal and lateral skin granular, smooth
ventrally.
Males with a pair of subgular vocal sacs, nuptial
pads, and humeral glands.
Ranidae
© A. Nöllert
Dorsal and lateral coloration black, with yellow or
orange red spots; a stripe of matching colour from
the tip of the snout up to above the eyes; dorsola-
teral stripes of the same colour continuous or in-
terrupted (see Taxonomic remarks above); ventral
side of males dark, lighter in females; limbs yel-
low or orange red above, with blackish brown cross-
bars.
Tadpoles relatively slender, oval in shape, slightly
compressed dorsoventrally; head-body-length up
to 13 mm, total length up to 41 mm; labial teeth
rows formula 3(2-3)/3(1) to 4(2-4)/3(1); first lower
row of denticles interrupted only to a minor ex-
tent; blackish brown dorsally, tail pale grey, un-
pigmented ventrally, covered with small, whitish
glands. These glands form a wide ring around the
oral field, a separated round spot behind it, a nar-
row ventrolateral series between insertion of legs
and below spiracle, and a dorsolateral row between
eye and base of tail. Rows of glands continued on
167
Ranidae

both fins over the latter’s full

length, but absent on caudal
muscle. Arrangement and num-
ber of these glands differ slightly
in the two forms.

Ecology/Ethology
Rana signata and R. picturata
are nocturnal inhabitants of
calm sidearms or respective
sections of small to moderately
large (up to about 10 m in
width) running water bodies.
Their occurrence is limited to
primary and old secondary for-
ests. The juveniles migrate
away from the sites of their lar-
val development and spend
their early life on the forest
floor, only to return to a calm
waters on reaching maturity.
They are usually found on the
ground, dead wood, or rocks,
more rarely climbing in low
vegetation (up to about 40 cm).
The males call individually at
first, but appear to trigger oth-
ers to do so as well, so that large Fig. LXIX Sonagram and oscillograms of Rana signata
calling communities establish
themselves at regular intervals
(INGER & STUEBING 1997). SANCHEZ-HERRAIZ et al. Call: (+24°C) It is at present extremely difficult to
(1995) reported on distances of several metres determine which call belongs to which form as both
between calling males with the calls not being have been treated as one and the same species in the
emitted in choruses. past. According to INGER & STUEBING (1997), the call
Oviposition between April and August, preferably of R. picturata sounds like a metallic burr (duration
in places without or only a weak current, e.g. in 620 ms, consisting of 4-6(8) pulsed notes of 50-80
rock basins or cut-off sidearms of streams or riv- ms each) with the intensity of the notes continually
ers. The larvae live in shallow, but also in deep, decreasing; frequency 1.1-204 kHz (S ANCHEZ -
sections of their water bodies where the bottom is HERRAIZ et al. 1995). There is, however, a deviant
muddy or consists of gravel, often in accumula- description of the call by GRANDISON (1972), who
tions of leaf litter (1d, f). Stomach analyses revealed characterized it as similar to human laughter. There
for a major part blue algae, besides about equal is a chance that the latter might refer to R. signata.
parts of plant particles, diatoms, and fungal hy-
phens. Distribution
The altitudinal range spans 0-750 m, whereby INGER Malay Peninsula, Sumatra, Borneo, Philippines
& STUEBING (1997) state R. picturata to be distrib- (with three subspecies in INGER, 1954). Mt. Kina-
uted throughout Borneo, while R. signata would balu: Sg. Kipungit, Sg. Mamut, Sg. Tananansad,
prefer the lowland. Sg. Sasapan.

168
 Ranidae

Genus Staurois COPE, 1865

The distribution of the genus Staurois

is mainly restricted to Borneo with
only S. natator also occurring on is-
lands of the Philippines.
It comprises small to medium-sized,
very slender frogs with long hind
limbs. Vomerine teeth generally ab-
sent, but often present in S. natator
at least on one side. Finger tips and
toes expanded into oval structures,
marginal disk folds and ventral cross
folds present. Toes fully webbed; 1st
finger always shorter than 2nd; dorso-
lateral folds wanting, tympanum vis-
ible, ventral skin coarsely granular.

© H. Sauer
All species share a distinct predilec-
tion for rapids and waterfalls where
they can usually be seen perched on Fig. 164 Foot flagging display in the genus Staurois; here: S. tuberi-
boulders on the banks or those pro- linguis from Sg. Liwago.
truding from the water. Besides of the
common acoustic advertisement, the
males have developed a particular optical display litter covering the bottoms of basins or small rivu-
in which the colourful webbings between the toes lets in or forking off from streams.
are flashed by stretching a hind limb and spread- Three species occur on Borneo, all of which are
ing the toes either unilaterally or alternatingly. As also found on Mt. Kinabalu.
this gives an appearance not dissimilar to naval flag
signals this behaviour is appropriately termed “foot
flagging display” (HARDING 1982, DAVISON 1984). Key to the Bornean species of Staurois
Only the larvae of S. natator (INGER & TAN 1990) (after INGER 1966)
and S. tuberilinguis (MALKMUS et al. 2000) are
known. Their external morphology and choice of
habitat is largely congruent, with the body being 1 Outer fingers webbed .................... latopalmatus
compressed, the tail long with reduced fins, the eyes • Fingers not webbed ........................................... 2
small and situated dorsally, and the skin hardly 2 Outer metatarsal tubercle present; no lingual pa-
pigmented. Differences are only evident in the body pilla ......................................................... natator
lengths at the various stages of development, and • No outer metatarsal tubercle; lingual papilla
in the numbers of denticle rows on the lower lip. present ........................................... tuberilinguis
The tadpoles lead a secretive life among the leaf

Tab. 17 Distribution of Staurois-species on Borneo.

169
 Ranidae
Staurois latopalmatus
(BOULENGER, 1887)
Ixalus latopalmatus BOULENGER, 1887b: 97 – Type locality:
Mount Kina Baloo, North Borneo
Ixalus latopalmatus, MOCQUARD 1890a: 152; WHITEHEAD 1893:
121
Sinomantis latopalmata, VAN KAMPEN 1923: 238; SMITH 1931:
17; INGER 1962/63: 46
Staurois latopalmatus, INGER 1966: 252, 1978: 313; MANTHEY
1983: 23; MALKMUS 1988c: 176, 1992a: 115, 1994b: 226,
1996c: 285, INGER & STUEBING 1992: 42; INGER , STUEBING &
TAN 1996: 363; INGER & TAN 1996a: 564; LAKIM et al. 1999:
32; INGER et al. 2000: 17
Diagnosis
A moderately slender terrestrial frog with substan-
tially expanded finger tips. The tips of all fingers
are wider in diameter than the tympanum, while in
all other Bornean species of this genus only the
tips of the outer two fingers are wider. In contrast
to the other species of Staurois the outer two fin-
Fig. 165 Staurois latopalmatus from Sg. Kipungit II,
640 m, Poring.
© R. Malkmus
170
gers are webbed, and the snout of S. latopalmatus
is very short and blunt, while it is very long and
pointed in the other two.
Description
Males 40-50 mm, females 60-70 mm; TL/SVL
0.58-0.71; habitus slender, but rather stout in com-
parison with the other two Bornean species of this
genus; head longer than wide; vomerine teeth ab-
sent; nostrils closer to tip of snout than to eyes;
canthus rostralis distinct; tympanum visible, 1/3
to 2/5 of eye diameter; supratympanic fold absent;
1st finger shorter than 2nd; finger tips expanded into
disk-like structures, diameters of all finger tips
are greater than diameter of tympanum; marginal
disk folds present; fingers not fully webbed; toe
tips expanded into disk-like structures, smaller
than finger tips; toes fully webbed; inner meta-
tarsal tubercle oval, no outer one; dorsolateral
folds absent; dorsal skin granular, smooth
ventrally.
Males with a pair of vocal sacs and whitish nuptial
pads on first metacarpal.
Dorsal coloration black with yellowish or white
dots, ventral side white to cream-coloured; toe
webbings of males whitish to pale grey, dark grey
in females.
The tadpole of Staurois latopalmatus will soon be
described by I. DAS.
Ecology/Ethology
Staurois latopalmatus is a diurnal and nocturnal
ground dweller which is found along running wa-
ters, often on boulders in or on the banks of streams,
but occasionally also in low vegetation. If disturbed,
it will leap from rock to rock, or it may even leap
into swift water and skip across the surface to an-
other rock. At night, juveniles are often encoun-
tered perched together in groups, whereas adults
are mostly found singly. Foot flagging displays can
only be observed between 6.30 and 8.00 a.m. dur-
ing which time the males often congregate in
groups. Their calls can be heard throughout the
night and the early morning hours (HARDING 1982,
DAVISON 1984). The vertical distribution of the spe-
cies ranges from 150 to 1000 m a.s.l.
Call: A short, rising trill.

Distribution
Endemic to Borneo with records from Sabah,
Sarawak, Kalimantan, and Brunei. Mt. Kinabalu:
Sayap, Sg. Panataran, Kiau, Sg. Liwago, Sg. Man-
tukungan, Sg. Mamut, Sg. Kipungit, Sg. Langanan,
Sg. Sasapan.
Staurois natator (GÜNTHER, 1859)
Ixalus natator GÜNTHER, 1859 (1858): 75 – Type locality:
Philippines
Ixalus guttatus GÜNTHER 1859 (1858): 76, Pl. 4: Fig. D – Type
locality: Borneo (syn. after BOULENGER 1882)
Ixalus natator, HANITSCH 1900a: 74
Staurois guttatus, BOULENGER 1918: 374; VAN KAMPEN 1923: 234
Rana guttatus, SMITH 1931: 30
Staurois natator, INGER 1962/63: 46, 1966: 245, 1978: 313;
MANTHEY 1983: 23; MALKMUS 1988c: 176, 1988d: 8, 1989:
188, 1992a: 115; INGER & TAN 1990: 4; INGER & STUEBING
1992: 42; WONG 1994: 32; HOFFMANN 1995b: 13; INGER,
STUEBING & TAN 1996: 363; LAKIM et al. 1999: 32; INGER et
al. 2000: 17
Fig. 166 Staurois natator from Sg. Kipungit II, 650 m, Poring.
Ranidae
Diagnosis
A very slender ground dweller with expanded fin-
ger tips and a pointed snout. The external morphol-
ogy corresponds with that of S. tuberilinguis, dif-
fering only in the greater SVL of adults and with
regard to the colour pattern. Olive green dorsally,
with large black spots; dorsal skin homogeneously
granular (in contrast to S. tuberilinguis which is
brown dorsally with irregular green spots and has
a heterogeneously granular skin). Ventral and lat-
eral sides of S. natator lemon-yellow to yellowish
green, S. tuberilinguis bluish white ventrally and
green with small white dots laterally.
Description
Males 29-37 mm, females 44-55 mm; TL/SVL
0.55-0.65; habitus very slender; head longer than
wide; vomerine teeth almost always present, but
not always on both sides; nostrils closer to tip of
snout than to eyes; canthus rostralis distinct; tym-
panum visible, its diameter about 1/3 to 1/2 eye
© P. Hoffmann
171
 Ranidae
diameter; supratympanic fold absent; 1 st finger
shorter than 2nd; finger tips expanded into disk-like
structures, outer two wider than tympanum diam-
eter; marginal disk folds present; 2nd and 3rd fingers
with dermal fringes; toe tips expanded into disk-like
structures, but smaller than finger tips; toes fully
webbed; inner metatarsal tubercle oval, outer one
round and small; dorsolateral folds present; skin on
dorsum and on posterior portion of belly granular,
chest and throat smooth.
Males with a pair of vocal sacs and yellowish nup-
tial pads on 1st finger; their webbings between toes
light greenish blue.
Dorsal coloration olive green, with large black
spots; ventral and lateral sides lemon-yellow to
yellowish green; a narrow, black stripe along can-
thus rostralis from eye to tip of snout.
Body of tadpoles oval in shape and slightly flat-
tened; length of tail more than twice the head-
body-length, e.g. at stage 41 head-body-length
10.5 mm, total length 34.5 mm; eyes small (di-
ameter 0.08 times head-body-length), set far
back, oriented dorsally, and thickly covered with
skin at early stages of development, formation
of a transparent “window” from stage 36; nos-
trils small and positioned subterminally; tail with
very low fins, ending in a broad tip; oral disk
situated ventrally, subterminally; labial teeth
Fig. 167 A pair of Staurois natator from Danum Valley,
Sabah, in amplexus.
© W.Hödl
172
rows formula 2(2)/8(1) to 2(2)/10(1); spiracle
sinistral, with a short, terminal tube; anal tube
long, situated medially; almost transparent, with
a reddish belly resulting from the visibility of
blood vessels.
Ecology/Ethology
A mainly diurnal to crepuscular inhabitant of the
rocks on the banks of streams and small rivers in
primary forests. The frog is also often found in low
banking vegetation, but rarely encountered farther
than 100 m from the nearest water. Especially
young specimens congregate at night to form “dor-
mitories” on larger leaves etc. The tadpoles live
predominantly in the thick layers of sediment con-
sisting of leaf litter and mud in sections of streams
where there are no currents (m: 1d, e).
The known altitudinal range extends from 110 to
650 m a.s.l.
Call: The call consists of a sharp chirp.
Distribution
Borneo and the Philippine Islands. Mt. Kinabalu:
Sayap, Sg. Mamut, Sg. Kipungit, Sg. Sasapan.
The record “Sg. Silau-Silau (1540-1580 m)” by
WONG (1994) is probably based on misidentified
S. tuberilinguis.
Staurois tuberilinguis BOULENGER, 1918
Staurois tuberilinguis BOULENGER, 1918: 374 – Type locality:
Mt. Kina Balu, North Borneo, altitude 4200 feet and Mt.
Batu Song, Sarawak, 1000 feet
Staurois tuberilinguis, VAN KAMPEN 1923: 237; INGER 1962/63:
48, 1966: 250, 1978: 313; MANTHEY & DENZER 1982: 12;
MANTHEY 1983: 23; MALKMUS 1985: 8, 1987: 281, 1988d: 8,
1988c: 176, 1989: 187, 1991a: 28, 1992a: 114, 1994b: 226,
1996b: 22, 1996c: 285, 1996e: 11; INGER & STUEBING 1992:
42; WONG 1994: 32; HOFFMANN 1995b: 13, 1998b: 86; INGER,
STUEBING & T AN 1996: 363; INGER & TAN 1996a: 564;
MALKMUS 1999: 15; MALKMUS, KOSUCH & KREUZ 1999: 17;
LAKIM et al. 1999: 33; INGER et al. 2000: 17
Rana tuberilinguis, SMITH 1931: 17
Diagnosis
A very slender ground dweller with expanded fin-
ger tips. Dorsum brown with irregular green spots,
heterogeneously granular; ventral sides bluish
 Ranidae

white; lateral sides green with small white dots.

Distinguished from S. natator (see above) on the
basis of the colour pattern, the absence of an outer
metatarsal tubercle, and the presence of a lingual
papilla (see key).

Description
Males 23-37 mm, females 29-43 mm; TL/SVL
0.49-0.57; habitus very slender; head longer than
wide; snout pointed; vomerine teeth absent; nos-
trils closer to tip of snout than to eyes; canthus
rostralis distinct; tympanum visible, about 1/3 to
2/5 the eye diameter; supratympanic fold present;
1st finger much shorter than 2nd; finger tips ex-
panded into disk-like structures, those of 3rd and
4th fingers wider than tympanum diameter; mar-
ginal disk folds present; fingers without dermal
fringes; toe tips expanded into disk-like structures,
but not as wide as tips of outer fingers; toes fully

© U. Manthey
webbed; inner metatarsal tubercle oval, outer one
round and small in about a third of all individu-
als, absent in the remainder; dorsolateral folds
wanting; dorsal skin heterogeneously granular, Fig. 168 Staurois tuberilinguis from Sg. Silau-Silau,
smooth ventrally. 1600 m.
Colour dorsally brown, with elongate, green spots;
green laterally, with small, white glands; bluish
white ventrally, usually with dark dots on the throat. tip of snout more than three times as large as
Males with a pair of vocal sacs and whitish nuptial interocular distance; oral field ventral, subtermi-
pads on the 1st metacarpal; webbings between the nal; marginal oral papillae small, in one row;
toes white. denticles 2(2)/6(1); proximal row of teeth on up-
Tadpoles morphologically very similar to those of per lip widely interrupted medially, inner row of
S. natator; body oviform, distinctly compressed
dorsoventrally; tail more than twice as long as head-
body portion, e.g. at stage 25, head-body-length Fig. LXXI Oral disc of Staurois tuberilinguis (Sg.
3.7 mm, tail length 8.1 mm; caudal fins low, tip Liwago, 1500 m).
blunt; spiracle sinistral, short, straight, directed
backward; anal tube median; eyes very small, dor-
sal; distance between anterior margin of eye and

Fig. LXX Tadpole of Staurois tuberilinguis.

R. Malkmus R. Malkmus

173
 Ranidae

predictably at another site. Foot flagging is shown

by the males often right after a series of calls. As the
white coloration of the vocal sacs is well visible dur-
ing the calling, they represent another optical sig-
nal. Suitable advertising sites are rocky banks and
solitary boulders on the banks. The males call dur-
ing the day between 6.00 a.m. and 6.30 p.m. Single
calls are often answered by antiphonic calls, but
never to such an extent that choruses might form.
The tadpoles have as yet only been described from
a basin on Sg. Liwago which was filled to the rim
with leaf litter (m: 1d, e). There they lived photo-
phobically in an environment with almost no cur-
© P. Hoffmann

rent.
The altitudinal distribution covers 200-1800 m a.s.l.

Fig. 169 Staurois tuberilinguis from Sg. Wariu, 950 m, Call: (+19°C) Note series of 8-12 s in duration,
Sayap. with 40-60 notes of 80-100 ms each, resembling
the trill of an insect. The frequency rises from an
initial 3.8 kHz to 4.1 kHz within a note series, with
lower lip with a small gap; lengths of denticle rows the note repetition rate increasing from 3 to 8 notes
decreases continually distally; edges of both jaws per second and the internote intervals decreasing
narrow, with black pigmentation, and homogene- from 400 to 50 ms.
ously serrated. Larvae light brown dorsally, fading
laterally so that they appear unpigmented lateroven- Distribution
trally. Tail with two central and two marginal Endemic to Borneo with records from Sabah,
brownish stripes. The transparent skin of the ven- Sarawak and Brunei. Mt. Kinabalu: Sayap,
ter permits to see the intestinal coils and the in- Kenokok, Sg. Kinateki, Marei Parei, Sg. Kada-
tensely red gills. maian above Kiau, Lumu Lumu, Lobang, Sg. Silau-
Silau, Sg. Liwago, Mesilau, Sg. Mantukungan, Sg.
Ecology/Ethology Mamut, Sg. Kipungit, Sg. Langanan.
This frog is a predominantly di-
urnal and crepuscular inhabitant
of the banks of fast-flowing Fig. LXXII Sonagram and oscillogram of Staurois tuberilinguis
mountain streams in primary for- (Sg. Liwago, 1500 m; 8.00 p.m. +19°C)
ests, particularly of waterfalls
and rapids. Here, the animals are
mostly seen perched on rocks
during the day while they spend
the night, well-camouflaged by
their green coloration, singly on
the upper side of leaves of bank-
ing bushes 50-100 cm above the
ground. S. tuberilinguis is very
shy during the day, escaping in
almost every instance by plung-
ing into the nearest rapid, only to
re-appear unexpectedly and un-

174

Rhacophoridae
Rhacophorid frogs are distributed in tropical Af-
rica and Madagascar, but have their main centre in
the Asian tropics and subtropics. Here, they range
from India to Japan, including the Greater Sunda
Islands and the Philippines.
Rhacophorids have a firmisternal pectoral girdle,
sacrum with cylindrical diapophyses, omosternum and
sternum ossified, eight procoelous presacral vertebrae
present. Maxillae and premaxillae dentate. Fingers
and toes dilated into large disks, short cartilaginous
intercalary element between penultimate and termi-
nal phalanges present; toes webbed over at least half
their lengths. Some species of Rhacophorus with ex-
tensive webbings on hands and feet enabling them to
perform paragliding flights. Pupil horizontal (except
in Nyctixalus). Mating takes place in axillary amplexus.
Key to the Bornean genera of Rhacophoridae (after MANTHEY & GROSSMANN 1997)
1 Pupil roundish or horizontal (A) ...................... 2
• Pupil vertical or diamond-shaped (B) .................
........................................................... Nyctixalus
2 Skin membrane between two outer metacarpals
present (C), terminal phalanges of fingers and toes
Y-shaped (D) ..................................................... 3
• No skin membrane between two outer metacar-
pals; terminal phalanges of fingers and toes not
Y-shaped, but partly slightly notched (E) ...........
............................................................ Philautus
3 One or more finger disks with a ring-shaped fold
(F) on lower side .......................... Rhacophorus
• No ring-shaped fold on lower sides of finger disks
........................................................................... 4
4 Upper side, particularly limbs, with numerous
smaller and larger warts (G) ........... Theloderma
• Skin of upper side smooth .............. Polypedates
Rhacophoridae
The species of all Bornean genera, save for
Nyctixalus and Philautus, produce foam nests in
which the eggs develop to tadpoles. The larvae are
aquatic, with terminal mouths with beaks and rows
of labial teeth; those of Nyctixalus and Theloderma
are non-feeding. Several lineages of Philautus are
known for their non-aquatic larvae – their eggs
developing directly into froglets.
All Bornean species are tree frogs, dwelling pri-
mary and secondary forests, some species appear
to prefer habitats modified by human activity.
Rhacophorids are very common on Mt. Kinabalu
at all altitudinal levels up to 3000 m above sea level.
The family is represented on Borneo by 5 genera
with 35 species, on Mt. Kinabalu by 4 genera and
22 species.
A B
E
D
C
G
F
175
 Rhacophoridae
Genus Nyctixalus BOULENGER, 1882
The genus Nyctixalus is represented on Borneo by
a single species, Nyctixalus pictus (PETERS, 1871),
which is known from Sabah, Sarawak, Brunei
Darussalam and Kalimantan.
Nyctixalus pictus (PETERS, 1871)
Ixalus pictus PETERS, 1871: 580 – Type locality: Sarawak, Bor-
neo
Philautus pictus, SMITH 1931: 19; INGER 1956: 422
Philautus pictus pictus, INGER 1966: 349
Hazelia picta, INGER 1978: 313; MANTHEY 1983: 23
Nyctixalus pictus, MALKMUS 1991: 30, 1992a: 123; INGER &
STUEBING 1992: 41-51; HOFFMANN 1995b: 13; INGER, STUEBING
& TAN 1996: 363; MALKMUS 1996c: 286; MANTHEY & GROSS-
MANN 1997: 124; LAKIM et al. 1999; INGER et al. 2000
Diagnosis
A slender, chestnut-brown frog (< 35 mm) with
white spots and dots on the back and sides; pupil
vertical-oval to diamond-shaped; fingers without
webbings; vomerine teeth absent.
Description
Males up to 33 mm, females up to 38 mm; TL/SVL
0.50-0.61; body and limbs slender; body flattened
Fig. 170 Nyctixalus pictus from Sg. Wariu, ca. 1000 m,
Sayap.
© P. Hoffmann
176
dorsolaterally; head longer than wide; snout obtusely
pointed; canthus rostralis distinct; lores nearly ver-
tical, feebly concave; tympanum distinct, two thirds
of eye diameter; tips of fingers expanded into round
or oval disks, no webbing; metacarpal tubercles
present; disks of toes smaller than those of fingers;
toe webbing 1(0.5), 2(0.5/0.5), 3(1/1), 4(2/2.5),
5(1.5-2); an oval inner metatarsal tubercle; dorsal
skin with spinose tubercles; throat smooth, chest
rugose, abdomen coarsely granular; no supratym-
panic fold; males without vocal sacs; a yellowish
nuptial pad on dorsal surface of 1st finger.
Dorsal sides of body and limbs dark chestnut-brown
to reddish brown, lighter laterally; lower side whit-
ish with a reddish tinge, belly sometimes blackish;
dark brown portions peppered with ivory-white dots
and spots which are reduced in numbers in speci-
mens from high altitudes. Elliptical shoulder spots,
a horizontal row of spots along canthus rostralis and
above eye, spots on knee and heel joints, and short
bars across hind limbs and lower arms are always
present, however. Upper third of iris greyish yellow
with a blackish network of veins, lower two thirds
dark brown.
Tadpoles: Head-body portion of tadpoles oval,
almost as wide as long, slightly flattened; snout
broadly rounded; eyes dorsal; mouth ventral; beaks
with black edges, serrated; labial teeth rows formula
Fig. 171 Ventral pattern of Nyctixalus pictus from Sg.
Silau-Silau, 1550 m.
© M. Schroth
 Rhacophoridae
© U. Manthey

© U. Manthey
Fig. 172 Tadpole (st 26) of Nyctixalus pictus from Pulau Fig. 173 Tadpole (st 43) of Nyctixalus pictus from Sin-
Pinang, Peninsular Malaysia. gapore.

5(2-5 or 3-5)/3; spiracle sinistral; tail slightly con- frequency 1.8-2.2 kHz; internote interval 80-90 ms)
vex, tapering to broadly rounded tip, 1.3-1.5 times at an increasing volume. This call resembles a muf-
the head-body-length. Tadpole entirely black to fled tweeting bird call. Harmonics at 4 kHz.
purplish brown, fins and ventral side of body
lighter; total length up to 53 mm; freshly metamor- Distribution
phosed froglets 17 mm. Malay Peninsula, Sumatra, Borneo, and Palawan.
Mt. Kinabalu: Poring (500 m), Kiau, Lumu Lumu,
Ecology/Ethology Sayap (Sg. Wariu), upper Liwago Valley (up to
Nyctixalus pictus inhabits the montane 1750 m), Headquarters (1540 m).
rain forests between 500-1750 m a.s.l.
on Mt. Kinabalu. It is found off from
Fig. LXXIII Sonagram and oscillograms of Nyctixalus pictus
streams in the lower stratum (shrub and
(Sg. Liwago, 1500 m; 8.00 p.m. +20,5°C)
bush level) up to 3 m above the ground,
but also in the leaf litter layer. The spe-
cies is nocturnal, leaving its shelters to
forage with the onset of dusk. This spe-
cies is predominantly a sit-and-wait
predator. Its diet comprises spiders, hair-
less caterpillars, and the larvae of bee-
tles.
Spawn was found in the form of clutches
containing 7-15 eggs hanging on the
walls of water-filled tree holes 1-1.6 m
above the ground, and in large empty fruit
casks 3-10 cm above the water level (m:
2d, g, h).

Call: (+20.5°C) The note series starts off

with 2-4 distinctly pulsed notes (dura-
tion 34-45 ms; frequency 1.9 kHz; inter-
note interval decreasing from 150 to
90 ms) which are followed by 4
weakly pulsed notes (duration 10-13 ms;

177
Rhacophoridae
Genus Philautus GISTEL, 1848
The species of this genus are distributed from In-
dia and Sri Lanka to China, the Greater Sunda Is-
lands, and the Philippines.
The genus Philautus comprises small to medium-
sized, usually stout tree frog species, many of which
are characterised by a confusing morphological
variability. The resulting problems are even en-
hanced by the fact that many species are able to
alter their colorations and patterns. The knowledge
of the specific advertisement calls is therefore of
fundamental importance for the field herpetologist.
An attempt to establish an identification key to the
species on the basis of morphological traits is for
the same reason rendered entirely dissatisfactory.
Under these circumstances we are much rather in-
clined to follow the proposal by DRING (1987) to
allocate the Bornean forms to four species groups
which are defined as follows:
- aurifasciatus group: small or medium-sized spe-
cies; vomerine teeth absent; nuptial pad small
and smooth; ova unpigmented; calls generally
pulsed, poorly tuned, with no frequency modu-
lation. Content: P. amoenus, longicrus, mjobergi,
petersi, refugii, saueri, umbra.
Tab. 18 Distribution of Philautus-species on Borneo.
178
- hosii group: large species; vomerine teeth present;
nuptial pad absent; ova with dark pigmentation;
calls not pulsed, well tuned, with frequency
modulation; stream-associated species. Content:
P. hosii, ingeri.
- tectus group: small species; vomerine teeth
present or absent; nuptial pad large and spinu-
lose; ova without pigmentation; call not yet de-
scribed; lowland species. Content: P. tectus.
- vermiculatus group: moderately large to large
species; vomerine teeth present or absent; nup-
tial pad absent; ova half-pigmented; call pulsed,
poorly tuned, sometimes with frequency modu-
lation. Content: P. acutus, aurantium, bunitus,
disgregus, kerangae.
It must be stated though that this categorisation has
meanwhile turned out to be in need of further revi-
sion.
The species of the genus Philautus are predomi-
nantly found in montane forests, between 700 and
3050 m on Mt. Kinabalu. The boundaries of the
individual species’ altitudinal ranges are usually
sharply defined at high montane levels and do nor-
mally not include wider transitory zones. Most spe-
cies inhabit the lower stratum (10-300 cm) which
is where the clutches are also deposited in cush-
ions of moss, rock crevices, leaf axils, and pitchers
 Rhacophoridae

of Nepenthes. Since it is known of P. kerangae,
mjobergi, petersi, and saueri that their larval de-
velopment up to metamorphosis is completed
within the jelly shell, it is to be presumed that this
mode of development is equally applicable in all
the other Bornean species of Philautus.
Altogether 15 species are known from Borneo, 8
of which from Mt. Kinabalu:
Remarks: It is at present unclear if P. hosii occurs
on Mt. Kinabalu. DRING (1987) re-identified a speci-
men described as “Rhacophorus hosii/Bundu
Tuhan” as being P. ingeri. Notwithstanding, P. hosii
appears in the check list by INGER, STUEBING & TAN
(1996) whereas P. ingeri does not.
Besides light brown specimens with almost no pat-
tern save for a dark interorbital bar, there are pale
brown ones with broad, dark dorsal stripes form-
ing a sand-glass like marking. Others irregularly
marbled with wide crossbars on light brown or
bright scarlet. Some species with uniformly red-
dish brown or moss-green dorsal sides framed with
black dorsolaterally and white flanks with fine
blackish brown reticulation; lower side usually
uniformly whitish grey to rich dark brown. All
specimens, however, show an oblique white bar
extending from posterior half of eye to upper lip;
From top to base: Philautus amoenus,
between Pondok Ubah and Pondok Lowii, 2100-2300 m.
Fig. 174 Pair Fig. 175 Male

Philautus amoenus SMITH, 1931

Philautus amoenus SMITH, 1931: 18 – Type locality: Kamboran-
gah
Philautus amoenus, INGER 1966: 340; MANTHEY 1983: 23; DRING
1987: 37; MALKMUS 1988c: 176, 1989: 193, 1992a: 124; INGER
& STUEBING 1992: 41-51; MALKMUS 1994b: 230, 1995a: 73,
1996b: 20-26, 1996c: 286; INGER, STUEBING & TAN 1996: 363;
INGER & TAN 1996a: 565; MALKMUS & RIEDE 1996a: 32; INGER
et al. 2000

© R. Malkmus
Diagnosis
A high montane species of the aurifasciatus group
with a hoarse, scraping voice; lingual papilla
present; nuptial pad oval and prominent; legs rela-
tively short; coloration and pattern extremely vari-
able.

Description
Males up to 24.2 mm, females up to 34.7 mm;
TL/SVL 0.47-0.60 in males, 0.45-0.57 in females;
habitus stocky; snout broadly rounded; canthus
rostralis distinct, a quarter of eye diameter; pupil
horizontal; fingers with disks, with or without ru-
dimentary webbings; disks of toes a little smaller
than those of outer fingers; toe webbing 1(1), 2(1-
1.5/0.5), 3(1.5/1), 4(2/2), 5(1); tibio-tarsal articu-
lation reaches up to tip of snout or a little beyond;
metatarsals with many small tubercles; skin above
© M. Schroth

and on flanks smooth to granular, weakly granular

ventrally; supratympanic ridge distinct; adult males
with a subgular vocal sac.

179
 Rhacophoridae
© R. Malkmus
Fig. 176 A pair of Philautus amoenus from Pondok Lowii, 2300 m in amplexus.
iris yellowish brown to greenish with golden gleam
and network of fine black veins.
Ecology/Ethology
Philautus amoenus is a typical inhabitant of the
lower stratum of the mossy forest at elevations from
1800 to 2400 m, occasionally up to 2500 m a.s.l.
Its calls can be heard already before the onset of
dusk, increasing in intensity to climax between 6.00
and 8.00 p.m. The males call from leaves and twigs
on bushes and shrubs, rarely higher than 150 cm
above the ground, occasionally also from roots and
tree stumps overgrown with mosses nearer to the
180
ground. During prolonged periods of drought the
frog is often found in cracks in the soil and rock
crevices on the banks of streams, as well as in cavi-
ties among rock slopes.
A clutch was discovered in Bambusa gibbsiae for-
est at 2300 m altitude. It comprised 15 eggs of
8-10 mm in diameter. They were hanging on the
face of a rock fissure.
Specimens kept in a terrarium presented themselves
as exceedingly voracious, trying to devour virtu-
ally everything that moved and did not exceed their
own body sizes, in particular butterflies and hair-
less caterpillars, spiders, flies, roaches, but also
 Rhacophoridae

wood lice, ants and earthworms. Fig. LXXIV Sonagram and oscillograms of Philautus amoenus
Suitable day shelters are such as (Pondok Ubah, 2100 m; 8.30 p.m. +12,5°C)
epiphytic cushions of moss, holes in
branches and roots, rock crevices,
empty pitchers of Nepenthes, and
fruit husks.

Call: (+12.5°C) The hoarse, scrap-

ing note series bears some resem-
blance to the call of P. bunitus and
consists of notes arranged in groups
of two or three (note duration 110 ms;
internote interval 150 ms) which are
emitted at intervals of 2-3 s. Each
group of notes is preceded by a short
grace note and is distinctly pulsed.
The carrier frequency lies at 1.8 kHz
with harmonics at 5 kHz.

Distribution
Philautus amoenus is as yet known
only from Mt. Kinabalu where it ap-
pears to be restricted to the high
montane zone between Powerstation
and Layang Layang and to the
Mesilau region.

Philautus aurantium INGER, 1989 Diagnosis

Philautus aurantium I NGER , 1989: 239 – Type locality:
Mendolong, Sipitang District, Sabah
Philautus aurifasciatus, (not Philautus aurifasciatus SCHLEGEL,
1837) MALKMUS 1985: 8
Philautus sp., MALKMUS 1989: 194, 1991: 30
Philautus “A”, MALKMUS 1992a: 125
Philautus “B”, MALKMUS 1992a: 125
Philautus “A/B”, MALKMUS 1994b: 231
Philautus aurantium, HOFFMANN 1995b: 14; INGER & T AN
1996b: 66; MALKMUS 1996b: 20-26, 1996c: 286; MALKMUS
& RIEDE 1996b: 21
Philautus aurantium montanum, MALKMUS 1996c: 286
Subspecies:
Philautus aurantium gunungensis, MALKMUS &
RIEDE 1996b: 22 – Type locality: “Fagaceenmisch-
wald oberhalb des Silaubaches, 1450 m, am
Südhang des Mt. Kinabalu”
A small species of the vermiculatus group with
sharp, short, single calls. Dorsal side light brown
with small black spots; in contrast to P. amoenus
with little variation in pattern; characteristic traits
are the unmarked, pale orange anterior and poste-
rior faces of the thigh; both vomerine teeth and lin-
gual papilla absent.
Description
Males up to 27.7 mm, females up to 28 mm; TL/SVL
0.50-0.58; body stocky; head wider than long; snout
broadly rounded; canthus rostralis distinct, slightly
curved; lores oblique, weakly concave; tympanum
obscure; pupil horizontal; tips of fingers expanded
into large disks, with very small rudiments of
webbings at the bases; thenar and palmar tubercles
distinct; disks of toes smaller than those of fingers;

181
 Rhacophoridae
toe webbing 1(1), 2(1/0), 3(1/0), 4(1-0.5/1-0.5),
5(0-0.5); a low inner metatarsal tubercle, no outer
one; dorsal and lateral skin smooth; tiny tubercles
on legs and upper eyelid; chest and belly coarsely,
throat softly granular; supratympanic fold weak,
but nevertheless distinct; males with a subgular
vocal sac.
Ground coloration light yellowish brown to red-
dish brown, flanks and sides of head lighter; lower
side uniformly whitish, partly tinged with rosy;
occasionally small dark dots on throat and belly;
inguinal region and inner sides of upper thighs in-
tensely yellow; pitch black dots of various sizes on
back, sides, and thighs, arranged in rows on back;
iris reddish brown to yellowish brown with a golden
shine and a network of very fine dark veins.
The montane populations on Mt. Kinabalu were
described as a separate subspecies, Philautus
aurantium gunungensis MALKMUS & RIEDE, 1996.
They exhibit a tendency towards longer tibia, a rela-
tively wider head, and a longer inter-narial distance.
Their advertisement calls differ distinctly from
Fig. 177 Philautus aurantium gunungensis from Sg. Liwago.
© R. Malkmus
182
those of the nominate form despite the similarities
in structure.
Their eggs are half-pigmented.
Ecology/Ethology
On Mount Kinabalu, this species inhabits the
montane forests between 700 and 1500 m a.s.l. The
males form calling communities off from running
waters. They call perched on the leaves of shrubs
and bushes 50-200 cm above the ground. Calling
activity begins with the onset of dusk, increases
rapidly – particularly after rain showers – to a high
intensity and continues throughout the night up to
the dawn of the following morning. The calling
communities are static to a remarkable extent: five
colonies were observed in exactly the same places
on the slopes of the Silau-Liwago valley between
1984 and 1998. This species is occasionally found
during the day resting with the limbs drawn up
against the body on leaves; such specimens are al-
most white with the dark dotted pattern suppressed
(bird dropping mimesis).
 Rhacophoridae

Fig. LXXV Sonagram and oscillograms of Philautus aurantium

gunungensis (Sg. Silau-Silau; 1450 m; 9.00 p.m. +21°C)

© M. Schroth
Fig. 178 A calling male of Philautus
aurantium aurantium from Sg. Langanan.

Call: (+21°C) The note series con-

sists of 14-30 sharply metallic notes
(duration 120-160 ms) at a fre-
quency of 2.9 kHz and pulse modu-
lation, and an internote interval of
560-700 ms. Males engaged in du-
ets increases the volume and beat of
the note series in the second half of
their calls often resulting in a final
crescendo.

Fig. LXXVI Distribution of Philautus aurantium and Philautus bunitus in the area of the Headquarters.

183
 Rhacophoridae
In P. a. gunungensis the call duration and internote
intervals are distinctly longer, the carrier frequency
lower. In contrast to the nominate form where the
call begins with a short pulse (5-6 ms), that of
gunungensis sets in abruptly and exhibits a weak
pulse modulation.
Distribution
This species has so far been recorded only from
the province of Sabah. Mt. Kinabalu: P. a. auran-
tium: Sg. Langanan (700-950 m). P. a. gunungensis:
Sg. Silau-Silau/Liwago (1400-1500 m). The taxo-
nomic status and allocation of specimens from
Sayap (1000-1100 m) is unclear at present.
Philautus bunitus
INGER, STUEBING & TAN, 1995
Philautus bunitus INGER, STUEBING & TAN, 1995: 127 – Type
locality: Mount Lumaku, Sipitang District, Sabah
Rhacophorus sp. “2”, MALKMUS 1991a: 29
Rhacophorus “A”, MALKMUS 1992a: 121
Philautus bunitus, INGER, STUEBING & TAN 1996: 363; INGER &
TAN 1996b: 66; MALKMUS 1996c: 286; MALKMUS & RIEDE
1996b: 24; MANTHEY & GROSSMANN 1997: 127; LAKIM et al.
1999; INGER et al. 2000
Fig. 179 Male of Philautus bunitus, upper Sg. Liwago.
© M. Schroth
184
Diagnosis
A large species of the vermiculatus group with a
hoarse, scraping call; vomerine teeth present; lin-
gual papilla absent; dorsum green, often with dark
markings; venter orange.
Description
Males up to 41 mm, females up to 46 mm; TL/SVL
0.52-0.54; habitus stocky; head wider than long;
snout broadly rounded, not projecting in profile;
canthus rostralis rounded, distinct; lores oblique,
slightly concave; tympanum visible, slightly ob-
scure, half diameter of eye; pupil horizontal-ellip-
tical; supratympanic fold strong, rounded; fingers
with truncate or slightly rounded disks, rudimen-
tary webbings at bases; 1 distinct, large subarticular
tubercle on each finger; inner metacarpal tubercle
large; toes with disks smaller than those of outer
fingers; inner metatarsal tubercle oval, no outer one;
toe webbing 1(1), 2(0.5/0.5), 3(0.5/0.5), 4(1/1),
5(0.5); dorsum, limbs, and flanks with widely scat-
tered, small tubercles; venter coarsely granular;
males with a median subgular vocal sac and no
nuptial pad.
Influenced by exogenous factors and depending on
the level of excitement the coloration of the upper
 Rhacophoridae
© P. Hoffmann

© P. Hoffmann
Fig. 180 Ventral pattern of Philautus bunitus from Sg. Fig. 181 Calling male of Philautus bunitus from Sayap,
Kemantis, ca. 1050 m, Sayap. ca. 1100 m.

side may vary from pale pastel green to deep dark which often begin almost with hesitation, but then
green, sometimes uniform, but usually with black- follow at ever decreasing pauses. They exhibit a
ish to lead-grey, wide stripes and spots; flanks slightly rising frequency modulation and harmo-
lighter; throat clouded with grey or uni-
formly yellowish; chest and belly orange,
darkening posteriorly; lower sides of limbs Fig. LXXVII Sonagram and oscillograms of Philautus bunitus
and webbings also orange; iris pale golden, (Sg. Liwago, 1450 m; 8.00 p.m. +19°C)
with a rough network of black veins.

Ecology/Ethology
In contrast to the other species of Philau-
tus inhabiting Mt. Kinabalu, Philautus
bunitus is exclusive to the middle stratum
and rarely encountered below 3 m above
the ground. It inhabits the oak-chestnut for-
ests between 1000 and 1800 (1900) m a.s.l.
Its diet comprises spiders, caterpillars of
butterflies, grasshoppers, and moths. The
hardly mistakable voice of this species can
be heard throughout the night, occasionally
even during the day, particularly at the on-
set of a rainshower. It does not form iso-
lated choruses like e.g. P. aurantium, but is
relatively evenly dispersed in the forests.
Duets and vocal trios are often heard.

Call: (+19°C) The note series consists of

15-25 hoarse, scraping notes (carrier fre-
quency 1.8 kHz, note duration 190-200 ms)

185
Rhacophoridae
nics at 5-6 kHz. The note is distinctly pulsed at its
early stages, but indistinctly so towards the end;
pulse duration 6 ms.
Distribution
P. bunitus is only known from western Sabah. Mt.
Kinabalu: On Mt. Kinabalu it occurs in the entire
Liwago-Silau region between 1400 and 1800 m (up
to 1900 m in places, see also map on page 183). It is
furthermore known from the Sayap region (1000 m;
Sg. Kemantis, Sg. Wariu), and from the eastern Sg.
Mesilau (1900 m).
Philautus ingeri DRING, 1987
Philautus ingeri DRING, 1987: 21 – Type locality: Camp three,
1300 m, Gunung Mulu, Fourth Division, Sarawak
Rhacophorus colletti, (not. Polypedates colletti BOULENGER
1890) INGER 1954a: 250
Rhacophorus hosei, (not Philautus hosii BOULENGER, 1895)
INGER 1966: 304 (part), 1978: 313 (part); MANTHEY 1983: 24
(part)
Philautus ingeri, INGER & STUEBING 1992: 41-51; INGER & TAN
1996b: 67; MALKMUS & RIEDE 1996b: 25
Diagnosis
A large member of the hosii group with a soft call;
dorsum brown; vomerine teeth present, lingual pa-
pilla absent; canthus rostralis and iris with disrup-
tive patterns; upper eyelid and heels with promi-
nent tubercles.
Description
Males up to 36 mm, females up to 50 mm; TL/SVL
0.57-0.58 (males), 0.63-0.64 (females); head slightly
longer than broad; snout depressed, pointed, and
projecting in profile; canthus rostralis angular and
nearly straight; lores almost vertical; tympanum dis-
tinct; pupil horizontal-elliptical; fingers with wide
disks, webbing rudimentary; disks of toes slightly
smaller than those of outer fingers; inner metatarsal
tubercle oval, no outer one; toe webbing 1(1), 2(0/
0), 3(0/0), 4(2/2), 5(0); dorsum shagreened; tuber-
cles on dorsum, limbs, upper eyelid, and heels; throat
wrinkled; belly and undersides of thighs areolate;
males with a subgular vocal sac, no nuptial pad.
Dorsum brown, with a dark inter-orbital triangle;
dark bars on limbs; a broad black canthal stripe;
186
venter cream-coloured, with fine black speckling;
iris golden in upper third, lower two thirds dark
brown.
Ecology/Ethology
Philautus ingeri is known from the upper montane
forests. Males call at night, from leaves and
branches 1-5 m above the ground.
Call: A series of about 5 soft “quoink” notes. A
call analysis was not yet performed.
Distribution
This species is known only from Gunung Mulu in
Sarawak and Mt. Kinabalu (see Remarks at the end
of the genus account). Mt. Kinabalu: Bundu Tuhan
(1375 m).
Philautus longicrus (BOULENGER, 1894)
Ixalus longicrus BOULENGER , 1894a: 88 – Type locality:
Palawan
Philautus aurifasciatus, (not Philautus aurifasciatus SCHLEGEL,
1837) INGER 1966: 341 (part); MANTHEY 1983: 24 (part)
Philautus longicrus, INGER & STUEBING 1992: 41-51; INGER,
STUEBING & TAN 1996: 363; MALKMUS & RIEDE 1996a: 35;
INGER et al. 2000
Diagnosis
A small brown species of the aurifasciatus group
with a rattling call; vomerine teeth and lingual pa-
pilla absent; similar to P. petersi, but females with-
out a tubercle on the tip of the snout.
Description
Males up to 23 mm, females up to 29 mm; TL/SVL
0.59-0.64; head wider than long; snout obtusely
pointed; canthus rostralis distinct, lores concave;
tympanum obscure, one third eye diameter; pupil
horizontal; tips of fingers with wide round disks;
disks on toes smaller than those on outer fingers; an
elongated inner, but no outer metatarsal tubercle; toe
webbing intermediate in extension between P. petersi
and P. mjobergi; skin shagreened above; tubercles
and ridges irregularly distributed on back, head, and
limbs; throat weakly, chest and belly coarsely granu-
lar; supratympanic fold present; males with a
subgular vocal sac; nuptial pad oval and prominent.
 Rhacophoridae

Coloration and pattern extremely variable in speci-

mens from Sarawak and Palawan; there are speci-
mens with a barred dorsal pattern as well as conco-
lor ones, but none with a white streak below the
eye or a dark network on a white ground. There is
no description of specimens from Mt. Kinabalu.

Ecology/Ethology
Almost nothing is known about the life history of
this species other than it was found in primary for-
est between 900 and 3000 m altitude. It is active at
night and arboreal, by day it rests under litter on
the forest floor.

Call: A harsh rattle, similar to the call of P. mjo-

bergi.

Distribution
Philautus longicrus is known from the Philippines
(1200-1550 m), Sarawak (900-1200 m), and Sabah
(up to 3000 m). Mt. Kinabalu: Layang Layang,
Sg. Tibabar.
INGER & STUEBING (1992) listed this species for an
elevation of 3000 m (Pakka). Bioacoustic studies,
however, promote the idea that it shares the habitat
on Mt. Kinabalu with P. amoenus at 1800-2400 m
a.s.l.
Philautus mjobergi SMITH, 1925
Philautus mjöbergi SMITH, 1925a: 11 – Type locality: Mt.
Murud, 7000 feet, Sarawak
Philautus petersi, (not Philautus petersi BOULENGER, 1900)
SMITH 1931: 20 (part)
Philautus aurifasciatus, (not Philautus aurifasciatus SCHLEGEL,
1837) INGER 1966: 341 (part), 1978: 313 (part); MANTHEY
1983: 24 (part), MALKMUS 1985: 8, 1987: 284, 1988c: 176
Philautus mjobergi, SMITH 1931: 21 (part); DRING 1987: 38;
MALKMUS 1989: 193, 1991: 30, 1992a: 124; INGER & STUEBING
1992: 41-51; MALKMUS 1994b: 230; INGER, STUEBING & TAN
1996: 363; LAKIM et al. 1999; INGER et al. 2000
Philautus mjoebergi, HOFFMANN 1995b: 14; MALKMUS 1996c:
286; MALKMUS & RIEDE 1996a: 29
Diagnosis
A highly variable species of the aurifasciatus group
with a sharp, irregular, rattling call; vomerine teeth
absent; lingual papilla present, sometimes absent;
females without tubercle on tip of snout.
© P. Hoffmann
Fig. 182 Philautus mjobergi from Sg. Silau-Silau, ca.
1550 m.
Description
Males up to 24 mm, females up to 32 mm; TL/SVL
0.52-0.62; head wider than long; snout rounded to
elliptical in outline; canthus rostralis distinct, lores
oblique and concave; tympanum obscure; pupil
horizontal; finger tips with wide oval disks; dis-
tinct rudiments of webbings; disks on toes smaller
than those of outer fingers; inner metatarsal tuber-
cle oval, no outer one; toe webbing 1(1), 2(1/1),
3(1/1), 4(2-2.5/2-2.5), 5(0.5-1); dorsum sha-
greened, sometimes with a subtle tubercle on up-
per eyelid or on heel; supratympanic ridge distinct;
throat granular, belly and undersides of thighs
areolate; males with a subgular vocal sac; nuptial
pad oval, prominent.
Dorsal ground coloration pale brown with a ten-
dency towards yellowish or reddish; inter-orbital
bar and dorsal stripes usually present, narrowing
at mid-dorsum to form an sand-glass like marking;
concolor specimens are occasionally found, or
those with a light, broad medial stripe; thighs with
3-4 dark bars; flanks lighter, sometimes spotted in
lower third; lower sides of body yellowish to soft
flesh-coloured, sometimes obscurely spotted with

187
 Rhacophoridae

grey on chest; iris golden with a

dark horizontal bar.

Ecology/Ethology
On Mt. Kinabalu, Philautus
mjobergi is a typical faunal ele-
ment of the oak-chestnut forests
at altitudes between 900 and
1700 m, occasionally up to
1800 m a.s.l. The males call
mainly at night with maximum
intensities between 6.30 and
8.00 p.m., and again between
4.30 and 5.30 a.m, but can occa-
sionally also be heard during the
day. They advertise from leaves
and twigs of shrubs and bushes,
but preferably from thorny rattan
thickets, 20-250 cm above the
© M. Schroth

ground. On Mt. Murud Mjöberg

found them depositing their eggs
in the pitchers of the Pitcher
Fig. 183 Philautus mjobergi from Sg. Silau-Silau. Plants (Nepenthes), no other wa-
Fig. LXXVIII Sonagram and oscillograms of Philautus mjobergi ter being available for the purpose
(Sg. Silau-Silau, 1450m; 6.30 p.m. +20°C) (SMITH, 1925).
A female captured on Sg. Silau-
Silau in August deposited shortly
afterwards 7 large, unpigmented
eggs (diameter of the jelly shell
6 mm) on a dead leaf in a ter-
rarium.

Call: (+20°C) The advertisement

call consists of a series of 4-8
metallic, sharp, pulsed notes (car-
rier frequency 2.7-3.5 kHz; note
duration 80-105 ms; internote in-
terval 70-80 ms) which is fol-
lowed by a pause of 0.8-1 s and a
short, marked final grace note. A
note series inclusive of the grace
note lasts 1.8-2.5 s and is re-
peated at irregular, often long in-
tervals.

Distribution
Philautus mjobergi is probably
the species of Philautus with the

188

widest range on Borneo. Mt. Kinabalu: Common
in the entire Headquarters region (1400-1800 m)
with additional records from Mesilau Cave, Marei
Parei, and Sayap.
Philautus petersi (BOULENGER, 1900)
Ixalus petersi BOULENGER, 1900: 185 – Type locality: Mts.
Penrissen, Dulit, and Kina Balu, Borneo and Great Natuna
Philautus aurifasciatus, (not Philautus aurifasciatus SCHLEGEL,
1837) INGER 1966: 341 (part), 1978: 313 (part); MANTHEY
1983: 23 (part)
Philautus petersi, SMITH 1931: 20 (part); MALKMUS 1991: 30,
1992a: 124; INGER & STUEBING 1992: 41-51; MALKMUS 1994b:
231, 1996c: 286; INGER, STUEBING & TAN 1996: 363; INGER
& TAN 1996a: 566; MALKMUS & RIEDE 1996a: 35; HOFFMANN
1998c: 15; LAKIM et al. 1999; INGER et al. 2000
Taxonomic remarks: The original type locality is
a composite based on specimens from Mt. Dulit,
Mt. Penrissen, and Mt. Kinabalu (Pakka). However,
Fig. 184 Female of Philautus petersi from Sayap.
Rhacophoridae
the Kinabalu syntype is “an unidentifiable juve-
nile” (DRING 1987).
Diagnosis
A stout species of the aurifasciatus group with a
very characteristic, single-pulse-train call; vomer-
ine teeth absent; lingual papilla present or absent;
dorsal surface tubercular; female with a tubercle
on tip of snout.
Description
Males up to 27 mm, females up to 37 mm; TL/SVL
0.5-0.66.
DRING (1987) noted marked differences in the mor-
phological ratios of specimens from various geo-
graphical regions. Similar differences are also
found on a smaller scale in specimens from vari-
ous elevations from Mt. Kinabalu (MALKMUS &
RIEDE 1996a).
Head broader than long; snout elliptical in outline,
rounded in profile; canthus rostralis rounded, lores
© M. Schroth
189
 Rhacophoridae
© M. Schroth

Fig. 185 Male of Philautus petersi from upper Sg. Liwago.

oblique and concave; tympanum obscure; pupil
horizontal; tips of digits with wide oval disks; dis-
tinct rudiments of webbings; disks on toes some-
Fig. 186 A pair of Philautus petersi in amplexus from
Sayap.
what smaller than those of outer fingers; inner
metatarsal tubercle oval, no outer one; toe web-
bing 1(1), 2(0.5-1/0.5-1), 3(0.5-1/0.5-1), 4(2/2),
5(0.5-1); tibio-tarsal articulation reaching up to eye;
dorsum distinctly tubercular, particularly on upper
eyelid, shoulders, upper side of legs, and below
jaws; heel with a small, spine-like tubercle; supra-
tympanic fold distinct; throat granular or wrinkled,
belly and undersides of thighs areolate; males with
a subgular vocal sac; nuptial pads prominent.
Dorsum brown to grey, pattern highly variable;
flanks lighter; groin and lower flanks flecked with
reddish brown; underside whitish to grey; iris
brown to golden, with a central horizontal dark bar.

Ecology/Ethology
On Mt. Kinabalu, P. petersi occurs in submontane
© P. Hoffmann

and montane forests between 800 and 1700 m a.s.l.

The males call from shrub thickets, 30-300 cm
above the ground, often adjacent to rivulets. The

190
 Rhacophoridae

day is spent hidden beneath dead leaves Fig. LXXIX Sonagram and oscillograms of Philautus petersi
and bark. The female’s keratinized tu- (Sg. Liwago, 1450 m; 7.30 p.m. +19°C)
bercle on the tip of the snout is possibly
used for the excavation of nests, perhaps
in epiphytic mosses. Amplexi were ob-
served in the month of August.
A clutch of this species was found on a
dead leaf in the leaf litter layer at Sayap.
It contained up to 25 unpigmented eggs.

Call: (+19°C) The call is a 1.2-1.5 s long

train of 16-56 rough pulses (pulse dura-
tion 20-25 ms; pulse rate from 27 (in
specimens from Sayap) to 46 per sec-
ond; dominant frequency 1.6-2.2 kHz;
weak harmonics between 5 and 7 kHz;
no frequency modulation. The calls are
emitted at intervals of several minutes.

Distribution
Malay Peninsula; widely distributed on
Borneo. Mt. Kinabalu: Langanan Val-
ley (800-900 m), Sayap (900-1100 m),
Marei Parei (1500 m), Headquarters re-
gion (1450-1700 m).
The record by SMITH (1931a), “Kambo-
rangah, 2200 m”, very likely refers to P.
amoenus. That the “unidentifiable juvenile” (DRING Philautus saueri
1987) from Pakka Cave at 3000 m altitude in the
series of syntypes represents P. petersi, can be dis- MALKMUS & RIEDE, 1996
missed with certainty.
Philautus saueri MALKMUS & RIEDE, 1996a: 29 – Type locality:
Pakka (3050 m), Mt. Kinabalu
Philautus cf. mjoebergi, (not Philautus mjöbergi SMITH, 1925)
Fig. 187 Eggs of Philautus petersi from Sayap, ca. MALKMUS 1995a: 70
1000 m. Philautus saueri, MALKMUS 1996c: 286

Diagnosis
A small, stocky, high-montane species of the
aurifasciatus group with a soft, hoarse, rattling call.
Vomerine teeth absent; dorsal side of body brown
with a whitish inter-orbital bar.

Description
Male 21.4 mm; TL/SVL 0.54; head slightly broader
than long; snout rounded in both outline and pro-
© P. Hoffmann

file; canthus rostralis rounded, lores slightly con-

cave; tympanum obscure; pupil horizontal; tips of
digits with large oval disks; webbing rudiments

191
 Rhacophoridae
distinct; disk on toes smaller than those on outer
fingers; inner metatarsal tubercle distinct, no outer
one; toe webbing 1(1), 2(1/0.5), 3(1.5/1), 4(2/2),
5(1); dorsum smooth; heel with a small tubercle;
supratympanic fold prominent, tightly bent, from
eye to axilla; 2 rows of ridges from eyelid to centre
of back; flanks, throat, and belly granular; males
with a subgular vocal sac.
Upper sides of body, tibiae, and forearms blackish
brown, peppered with small whitish dots; remain-
ing portions of limbs and sides of body white to
whitish grey, reticulated with black; a whitish or-
bital bar separates reddish brown of snout from
blackish brown of dorsum; a broad white area be-
tween eye and upper lip, bordering blackish brown
of tympanum area; supratympanic fold whitish;
belly white, with a umber-coloured, reticulated
pattern; throat yellowish brown; lower sides of feet
and hands, and upper sides of first two fingers
whitish yellow; central portion of iris brownish
Fig. 188 Philautus saueri from Pakka Cave region, 3000 m.
© R. Malkmus
192
grey; with wider upper and narrower lower mar-
gins in yellowish green.
Ecology/Ethology
Calls of P. saueri can be heard at altitudes between
2200 and 3050 m a.s.l., very rarely though below
2400 m. In contrast to all other species of Philautus
occurring on Mt. Kinabalu, the males of this spe-
cies call also during daylight hours, even in bright
sunshine. Their diurnality must certainly be re-
garded as an adaptation to the fact that the tem-
peratures at these altitudes not rarely drop to just
5°C at night. This point of view is supported by the
observation that calls could still be recorded at
10.00 p.m. in the less elevated mossy forest (be-
low 2600 m). The males call out of low, very dense
bush vegetation.
Four pitchers of Nepenthes villosa were discovered
in the Leptospermum-Dacrydium forest near Pakka,
in August 1994, which each contained 9-17 gelati-
 Rhacophoridae

nous capsules of 11-15 mm in diameter of Fig. LXXX Sonagram and oscillograms of Philautus saueri
P. saueri. 20-30% of these capsules did nei- (Pakka, 3000 m; 11.00 a.m. +15°C)
ther contain eggs nor embryos, but were
merely filled with liquid. The other shells
held larvae at various stages of development.
All these pitchers were dead and leaking,
although their well-shaded locations and
contact to the ground secured the necessary
moisture for the development of the larvae.
It remains unclear, though, whether the con-
tents of the individual pitchers were the prod-
uct of one or several pairs of these frogs.

Call: (+15°C) A note series consists of 2-4(5)

pulsed notes (note duration 30-40 ms; pulse
rate 150 per second) at intervals of 70-100 ms
at a carrier frequency of 1.9-2.6 kHz. Spaced
by a pause of often several seconds, a drawn-
out “gweeh” sound (duration 90-120 ms)
may follow, which may also be emitted
alone. The structure of the note series ex-
hibits certain similarities with that of P.
mjobergi.

Distribution
Philautus saueri is as yet known only from the
montane forest between Layang Layang and Pakka
Cave.
Genus Polypedates TSCHUDI, 1838

The genus Polypedates is distributed from China

Fig. 189 Tadpole of Philautus saueri. and Japan, through tropical Southeast Asia, up to
the Philippines, Borneo, and Java.
Although CHANNING (1989) demonstrated in a
phylogenetic study that Polypedates is a genus
separate from Rhacophorus, its validity has re-
mained a subject of debate. In contrast to Rhaco-
phorus, a portion of the scalp is fused to the skull
in Polypedates (except in P. colletti), and the ring-
fold on the lower side of the digital disks typical
for Rhacophorus is missing in Polypedates. While
Rhacophorus possess webbings between the fin-
gers, such are absent in Polypedates.
The Bornean species are relatively large tree frogs
with compressed bodies and long limbs. Vomerine
teeth present, tympanum visible, pupil horizontal.
© H. Sauer

Fingers and toes with enlarged pads at their tips,

toes webbed over half to almost full length. Males
with subgular vocal sac and nuptial pads.

193
Rhacophoridae
Tab. 19 Distribution of Polypedates-species on Borneo.
The eggs lack pigmentation and are deposited in a
foam nest which is attached to the leaves of bushes
in a manner that hangs freely above a water body.
Tadpoles with almost spheroidal bodies, eyes on
sides of head, oral disk ventral, strong black beaks
and labial teeth rows present; tail with high fins.
Key to the Bornean species of Polypedates
1 Skin of head not fused to skull; conspicuous X-
shaped mark on dorsum .......................... colletti
• Skin of head fused to skull in adult specimens . 2
2 A serrated bony crest projecting above the tym-
panum .................................................. otilophus
• No such crest ..................................................... 3
Polypedates leucomystax
(GRAVENHORST, 1829)
Hyla leucomystax GRAVENHORST, 1829: 36 – Type locality: Java
Rhacophorus maculatus, (not Polypedates maculatus GRAY,
1834) BOULENGER 1887b: 95; MOCQUARD 1890: 149
Rhacophorus leucomystax var. quadrilineatus, (syn. after
STEJNEGER 1907: 157) MOCQUARD 1890: 150
Rhacophorus leucomystax sexvirgata, (syn. after STEJNEGER
1907: 157) VAN KAMPEN 1923: 249
Rhacophorus leucomystax, VAN KAMPEN 1923: 246; SMITH 1931:
9; INGER 1954a: 250; MALKMUS 1996c: 286
Rhacophorus leucomystax leucomystax, INGER 1966: 307
Polypedates leucomystax, INGER 1978: 313; MANTHEY & DENZER
1982: 15; MANTHEY 1983: 24; MALKMUS 1985: 11, 1987: 283,
1988d: 7; HOFFMANN 1995b: 14; INGER, STUEBING & TAN 1996:
363; HOFFMANN 1998b: 89
Polypedates leucomystax leucomystax, MATSUI 1979: 341;
LAKIM et al. 1999; INGER et al. 2000
Taxonomic remarks: NARINS et al. (1998) pointed
out the morphological, genetical, bioacoustical, and
194
The Bornean species live in the primary and sec-
ondary forests of the lowlands. P. leucomystax and
P. otilophus, appear also in cleared forests, agri-
cultural fields, villages, and towns.
Four species are known from Borneo, three of
which also from Mt. Kinabalu:
3 A broad black temporal stripe ............. macrotis
• Temporal stripe absent or weak; often 4 narrow
dorsal stripes .................................. leucomystax
ethological differences between the striped and
stripeless forms which they observed sympatrically
on the Malay Peninsula and considered it justified
to grant both species ranks.
Diagnosis
A small to moderately large, slender frog with long,
slender limbs; specimens from Mt. Kinabalu usu-
ally with 4 narrow dorsal stripes; no bony crest
above tympanum.
Description
Males up to 50 mm, females up to 75 mm; TL/SVL
0.47-0.60; head longer than broad; snout obtusely
pointed; canthus rostralis sharp, lores not concave;
tympanum distinct, 75% or more of eye diameter;
finger tips with large round disks, webbing rudimen-
tary; toe tips not as wide as those on outer fingers;
toe webbing 1(0), 2(0/0), 3(0/0), 4(2/2), 5(0); an oval
inner metatarsal tubercle, outer one small or absent;

© P. Hoffmann
Fig. 190 Polypedates leucomystax from Poring, 550 m.
dorsal skin smooth, venter coarsely granular; su-
pratympanic fold present, curved.
Light beige, yellowish, or brown above, with dark
flecks or four narrow stripes running down length
of back; limbs with dark crossbars; undersides
whitish to whitish grey; upper lip white; nuptial
pads yellowish; iris light golden with a fine black
network.
Tadpoles: Body spheroidal, flattened above,
rounded below; eyes dorsolateral; oral disk ven-
tral, subterminal; labial teeth rows formula 4(2-4)/3;
beaks black, strong, finely serrated; spiracle sinis-
tral; tail tapering abruptly in the last quarter to a
slim, drawn-out tip; total length up to 50 mm. Dor-
sum brown to yellowish, with dark stripes at stages
40-41; whitish ventrally; tail grey with dark dots;
fins transparent, dark spotted (YORKE 1983).
Ecology/Ethology
Polypedates leucomystax inhabits the marginal
zones of primary and secondary forests, but also
cultivated areas, road sides, and villages (where it
may even be encountered inside houses). It is a
nocturnal frog which may, however, also forage
during rain showers in the day. Its diet comprises
insects, spiders, and even small lizards. The males
congregate at stagnant and slowly flowing water
bodies and form choruses. It is not rare that a fe-
Rhacophoridae
© U. Manthey
© U. Manthey
© U. Manthey
From top to base: Polypedates leucomystax.
Fig. 191 Typical foam nest from West Malaysia.
Fig. 192 Tadpole from West Malaysia.
Fig. 193 Tadpole from Pulau Bintan, Indonesia.
male assumes the active advertising by ascending
to a branch, drumming against it for minutes and
occasionally emitting a croak. It does usually not
take long for some males to arrive. Once a pair is
united in an amplexus, it chooses a site for build-
ing a nest in the branchwork of a bush 10-70 cm
above a water surface or along the bank of pools
on the ground (earth, rocks). The female then ex-
pels from its cloaca a liquid produced by foam
glands which it whips to a primary foam with its
hind limbs. The subsequent thrust contains the eggs
(150-900) which are mixed with the male’s sperm
and the foam. The female does most of the nest
building and is supported by the male only occa-
sionally (KABISCH et al. 1994).
195
Rhacophoridae

Five to six days after oviposition the tadpoles

emerge and drop from the foam nest into the water
beneath (m: 2b).

Call: NARINS et al. (1998) identified three separate

call types for each of the two forms (striped and
stripeless, respectively). In the striped form – only
this one has so far been found on Mt. Kinabalu –
there are two pulsed call types with dominant fre-
quencies at 2238 Hz (call duration 174 ms, 13
pulses), and 934 Hz (call duration 56 ms, 4-5 pulses),
respectively, and staccato calls (dominant frequency
997 Hz, call duration 527 ms). Analyses of the
vocalisations produced by Kinabalu specimens are

not available (MATSUI 1982a, MATSUI et al. 1986,
SANCHEZ-HERRAIZ et al. 1995).
Distribution
This species has a vast distribution, ranging from
eastern India and southern China to Java, Borneo,
and the Philippine Islands, from sea level up to
1000 m, on Mt. Kinabalu up to 1500 m a.s.l. Mt.
Kinabalu: Sayap, Kiau, Tenompok, Bundu Tuhan,
Lobang, Kundasang, Poring.
Whether the “egg mass” reported from Kamboran-
goh by MATSUI (1979) really belongs to P. leucomys-
tax is questionable.
Polypedates macrotis (BOULENGER, 1891)
Rhacophorus macrotis BOULENGER, 1891: 282 – Type locality:
Baram District, Sarawak
Rhacophorus leucomystax linki, (syn. after INGER 1966: 308)
INGER 1956: 409
Rhacophorus macrotis, INGER 1966: 313
Polypedates macrotis, I NGER & S TUEBING 1992: 41-51;
HOFFMANN 1995b: 14; INGER, STUEBING & TAN 1996: 363;
LAKIM et al. 1999; INGER et al. 2000
© T. Eltz
Fig. 194 Polypedates macrotis from Deramakot, Sabah.
heads and finger tips, but narrower tympani than
males; skin smooth, some specimens with tiny
bumps on back; often a low, whitish dermal ridge
of skin on outer edge of forearm.
Upper side of head, back, and limbs tan to brown;
flanks sometimes with black dots; limbs with dark
crossbars; a broad dark stripe from eye to arm;
undersides whitish, with or without dark dots; throat
mottled with brown; iris light golden.
Tadpoles: Head-body portion oval; snout broadly
rounded; body flattened above; eyes lateral, vis-
ible from below; oral disk ventral, subterminal; la-
bial teeth rows formula 5(2-5/3(1) or 6(2-6)/3;
beaks weakly serrated; tail convex, 1.2-1.7 times
Fig. 195 Polypedates macrotis from Poring.

Diagnosis
A medium-sized to large tree frog with a triangu-
lar head and a characteristic, broad, blackish brown
temporal stripe.

Description
© P. Hoffmann

Males up to 57 mm, females up to 90 mm; TL/SVL

0.49-0.60; very similar in its morphological details
to P. leucomystax; females have relatively wider

196

head-body-length; total length 55-60 mm. Head and
body dark green, dorsally and laterally with light
spots, below silvery grey; caudal muscle with a dark
reticulated pattern; dorsal fin green, ventral fin
white with small black dots (INGER 1985).
Ecology/Ethology
Polypedates macrotis appears in lowland primary
and secondary forests. It is nocturnal, preying upon
insects (particularly roaches and Hemiptera) and
spiders. Males form large calling communities on
the ground or in low vegetation (up to 3 m) near
rainwater-filled pools.
The females attach large foam nests to vegetation
(shrubs, small trees, herbs, grasses) overhanging
ditches and rainwater-filled pools (m: 2b, c) at the
edges of forests.
Call: A long, drawn-out chuckle, the first note dis-
tinctly pronounced. The advertisement calls con-
sist of main and short notes, with the former last-
ing 0.1-0.79 s and consisting of 1-3 notes (dura-
tion 0.09-0.12 ms, 5-6 pulses per note). The short
notes (duration 0.005-0.01 s) are emitted before or
after the main notes. The frequency range is wide,
covering 0.4-2.6 kHz (+22°C) (MATSUI 1982b).
Distribution
This frog is known from the Malay Peninsula,
Sumatra, Borneo, Palawan, and the Sulu Archi-
pelago (200-1350 m a.s.l.). Mt. Kinabalu: Sayap,
Poring: Sg. Kipungit.
Polypedates otilophus
(BOULENGER, 1893)
Rhacophorus otilophus BOULENGER, 1893: 527 – Type locality:
Bongon, N. Borneo
Rhacophorus otilophus, MALKMUS 1989: 192
Polypedates otilophus, INGER & STUEBING 1992: 41-51; INGER,
STUEBING & TAN 1996: 363; LAKIM et al. (1999); INGER et al.
(2000)
Diagnosis
A large tree frog with a triangular head and a pro-
truding, sharp point at the angle of the jaws; a ser-
rated crest overhanging the tympanum; toes webbed
over only half their lengths.
Rhacophoridae
Description
Males up to 80 mm, females up to 97 mm; TL/SVL
0.47-0.55; head longer than broad; snout rounded
in profile, projecting; tympanum distinct, oval, 3/4
of eye diameter; tips of fingers expanded into large
disks; rudimentary webbing between 1st and 2nd fin-
gers; disks of toes much smaller than those of outer
fingers; an oval inner metatarsal tubercle, no outer
one; toe webbing 1(1), 2(1/0.5), 3(0.5/0.5), 4(2/2),
5(0.5); skin above smooth, flanks and undersides
coarsely granular; a pointed projection on heel and
often another, smaller one on elbow; a row of 4-6
white tubercles below vent.
Yellowish brown above, with numerous thin, dark
stripes running down back from the head; inguinal
region white with black lines and dots; thighs with
7-11 narrow black bars on the anterior and poste-
rior faces; undersides dirty white; nuptial pads grey-
ish or yellowish; iris yellowish grey.
Tadpoles: Head-body section spherical, top of head
flattened; very similar to the larvae of P. macrotis;
labial teeth rows formula 4(2-4)/3 or 5(2-5)/3; to-
tal length 63 mm, maximum up to 80 mm. Yellow-
ish green above, below silvery white; dorsal fin with
golden dots.
Ecology/Ethology
Polypedates otilophus dwells the edges of pri-
mary forests as well as habitats modified by man
Fig. 196 Polypedates otilophus from Poring.
© P. Hoffmann
197
 Rhacophoridae
© M. Veith
Fig. 197 Polypedates otilophus from West Kalimantan.
Genus Rhacophorus
KUHL & VAN HASSELT, 1822
Species of Rhacophorus are known from the re-
gion from India, Sri Lanka and China to Japan, the
Philippines, and the Greater Sunda Islands.
This genus includes small to moderately large, for-
est-dwelling tree frogs with robust bodies and long
limbs. Most species possess vomerine teeth, but
this trait is not constant in all species. There is no
ossification of the scalp which distinguishes it from
Polypedates. Tympanum visible, eyes large with a
horizontal pupil; tips of fingers and toes expanded
into large, round or oval disks, those of toes not as
wide as those of outer fingers; webbings between
fingers and toes present; undersides of fingers with
circum-marginal grooves. Some species are able
to perform paragliding flights by spreading their
excessively webbed hands and feet which thus de-
198
(secondary forests, tree planta-
tions, logged areas, even villages).
It is a lowland species, occurring
from sea level up to 500 m, rarely
higher (up to 1000 m a.s.l.). Its
diet consists of insects (especially
arboreal crickets) and spiders.
Captured specimens emanate a
characteristic, strong odour.
Males call at night from low veg-
etation (< 2 m) near small pools
(m: 2b, c) where the females, simi-
lar to P. macrotis, attach their
foam nests to low herbs about 30 cm
above the water surface.
Call: A series of slurred, rasping
notes followed by some loud
“chucks” (I NGER & S TUEBING
1997). A call analysis is not avail-
able.
Distribution
Borneo, Sumatra. Mt. Kinabalu:
Only known from Poring: Sg.
Kipungit.
celerates their descent (angle of descent less than
45°) due to the enlarged surface area. Males with a
subgular, median vocal sac and nuptial pads.
Rhacophorus construct foam nests which are at-
tached to vegetation above flowing (R. anguliros-
tris, cyanopunctatus, gauni) or stagnant water bod-
ies (all other species), sometimes next to water.
Tadpoles are free-swimming and actively feeding;
mouth ventral, with jaw beaks and labial teeth rows;
eyes dorsolateral; spiracle sinistral; anal tube me-
dian or lateral. There are two basically different
morphotypes of Rhacophorus larvae on Borneo:
type I is associated with stream microhabitats (m:
1b, c, d, e, g) and has cup-like oral disks, while
type II dwells stagnant water pools or water-filed
holes in phytothelmes (m: 2b-g).
Bornean species occur in all strata of primary and
old secondary forests.
A total of 14 species are known from Borneo, 10
from Mt. Kinabalu.

Preliminary key to the Bornean species of Rhacophorus
(partly after INGER & STUEBING 1997)
1 Webbing of foot green; a dermal projection above
the anus ................................................ dulitensis
• Webbing of foot at least partially black ........... 2
2 Body green ......................................................... 3
• Body not green ................................................... 6
3 Outer fingers webbed for only 1/3; SVL < 25 mm
.................................................................... kajau
• Outer fingers webbed for more than 1/2; SVL >
30 mm ................................................................. 4
4 Outer fingers fully webbed ................................ 5
• Outer fingers not fully webbed ............................
....................................... angulirostris (females)
5 A wide skin flap along arm and tarsus; flanks yel-
low or greenish ........................... nigropalmatus
• Flap of skin along arm and tarsus; flanks black
with blue and yellow spots ............... reinwardtii
6 Webbings of foot orange.................................... 7
• Webbings of foot not orange ............................. 8
7 Orange network on belly; forearm with flap of
skin along outer edge ............................ pardalis
• No such network on belly; no skin flaps .. rufipes
8 A pointed projection on heel .............. baluensis
• No such projection ............................................ 9
Tab. 20 Distribution of Rhacophorus-species on Borneo.
Rhacophoridae
9 Numerous conical projections on back head, arms,
and legs ............................. everetti macroscelis
• Without conical projections .......................... 10
10 White spot(s) below eye ............................... 11
• Without spot(s) ............................................... 13
11 Upper eyelid with dermal spike .............. gauni
• No such dermal spike .................................... 12
12 Flanks, groin, and inner sides of thighs with blue
spots ........................................ cyanopunctatus
• No blue spots; sides yellow with black or brown
spots .................................. angulirostris (part.)
13 Lower arm and tarsus with wavy fringes of skin
................................................. appendiculatus
• Without wavy fringes of skin ........................ 14
14 Inner side of 2nd finger webbed up to disc .........
......................................................... harrissoni
• Webbing smaller ............................................ 15
15 Snout pointed, canthus rostralis sharp ...............
........................................... angulirostris (part.)
• Snout blunt, without sharp ridge ....... fasciatus
199
Rhacophoridae

Fig. LXXXI Distribution of Rhacophorus-species

on Mt. Kinabalu

200

Rhacophorus angulirostris AHL, 1927
Rhacophorus acutirostris MOCQUARD , 1890a: 151 – Type
locality: Kina Balu
Rhacophorus acutirostris BOULENGER 1894a: 81; HANITSCH
1900a: 73; VAN KAMPEN 1923: 260; SMITH 1931: 17; INGER
1966: 283, 1978: 313; MANTHEY 1983: 13
Rhacophorus angulirostris AHL, 1927: 45 (substitute name for
Rhacophorus acutirostris), 1931: 150; INGER & TAN 1990: 4;
MALKMUS 1991a: 30, 1992a: 122; INGER & STUEBING 1992:
41-51; MALKMUS 1993c: 35, 1994b: 229; WONG 1994: 29-
37; HOFFMANN 1995b: 15; MALKMUS 1995d: 245, 1996b: 20-
26, 1996c: 286; INGER , STUEBING & TAN 1996: 363; MANTHEY
& GROSSMANN 1997: 131; MALKMUS 1999: 13-19; LAKIM et
al. 1999; INGER et al. 2000
Philautus bimaculatus, (not Rhacophorus bimaculatus PETERS,
1867) MALKMUS 1989: 194
Rhacophorus sp. 1, MALKMUS 1991a: 28
Rhacophorus “B”, MALKMUS 1992a: 122
Diagnosis
A small rhacophorid species with relatively short
limbs and a short, pointed snout; only the outer two
fingers are fully webbed; upper portion of iris blue.
Fig. 198 Female of Rhacophorus angulirostris from Sg. Silau-Silau, 1550 m.
Rhacophoridae
Description
Males up to 34.5 mm, females up to 51 mm; TL/SVL
0.48-0.67; head broader than long; snout pointed,
projecting; canthus rostralis sharp, lores weakly
concave; tympanum distinct; vomerine teeth
present; finger webbing 1(1), 2(1/1), 3(1.5/1.5),
4(0.5-1); toe webbing 1(0), 2(0-0.5/0-0.5), 3(0-0.5/
0-0.5), 4(1/1), 5(0-0.5); a small inner metatarsal
tubercle, no outer one; skin smooth above, coarsely
granular below except on throat; supratympanic
fold curved, from eye to axilla.
Males light brown to yellowish brown above, of-
ten with dark flecks, an X-mark, or crossbars on
back and crossbars on limbs; flanks and rear of
thighs yellow with brown or black dots, particu-
larly in groin region; ivory-white flecks below eye
(up to 9 on one side) or not; belly whitish, throat
clouded with greyish; iris dark golden brown, up-
per portion blue.
Females above dark green to yellowish green with
obscure dark markings; reddish grey below line
© P. Hoffmann
201
 Rhacophoridae
© R. Malkmus
Fig. 199 Male of Rhacophorus angulirostris from Sg.
Liwago.
canthus rostralis – eye – supratympanic fold, some-
times with white spots; venter yellow; flanks and
iris similar to those of males.
Tadpoles: Head-body oval, slightly flattened above,
rounded below; snout rounded; oral disk ventral, sub-
terminal, cup-like; labial teeth rows formula 4(3-4)/
4-5; tail convex, tapering gradually to tip, twice as
long as body; total length up to 36 mm. Coloration
Fig. 201 Female of Rhacophorus angulirostris from Sg.
Liwago, 1500 m.
© R. Malkmus
202
© H. Sauer
Fig. 200 Oral disc of Rhacophorus angulirostris, same
specimen as below.
dark brown or greyish brown above and laterally,
pigmentless below, with a silvery sheen; tail light
brown with dark spots; fins dusky with black spots.
Ecology/Ethology
Rhacophorus angulirostris was found along rocky
streams in primary forests, between altitudes of 500
and 1650 m. Calling males perch on vegetation
overhanging such streams (1-4 m above the water
surface). Calls are emitted between dusk (6.00-
6.30 p.m.) and dawn (5.30 a.m.) throughout the
night, sometimes with large breaks of 5-20 min-
utes. To the human ear the calls are very similar to
those of large arboreal crickets (Pseudophyllinae).
Captured specimens occasionally show a catalep-
tic behaviour.
Fig. 202 Tadpole of Rhacophorus angulirostris.
© H. Sauer
 Rhacophoridae
© M. Schroth

Fig. 203 Tadpole of Rhacophorus angulirostris.

The larvae live in montane streams with rocky bot-

toms (m: 1b, c).
The diet of the adults comprises insects and spi-
ders. Parts of a prey item that may protrude later-
ally from the mouth are taken with the hands and
straightened out forward.

Call: (+19°C) The call consists of a single, very

sharp, and shrill “tshirk”-note. This note (note du-
ration 90-100 ms) is composed of two major pulses
with a distinct intensity modulation; dominant fre-
quency 3.4-4.8 kHz; call intervals irregular, often
very long (5-10 min.).
Fig. LXXXIII Distribution of Rhacophorus-species in
Distribution
the Poring area.
This species is known only from Sumatra
and the Bornean province of Sabah. Mt.
Kinabalu: Common along Sg. Luidan, Sg.
Kinateki, Kiau region, Kenokok, Marei
Parei, Sayap: Sg. Kemantis, Poring: Sg.
Langanan (800-900 m), Sg. Kipungit (500-
600 m; questionable), Sg. Silau-Silau/Sg.
Liwago and tributaries (1400-1550 m), see
also map on page 200 and 204.

Fig. LXXXII Sonagram and oscillogram of

Rhacophorus angulirostris
(Sg. Silau-Silau, 1450 m; 7.00 p.m. +19°C)

203
 Rhacophoridae

Fig. LXXXIV Distribution of Rhacophorus-species in the area of the Headquarters.

Rhacophorus appendiculatus
(GÜNTHER, 1859)

Polypedates appendiculatus GÜNTHER, 1859 (1858): 79 – Type

locality: Philippine Islands
Polypedates appendiculatus INGER et al. 2000

Diagnosis
A small lowland tree frog with a robust body and
slender limbs; snout with a distinct, conical pro-
jection; outer two fingers partially webbed, others
unwebbed; wavy skin flap along arm and 4th finger
and another along tarsus; narrow flap on heel and
crenulate dermal projection on vent.

Description
Males up to 37 mm, females up to 50 mm; TL/SVL
0.45-0.57; head longer than broad; canthus rostralis
rounded; lores oblique; tympanum distinct, half the
eye diameter; vomerine teeth present; finger web-
© M. Schroth

Fig. 204 Rhacophorus appendiculatus from Mulu NP,

Sarawak.

204
 Rhacophoridae
© J.C. Murphy

© J.C. Murphy
Fig. 205 Male of Rhacophorus appendiculatus from Fig. 206 Female of Rhacophorus appendiculatus from
Danum Valley, Sabah. Danum Valley, Sabah.

bing 1(1-1.5), 2(1-1.5/1), 3(1/1), 4(1); toe webbing in flat areas of small, intermittent streams (m: 1i,
1(0-0.5), 2(0-0.5/0), 3(0/0), 4(0-0.5/0-0.5), 5(0); an 2b), 2-5 m above the water surface.
oval inner metatarsal tubercle, no outer one; skin
above with irregular low ridges; supratympanic fold Call: (+22°C) A series of soft, clicking, pulsed notes;
between eye and axilla. For the other cutaneous note duration 200-300 ms; each note increases from
appendages see diagnosis above. 1.9 to 2.9 kHz; note interval 600 ms; separated by the
Greyish brown above with obscure, highly variable, length of 2-4 notes a short (60 ms) click (1.8-2.9 kHz;
dark markings; limbs spotted, barred, or uni-
form; undersides whitish with slight yellow-
ish tinge; nuptial pads yellow; iris golden. Fig. LXXXV Sonagram and oscillograms of Rhacophorus
Tadpoles: The larvae belong to the morphotype appendiculatus (Danum region, 200 m; 8.00 p.m. +24°C)
II. Labial teeth rows formula 4-5(2-4 or 5)/3(1);
tail slender, pointed, twice body length; total
length 30 mm; head-body greyish brown with
darker crossbars; ventral sides whitish; caudal
muscle brown with light spots; dorsal fin dusky;
ventral fin dusky near its tip.

Ecology/Ethology
Rhacophorus appendiculatus lives in both pri-
mary and old secondary forests where it oc-
curs at all strata between low vegetation and
the canopy level. Its range is restricted to alti-
tudes between sea level and hilly regions up
to 500 m. It is active at night, hunting soft-
bodied insects including their larvae, and spi-
ders. Males form large calling groups on the
twigs and leaves of low vegetation 1-3 m above
the ground).
Foam nests can be found attached to the
branches of low vegetation on the edges of
rainwater-filled pools on the forest floor and

205
 Rhacophoridae
harmonics at 6.6-7.9 kHz); intensity modulation
present.
Distribution
This species is widely distributed on the Malay
Peninsula, Sumatra, the Philippine Islands, and over
all parts of Borneo. Mt. Kinabalu: Only recorded
from the Sg. Kipungit in the Poring region.
Rhacophorus baluensis INGER, 1954
Rhacophorus baluensis INGER, 1954a: 250 – Type locality:
Bundu Tuhan, 4,500 feet, Mount Kina Balu, North Borneo
Rhacophorus baluensis, INGER 1966: 298, 1978: 313; MANTHEY
1983: 23; MALKMUS 1988c: 176; INGER & STUEBING 1992:
41-51; INGER, STUEBING & TAN 1996: 363; INGER & TAN 1996a:
567,1996b: 69; LAKIM et al. 1999; INGER et al. 2000
Diagnosis
A brown, medium-sized, montane frog with a short,
pointed snout; outer fingers webbed over half their
lengths; dermal ridges along outer fingers and edge
of forearm up to elbow, and between outer toes and
Fig. 207 Rhacophorus baluensis from Sinsuron Ridge, Sabah.
© R.B. Stuebing
206
heel; a pointed dermal flap on heel, and a narrow
crenulate ridge above vent.
Description
Males up to 55 mm, females up to 65 mm; head
as long as broad; canthus rostralis sharp; lores
oblique, slightly concave; tympanum 3/5 the eye
diameter; vomerine teeth present; finger webbing
1(1), 2(1/0), 3(1/0-0.5), 4(0); toe webbing 1(0),
2(0/0), 3(0/0), 4(1/1), 5(0); an oval inner meta-
tarsal tubercle, no outer one; skin smooth or
weakly pebbled above, granular below; a feebly
curved supratympanic fold between eye and
shoulder; for dermal appendages see diagnosis
above.
Coloration highly variable, depending on ambient
temperatures, intensity of light, and endogenous fac-
tors; above sand brown to chocolate-brown, with
dark transverse bars and irregular light or dark
blotches; ridge above vent white; limbs with dark
crossbars; flanks with white spots; undersides whit-
ish, with dark spots on throat; iris golden; males with
grey nuptial pads.

Tadpoles: The larvae belong to the morphotype II.
Labial teeth rows formula 6-7(2-6 or 7)/3(1); beaks
black for less 50 %, serrated; tail elongate, taper-
ing to an obtusely pointed tip, twice as long as body;
total length up to 75 mm; head-body brown, belly
and posterior half of tail lighter.
Ecology/Ethology
This species occurs in primary as well as in dis-
turbed forests. It is restricted to montane zones with
records from between 900 and 1800 m a.s.l. The
males call at night, forming groups around poten-
tial spawning sites.
The larvae can be found in small pools formed by
accumulating rainwater on the forest floor (m: 2b),
often in the marginal zones of forests. They are
bottom suspension feeders
Call: The call is a single deep rattle consisting of
12-15 pulses at a constant rate of about 20 pulses
per second. The first 2 pulses are very soft; dura-
tion of the rattle 750-800 ms; dominant frequency
1-2.7 kHz; intensity modulation present.
Fig. LXXXVI Sonagram and oscillograms of Rhacopho-
rus baluensis (Gg. Mulu, 1300 m; 9.30 p.m. +20°C)
Rhacophoridae
Distribution
Rhacophorus baluensis is only known from a few
localities in western Sabah and north-eastern
Sarawak. Mt. Kinabalu: Bundu Tuhan (1375 m),
Marei Parei, Headquarters (1500 m), Mesilau (see
also map on page 200 and 204).
Rhacophorus cyanopunctatus
MANTHEY & STEIOF, 1998
Rhacophorus cyanopunctatus MANTHEY & STEIOF, 1998: 37 –
Type locality: “Südlicher Rand des Khao Sok National Parks
(ca. 200 m ü NN), Provinz Surat Thani, Distrikt Phanom,
Thailand”
Philautus bimaculatus, (not Rhacophorus bimaculatus PETERS,
1867) I NGER 1954a: 250, 1966: 344 (part), 1978: 313;
MANTHEY 1983: 23
Rhacophorus bimaculatus, (not Rhacophorus bimaculatus PE-
TERS, 1867) INGER, STUEBING & TAN 1996: 364
Diagnosis
A brown, small tree frog with a slender body, long
limbs and large eyes; one or more white spots below
eye, similar to R. gauni. The only species of
Rhacophorus in the Sunda region with blue spots on
flanks, on posterior and anterior faces of thighs, on
underside of tibia, and on axillae; no dermal projec-
tions; the tibio-tarsal joint reaches beyond tip of snout.
Description
Males up to 35 mm, females up to 43 mm; TL/SVL
up to 0.63; head as wide as long; snout pointed,
sloping backward to mouth in profile; canthus
rostralis sharp, curved; lores not concave; tympa-
Fig. 208 Holotype (female) of Rhacophorus cyano-
punctatus.
© U. Manthey
207
 Rhacophoridae
© U. Manthey
Fig. 209 and 210 Holotype (female) of Rhacophorus cyanopunctatus from South Thailand.
num half eye diameter; vomerine teeth absent; fin-
ger webbing 1(1), 2(1/0), 3(1.5/0), 4(0); toe web-
bing 1(0), 2(0/0), 3(0/0), 4(1/0.5), 5(0); an oval in-
ner metatarsal tubercle, no outer one; skin of dor-
sum, throat, and chest smooth, belly granular;
supratympanic fold curved, between eye and ax-
illa; a narrow dermal fringe along outer edge of
both 4th finger and the 5th toe; white tubercles on
vent; nuptial pads absent.
Light brown above, with a narrow dark inter-or-
bital bar; dark markings (often as crossbars) on
dorsum; sides of body and head below line can-
thus rostralis - eye - supratympanic fold dark brown;
blue spots on blackish brown ground (see diagno-
sis above); undersides whitish, often dusted with
brown; belly yellowish; iris golden brown. The
208
dorsal pattern is highly influenced in its intensity
by the time of the day and the prevailing mood.
The white spots below the eye and blue spots vary
very much with the individual.
Tadpoles: The larvae have not been identified with
certainty (INGER & STUEBING 1997), but presumably
belong to the morphotype I.
Ecology/Ethology
Rhacophorus cyanopunctatus occur in lowland
primary forests, up to an altitude of 600 m, but are
actually rare above 300 m. Males call at night, from
vegetation 60-250 cm above the ground near small
streams.
Larvae presumably living in forest streams with
moderate to strong currents (m: 1a, b, c).

Fig. LXXXVII Sonagram and oscillogram of Rhacopho-
rus cyanopunctatus (Gunung Lawit, 250 m; 9.00 p.m.)
Call: (+21°C) The call is composed of 2 short,
pulsed rattles, separated by an interval of about 1
second; the first signal (duration 80 ms) consists
of 3-4 pulses, the second (duration 110 ms) of 2-3
pulses; dominant frequency 2.3-4.8 kHz.
Distribution
This species is known from the Malay Peninsula,
and from northern Sumatra and Borneo. Mt. Kina-
balu: along Sg. Mamut, Sg. Kipungit (500-600 m).
Fig. 211 A pair of Rhacophorus dulitensis in amplexus from Deramakot, Sabah.
Rhacophoridae
The record from “Lumu Lumu (1600 m)” by INGER
(1966, 1978) is probably based on a misidentifica-
tion.
Rhacophorus dulitensis
BOULENGER, 1892
Rhacophorus dulitensis BOULENGER, 1892: 507 – Type locality:
Mt. Dulit, Borneo
Rhacophorus dulitensis, WONG 1994: 29-37; I NGER et al. 2000
Diagnosis
A small tree frog with a sharply pointed snout, a
slender body, and slim limbs; fingers almost fully
webbed; a white dermal ridge along outer edges of
arm and tarsus, respectively; small conical or round
skin flap on heel; transverse, two-lobed skin fold
above anus; no supratympanic fold.
Description
Males up to 36 mm, females up to 50 mm; TL/SVL
0.47-0.49; head longer than wide; canthus rostralis
sharp; lores vertical, not concave; tympanum 2/3
eye diameter; vomerine teeth present; finger
© T. Eltz
209
 Rhacophoridae
© W. Hödl
Fig. 212 Rhacophorus dulitensis from Danum Valley
with its foam nest.
webbing 1(1), 2(1/0), 3(0/0), 4(0); toes fully
webbed up to disks; an oval inner metatarsal tu-
bercle, no outer one; skin above smooth, granular
below; for cutaneous appendages see diagnosis
above.
Bright jade-green above with a brownish tinge
down middle of the back, and tiny white spots; a
yellowish canthal stripe; upper eyelids and snout
brown; undersides whitish green; webbing pale
green, some specimens with reddish brown spots
on webbing between outer two toes; nuptial pads
yellowish. The bones of the limbs are green and
visible through the tissue from below.
Tadpoles: The larvae belonging to the morphotype
II; labial teeth rows formula 6(2-6)/3(1); beaks
blacks in their marginal thirds, finely serrated; tail
a little less than twice body length; total length up
to 50 mm; head-body and anterior half of tail grey-
ish brown; undersides whitish; a wavy, vertical,
black line on mid-line of tail.
210
Ecology/Ethology
Nearly nothing is known about the life history of
this species, other than it lives in lowland primary
forests, and that males form large groups around
forest pools.
Spawning sites are on the forest floor (m: 2b, c).
The tadpoles are large bottom suspension feeders
Call: The call is a short, weak trill, similar to the
song of a cricket. A call analysis is not described.
Distribution
This frog is known from Sumatra and Borneo. Mt.
Kinabalu: There is only a single record from Sg.
Kipungit, see also map on page 200 and 203.
Rhacophorus everetti macroscelis
BOULENGER, 1896
Rhacophorus macroscelis BOULENGER, 1896a: 403 – Type
locality: Mount Kina Balu, N. Borneo
Rhacophorus macroscelis, VAN KAMPEN 1923: 252; SMITH 1931: 31
Philautus spiculatus SMITH, 1931: 20 – Type locality: Kenokok,
Mount Kina Balu, North Borneo (syn. after INGER 1966)
Rhacophorus spiculatus, INGER 1954a: 251
Rhacophorus everetti macroscelis, INGER 1966: 298, 1978: 313;
MANTHEY 1983: 23; MALKMUS 1985: 8, 1986: 3, 1987: 282,
1988c: 176, 1989: 192, 1991a: 28, 1992a: 118, 1994b: 228,
1994c: 87; HOFFMANN 1995b: 15; MALKMUS 1996c: 286
Rhacophorus everetti, INGER & STUEBING 1992: 41-51; INGER,
STUEBING & TAN 1996: 364; INGER & TAN 1996a: 567, 1996b:
69; LAKIM et al. 1999; INGER et al. 2000
Taxonomic remarks: Based upon morphological
differences, INGER (1966) considered the Bornean
specimens a separate subspecies, R. e. macroscelis,
thus restricting the nominate subspecies to the Phil-
ippine island of Palawan. The present subspecies
has longer tubercles, a wider inter-orbital marking,
and a slightly smaller tympanum. In contrast to R.
e. everetti, it possesses tubercles on the dorsal sur-
faces of the head and body.
Diagnosis
A small to medium sized, stocky, montane tree frog
with slender, long legs and a short blunt snout; fin-
gers with rudiments of webbings confined to bases;
many pointed tubercles on head and neck; cone-
 Rhacophoridae

© M. Schroth
Above: Fig. 213 Rhacophorus everetti macroscelis from
Sg. Kemantis, Sayap, 900 m.

shaped dermal projections along outer edge of fore-

arm and leg, respectively; females with larger,
spiculate dermal appendages.

Description

© R. Malkmus
Males up to 32 mm, females up to 55 mm; TL/SVL
0.44-0.68; head wider than long; canthus rostralis
sharp, curved; lores concave; tympanum half eye
diameter; vomerine teeth present; webbing of toes
1(1), 2(1/0.5), 3(1.5/0.5), 4(1-1.5/1-1.5), 5(0); a small
inner metatarsal tubercle, no outer one; skin of the
back shagreened with white-pointed tubercles on
forehead and eyelids, forearms, and on tarsi; flanks
and ventral sides coarsely granular; some white tu-
bercles below vent; nuptial pads yellowish.
The coloration is significantly influenced by ex-
citement and exogenous factors and as a conse-
© M. Schroth

Right from top to base: Rhacophorus e. macroscelis

Fig. 214 Female from Sg. Liwago, 1450 m.
Fig. 215 Juvenile from Sg. Silau-Silau, 1500 m.

211
 Rhacophoridae
© R. Malkmus
Fig. 216 Fermale of Rhacophorus everetti macroscelis
from Sg. Liwago, 1450 m.
quence is highly variable. Relatively constant traits
are an inter-orbital bar, a large W-shaped to trap-
ezoid marking on the occiput, and dark crossbars
on the hind limbs. Upper side of body spotted in a
variety of shades of brown, lighter laterally; groin
region and inner sides of thighs bright yellow with
mocha-brown spots; lower sides of body lemon-
yellow, occasionally peppered with brown dots;
insides of mouth and tongue green. Exposed to
daylight some of the frogs change colour to a whit-
ish green with a metallic bronze sheen. Iris whitish
yellow with a fine brown network. Females green
above, with a large, W-shaped, black dorsal mark-
ing; flanks light green with black spots and no yel-
low inguinal region; lower sides yellow.
The surface structure and the coloration award per-
fect camouflage to the frogs, especially when
perched on a surface which is overgrown by lichens
(lichen mimesis).
Tadpoles presumably stream-dwelling and of
morphotype I.
Ecology/Ethology
This frog is an inhabitant of montane primary for-
ests at elevations between 900 and 1840 m. It is
nocturnal. The males call in colonies established
in the vicinity of small streams perched on
branches and leaves of bushes and shrubs 150 to
600 cm above the ground. Their call activity starts
often before the onset of dusk (from 5.30 p.m.) if
it rains. Specimens kept in a terrarium always re-
turned from their nightly excursions to their rest-
212
ing-place. Their diet includes insects (moths, flies,
small grasshoppers) and their larvae, as well as
spiders. Parts of a prey item that may protrude
laterally from the mouth (e.g. the legs of Opiliones)
are taken with the hands and straightened out for-
ward. We found freshly metamorphosed juveniles
on Sg. Silau-Silau during the months of March
and August.
Call: The note series consists of a rasping sound
which is repeated 3-10 times at continually increas-
ing volume. This series is occasionally followed
by a sharp “di di di di dit”. A call analysis does not
exist.
Distribution
This subspecies is only known from a few locali-
ties in western Sabah, north-eastern Sarawak, and
Brunei. Mt. Kinabalu: Kenokok, Marei Parei,
Kiau, Bundu Tuhan, Sayap: Sg. Kemantis (1000 m),
common in the region between Sg. Silau-Silau and
Liwago including the tributaries (1400-1650 m, Sg.
Tibabar up to 1800 m), Sg. Mesilau (up to 1840 m),
see also maps on page 200 and 204.
Rhacophorus gauni (INGER, 1966)
Philautus gauni INGER , 1966: 346 – Type locality: Mengiong
River, Kapit District, Sarawak
Rhacophorus gauni, INGER 1985: 81; INGER & TAN 1990: 6;
INGER & STUEBING 1992: 41-51; WONG 1994: 29-37; INGER,
STUEBING & TAN 1996: 364; MALKMUS 1996c: 287; LAKIM et
al. 1999
Diagnosis
A small, brown, slender lowland species with a very
short snout; conical projections on heel and on
upper eyelids, respectively; outer finger almost fully
webbed; a white spot below large eyes; tibio-tarsal
joint not reaching beyond eye; nuptial pads absent.
Description
Males up to 32, females up to 38 mm; TL/SVL
0.51-0.58; head equal in length and width; snout
broadly rounded; canthus rostralis curved, lores
sloping; tympanum 1/3 eye diameter; vomerine
teeth absent; finger webbing 1(1-1.5), 2(1-1.5/0),
3(01/0.5), 4(0-0.5); toe webbing 1(0), 2(0/0), 3(0/0),
 Rhacophoridae

© R. Malkmus
Fig. 217 Rhacophorus gauni from Sg. Langanan, 700 m.

4(1/1), 5(0); an oval inner metatarsal tubercle, no

outer one; skin smooth above; a pointed, conical
tubercle on upper eyelid and another one on heel;
3-4 small projections between heel and base of
outer toe; curved, weakly expressed supratympanic
fold; throat smooth, chest and belly coarsely granu-
lar.
Upper sides of the body brown with large dark spots
and transverse bands; crossbars on arms and legs;
groin and rear of thighs yellow to reddish orange;
ventral sides white; iris golden brown, lighter on
margins.
Tadpoles: The larvae belong to the morphotype I.
Labial teeth rows formula 4(3-4)/3(1); beaks V-
shaped, heavy, black, coarsely serrated; tail con-
vex, tapering gradually to a pointed tip; total length
25 mm; head-body portion yellowish brown
dorsally and laterally, unpigmented ventrally; cau-
© H. Sauer

Fig. 218 Ventral pattern of Rhacophorus gauni from Sg.

Kipungit, 600 m.

213
Rhacophoridae

dal muscle and dorsal fin with large dark spots;

ventral fin pigmentless.
Indications are that the tadpoles of this species have
developed a bimodal feeding strategy as there are
non-feeding as well as exotrophic larvae (INGER
1992a).

Ecology/Ethology
Rhacophorus gauni is restricted to primary for-
ests in lowland and hilly regions (150-950 m

a.s.l.). At night, males are found on vegetation
(30-800 cm above the ground) overhanging rif-
fles of rocky streams, forming loose calling
groups. We found specimens of this species in
the midst of a calling community of Philautus
aurantium in a bamboo-rattan mixed forest, some
300 m away from the nearest stream in the
Langanan valley. The small foam nest is attached
to leaves of vegetation overhanging forest
streams. The larvae live among gravel and rocks
(m: 1a, b, d, e).
Call: A single high-pitched chirp. A call analysis
is not yet described.
Distribution
Rhacophorus gauni is only known from a few
localities in Sarawak and Sabah. Mt. Kinabalu:
Sg. Kinateki, Poring region: Sg. Mantukungan,
Sg. Kipungit I and II (500-640 m), Sg. Langanan
(750-950 m), see also maps on page 200 and 203.
© J.C. Murphy
Fig. 219 Rhacophorus harrissoni from Sabah.
Description
Males up to 56 mm, females up to 70 mm; TL/SVL
0.52-0.58; head longer than wide; canthus rostralis
sharp, lores almost vertical, weakly concave; tym-
panum distinct, larger than half eye diameter;
vomerine teeth present; all toes webbed up to disks;
dermal flap along outer edge of 1st toe; a small,
oval inner metatarsal tubercle, no outer one; dorsal
skin smooth and finely pebbled; supratympanic fold
straight, from eye to tympanum and shoulder; un-
dersides granular.
Back and upper side of head reddish brown; dark
crossbars on limbs; side of head sometimes with
white spots; ventrum white, with dark spots; nup-
tial pads yellowish; iris light brown in upper half,
dark reddish brown in lower.

Rhacophorus harrissoni Fig. 220 Rhacophorus harrissoni from Marak Parak, Sabah.
INGER & HAILE, 1959

Rhacophorus harrissoni INGER & HAILE, 1959: 270 – Type

locality: “Ulu Patah, a tributary of the Baram River, Fourth
Division, Sarawak”
Rhacophorus harrissoni, INGER & STUEBING 1992: 41-51; INGER,
STUEBING & TAN 1996: 364; INGER et al. 2000

Diagnosis
A moderately large, brown tree frog with a trian-
© R.B. Stuebing

gular head and an obtusely, pointed, sharp-edged

snout; three outer fingers fully webbed; webbing
between fingers and toes brown or black; no der-
mal projections on limbs.

214

R. Malkmus
Fig. LXXXVIII Oral disc of Rhacophorus harrissoni.
Tadpoles: The larvae belong to the morphotype II.
Labial teeth rows formula 4-5(2-4 or 5)/3; beak
finely serrated; tail slightly convex, tapering near
end to narrow rounded tip; tail about twice body
length; total length up to 41 mm; head-body por-
tion and tail dark grey.
Ecology/Ethology
Rhacophorus harrissoni lives in primary and old
secondary forests covering hilly terrains at low el-
evations. Very little is known about its life history.
Specimens were found in tree holes, among epi-
phytes, or on the leaves of vegetation, 30-900 cm
above the ground.
Foam nests are attached to the bark above water-
filled tree holes, tree trunks, or water tanks formed
by anastomosing buttresses, 1-4 m above the ground
(m: 2e, f, g).
Distribution
This frog is widely distributed in all parts of Borneo.
Mt. Kinabalu: Known only from the Poring region
(Sg. Kipungit), see also maps on page 200 and 203.
Rhacophorus nigropalmatus
BOULENGER, 1895
Rhacophorus nigropalmatus BOULENGER, 1895: 170 – Type
locality: “Akar River”
Diagnosis
A green tree frog, very large by comparison, with
long, slender limbs and large hands and feet; all
Rhacophoridae
fingers and toes fully webbed; webbings black,
yellow towards tips of toes; broad skin flaps on
forearm, tarsus, and heel.
Description
Males up to 90 mm, females up to 100 mm; TL/SVL
0.54-0.58; head as wide as long or slightly broader;
canthus rostralis distinct, lores sloping, a little con-
cave; tympanum half eye diameter; inner edge of
1st finger and toe, and outer edges of 4th finger and
5th toe, with flaps of skin; an oval inner, no outer
metatarsal tubercle; skin above finely pebbled,
granular below, except on the throat; no supra-
tympanic fold.
Emerald green above, often with tiny white spots;
flanks yellow at insertions of arm and leg; under-
sides whitish, with a yellowish tinge on chest and
belly; iris pale golden; nuptial pads yellowish.
Tadpoles: The larvae belong to the morphotype II.
Labial teeth rows formula 4-6(1-4, 5, or 6)/2(1) or
3-7(2-3, 4, 5, 6, or 7)/3(1) or 6(3-6)/3(1); beaks
finely serrated, black in their marginal thirds; tail
length nearly twice body length; head-body pale
grey above, white below; tail grey with dark spots.
Ecology/Ethology
Rhacophorus nigropalmatus is a poorly known low-
land species occurring from sea level up to 500 m
altitude. It dwells the canopy of primary forests
leaving the trees only for breeding. WALLACE (1869)
Fig. 221 Rhacophorus nigropalmatus from Sabah.
© J.C. Murphy
215
 Rhacophoridae

Rhacophorus pardalis GÜNTHER, 1859

Rhacophorus pardalis GÜNTHER , 1859 (1858): 83 – Type

locality: Philippines, Borneo, and East Indies
Rhacophorus pardalis, INGER & STUEBING 1992: 41-51; WONG
1994: 29-37; HOFFMANN 1995b: 15; INGER et al. 2000

Diagnosis
A small to medium-sized, slender to robust, brown
tree frog with long legs; the outer three fingers are
fully webbed; webbings on hands and feet orange-
red; an orange network on the belly.

Description
Males up to 55 mm, females up to 75 mm; TL/SVL
0.47-0.49; head longer than wide, snout rounded;
canthus rostralis distinct; lores sloping, weakly
concave; tympanum half eye diameter; vomerine
teeth present; finger webbing 1(0.5-1), 2(0.5-1/0,
3(0/0), 4(0); toes fully webbed; a small, oval inner

Fig. 223 Rhacophorus pardalis from Poring, 550 m.

© M. Pfeiffer

Fig. 222 Rhacophorus nigropalmatus from Sg. Kipun-

git I, 550 m, Poring.

and HANITSCH (1900b) were the first to describe the

gliding flight of this frog, and INGER (1966) ob-
served: “One frog was thrown into the air from a
height of five meters above ground. It immediately
righted itself and held the hands and feet in the
plane of the body with the digits widely spread”;
see also EMERSON & KOEHL (1990).
The large foam nests are attached to low vegeta-
tion or deposited on the muddy banks of small pools
and wallows of rhinos and wild boars on the forest
floor (m: 2b, c). The larvae often develop in ex-
tremely murky, turbid water (INGER & STUEBING
1997) and are bottom suspension feeders.

Distribution
© P. Hoffmann

Malay Peninsula, Sumatra, and Borneo. Mt. Kina-

balu: There is only one record from Sg. Kipungit I
(500 m).

216
 Rhacophoridae

metatarsal tubercle, no outer

one; dorsal skin shagreened,
coarsely granular below
(smooth on throat of females);
dermal flaps along outer edges
of forearm and hand and on
tarsus, round flap on heel;
supratympanic fold low,
curved.
Coloration very variable; up-
per sides tan to reddish brown
with dark markings, often ir-
regular white, blue, red, or yel-
low spots on back or legs;
limbs with dark crossbars;
flanks yellow with black
flecks; chest and belly pale
yellow with an orange net-
work; throat whitish; iris light
golden.
Tadpoles: The larvae belong
to the morphotype II. Labial
teeth rows formula 6(3-6)/3 or
7(3-7)/3; tail tapering to a nar-
row tip; tail length slightly less
than twice body length; total
length up to 45 mm; head-
body and tail grey to light
brown, sometimes with sev-
eral dark spots; often with
only one spot on side of head.

© J.C. Murphy
Ecology/Ethology
Rhacophorus pardalis appears
to be a resident of the canopy Fig. 224 Rhacophorus pardalis from Sabah.
stratum. It can be found in pri-
mary and secondary forests
and is rather common along logging roads. It is Call: A short, raspy chuckle. A call analysis is not
able to perform paragliding flights by spreading described.
its webbed digits to break the fall. At night, the
males form calling groups on low vegetation (1-3 m Distribution
above the ground) around spawning sites. Foam nests R. pardalis occurs in Peninsular Malaysia, on
can be found on herbs and overhanging vegetation Sumatra, Borneo, and the Philippines. Mt. Kina-
at the edges of small streams with reduced gradi- balu: Locality records exist only from the Poring
ents (m: 1d, f, g, i) or rainwater-filled pools on the region (Sg. Kipungit), see also map on page 200
forest floor (m: 2b, c). and 203. The record of this lowland species from
R. pardalis is a lowland species occurring between Sg. Silau-Silau (1500 m) by WONG (1994) is prob-
100 and 500 m a.s.l. ably based on a confusion with another species.

217
Ichthyophiidae

Gymnophiona
Caecilians are pantropical, elongate, limbless amphibians of a worm-like appearance. They lack sternal
and pelvic girdles, and the columella is either massive or absent. The body is segmented by annular
grooves with the number of annuli corresponding to that of the vertebrae (up to 300). In the more primi-
tive genera the lower layers of the skin encase ossified scales. The tail, if present, is very short and
pointed.
Caecilians are adapted to drill-burrowing with their heavily ossified skulls. For the same reason the eyes
are rudimentary and obscure, although they still possess a lens and a retina with photoreceptors which
can distinguish between light and dark. Between the eye and the nostril there is a protrusible sensory
tentacle. The teeth are curved, present on premaxillae, maxillopalatines, vomers, dentaries, and splenials.
In contrast to all other vertebrates which possess only a single pair of jaw muscles, caecilians have
another, second pair. Lungs are present (in except of Atretochoana) with the left lung usually being
rudimentary.
Caecilians reproduce by internal fertilization for which the males are equipped with a single, median,
protrusible penis (phallodeum). Females either lay a clutch of eggs or give birth to live youngs. In the
case of egg-laying species, the female guards her clutch until it hatches. The larvae are amphibic with
gill slits (spiracle) and labial lobes. They live in shallow water and feed at night. Caecilians are car-
nivorous.
While numerous works exist on systematic-taxonomic and anatomic questions, only very little is known
on the natural history of the secretive gymnophionians which are also only rarely kept in captivity.
Borneo is home to one family, the Ichthyophiidae.

Ichthyophiidae
This family is widespread throughout Southeast Asia ranging from India and Sri Lanka to Borneo and
the Philippines.
It contains medium-sized caecilians of up to 500 mm in length with pointed tails and subterminal mouths.
The tentacular openings are situated mid-way between the eyes and the nostrils. Their primary annuli are
subdivided into secondary annuli (2-4 secondaries per primary), both of which form chevrons pointing
toward the tail on the ventral side. The dermis includes small ossified scales.
The eggs produced by the species of this family are very rich in yolk. They are deposited in a string in moist
soil and guarded and defended by the female. Eggs attacked by fungal infections or which have perished
otherwise are consumed by the female. The period of development from oviposition to the emergence of the
larvae takes 70-80 days. The larvae lead an amphibic life and are equipped with well-developed lateral line
organs (neuromasts) and ampullary organs. While the former serve as mechanoreceptors, sensitive to water
currents and pressures, the latter are electroreceptors. The larvae have one or two gill slits, and low caudal fins.
The tentacles do not appear until shortly before the completion of the larval development, that is after 12-14
months.
Adults live in primary and old secondary forests in earth holes, under leaf litter, or beneath logs. Their food
comprises arthropods, earth worms, and small vertebrates. The life histories of all species are nearly unknown.
This family is represented on Borneo by 2 genera (Caudacaecilia and Ichthyophis) and 6 species, on Mt.
Kinabalu by one genus (Ichthyophis) and one species.
These two genera can hardly be distinguished on the basis of their external morphologies - Cauda-
caecilia has one row of teeth on the lower jaw, Ichthyophis two.

218

gill slit
Fig. LXXXIX Larva of Ichthyophis.
ampullary organs
Genus Ichthyophis FITZINGER, 1826
The species of this genus are almost as widespread
as those of the entire family. They exhibit all the
above mentioned traits typical for the family.
Four species are known from Borneo, one from Mt.
Kinabalu.
Ichthyophis monochrous BLEEKER, 1859
Ichthyophis monochrous BLEEKER, 1859 (1858): 188 – Type
locality: Singkawang, Borneo
Ichthyophis monochrous, INGER 1962/63: 46, 1966: 355
Diagnosis
Purplish black with tiny, diffuse, yellow spots on
the throat; tentacles at upper lip, much closer to
the eye than to the nostril.
Tab. 21 Distribution of Ichthyophis-species on Borneo.
Ichthyophiidae
tail fin
neuromasts ampullary organs
Fig. XC Lateral-line organs in Ichthyophis.
Description
Total length up to 280 mm; head as wide as body;
snout rounded; small eyes visible through the skin;
body with 247 annuli, tail with 4-6 annuli; 18-25
maxillary-premaxillary teeth, 19-21 vomero-
palatal teeth, 19-23 mandibular teeth, 2-4 sple-
nial teeth.
Eggs: unknown.
Larvae: head depressed; caudal fin distinct; total
length up to 192 mm.
Ecology/Ethology
A forest species ranging from sea level up to 600 m
altitude, burrowing beneath the duff of logs.
The larvae live in small, clear forest streams.
Distribution
Widespread on Borneo. Mt. Kinabalu: Mamut area/
Ranau District; Sg. Kipungit.
219
 Reptilia (reptiles)
Reptiles are marked by a skin covered with scales and usually have a creeping
mode of locomotion. The conservative systematic considers the class Reptilia a
group of representatives of the phylum Vertebrata or vertebrates. These are animals
breathing with lungs whose body temperatures largely depend on the temperature
of their immediate environments. All of today’s reptiles evolved from ancestors
which originally possessed two pairs of limbs, ending in clawed fingers and
toes, and a tail. The skin incorporates but a small number of glands and is usually
covered with a protective layer of horny scales. The number, arrangement, and
individual shapes of the latter are, within certain limits of natural variability,
indicative of species and genera. Reptiles reproduce under the principles of internal
fertilization, but there are a few species known which multiply parthenoge-
netically, i.e. without the necessity of a mating of two different sexes (e.g. within
the family Gekkonidae). Some species give birth to fully developed young, but
the vast majority of reptiles lay relatively large eggs with hard and calcified or
parch-like shells which contain substantial amounts of yolk. As a rule, these are
deposited on land. The clutches are usually neither guarded nor actively incubated
by the parents. The young hatch after some time and are immediately self-
supporting.
There are in excess of 6000 species of reptiles, systematically grouped in 4 orders.
Borneo is inhabited by reptiles of three orders, namely Crocodylia (crocodiles),
Squamata (scaled reptiles), and Testudines (turtles, tortoises, and terrapins).
Crocodiles are easily identified as they are the only reptiles whose teeth are
visible when the mouth is closed. Equally easy to recognize are turtles and tortoises
whose bodies are protected by a shell. The latter may be domed or low, and hard
or soft to the touch. All the remaining reptiles of Borneo belong to the order
Squamata and comprise the lizards and the snakes.

Squamata (scaly reptiles)

In contrast to crocodiles, squamates all have a transversally arranged anal slit,
and no teeth are visible when the mouth is closed. Males possess a pair of
copulation organs (penes) of which but one is employed for the actual mating.
The skin is covered by a protective layer of horny scales which does not grow
with the animal and has to be shed from time to time. The underlying new scale
cover then is slightly larger than the previous one. The hypodermis (lower layer
of skin) contains pigment cells which determine the colouration and colour pattern
of a reptile depending on the type and arrangement of these cells.
This order comprises two suborders, named Sauria (lizards) and Serpentes
(snakes), respectively. The latter have evolved from the lizards and therefore
represent a much younger stage of evolution. Lizards are usually, but not generally,
characterized by four limbs and movable eyelids. Snakes, on the other hand, do
neither have limbs, nor movable eyelids, nor ear openings. Certain fossorial groups
bring about complex problems as to their proper allocation to one group or another.
The members of the snake family Typhlopidae (blind snakes) for example exhibit
skull structures which are very similar to those of lizards, while the lizard family
Dibamidae (blind skinks), whose members do not possess limbs (males with
rudiments) or tympani, share many traits with the snakes.
 ey
th
an
.M
U
©
Sauria (lizards)
Lizards have conquered almost all types of habitats. Some live
high up on trees and have developed the ability to perform a
controlled gliding flight, others dwell on or even in the ground.
Several of them are excellent swimmers, and a few do not
even avoid the ocean. Lizards feed mainly on live insects, but
some species also prey upon molluscs, worms, small
vertebrates, crustaceans, and carrion, or have specialized on a Agamidae
vegetarian diet.
Our knowledge about how lizards live in the wild is still
relatively limited. Observations are usually superficial and
restricted to brief moments before a lizard takes to flight and
Gekkonidae

©
M
disappears into the layer of leaf litter (like many skinks for

.S
ch
example), or dashes up a tree trunk with tremendous speed

ro
th
and thus withdraws from the curious eyes of an observer. Other
lizards rely on the camouflage effects of their colours and colour
patterns which blend in with their surroundings and may not
give themselves away by movements for hours on end. To locate
them in the semishade of a forest is extremely difficult and
very much dependent on incidental discovery. Very few lizards
are true to a locality for longer periods of time so that a purpose-
driven search is rendered extremely difficult if not impossible.
Some 110 species of lizards are represented on Borneo which
are systematically grouped in eight families and thirty-seven
genera. Five of these families and twenty-five genera have
been recorded from Mt. Kinabalu (see Fig. XII). Using the

©
S.
identification key (page 223), they can easily be allocated to

M
an
th
their proper families. Considerably more difficult, however,

ey
is the correct allocation of representatives of the families
Scincidae, where generic traits are not always discernible
externally, and Gekkonidae, where indicative morphological
traits can sometimes only be evaluated with the aid of at least
a magnifying glass. This is even more true when it comes to
the identification of the individual species. The latter often
requires the use of a binocular microscope which facilitates Scincidae
the counting of scales etc., but a layman may still be faced
with insurmountable problems.
As far as the agamids are concerned, the generic identification
is somewhat easier because it can be based on relatively well-
discernible external traits. When it comes to species level,
©
U
.M

however, another kind of complication may make itself felt as Varanidae

an
th
ey

females often exhibit an appearance which is entirely dissimilar

to that of the males of the same species.
As the pigment cells embedded in the hypodermis can alter
the colouration and the distribution of colours to a sometimes
substantial extent by changes in cell size and arrangement,
colour and pattern traits have only a limited suitability for
identification purposes.

Background: typical stream on Mt. Kinabalu © A. Nöllert

Sauria

Distinguishing features in lizards

rostral
supranasal mental
(internasal)
frontonasal postmental
prefrontal
frontal submaxillaries
supraoculars

frontoparietal
parietal gulars
interparietal
nuchal

Fig. XCI Head scales of a skink

temporals
supraciliaries

preoculars
postnasal loreal

nasal

rostral

Fig. XCII Toe mental

infralabials

supralabials
subdigitals tympanum

tail length (TL) snout-vent length (SVL) femoral pores

preanal pores

Fig. XCIII Place of measurements in lizards Fig. XCIV Anal region of a gecko

222
 Sauria

Fig. XCV Rhombic imbricate Fig. XCVI Inhomogeneous Fig. XCVII Granular scales of a
scales of an agamid lizard, scales of an agamid lizard, gecko, intermixed with larger
homogeneous in shape, size, intermixed with larger ones. roundish tubercles.
and orientation.
supraciliary edge nuchal crest
dorsal crest
canthus rostralis

Fig. IC Some external

morphological charac- fold in front of
ters of agamid lizards. the shoulder

Fig. XCVIII Keeled scales gular sac

of an agamid lizard.

Key to the families of Bornean lizards (after MANTHEY & GROSSMANN 1997)
1 Eyes visible ....................................................... 2
• Eyes hidden beneath scales (A) ........ Dibamidae A B C
2 Head scales large, symmetrical (B) .................. 3
• Head scales small, usually asymmetrical (C) .... 5
3 Lateral fold present (D), limbless ..... Anguidae
• Lateral fold absent, limbs present or absent .... 4
4 Scales shiny, tail length < 3 × SVL ... Scincidae D E F
• Scales dull, tail length > 4,5 × SVL (E) ..............
........................................................... Lacertidae
5 Body and tail with distinct rows of large tubercles
(F) ................................................ Lanthanotidae
• Body and tail without rows of tubercles .......... 6 G
H I
6 Body compressed laterally, gular sac and nuchal/
dorsal crests usually present (G) ...... Agamidae
• Body compressed dorsoventrally, no gular sac or
crests .................................................................. 7
7 Pupil vertically oval (H); if the pupil is round,
fingers and toes are filigrane (I); adult specimens J K
< 40 cm in total length ................... Gekkonidae
• Pupil round, fingers and toes sturdy (J); snout and
neck conspicuously long (K); adult specimens >
100 cm in total length ....................... Varanidae

223
Agamidae

Agamidae
The distribution of agamid lizards is confined to the Old World. They are represented in south-eastern
Europe, on most of the African continent, in southern and central Asia as far north as Japan, the entire Indo-
Australian Archipelago, and in Australia.
On Borneo and the surrounding region, agamid lizards are characterized by laterally compressed bodies with
four well-developed limbs, each with five clawed fingers and toes. Their dull scales do not form a symmetrical
or clearly differentiated arrangement on the head. The pupils are always round. The majority of agamids
possess a vertebral crest and/or a gular sac. In some species, females differ little from males, in others the sexes
are so dissimilar in their appearances that it is difficult to believe they belong to the same species.
These lizards are almost exclusively diurnal and live more or less strictly on trees and shrubs. Many species
are able to alter their coloration and colour pattern substantially. Agamids feed on a large variety of insects,
but may occasionally predate on small bird, fish, or rodent.
Almost all agamids are oviparous. Some time after successful mating the female descends to the ground in
order to dig a hole for egg deposition. Afterwards the hole is refilled with soil, which is solidified by forceful
head butts. Due to the absorption of moisture the egg size increases steadily until the young will hatch.
On Mt. Kinabalu the vertical distribution of agamid lizards extends to an altitude of approx. 2100 m a.s.l.
(Phoxophrys cephalum).
This family is represented on Borneo by nine genera and 27 species, with seven genera (two of which are
monotypic and endemic) and 14 species occurring on Mt. Kinabalu.

Key to the Bornean genera of Agamidae (in part after MANTHEY & GROSSMANN 1967)

A C
1 Sides of body with large, spreadable skin mem-
branes and neck lappets (A) .................... Draco
• Sides of body without spreadable skin membranes
........................................................................... 2
2 Sides of head covered with extremely large shields
below the tympanum (B) .............. Hypsicalotes B
• Sides of head without large shields .................. 3 D
3 Tympanum clearly visible (C) .......................... 4 E
• Tympanum invisible .......................................... 8
4 Males with a horn-like protrusion (D), females
with a small hump (E) on the tip of the snout ....
....................................................... Harpesaurus F G
• Both sexes without horn-like protrusions or humps
on the tip of the snout ........................................ 5
5 Lateral skin folds on the neck present (F), homo-
geneous rhombic dorsals (G), TL/SVL > 3; eggs
spindle-shaped (H) ........................ Bronchocela
• No lateral skin folds on the neck, TL/SVL < 3; H I
eggs oval (I) ....................................................... 6

224
 Agamidae

6 Ventrals > dorsals; dorsals usually intermixed with J

larger, tubercular or rhombic scales (J); if scala-
tion homogeneous, dorsals neither rhombic nor
keeled ......................................... Gonocephalus
• Ventrals @ dorsals .............................................. 7
7 Gular sac of males with lateral pockets (K), dorsals K
of various shapes, sizes, and orientations, rhombic,
tips mainly rounded ....................... Complicitus
• Gular sac of males without lateral pockets, dorsals L
vary slightly in size, rhombic, tips truncate ........
..................................................... Pseudocalotes
8 A small protrusion usually present (L), TL/SVL >
1,9, base of the tail rotund in males ... Aphaniotis
• No protrusion, base of the tail in males flattened
above, TL/SVL < 1,9; if TL/SVL > 1,8, supraci-
liary edge and nape with spines (M) ...................
M
......................................................... Phoxophrys

small and heterogeneous, ventrals always keeled

Genus Aphaniotis PETERS, 1864
Species of the genus Aphaniotis are distributed on
the Malay Peninsula, Sumatra, Borneo, and on
many small offshore islands.
Small agamids (SVL up to 70 mm max.), with slen-
der bodies and long, delicate limbs; enlarged,
hump-like scales on the tip of the snout or a scaly
protrusion in two of the three species of Aphaniotis.
Lining of the mouth blue. Tympanum hidden, some-
times indicated by enlarged scales. Dorsal scales
and larger than dorsals. Nuchal and dorsal crest
absent or weakly developed.
These tree- and bush-dwellers inhabit primary
lowland forests and hilly regions in the general
vicinity of running water. They feed on small in-
sects.
All three species of this genus occur on Borneo,
although presence of Aphaniotis acutirostris is sub-
ject to some debate. Mount Kinabalu is home to
only one species, A. ornata, which has been re-
corded up to elevations of 900 m a.s.l.

Key to the Bornean species of Aphaniotis

1 A distinct protrusion on the snout ........... ornata 2 Flanks with transverse bands, iris of males brown
• No or only weakly developed protrusion on the .......................................................... acutirostris
snout .................................................................. 2 • Flanks without transverse bands, iris of males blue
or green ....................................................... fusca

Tab. 22 Distribution of Aphaniotis-species on Borneo.

225
 Agamidae

Aphaniotis ornata Diagnosis

(LIDTH DE JEUDE, 1893)
Japalura ornata LIDTH DE JEUDE, 1893: 251 – Type locality:
“in the neighbourhood of the Sandakan-Bay (N. Borneo)”
Japalura ornata, BOULENGER 1894d: 723; DE ROOIJ 1915: 92,
fig. 47; SMITH 1931: 22, pl. II: fig. 2; MANTHEY 1983: 26;
EBENHARD & SJÖGREN 1984: 24
Aphaniotis ornata, MALKMUS 1994b: 245; TAN 1993: 1
Japalura nasuta DE J ONG , 1930 (syn. after M ANTHEY &
GROSSMANN 1997 and OTA & HIKIDA, 2000)
Japalura nasuta, MANTHEY 1983: 26; MALKMUS 1988c: 174
Aphaniotis nasuta, MALKMUS 1994b: 245
Smallest species of the genus Aphaniotis (SVL less
than 60 mm). Inconspicuously brown; both sexes
with a distinct protrusion.
Description
SVL males 54-55, females 52-57 mm; TL males
110-115, females about 102 mm; tip of the snout
with a flexible, often backward-pointing protrusion;
small gular sac present; low nuchal crest consist-
ing of triangular scales; dorsal crest separated from
the former, consisting of small, serrated scales,

Fig. 230 Male of Aphaniotis ornata from Kutai, Kalimantan.

© D. Knowles

226
 Agamidae

paralleled by two rows of tubercular

scales; conical laterals smaller than
dorsals, intermixed with larger
scales.
Upper side uniformly brownish, head
and body sometimes with a faint ol-
ive tinge; occasionally with a light,
indistinct vertebral stripe.
Females with dark radial pattern
around the eye and with two light
bands between the eyes; rostral pro-
trusion slightly conical as opposed
to flattened in males; males with a
small nuchal sail and distinctly
broaden tail base.
Two eggs (7 × 15 mm) per clutch.

Ecology/Ethology

© C. Brühl
An inhabitant of primary forests.

Distribution Fig. 231 Female of Aphaniotis ornata from Deramakot, Sabah.

Two localities are known from Ka-
limantan, i.e. upper Mahakkam river
(LIDTH DE J EUDE 1905) and Res. Kutai (photo- Sabah and Brunei (DAS, pers. comm.). Mt. Kina-
graphic record K NOWLES), all the others lie in balu: Kiau.

Genus Bronchocela KAUP, 1827 its base, then roundish or slightly laterally com-
pressed.
In contrast to all other similar agamids, females of
Of the agamid genera occurring on Mt. Kinabalu, this genus deposit spindle-shaped eggs (not con-
Bronchocela has the widest distribution. Its range firmed for B. celebensis, smaragdina, and danieli).
extends from the Nicobar Islands through Myanmar
as the western limit on the mainland, to Vietnam in
the east, and southeast through the Malay Penin- Fig. 232 For Bronchocela typical spindle-shaped eggs.
sula, to New Guinea and to the Philippines, includ-
ing numerous smaller islands in the whole region.
Medium-sized to large species (SVL 80-130 mm)
with long, slender limbs and tail. Dorsals and
laterals homogeneous in shape, size, and orienta-
tion. Skin folds on the sides of the neck supported
by the hyoid apparatus. Nuchal crest usually present
or at least indicated, dorsal crest present, indicated,
or absent. If dorsal crest present, continuous with
© U. Manthey

nuchal crest. Gular sac absent or only weakly de-

veloped. Gular scales and scales covering the gu-
lar sac rhombic in shape. Tail slightly triangular at

227
 Agamidae
Borneo is home to Bronchocela jubata (D AS ,
HALLERMANN, pers. comm.) and B. cristatella. The
latter occurs both within forested habitats and in
heavily disturbed, even urban, setting. but seems
to prefer open areas. In the Headquarters region of
Mt. Kinabalu, these lizards can be found in forests
up to altitudes of approx. 1700 m a.s.l. In the Po-
ring region it was found outside the forest
(MANTHEY & DENZER 1982).
Key to the Bornean species of
Bronchocela
1 6-10 (rarely 4-5 or 12) upper rows of dorsals point-
ing upwards; dorsals/ventrals > 1,5 .....................
............................................................ cristatella
• 2-4 upper rows of dorsals pointing upwards;
dorsals/ventrals £ 1,2 ............................... jubata
Fig. 233 Bronchocela cristatella from the Headquarters.
© T. Ulber
228
Bronchocela cristatella (KUHL, 1820)
Agama cristatella KUHL, 1820: 108 – Type locality: not
indicated
Calotes cristatellus, MOCQUARD 1890: 129, pl. VII: fig. 3, 3a;
BARTLETT 1895a: 85; DE ROOIJ 1915: 121; SMITH 1931: 10,
31; MANTHEY & DENZER 1982: 18; MANTHEY 1983: 25;
EBENHARD & SJÖGREN 1984: 24; MALKMUS 1985: 9; OTA &
HIKIDA 1991: 180
Bronchocela cristatella, MALKMUS 1987: 285, 1988c: 175,
1991a: 33, 1992a: 130; TAN 1993: 2; MALKMUS 1994b: 245;
MANTHEY & GROSSMANN 1997: 161
Bronchocela sp., MALKMUS 1994b: 234, figs. 12, 13, 1996c: 288
Diagnosis
Slender, long-tailed, long-limbed, and narrow-
headed agamids with a low nuchal crest and dorsal
crest indicated. Dorsals diamond-shaped, homog-
enous, imbricate, smaller than the similarly shaped
ventrals.
Description
SVL males 87-130, females 82-96 mm; TL males
300-440, females 290-330 mm. A row of enlarged
 Agamidae

scales of variable lengths

behind the supraciliary
edge. Small nuchal crest of
lanceolate, erect scales;
dorsal crest indicated by a
low, serrated ridge; 6-10
(rarely 4-5 or 12) rows of
dorsals directed up- and
backwards, laterals back-
and downwards; ventrals
about 1,5 to 2 times as large
as dorsals; scales on the gu-
lar sac smaller than ven-
trals.
Upper side usually green;
whitish or light blue spots
visible on the sides, often
fusing to form vertical
bands; flanks sometimes
reticulated with brown;
sides of the head green to
light blue, occasionally
with a reddish tinge; orbit,
or only orbit edge, and
tympanum usually black;
ventral surfaces completely
greenish yellow, gular sac
occasionally yellowish red,
yellow, or whitish. Entire
coloration subject to colour
changes towards brown,
dark green to blackish, yel-
lowish, or reddish.
1-2 spindle-shaped eggs
(9-10 × 40,5-42 mm) per
clutch.

Ecology/Ethology
Bronchocela cristatella in-
habits trees and bushes on
the edge of forests, and ur-
ban areas such as parks.Its
vertical distribution ranges
© R. Malkmus

Fig. 234 Bronchocela crista-

tella from the Headquarters
in a blackish colour phase.

229
Agamidae
from sea level up to approx. 1700 m a.s.l. On Mt.
Kinabalu this agamid has also been found in pri-
mary forest (M ALKMUS 1994b). Their slightly
spreadable ribs enables it to parachute from a tree
to the ground (MANTHEY & GROSSMANN 1994).
Distribution
Myanmar to New Guinea, and Philippines. Mt.
Kinabalu: Poring, Kiau, Kenokok, Sayap, Lobang,
Bundu Tuhan, Headquarters region including the
Liwago Trail.
Genus Complicitus
MANTHEY in MANTHEY & GROSSMANN, 1997
Monotypic genus endemic to Mt. Kinabalu, de-
scribed on the basis of a single male specimen with
a habitus similar to species of the genus Pseudo-
calotes. Unique among agamid lizards due to the
combination of lateral gular sac pockets and almost
granular scutellation of the gular sac.
Fig. C Male of Complicitus nigrigularis, drawing after the holotype.
230
Complicitus nigrigularis
(OTA & HIKIDA, 1991)
Calotes nigrigularis OTA & HIKIDA, 1991: 185-187, fig. 4 –
Type locality: Kinabalu Park (alt. ca. 1500 m)
Bronchocela nigrigularis, MALKMUS 1994b: 235
Calotes (inc. sed.) nigrigularis, MALKMUS 1994b: 245
Complicitus nigrigularis, MANTHEY in MANTHEY & GROSSMANN
1997: 165, fig. LXII
Diagnosis
Small agamids (SVL less than 80 mm) with slightly
heterogeneous dorsals regarding shape, size, and
orientation. Males with lateral gular sac pockets,
nuchal crest, and dorsal crest indicated; limbs and
tail comparatively short.
Description
SVL male 70 mm, TL male 155 mm; nuchal crest
composed of 5 high, lanceolate scales; dorsal crest
indicated by erect, distinctly spaced scales, con-
tinued on the anterior portion of the tail; dorsals of
various sizes, mainly smooth; laterals usually dia-
mond-shaped, pointed or broadly rounded, differ-
V. Heinrich

ing in size, arranged in irregular rows; ventrals
slightly heterogeneous in size, keeled; scales on
the gular sac very small, predominantly granular;
shoulder fold weakly developed; tail substantially
laterally compressed laterally behind its base.
Ground colour of males dark greyish brown above;
several white spots below the eye, on shoulder and
nuchal region; two wide, whitish cross bands on
back and upper flanks; lower side predominantly
whitish; gular sac black with a white margin and
white spots.
Ecology/Ethology: Unknown.
Distribution
Mt. Kinabalu: Only known from the type locality.
Genus Draco LINNAEUS, 1758
The distribution of this genus ranges from India
east to China, the Philippines, and the Indo-Aus-
tralian Archipelago.
Very slender, petite build lizards up to a SVL of
140 mm. Two large, spreadable, often colourful
wings (patagia) supported by extended, movable
ribs. Additional skin sails on the sides of the neck
(neck lappets) and dewlaps in various shapes and
colours, indicative of the species and often also of
the sex. Nostrils oriented upwards or laterally; tym-
panum visible or covered with scales.
Tab. 23 Distribution of Draco-species on Borneo.
Agamidae
Shape and coloration of dewlaps and inner side of
the neck lappets are involved in the recognition of
conspecifics and are therefore of great value for
the taxonomist. The patagia are folded back along
the sides of the body when the lizard rests. Only
when a flying dragon jumps off a tree these “wings”
are opened and permit controlled gliding. Depend-
ing on their starting point and the available unob-
structed space, up to 50 metres may be covered in
a single flight. These agamids use this mode of trav-
elling to change from one tree to another and al-
ways land in an upward-oriented position. In dis-
plays of threatening behaviour and advertisement,
they do not only use their dewlaps, but also often
employ their patagia.
They are strictly arboreal and inhabit sparse for-
ests as well as dense lowland and montane rain
forests. Some species are even found in coconut
plantations and in parks of large cities. Their diet
consists mainly of large quantities of ants and ter-
mites. Almost all species lay 1-4, in exceptional
cases 6 eggs.
Ten (9) species are known to inhabit Borneo, 7 (6, if
D. affinis is not a good species) of which live on Mt.
Kinabalu.
Taxonomic comments: Following recent studies
(HONDA, OTA, KOBAYASHI, NABHITABHATA, YONG &
HIKIDA 1999; MCGUIRE & KIEW 2001), the revision
of this genus by MUSTERS (1983) is no longer ten-
able in all its details. Here the results of their in-
vestigations are taken into consideration.
D. affinis BARTLETT, 1895 is not included in the key
because its validity is currently uncertain.
231
Agamidae
Key to the Bornean species of Draco
1 Five patagial ribs ............................................... 2
• Six patagial ribs ................................................ 6
2 Nostril directed upwards, supraciliary edge with-
out a thornlike scale .......................................... 3
• Nostril directed laterally, supraciliary edge with
or without a thornlike scale .............................. 5
3 Patagium dorsally yellowish to greenish or bronze
to golden brown with 5-6 pairs of dark narrow
bands, dewlap smoky grey (males) or pale yellow
(females), lower portion of dewlap with greatly
enlarged scales .................... obscurus obscurus
• Patagium dorsally black or blackish, dewlap
without enlarged scales ..................................... 4
4 Patagium dorsally black with orange-yellow spots,
in males dewlap jet black with a white portion on
posterior part of base; in females dewlap black to
dark reddish, base white; in both sexes front por-
tion of the inner side of neck lappets black, basal
portion pure white ................................................
................... melanopogon nigriappendiculatus
• Patagium dorsally with black marbling sometimes
forming five ill-defined bands or similar as in 4,
in males dewlap bright yellow with a large black
spot on base, in females dewlap reddish to lightly
washed with orange, same also on inner side of
neck lappets in both sexes ...................................
.................................. haematopogon microlepis
? Draco affinis BARTLETT, 1895
Draco affinis BARTLETT, 1895a: 80 – Type locality: Borneo
Draco cornutus, SMITH 1931: 22 (part., females)
Diagnosis (after BARTLETT 1895a)
“Similar to D. cornutus, but without large spine-
like scale above the eye. Gular appendage very
small; in females almost absent.”
Description (after BARTLETT 1895a)
“Male – Back dull brown, tinged with green; three
distinct greyish white transverse bands on the back;
232
5 Supraciliary edge with a thornlike scale, body with
lateral row of enlarged scales, in both males and
females dewlap and inner side of neck lappets pale
reddish to yellowish, tail without crest ...............
......................................... fimbriatus fimbriatus
• Supraciliary edge without a thornlike scale, body
without enlarged scales; dewlap in males golden-
yellow with brown anterior edge, tail of males with
crest .................................................. cristatellus
6 Nostril directed upwards, supraciliary edge with-
out a thornlike scale .......................................... 7
• Nostril directed laterally, supraciliary edge with
a thornlike scale ................................................ 8
7 Patagium dorsally with a pattern of alternating or-
ange red and black bands .....................................
............................ quinquefasciatus longibarba
• Patagium dorsally with a pattern of alternating
green and black bands ......................... maximus
8 Patagium usually reddish dorsally in both sexes
with black spots, or dark with green stripes (some
males); upper side of body predominantly green
(males) or brownish (females) ............. cornutus
• Patagium black dorsally with lighter spots, upper
side of body bronze coloured or light brownish to
light greyish ......................... volans sumatranus
wing membranes above bright brick red spotted with
black, with broad black outer margins, a pale bluish
grey line down the centre of the belly; underside of
wing membranes dull brick red tinged with blue,
and spotted with blackish brown, margined with
blotches of black and greyish white.
Adult female – Back grey mottled with dark brown
and tinged with green; wing membranes bronze
green spotted with black, with a broad black band
on the outer margin; gular appendage small, yel-
lowish green; chest blue; a line down the centre of
belly and under part of hind limbs pale blue, sides
of body greyish white mottled with black, under-
 Agamidae

side of wing membrane yellowish green, outer

margin blue.”
Remarks: The above quoted description of the
dewlap of the female is based on an error (com-
pare Diagnosis); it actually refers to males where
such description is missing. The allocation of the
D. cornutus females to the taxon D. affinis by SMITH
(1931) is based on the mentioned absence of a dew-

© T. Hikida
lap. Of all species of Draco inhabiting the Sunda
region, the absence of a dewlap in females is unique
to D. affinis. The allocation of females referred to
by SMITH (1931) to D. affinis is done here with res-
ervations, as neither the types of D. affinis nor the
specimens originally available to SMITH could be
examined. The validity of D. affinis is currently
uncertain.

Ecology/Ethology: Unknown.

© T. Hikida
Distribution
Borneo. Mt. Kinabalu: ? Kiau.

Draco cornutus GÜNTHER, 1864

Draco cornutus GÜNTHER, 1864: 125 – Type locality: Borneo

Draco cornutus, MOCQUARD 1890: 128; HANITSCH 1900a: 70,
1900c: 2; SMITH 1931: 22 (part. (?)); DE ROOIJ 1915: 72; MUS-
TERS 1983: 19; EBENHARD & SJÖGREN 1984: 24; TAN 1993: 2;
MALKMUS 1994b: 245; HONDA et al. 1999: 198, fig. 2A, B, C.
Draco spilopterus cornutus, HENNIG 1936: 183; MANTHEY 1983:
25

Diagnosis

© T. Hikida
A small to medium-sized species (SVL max. 85 mm)
with 6 patagial ribs and a thornlike scale on the
supraciliary edge. Nostrils oriented laterally; dew-
lap triangular, tip orange yellow in males, in fe-
males dewlap white; inner side of neck lappets and
dewlap similarly coloured.

From top to base: Draco cornutus

Fig. 235 Female (left dorsal pattern, right ventral pat-
tern of patagium).
Fig. 236 Male (left dorsal pattern, right ventral pattern
© T. Hikida

of patagium).
Fig. 237 Male (ventral pattern of patagium).
Fig. 238 Dewlap of a male.

233
 Agamidae
© S. Wilson

Fig. 239 Male of Draco cornutus from Niah Caves NP, Sarawak.
© S. Wilson

Fig. 240 Female of Draco cornutus

from Niah Caves NP, Sarawak.

234
 Agamidae

Description Draco fimbriatus fimbriatus KUHL, 1820

SVL males up to 78, females up to 85 mm; 6 patagial
ribs; nostrils oriented laterally; a thornlike scale on
Draco fimbriatus KUHL, 1820: 101 – Type locality: “India
supraciliary edge; tympanum usually not covered
orientali” – Restricted type locality: Malay Peninsula
with scales; dewlap triangular, with small scales; (HENNIG 1936)
neck lappets with larger scales; males and females Draco fimbriatus, SMITH 1931: 10, 31; EBENHARD & SJÖGREN
with a small nuchal crest composed of triangular 1984: 24; MALKMUS 1987: 285, 1992a: 130; TAN 1993: 3;
scales; dorsal crests absent in either sex; dorsals vary- MALKMUS 1994b: 245;
Draco fimbriatus fimbriatus, MANTHEY 1983: 25
ing in size, smooth, sometimes intermixed with some
larger scales; keeled scales on bases of patagium;
Diagnosis
fringe-like scales along the rear edges of the thighs;
A moderately large species (SVL 90-117 mm) with
ventrals keeled, about size of dorsals.
5 patagial ribs and a thornlike scale on the supra-
Dorsal surface of males green or greenish brown,
ciliary edge. Nostrils oriented laterally; dewlap tri-
occasionally with indistinct, dark bands; dorsum of
angular; inner side of neck lappets and dewlap or-
females light brownish with extensive darker inclu-
ange (yellow in specimens from Poring, MCGUIRE,
sions; both sexes with dark spots on head, nape, and
pers. comm.)
on sides of the neck; lower surface of males and fe-
males whitish with a turquoise to bluish centre and
Description
small ventrolateral dark (males) or yellowish brown
SVL males 90-112, females 102-117 mm; 5 pata-
(females) spots; patagium reddish above, more
gial ribs; nostrils directed laterally; a thornlike
greenish towards the body (males), or brownish (fe-
males) with dark spots occasionally converging to
form bands; a wide dark marginal band sometimes Fig. 241 Draco fimbriatus fimbriatus from Poring.
with very light inclusions; some males dark with
indistinct, greenish longitudinal lines; in males lower
surface of patagium bluish or reddish with a blue
marginal band and large black blotches fusing on
the margins; in females lower surface of patagium
yellowish white proximally, then sometimes reddish,
and bluish with dark spots distally, dark spots occa-
sionally fusing to form a marginal band; dewlap in
males with a yellowish to orange yellow tip and a
greenish white base, inner side of neck lappets with
a corresponding coloration; inner side of neck lappets
and dewlap in females white with very pale yellow-
ish brown speckles.
3-4 eggs per clutch.

Ecology/Ethology
Draco cornutus usually inhabit lowland forests and
hilly regions. Probably this is a forest edge and
canopy species and can be found in totally disturbed
urban areas, such as the grounds of the Sabah Mu-
seum.
© R. Malkmus

Distribution
Borneo, Sumatra, Java, and Bunguran Islands. Mt.
Kinabalu: Kiau, Poring.

235
Agamidae

scale on supraciliary edge; tympanum usually with-
out scales; dewlap triangular with small scales;
neck lappets with larger scales; dorsals heteroge-
neous, predominantly smooth, with only a few
keeled scales intermixed; lateral row of enlarged
scales; fringe-like scales on posterior edges of
thighs and base of tail; ventrals keeled, larger than
dorsals.
Dorsally coloration like bark of tropical trees,
brown, grey, olive, and pale greenish, interspersed
with darker patches; patagium indistinctly striped
with black and greyish brown; lower surface of
patagium light grey, spotted with dark brown
(males), or light orange with black spots (females);
inner side of neck lappets orange (yellow in Por-
ing specimens); dewlap reddish orange (yellow in
Poring specimens), anterior and lower margins
lighter (females); similar coloration in males.
Males with low nuchal sail and caudal crest of
small, triangular scales, dewlap comparatively
larger and wider. Nuchal crest of females consist-
ing only of slightly enlarged scales.
3-4 eggs (10 × 17 mm) per clutch.
Ecology/Ethology
Draco f. fimbriatus inhabits predominantly lowland
forests and hilly regions.
Distribution
Malay Peninsula, Borneo, Sumatra including the
surrounding smaller islands, and Java. Mt.
Kinabalu: Kiau; Poring.

Draco maximus BOULENGER, 1893 Fig. 242 Female of Draco maximus from Poring.

Draco maximus BOULENGER, 1893: 522 – Type locality: Mt.

Dulit, 2000 feet [Sarawak]
Draco maximus, WERNER 1910: 16; DE ROOIJ 1915: 81; SMITH
1931: 31; MUSTERS 1983: 57; EBENHARD & SJÖGREN 1984:
24; TAN 1993: 3; MALKMUS 1994b: 245, 1996c: 289, fig. 9;
MANTHEY & GROSSMANN 1997: 170, fig. 113
Draco maximus maximus, HENNIG 1936: 188; MANTHEY 1983:
25

Diagnosis
The largest species of Draco on Borneo (SVL up
to 139 mm) with 6 patagial ribs, and no thornlike
scale on supraciliary edge. Nostrils oriented up-
ward; dewlap triangular, lower portion reddish
brown (males) or greenish grey (females); inner side
of neck lappets grey to dark grey.

Description
SVL males up to 139, females up to 137 mm; 6
patagial ribs; nostrils oriented upward; no thorn-
like scale on the supraciliary edge; tympanum par-
tially or completely covered with scales; dewlap
triangular, with small scales; neck lappets with
slightly enlarged scales along the edges; dorsals
very small, primarily homogeneous with just a
small series of enlarged scales along base of
© A. Nöllert

patagium; fringe-like scales on the rear margins of

thighs and on base of tail; ventrals keeled, larger
than dorsals.

236

Dorsally predominantly green with a brownish
black pattern; patagium black with numerous light,
brownish green, discontinuous longitudinal lines;
lower side of patagium almost colourless; inner side
of neck lappets grey (males) or dark grey (females);
dewlap black at its base, with white dots, remain-
ing portions reddish brown (males) or greenish grey
(females).
Males with nuchal sail and a much larger dewlap.
1-5 eggs (11 × 18 mm) per clutch.
Ecology/Ethology
Draco maximus prefers primary rain forests up to
altitudes of approx. 600 m a.s.l.
Distribution
Peninsular Malaysia, Sumatra, Natuna Islands, and
Borneo. Mt. Kinabalu: Poring.
Fig. 243 Female of Draco melanopogon nigriappen-
diculatus from Sumatra.
© U. Manthey
Agamidae
Draco melanopogon nigriappendiculatus
BARTLETT, 1895
Draco nigriappendiculatus BARTLETT , 1895a: 92 – Type
locality: Kuching, Sarawak
Diagnosis
A small to moderately large species (SVL up to
93 mm) with 5 patagial ribs, no thornlike scale on
the supraciliary edge. Nostrils directed upward;
dewlap triangular, black; inner side of neck lappets
black and white.
Description
SVL males up to 89, females up to 93 mm; 5 pata-
gial ribs; nostrils oriented upward; no thornlike
scale on supraciliary edge; tympanum partly or
completely covered with smooth skin; dewlap long,
Fig. 244 Male of Draco melanopogon nigriappendi-
culatus from Sumatra.
© U. Manthey
237
 Agamidae

triangular; neck lappets with enlarged scales; dor-

sals heterogeneous, smooth or faintly keeled; a lat-
eral row of enlarged scales; fringe-like scales on
posterior part of thighs and on base of tail; ventrals
keeled, larger than dorsals.
Dorsally mostly green, with brownish bands or dia-
mond shaped spots and lighter inclusions; patagium
dorsally black with numerous yellow-orange spots,
these sometimes with small dark centre; dorsal
coloration and pattern of patagia shimmering
through, visible on the underside. Front portion of
inner side of neck lappets black, basal portion pure
white; in males dewlap jet black with a white por-
tion on posterior part of base, in females dewlap
black to dark reddish, base white.
1-2 eggs per clutch.

Ecology/Ethology
Draco m. nigriappendiculatus inhabits usually sec-
ondary and primary lowland forests up to eleva-
tions of approx. 600 m a.s.l.
© U. Manthey

Distribution
Sumatra and Borneo. Mt. Kinabalu: Poring
Fig. 245 Male of Draco melanopogon nigriappendicu- (MCGUIRE, pers. comm.).
latus from Sumatra.

Fig. 246 Male of Draco melanopogon nigriappendicu-

latus from Santubong, Sarawak. Draco obscurus obscurus
BOULENGER, 1887
Draco obscurus BOULENGER, 1887b: 95 – Type locality: “Mount
Kina Baloo”
Draco obscurus, MOCQUARD 1890: 128, pl. VIII: fig. 1, 1a, 1b;
DE ROOIJ 1915: 85; SMITH 1931: 10, 31; TAN 1993: 3; MALKMUS
1994b: 245;
Draco formosus, (not Draco formosus BOULENGER, 1900) WER-
NER 1910: 12; SMITH 1931: 10, 31
Draco formosus obscurus, HENNIG 1936: 218; MANTHEY 1983:
25; EBENHARD & SJÖGREN: 24
Draco obscurus obscurus, MUSTERS 1983: 65
Draco volans, (not Draco volans LINNAEUS, 1758) MANTHEY & DEN-
ZER 1982: 16; EBENHARD & SJÖGREN: 24; MALKMUS 1994b: 245
Draco volans volans, (not Draco volans L INNAEUS , 1758)
MANTHEY 1983: 25

Diagnosis
A moderately large species (SVL up to 100 mm)
© W. Denzer

with 5 patagial ribs; no thornlike scale on the su-

praciliary edge. Nostrils oriented upward; dewlap
tongue-shaped, smoky grey (males) or pale yellow

238

© U. Manthey
Fig. 247 Male of Draco obscurus obscurus from Por-
ing Hot Springs.
(females); inner side of neck lappets maroon
(males) or pale pink (females).
Description
SVL males up to 100, females up to 94 mm; 5
patagial ribs; nostrils oriented upward; no thorn-
like scale on supraciliary edge; tympanum entirely
or partially covered with smooth skin; dewlap long,
lingulate; slightly constricted at its base, lower
portion with greatly enlarged scales; neck lappets
with enlarged scales; dorsals heterogeneous, mainly
smooth; a lateral row of enlarged scales; fringe-
like scales on posterior part of thighs and on tail
base; ventrals varying in size, keeled, slightly larger
than dorsals.
Dorsally mostly grey, sometimes pale turquoise
with numerous brownish inclusions; patagium yel-
lowish to greenish, also bronze coloured to golden
brown (females with more brilliant colours); 5-6
pairs of dark narrow bands; rear margin of patagium
Agamidae
chestnut coloured; lower side of patagium grey to
whitish, with indistinct bands and maroon margins
often fragmented into spots in females; inner side
of neck lappets maroon (males) or pale pink (fe-
males); dewlap smoky grey (males) or pale yellow
(females).
1-5 eggs per clutch.
Ecology/Ethology
Draco o. obscurus inhabits primary forests up to
elevations of 900 m a.s.l.
Distribution
Sumatra and Borneo. Mt. Kinabalu: Kiau, Po-
ring.
Draco quinquefasciatus longibarba
HENNIG, 1936
Draco quinquefasciatus longibarba HENNIG, 1936: 193 – Type
locality: Baram, Sarawak, Borneo
Draco quinquefasciatus, SMITH 1931: 10, 31; MUSTERS 1983:
70; EBENHARD & SJÖGREN 1984: 24; TAN 1993: 3; MALKMUS
1994b: 245
Draco quinquefasciatus longibarba, MANTHEY 1983: 25
Diagnosis
A medium-sized species (SVL up to 106 mm) with
6 patagial ribs, no thornlike scale on the supracili-
ary edge. Nostrils pointing upward, dewlap cres-
cent-shaped, mustard-yellow (males) or dull yel-
low (females), same colours on inner side of neck
lappets.
Description
SVL males up to 106, females up to 108 mm; 6
patagial ribs; nostrils oriented upward; no thorn-
like scale on supraciliary edge; tympanum fully
covered with scales; dewlap long, slender, crescent-
shaped, with small scales; neck lappets with en-
larged scales; dorsals heterogeneous, predomi-
nantly smooth; a lateral row of enlarged scales;
fringe-like scales on the rear margins of thighs and
on base of tail; ventrals keeled, about same size as
dorsals.
Bright green (males) or greenish to olive (females)
above, speckled with reddish to dirty brown all over,
239
 Agamidae
© I. Das
Fig. 248 Draco quinquefasciatus longibarba, juv. from Batu Apoi, Brunei.
Genus Gonocephalus KAUP, 1825
The distribution of this genus is limited to the Sunda
region and the Philippine Islands.
Moderately large to large agamids (SVL 90-170 mm,
TL 430 mm max.) with a distinct angular bony ridge
between eye and nose (Canthus rostralis) and occa-
sionally strongly raised angular supraciliary border.
Both sexes with gular and shoulder folds, gular sac,
nuchal crest, and visible tympani. Males and the
majority of females have a dorsal crest (variable in
height even within a population, independent of body
size). Nuchal crest usually situated on a nuchal sail,
the latter covered with rows of large scales. Similar
rows of scales on the base of the dorsal crest, differ-
ent from the remaining dorsal scales. Head covered
with small, granular, heterogeneous scales. Dorsals
240
with a pattern of broad brown
bands (males and females);
patagium yellowish orange to
reddish, with 5 dark brown to
black bands, each with a mid-
line of fine white dots; colora-
tion and pattern of females more
contrasting and brilliant; lower
surface of patagium yellowish
white, with 4 black bands (males
and females); inner side of neck
lappets mustard-yellow (males)
or dull yellow (females) at base,
tip white; dewlap mustard-yel-
low with greenish stripes on base
(males) or dull yellow (females).
Males with low nuchal sail and
larger dewlap than females.
1-4 eggs per clutch.
Ecology/Ethology
D. q. longibarba live in lowland
forests and hilly regions.
Distribution
Sumatra and Borneo. Mt. Kina-
balu: Kiau and Poring.
and laterals sometimes almost equal in size, usually
intermixed with larger ones. Ventrals diamond-
shaped, imbricate, and larger than dorsals.
Angle-headed lizards often inhabit the canopy of
rainforests at low and medium elevations, where
they are difficult to locate. Favourite resting places
at night are the outer ends of thin branches. Field
observations seem to indicate that they do not wan-
der too far away from running waters.
Borneo is home to four species, two of which have
also been recorded from Mt. Kinabalu.
Taxonomic comments: After examination of a
larger number of specimens of Gonocephalus
bornensis it became clear that Gonocephalus
denzeri MANTHEY, 1991 is a junior synonym of G.
bornensis (M ANTHEY , unpublished, see also
MANTHEY & GROSSMANN 1997: 191).

Key to the Bornean species of Gonocephalus
1 Supraciliary border strongly raised and angular
posteriorly ................................... doriae doriae
• Supraciliary border round ................................. 2
2 Dorsal scales almost of equal size, not intermixed
with larger scales; dorsal and nuchal crest discon-
tinuous in males, females with nuchal sail only .
................................................................ grandis
• Dorsals intermixed with larger scales; dorsal and
nuchal crests continuous in both males and
females .............................................................. 3
Remark: Because of G. mjobergi there is only one female specimen known; it is not included in the key.
Tab. 24 Distribution of Gonocephalus-species on Borneo.
Gonocephalus bornensis
(SCHLEGEL, 1848)
Lophyrus bornensis (sic!) SCHLEGEL, 1848: 4, pl. III – Type
locality: Borneo
Gonocephalus borneensis, S MITH 1931: 10, 31; MALKMUS
1994b: 245; MANTHEY & GROSSMANN 1997: 179, fig. 122,
123
Tiaris miotympanum GÜNTHER, 1872 (syn. after MANTHEY &
DENZER 1992: 12)
Gonocephalus borneensis, MANTHEY & DENZER 1982: 18;
MANTHEY 1983: 26; MALKMUS 1988c: 175, 1992a: 130
Gonocephalus bellii (borneensis), MANTHEY & DENZER 1982:
18
Gonocephalus bellii, (not Lophyrus bellii DUMÉRIL & BIBRON,
1837) MANTHEY 1983: 25; EBENHARD & SJÖGREN 1984: 24;
MALKMUS 1988c: 175; TAN 1993: 4
Gonocephalus liogaster, (not Tiaris liogaster GÜNTHER, 1872)
TAN 1993: 4
Agamidae
3 A row of enlarged scales below the eye ending
above the tympanum (tympanum/eye > 0,7); sides
of body with or without light roundish spots; iris
of males bright blue ............................. liogaster
• No row of enlarged scales below the eye,
tympanum/eye < 0,7; sides of body usually with
light oval spots; iris of males brown or light blue
............................................................ bornensis
Diagnosis
A moderately large Gonocephalus (SVL up to
136 mm) with a round supraciliary border and a
small tympanum; small conical to weakly granu-
lar dorsals usually intermixed with larger ones;
males and females with a continuous vertebral crest
composed of lanceolate scales.
Description
SVL males 118-136, females 90-130 mm; TL
males 261-310, females 215-275 mm; body mus-
cular, with an almost round tail and limbs of mod-
erate lengths; supraciliary border round; continu-
ous, high crest of closely arranged, lanceolate
scales (fastened to one another in young males),
decreasing in height towards the terminal third of
the body, and ending abruptly on the base of the
241
 Agamidae

Fig. 249–252
Gonocephalus bornensis

© U. Manthey
Fig. 249 Adult male Fig. 250 Semiadult male
© C. Brühl

from Deramakot, Sabah. from Poring Hot Springs.

Fig. 251 Female from Kipungit II. Fig. 252 Female from Kipungit I.

© R. Malkmus
© U. Manthey

242
 Agamidae

tail; tail with low, serrated scales (old males); fe-

males with a high nuchal crest of lanceolate scales,
dorsal crest considerably lower, decreasing in
height substantially in the anterior half of the body,
reduced to small triangular scales in the posterior
portion. Dorsals and laterals small, conical, usu-
ally intermixed with some larger scales on the
flanks; often with a row of enlarged scales paral-
lel to the dorsal crest; upper sides of limbs with
keeled scales; scales on throat and gular sac con-
vex, not keeled; ventrals smooth but more or less
distinctly convex on the chest (young specimens

© C. Brühl
up to about 65 mm SVL with keeled ventrals);
diameter of tympanum small, ratio to eye diam-
eter 0,37-0,59.
Semiadult males brown, olive, or green dorsally
and laterally, with a dark, varying, reticulated pat-
tern; adult males predominantly brownish, reticu-
lated pattern often indistinct; iris brown or a faded
light blue; females rust-red dorsally and dorso-
laterally; with more or less distinct, light, oval spots;
dirty white ventrolaterally, with dark reticulation,
or dorsal side greenish with an almost black re-
ticulated pattern; iris brown; gular sac light, with
dark, interrupted stripes; limbs and tail banded light
and dark. Hatchlings green to olive above with a

© A. Nöllert
dark reticulated pattern and short light brown or
rust-brown bars on the back.
Four eggs per clutch of 22 mm length; hatchlings
35-37 + 63-65 mm (SVL + TL).

Ecology/Ethology
G. bornensis inhabits primary rain forests up to
altitudes of approx. 700 m a.s.l. They live on tree
trunks and vines, very often in the vicinity of run-
ning waters. In case they are disturbed, they may
also retreat into hiding places on the ground.
Eggs are deposited in intervals of about 3 months.

Distribution
Endemic to Borneo. Mt. Kinabalu: Poring, Kiau.

Right from top to base Gonocephalus bornensis

Fig. 253 A female from Sg. Kipungit I, 500 m, near
© A. Nöllert

Poring is laying eggs.

Fig. 254 Hatchling from Sg. Kipungit I, Poring.
Fig. 255 Juvenile male from Poring.

243
 Agamidae

Gonocephalus grandis (GRAY, 1845)

Dilophyrus grandis GRAY, 1845: 239 – Type locality: Rangoon

[in error]
Gonocephalus grandis, MANTHEY & DENZER 1982: 18; MANTHEY
1983: 26; EBENHARD & SJÖGREN 1984: 24; MALKMUS 1988c:
175; MANTHEY & DENZER 1991: 3; TAN 1993: 4; MALKMUS
1994b: 237, 245

Diagnosis
A large angle-headed dragon (SVL up to 160 mm)
with a round supraciliary border; dorsals homog-
enous, very small, not intermixed with larger scales;
dorsal crest absent (females), or distinctly separated
from the nuchal crest (males).

© M. Schroth
Description
SVL males 134-160, females 115-137 mm; TL males
345-427, females 305-365 mm. Body more or less Fig. 257 Female of Gonocephalus grandis from Poring.
strongly compressed, robust; males slimmer than
females; limbs relatively short; supraciliary border Fig. 258 Gonocephalus grandis, some days/weeks old
from Sg. Kipungit I, Poring.
round; nuchal crest of males composed of long, im-
bricate, lanceolate scales on an arched sail; dorsal

Fig. 256 Male of Gonocephalus grandis from Sumatra.

© P. Hoffmann
© U. Manthey

244

crest clearly separated from the nuchal crest, also
consisting of long, imbricate, lanceolate scales, de-
creasing in height at both ends; caudal crest lim-
ited to small, triangular scales; females without
dorsal crest, nuchal sail with irregularly arranged
larger scales; dorsals and laterals very small, ho-
mogenous, smooth, without intermixed enlarged
scales; limbs covered with small, smooth or slightly
keeled scales; scales on the gular sac and ventrum
smooth.
Back of males in various shades of green to almost
black; flanks blue, with yellow spotting; gular sac
yellow to red, striped with blue; females exhibit a
dark band with light borders between eye and shoul-
der; back brownish, greenish, anthracite, or almost
black; light, chevron-shaped bands; coloration of
flanks similar to that of males; colour of gular sac
light with dark stripes. Juveniles similar to females,
but flanks pale and reticulated dark.
3-6 (most 4-6) eggs (10-11 × 21-26 mm) per clutch;
hatchlings 32-35 + 48-65 mm (SVL+TL).
Genus Hypsicalotes
MANTHEY & DENZER, 2000
Monotypical genus known only from three males
from Mt. Kinabalu, differing from all other Asian
species by extremely large plates on the sides of
the head in combination with a heterogeneous
scutellation of throat and gular sac.
Hypsicalotes kinabaluensis
(DE GRIJS, 1937)
Calotes kinabaluensis DE GRIJS , 1937: 136 – Type locality:
“Kina-Balu Mountains, North Borneo”
Calotes kinabaluensis, WERMUTH 1967: 39; TAN 1993: 2; OTA
& HIKIDA 1996: 188; INGER & LAKIM 1998: 143
Calotes kinabaluensis, (inc. sed.) M ALKMUS 1994b: 245,
MANTHEY & GROSSMANN 1997: 152
Bronchocela kinabaluensis, MALKMUS 1994b: 235
Hypsicalotes kinabaluensis, MANTHEY & DENZER 2000:13-20
Agamidae
Ecology/Ethology
Gonocephalus grandis is native to primary rain
forests up to elevations of 1400 m a.s.l., on Mt.
Kinabalu up to approx. 700 m a.s.l. These agamids
are often found on trees or shrubs immediately ad-
jacent to small rivulets, rapidly flowing streams,
or wide rivers. Males usually seek the vicinity of
females. Females, but also juveniles, like to perch
on rocks in, or next to a waterbody, particularly
during morning hours. If in danger, they leap into
water and dive out of sight. Defecation also takes
place in water.
Females mature at ca. 20 months and begin laying
clutches of eggs every 1-3 months. Incubation takes
75-90 days.
Distribution
Malay Peninsula as well as Pulau Pinang and
Tioman, Sumatra and Pulau Nias, Nako, and
Sipora, Borneo. Mt. Kinabalu: Poring, Sg.
Manggis.
Diagnosis
Relatively large agamid lizard (SVL up to 145 mm)
with extremely large scales on the sides of the head
in combination with a heterogeneous scutellation
of throat and gular sac.
Fig. 259 Head of the neotype of Hypsicalotes kinaba-
luensis (male).
© U. Manthey
245
Agamidae
V. Heinrich
Fig. CI Male of Hypsicalotes kinabaluensis, drawing after the neotype.
Description
Relatively large agamid lizard (SVL up to 145 mm)
with moderately long limbs and tail; dorsals scales
equal in size and orientation; laterals varying in
size, shape, and arrangement; sides of head with
an extremely large scale below the tympanum
(about the size of the ocular cavity), probably
smaller in females; nuchal and separate dorsal
crests present at least in males, the latter proceed-
ing on the tail. Males with a distinctly developed
gular sac, anterior edge with long, lanceolate
scales; scales on gular sac very small, ranging in
shape from oval to rhombic and varying in size;
throat with scales differing extraordinarily in
shape and size. Distinct fold on the shoulder; tail
246
swollen behind its base, then substantially com-
pressed laterally.
Head and body green, the latter with a banded
pattern of chocolate-brown and black spots; tail
predominantly brown with small green inclu-
sions.
Ecology/Ethology
One specimen was discovered at night sleeping on
the leaf of a bush 20 cm above the ground and 1.5 m
from a small stream.
Distribution
Endemic to Mt. Kinabalu, the only precise locality
known being Sayap, 900 m a.s.l.

Genus Phoxophrys HUBRECHT, 1881
The distribution of this genus is limited to the is-
lands of Sumatra and Borneo.
Small (SVL up to 84 mm, but usually smaller),
poorly known species without horn-like protru-
sion; tympanum not visible. Gular sac absent or
Key to the Bornean species of Phoxophrys (in part after INGER 1960)
1 Supraciliary border with spines ............ spinipes
• Supraciliary border without spines .................. 2
2 Continuous dorsal crest of low, closely arranged
vertebral scales .................................. nigrilabris
• Dorsal crest indicated by several widely spaced
vertebral scales .................................................. 3
Tab. 25 Distribution of Phoxophrys-species on Borneo.
Phoxophrys borneensis INGER, 1960
Phoxophrys borneensis INGER, 1960: 222, figs. 1, 2A – Type
locality: Bundu Tuhan, Mount Kinabalu, North Borneo
Phoxophrys borneensis, MANTHEY 1983: 26; E BENHARD &
SJÖGREN 1984: 24; MALKMUS 1988c: 174; TAN 1993: 7;
MALKMUS 1994b: 237, 245, 1995e: 67, fig. p. 66, 1996c: 289;
MANTHEY & GROSSMANN 1997: 195, fig. 141
Diagnosis
A small agamid (SVL less than 79 mm) without a
supraciliar spine; vertebral crest composed of
widely spaced scales; base of the tail with keeled
scales laterally.
Description
SVL males 63-66, females 61 mm; TL males 92-104,
females 90 mm; occipital region and sides of head
with conical scales; nuchal crest short, consisting
Agamidae
very small; low nuchal crest usually present. Dor-
sal crest absent or indicated. Dorsals small, het-
erogeneous, and intermixed with larger tubercu-
lar scales. Base of tail in males substantially thick-
ened, distinctly elongate and dorsally flattened in
shape.
Borneo is home to four species, two of which oc-
cur on Mt. Kinabalu.
3 Gular scales and lateral scales on the tail base
keeled, head of males “normal” in size ...............
........................................................... borneensis
• Gular scales and lateral scales on the tail base
smooth, head of males conspicuously large .......
............................................................ cephalum
of roundish (males), or flattened (females), coni-
cal scales; gular scales keeled; dorsal crest indi-
cated by some raised scales; males with a small
gular sac; dorsals and laterals heterogeneous,
mainly smooth, intermixed with larger, keeled, and
some tubercular scales; ventrals homogeneous on
the chest, keeled, more heterogeneous posteriorly
and almost smooth; base of the tail with keeled
scales on the sides.
Males brownish to almost black dorsally, with yel-
low, oblique bands and a yellow radial pattern
around the eye; throat yellow with a brown striped
pattern; ventral sides whitish, marbled with brown.
Young females dark brown dorsally with narrow,
green, oblique bands; some enlarged scales whit-
ish; upper sides of limbs same coloured; two light
stripes between the eyes, another from the eye to
the angle of the moth; sides of head and neck with
247
 Agamidae

some green markings; throat whitish, marked with

a network of brown stripes. Youngs rust-red dorsally
with a pattern of light brown to light olive bands,
upper sides of limbs same coloured; two whitish
bars between eyes, another one from the eye to the
angle of the mouth; great parts of neck and sides of
head whitish to light greenish.
In males tail base substantially widened with 4 rows
of keeled subcaudals.
Two eggs per clutch.

Ecology/Ethology
P. borneensis inhabits preferably shrubs. These
small agamids have been recorded from altitudes
between 1300 and 1500 m a.s.l. on Mt. Kinabalu.

Distribution
Borneo, Sabah: Crocker Range (I. DAS pers. comm.)
Sarawak: region of the Trusan River. Mt. Kinabalu:
Bundu Tuhan, Sg. Liwago, Sayap (Sg. Kemantis).
© H. Sauer

Phoxophrys cephalum
Fig. 260 Young female of Phoxophrys borneensis from
(MOCQUARD, 1890)
the Headquarters.
Pelturagonia cephalum MOCQUARD, 1890: 130, pl. VII: figs. 4,
Fig. 261 Male of Phoxophrys borneensis from Kiau, 4a – Type locality: “Kina Balu” (syn. of Otocryptis
900 m. (Japalura) nigrilabris PETERS, 1864 after BOULENGER 1891)
Pelturagonia cephalum, WHITEHEAD 1893: 183
Phoxophrys cephalum, I NGER 1960: 223 (valid species);
MANTHEY 1983: 26; EBENHARD & SJÖGREN 1984: 24; TAN 1993:
7; MALKMUS 1994b: 245, 1995e: 68, figs. p. 66, 67, 68, 1996c:
289, fig. 8; MANTHEY & GROSSMANN 1997: 196, fig. 142
Japalura nigrilabris, (not Otocryptis (Japalura) nigrilabris
PETERS, 1864) BOULENGER 1891: 342; HANITSCH 1900a: 70,
1900c: 2; DE ROOIJ 1915: 93; SMITH 1931: 22
Phoxophrys nigrilabris, (not Otocryptis (Japalura) nigrilabris
PETERS, 1864) TAN 1993: 8; MALKMUS 1994b: 245

Diagnosis
A relatively small agamid (SVL up to 84 mm) with-
out a supraciliar spine; vertebral crest of widely
spaced scales; base of the tail with smooth scales
laterally.

Description
SVL males 71-84, females 59-74 mm; TL males
© U. Kuch

126-140, females 111 mm; terminal portion of su-

praciliary edge, occipital region, sides of head, and

248

lower jaw covered with conical scales; gular scales
varying in size, nearly diamond-shaped, smooth to
granular, sometimes intermixed with almost tuber-
cular scales; nuchal crest of 5-8 separated, thick,
triangular scales of various size; vertebral crest in-
dicated by a few, widely spaced and erected scales;
dorsals heterogeneous, intermixed with some
larger, conical scales; ventrals smooth to feebly
convex in adult specimens, sometimes keeled in
juveniles and semi-adults, larger than dorsals; base
of tail with smooth scales laterally.
Greenish to azure dorsally, with dark green, ob-
lique bands varying in width; head often paler
than back; throat whitish with some dark green
spots; ventral sides light, marbled with dark. Fe-
males dull brown (WHITEHEAD 1893) with a yel-
low throat.
Head disproportionately large in males relative to
body size; base of tail substantially widened,
dorsally strongly flattened with a series of hard,
mucronate scales along the edges, laterally with
smooth scales; two rows of keeled subcaudals.
Fig. 262 Male of Phoxophrys cephalum from Kamborangoh Road, 1750 m.
Agamidae
Ecology/Ethology
Phoxophrys cephalum appears to be a slow-mov-
ing lizard. When disturbed this species will remain
completely motionless. They may exhibit a cata-
leptic behaviour over a period of several minutes
which includes a half-open mouth and a vaulted
tongue. During this stage individuals can be ma-
nipulated at will.
These lizards inhabit shrubs and trees in rain for-
ests (< 500 up to approx. 2100 m a.s.l.).
Remark: Interestingly, the two specimens which
subsequently became the type specimens were
caught by natives in rat-traps positioned on the
ground. This may indicate a more terrestrial life
than most other Bornean agamid lizards lead.
Distribution
Sabah: Sunsuron, Sg. Mesilau, Ranau. Mt. Kina-
balu: Kiau, Kenokok, Tenompok, Lumu Lumu,
Bundu Tuhan, Kamborangoh Road, Kamborangoh,
Sayap (Sg. Kemantis).
© H. Sauer
249
Anguidae

Anguidae
The family of the anguine lizards is organized in three subfamilies which inhabit mainly the temperate
and subtropical regions of the northern hemisphere, but are also represented by some species in south
America. Only a few species are known from Southeast Asia.
Apart from primitive four-legged species, various transitional stages lead to more advanced legless forms
with hardly any remains of a girdle skeleton. All species are considered to be carnivorous. Most of them
are oviparous, but some give birth to fully developed young.
As far as Borneo is concerned, the family is represented only by the genus Ophisaurus (glass lizards) of
the subfamily Gerrhonotinae whose members are characterized by the absence of limbs. They are distin-
guished from all other legless reptiles by the presence of lateral folds.

sals in 16-18 longitudinal and 98-105 transverse

Genus Ophisaurus DAUDIN, 1803
The distribution of this genus is disjunct with rep-
resentatives in the southeastern USA, parts of north-
ern Africa, southeastern Europe, West Asia, and
Southeast Asia.
Comparatively rigid body with strong lateral folds;
no limbs. Head slightly pointed anteriorly and hardly
distinct from the neck. Eyes with movable eyelids,
ear-opening present. Tail fragile with the ability to
regenerate. Large, rhombic scales in regular longi-
tudinal and transverse rows, attached to ossified
plates. Lateral folds extensible, with small,
unossified scales adding to the animal’s mobility.
Only one species is known from Borneo and has
also been recorded from Mt. Kinabalu.
Ophisaurus buttikoferi
LIDTH DE JEUDE, 1905
Ophisaurus Büttikoferi LIDTH DE JEUDE , 1905: 192 – Type
locality: Mount Liang Koeboeng, Central-Borneo (= Bukit
Liang Kubung, on the Sungai Kapuas, West Kalimantan)
Ophisaurus buettikoferi, INGER 1958d: 479; MANTHEY 1983: 27;
EBENHARD & SJÖGREN 1984: 24; MALKMUS 1994b: 246
Diagnosis
A moderately large lizard of up to 125 mm SVL,
characterized by almost equally sized frontal and
interparietal shields which form a wide suture.
rows, central 12 rows of dorsals keeled; ventrals
smooth, in 10 longitudinal rows; all caudals
keeled.
Colour in alcohol (after DE ROOIJ 1915):
“Brown above; a dark lateral band edged above
with a light band; this lateral band is continued
on the tail. A small blue spot on the interparietal,
indicating the location of the parietal-eye. On the
anterior part of the back irregular transverse se-
ries of bluish spots, anteriorly edged with black.
Lips and lower parts pale yellowish; five oblique
dark lines on the side of the head, running across
the lips, the first behind the nostril, the second
below the eye, the third at the corner of the mouth,
the fourth below the ear-opening and the fifth at
the nape; the latter is continued on each side of
the belly.”
Ecology/Ethology
This anguine possibly leads a secretive life be-
neath leaf litter or in the soil. Only three speci-
mens have been found so far, of which one was
discovered on a path through primary rain forest
at about 300 m altitude. The stomach of another
specimen contained the remains of a cockroach
and fragments of the leg of a rat. Such rodent
would, however, be much too large for this lizard
so it may be safe to assume that the leg was torn
off a dead rat.

Description Distribution
SVL 119-125 mm, TL 375 mm; frontal and inter- This species is endemic to Borneo. Mt. Kinabalu:
parietal form a wide suture; 5 supraoculars; dor- without precise locality data.

250
 Gekkonidae

Gekkonidae
Geckos have a cosmopolitan distribution inhabiting all temperate and tropical regions. Often they are the
first reptiles to invade newly formed islands (e.g. from volcanic eruptions).
They are small to moderately large (total length up to about 400 mm), mainly crepuscular and nocturnal
lizards, typically with a granular skin structure. Usually they possess vertical-oval pupils as well as
adhesive lamellae underneath their digits (subdigital lamellae). The arrangement and shape of these
lamellae are important features for the identification of several genera. Adhesive lamellae enable geckos
to climb on seemingly smooth, vertical surfaces. Geckos are known for a highly evolved autotomy and
the ability to regenerate lost tails. Regenerates are however different in skin structure and shape. Geckos
inhabiting Borneo usually posses (original) tails which are longer than the snout-vent-length.
In contrast to most other groups of lizards, many gekkonid lizards are able to produce clearly audible
sounds which often are indicative of a species. They feed on a large variety of invertebrates and several
species exploit the abundance of insects attracted at night by electric light. Many geckos adapted to man-
made alterations of nature and live in and around human habitations. All species in the range reproduce
by laying eggs which may be attached to a surface. Production of several fertilized clutches after a single
mating process is (Amphigonia retardata) is not uncommon among geckos.
Borneo is home to 11 genera comprising 29 geckonid species with 8 genera and 10 species occurring on
Mt. Kinabalu.

Key to the Bornean genera of Gekkonidae (in part after MANTHEY & GROSSMANN 1997)
1 True eyelids present (A), body compressed
laterally .................................... Aeluroscalabotes
• Without eyelids, body compressed dorsoventrally A
........................................................................... 2 B
2 Pupil vertical (B) ............................................... 3
• Pupil round ........................................ Cnemaspis
3 Fingers dilated (C) ............................................ 4
D
C
• Fingers slender, slim, curved (D) ........................
..................................................... Cyrtodactylus
4 Inner finger/toe well-developed ....................... 5
• Inner finger/toe rudimentary (E) ......................... E
........................................... Hemiphyllodactylus F G
5 Inner finger/toe clawed (F) ............................... 6
• Inner finger/toe without claw (G) ..................... 7
6 Body and tail with lateral skin fringes (H) .........
......................................................... Cosymbotus
• Body and tail without lateral skin fringes ...........
...................................................... Hemidactylus
7 Body and tail with lateral skin fringes (H) .........
......................................................... Ptychozoon H
• Body and tail without lateral skin fringes ........ 8

251
Gekkonidae
8 Subdigital lamellae divided (I) ...................... 10
• Subdigital lamellae not divided (J) ................. 9
9 Paired submaxillars present (K), median row of
subcaudals enlarged (L) ........................... Gekko
• Submaxillars not in pairs (M), median row of
subcaudals not enlarged .............. Luperosaurus
10 Terminal ends of outer 4 finger free and raised at
an angle from the dilated portion of the finger
(N); skin tears easily ............................. Gehyra
• Terminal ends of outer 4 finger continuous with
the dilated portion of the finger (O); skin not
easily torn ................................. Lepidodactylus
Genus Cyrtodactylus GRAY, 1827
Geckos of the genus Cyrtodactylus are distributed
from Southeast Asia to New Guinea but also occur
in northeastern Australia and on the Solomon Is-
lands.
Mainly moderately large lizards (SVL of Bornean
species up to 125 mm); body compressed dorsoven-
trally, with ground colour usually being some shade
of brownish; dorsal surface marked with highly vari-
able, dark spotted or banded patterns. Head acute-
triangular in shape when viewed from above, and
usually very distinct from the neck. Pupil vertical-
oval with crenate edges. Fingers and toes clawed,
slender, not dilated, their arched terminal phalanges
compressed laterally. Ventral surface of basal
phalanges with enlarged plates or lamellae in rows.
Males usually with preanal and/or femoral pores,
some species with preanal grooves.
Tab. 26 Distribution of Cyrtodactylus-species on Borneo.
252
I J K L
M
N
O
Species of Cyrtodactylus occurring in the Sunda
region inhabit predominantly primary rain forests,
but some have also adapted to secondary and bush
forests. They normally live at altitudes between
sea-level and approx. 1300 m, on Mt. Kinabalu
up to 2500 m a.s.l. These are strictly nocturnal
geckos which often leave their shelters only a few
hours after sundown. They are found on the for-
est floor, on rocks, in caves, or on and in tree
trunks, very often in the vicinity of running wa-
ter. Many species are able to curl up their tails
laterally. They feed on a variety of invertebrates.
Females produce clutches of usually two roundish
hard-shelled eggs, in some species the eggs are
spindle shaped.
Of the eight species known to inhabit Borneo,
four have been recorded from Mt. Kinabalu.
Cyrtodactylus is therefore the gekkonid genus
with the greatest specific diversity on this moun-
tain.

Key to the Bornean species of Cyrtodactylus (in part after HIKIDA 1990)
1 Light narrow network pattern on head and light
narrow bands on the back ................................. 2
• Colour pattern different from above ................. 3
2 With enlarged femoral scales, males and females
with 9-14 preanal pores forming a narrow angu-
lar series, males with 0-6 femoral pores; subdigital
lamellae beneath fourth toe 22-28, supralabials 10-
16; 58-65 ventrals (at mid-body between ventro-
lateral folds) ................................... consobrinus
• Without enlarged femoral scales, males with 8-
10 preanal pores forming a wide angular series,
without femoral pores; subdigital lamellae 21-23;
supralabials 8-11; 58-62 ventrals ... malayanus*
3 Supralabials 8-9, no femoral pores or enlarged
femoral scales, males with preanal groove con-
taining two pairs of pores; subdigital lamellae 22-
26; supralabials 8-9; 51-58 ventrals ....................
........................................................ cavernicolus
• More than 9 supralabials .................................. 4
4 Males with femoral pores (4-9 on each side), en-
larged femoral scales and 9-11 preanal pores form-
ing a narrow angular series; subdigital lamellae
20-23; supralabials 10-12; 40-45 ventrals ..........
............................................................. baluensis
• Males without femoral pores or enlarged scales
........................................................................... 5
Cyrtodactylus baluensis
(MOCQUARD, 1890)
Gymnodactylus Baluensis MOCQUARD, 1890: 125, pl. VII: fig.
1, 1a, 1b, 1c – Type locality: Kina Balu
Gymnodactylus baluensis, BOULENGER 1894d: 722; DE ROOIJ
1915: 14; SMITH 1931: 10, 11, 21, 31; MANTHEY 1983: 24
Cyrtodactylus baluensis, EBENHARD & SJÖGREN 1984: 23, 24;
MALKMUS 1988a: 29, 1989: 195; HIKIDA 1990: 91; MALKMUS
1991a: 33, 1991c: 6, fig. 1, 1a, 1992a: 126, 1994b: 234, 245,
1996c: 288; MANTHEY & GROSSMANN 1997: 219, fig. 155
Gonydactylus baluensis, TAN 1993: 5
Gymnodactylus marmoratus, (not Cyrtodactylus marmoratus
GRAY, 1831) HANITSCH 1900a: 70, 1900c: 1; DE ROOIJ 1915:
13 (part.); SMITH 1931: 31; MANTHEY 1983: 24
Cyrtodactylus marmoratus, (not Cyrtodactylus marmoratus
GRAY, 1831) EBENHARD & SJÖGREN 1984: 24
Lymnodactylus sp., JACOBSON 1985: 42 fig. 42a
Gekkonidae
5 Males with preanal groove containing 3-5 pairs
of pores (7-9 total); subdigital lamellae 17-22;
supralabials 10-11; 43-55 ventrals ... pubisulcus
• Males without a preanal groove ....................... 6
6 Back with a pattern of dark blotches; males with
7-8 preanal pores forming a narrow angular se-
ries; subdigital lamellae 23-28; supralabials 10-
12; 40-43 ventrals ..................................... ingeri
• Back with dark cross bands .............................. 7
7 Males with 7-8 preanal pores forming a wide an-
gular series; subdigital lamellae 22; supralabials
10-12; 48-51 ventrals ............................. matsuii
• Males with 8-12 preanal pores forming a narrow
angular series; subdigital lamellae 25-30;
supralabials 10-14; 50-58 ventrals ........... yoshii
* The occurrence of C. malayanus on Borneo is ques-
tionable.
Remark: A satisfactory identification key which
takes both males and females into consideration is
currently out of question since females of some spe-
cies are still unknown.
Diagnosis
A moderately large Cyrtodactylus with a maximum
SVL of 86 mm; no preanal groove; males with 9-
10 preanal pores arranged at an acute angle in a
shallow depression, and 6-9 separated femoral
pores on either side of the former; 40-45 rows of
ventrals at mid-body between ventrolateral folds;
21-23 subdigital lamellae beneath fourth toe.
Description
SVL up to 86 mm; tail slightly longer; body slen-
der; dorsal scales fine, intermixed with larger tu-
bercles in 21-24 irregular rows; ventrolateral folds
weakly developed; 4-45 rows of small, roundish,
imbricate ventrals between ventrolateral folds;
rostral broad with a central groove, in contact with
253
 Gekkonidae

nostrils; 10-12 supralabials, 9-10 infralabials; men-
tal triangular, followed by 2-3 pairs of submaxillars;
fourth toe with 20-23 subdigital plates, largest on
basal phalange; tail dorsally with small scales, in-
termixed with tubercles; its basal portion with 8
scales and 2 tubercles per segment.
Males with 9-11 preanal pores arranged at an acute
angle in a shallow depression (4-6 per side); preanal
pores widely separated, in large femoral scales, 4-9
on each thigh; females without pores, but with a se-
ries of enlarged scales arranged at an obtuse angle
in a flat preanal region; males with 3-4 pairs of large,
females with 2-3 pairs of small postanal tubercles.
Beige to yellowish brown dorsally, with irregular,
dark brown spots, waved or crenated bands or short,
broad, longitudinal dashes; head often with a dark
brown, almost V-shaped lateral band from the tip
of snout to the nape; limbs and tail banded in light
and dark brown above; ventral surface almost uni-
formly violet grey.
Fig. 263 Cyrtodactylus baluensis from Sg. Silau-Silau.
Two hard-shelled, roundish eggs (12 × 15 mm) per
clutch; hatchlings 31-32 + 34-35 mm (SVL + TL)
[HIKIDA 1990].
Ecology/Ethology
Cyrtodactylus baluensis usually inhabits montane
forests at altitudes of 900 to 1800 m, up to 2500 m
a.s.l. on Mt. Kinabalu (photographic records
KNOWLES), but populations also exist in the low-
lands at 160 m a.s.l. (Melinau George, Mulu Na-
tional Park, Sarawak). This gecko finds suitable day
shelters, among others, under loose bark and be-
comes active only after dark when humidity has
risen sufficiently. It can be found on tree trunks up
to heights of 4 metres above the ground, between
plank roots, fallen logs, in the foliage of young
trees, branchwork of bushes, and more rarely on
the ground. C. baluensis is rarely encountered dur-
ing extended dry periods. When touched, the gecko
raises high on its legs, bends its body into an S,
turns the head to one
side, and hisses. If given
a chance, it will bite with
vigour.
In parts of its distribution
it occurs syntopically
with C. consobrinus and
C. yoshii (Danum Val-
ley), and with C. conso-
brinus and C. pubisulcus
(Mulu).
Females lay the eggs in
humus beneath the roots
of trees.

Distribution
Cyrtodactylus baluensis
is endemic to Borneo
(Sabah, Brunei). Mt.
Kinabalu: Poring, Kiau,
Sg. Kemantis, Kenokok,
Bundu Tuhan, Lumu
Lumu, Power Station,
Kamborangoh, Sg. Mesi-
lau, Headquarters re-
© M. Maronde

gion, and on the Summit

Trail up to 2500 m a.s.l.
(KNOWLES pers. comm.).

254

Cyrtodactylus consobrinus
(PETERS, 1871)
Gymnodactylus consobrinus PETERS (1871): 569 – Type local-
ity: Sarawak, Borneo
Cyrtodactylus consobrinus, MALKMUS 1988d: 9, fig. 5, 11;
HIKIDA 1990: 93; MALKMUS 1991c: 8, fig. 3, 1992a: 129,
1994b: 245
Cyrtodactylus consobrinus kinabaluensis MALKMUS, 1989: 196
– Type locality: “Waldbach hinter den Hot Springs (600 m)”
(syn. after HIKIDA 1990: 93)
Gonydactylus consobrinus, TAN 1993: 5
Diagnosis
A large species of Cyrtodactylus with a maximum
SVL of 125 mm; no preanal groove; males with 9-
11 preanal pores arranged in an obtuse angle, and
0-6 femoral pores per thigh; 65-70 rows of ventrals
at mid-body between ventrolateral folds; 22-28
subdigital lamellae beneath fourth toe.
Description
SVL up to 125 mm, TL up to 163 mm; body slen-
der, but muscular; body scales small, intermixed
Fig. 264 Cyrtodactylus consobrinus from Poring.
Gekkonidae
with roundish, weakly keeled tubercles in 18-20
irregular rows; ventrolateral fold with larger tuber-
cles; small, roundish, imbricate ventrals in 65-70
rows at mid-body between ventrolateral folds.
Rostral large with a central groove, quadrangular, in
contact with the nostrils; 10-16 supralabials, 9-13
infralabials; mental triangular, followed by one pair
(more rarely two pairs) of large submaxillars.
Fourth toe with 22-28 subdigital lamellae, largest
on basal phalange; tail roundish in cross-section,
tapering to a point, with small flat scales above and
4-8 rows of keeled tubercles at its base.
Males with 9-11 preanal pores arranged at an ob-
tuse angle (4-6 per side), and 0-6 femoral pores on
each thigh, and 6-7 enlarged femoral scales; no
preanal groove; 2-3 pairs of large postanal tuber-
cles; females with enlarged preanal and femoral
scales, but no pores, and 2 pairs of postanal tuber-
cles.
Dorsal surfaces rich chocolate-brown with darker
sections and scattered light dots; 4-8, usually indi-
vidually shaped, cream-coloured, white or yellow
bands or cross lines; dorsal surface of head with a
© M. Pfeifer
255
 Gekkonidae
characteristic, individual, light network of lines;
labials spotted with lighter colour; limbs and tail
with narrow light bands; ventral side dirty white,
beige, or reddish grey; juveniles coloured and pat-
terned like adults, but substantially more contrasted.
Two eggs per clutch; hatchlings 31 + 31 mm
(SVL+TL).
Ecology/Ethology
These geckos live in primary rain forests of the low-
lands. They prefer to stay near the ground (up to
approx. 1 metre) on tree trunks, between plank
roots, or in holes in the trunks of large, smooth-
barked trees in the vicinity or immediately next to
running waters. They usually become active only
well into the night. They react very sensitively to
light and will escape immediately when disturbed
by a torchlight.
Females produce clutches several times per year.
Distribution
Peninsular Malaysia, Sumatra, and Borneo. Mt.
Kinabalu: Poring: Hot Springs, Sg. Kipungit I
and II.
Fig. 265 Cyrtodactylus ingeri from Poring.
© K.E. Linsenmair
256
Cyrtodactylus ingeri HIKIDA, 1990
Cyrtodactylus ingeri HIKIDA, 1990: 96, fig. 7, 8 – Type local-
ity: “near Poring Hot Spring, Ranau District, Sabah”
Cyrtodactylus ingeri MALKMUS 1991c: 8, fig. 1b, 4-6, 1992a:
128, 1994b: 234, 245, 1996c: 288
Gonydactylus ingeri TAN 1993: 5
Cyrtodactylus pubisulcus (not Cyrtodactylus pubisulcus INGER,
1958) MALKMUS 1989: 195
Diagnosis
A smaller species of Cyrtodactylus with a maxi-
mum SVL of 80 mm; no preanal groove; males with
7-8 preanal pores arranged at an acute angle in a
depression; no femoral pores; 40-43 rows of
ventrals at mid-body between the ventrolateral
folds; 23-28 subdigital lamellae beneath fourth toe.
Description
SVL up to 80 mm, TL up to 100 mm; body slen-
der, slightly compressed dorsoventrally; scales
small, intermixed with 17 irregular rows of larger,
conical tubercles; ventrolateral folds weakly indi-
cated by a low, rounded tubercles; 40-43 rows of

© R. Malkmus
Fig. 266 Underside of Cyrtodactylus ingeri from Por-
ing.
small, roundish, imbricate ventrals at mid-body
between ventrolateral folds.
Rostral large, quadrangular, in contact with nos-
trils; 10-12 supralabials, 8-10 infralabials; mental
triangular, followed by 2 pairs of submaxillars.
23-29 subdigital plates beneath fourth toe, largest
on basal phalange. Tail round in cross-section, with
small scales above and rings of tubercles, 11-12
rows of granular scales between rings at tail base.
Males with 7-9 preanal pores (3-4 per side), ar-
ranged at an acute angle in a depression; no femo-
ral pores, no large femoral scales, no preanal
groove.
Dorsal surface ochre to sand-coloured or light grey
with a slight tinge of reddish, with 5-6 dark brown
bands sometimes fragmented to form large para-
vertebral blotches; neck often with a dark Y- or V-
shaped marking; dark stripe from posterior edge
of eye often extending to the insertion of the arm;
dorsal surface of limbs light brownish with dark
speckles; dorsal surface of tail with more or less
wide, dark bands; ventral surfaces whitish grey,
sometimes greyish violet.
One egg 9.5 × 12 mm per clutch.
Ecology/Ethology
This gecko leaves its day-shelter already with the
onset of dusk. It appears to prefer dwelling on leafs
Gekkonidae
and on trunks of young trees in the vicinity of run-
ning water and is often seen on branches overhang-
ing a stream. It can also be encountered on bushes,
rocky cliffs, and on the lower portions of bigger
trees up to heights of approx. 1,5 metres. When
disturbed, individuals produce a hissing sound
while fleeing. They have also been observed flip-
ping over the entire anterior body including the
arms in a way that the body is hanging down from
the tree trunk at an angle of approximately 60°.
They cling to the surface only with their hind limbs
and may maintain this cataleptic position for sev-
eral minutes.
Distribution
Endemic to Borneo. Mt. Kinabalu: Poring, Sg.
Langanan, Sayap, Sg. Wariu, Sg. Sasapan.
Cyrtodactylus matsuii HIKIDA, 1990
Cyrtodactylus matsuii HIKIDA, 1990: 100, fig. 11, 12 – Type
locality: “in the Park Headquarters of Kinabalu National Park
at 1600 m in Mt. Kinabalu, Sabah”
Cyrtodactylus matsuii, MALKMUS 1991c: 7, fig. 2, 1992a: 127,
1994b: 245, 1996c: 288
Gonydactylus matsuii, TAN 1993: 5
Diagnosis
A large species of Cyrtodactylus, SVL about 105 mm;
no preanal groove; males with 7 preanal pores ar-
ranged at an obtuse angle; no femoral pores; 51
rows of ventrals at mid-body between ventrolateral
folds; 22 subdigital plates beneath fourth toe.
Description
SVL up to 105 mm, TL 82 mm (regenerated); body
slender, slightly compressed dorsoventrally; scales
small, intermixed with larger tubercles in 18 irregu-
lar rows; a weakly indicated skinfold consisting of
low, round tubercles between axilla and groin; 51
rows of small, roundish, imbricate ventrals between
ventrolateral folds at mid-body. Large, notched,
quadrangular rostral, in contact with nostrils; 10-11
supralabials, 10-11 infralabials; mental triangu-
lar, followed by 2 pairs of submaxillars. Fourth toe
with 22 subdigital plates, largest on basal phalange.
Tail round in cross-section, with small scales
257
 Gekkonidae
© R. Malkmus
Fig. 267 Cyrtodactylus matsuii from Sg. Silau-Silau.
above, and rings of low tubercles; 4-7 rows of
granular scales separating the rings at base of tail.
Males with 7 preanal pores (3-4 per side), arranged
at an obtuse angle; no femoral pores, no preanal
groove.
Dorsal surface yellowish brown with highly irregu-
larly arranged and shaped dark brown spots and
bands on back; head with small dark spots; a dark
band from the posterior edge of the eye sometimes
meeting its counterpart on the nape; limbs and tail
with dark bands; ventral surfaces without any pat-
tern, lighter than above.
Ecology/Ethology
The few specimens found as yet were discovered
more than 2,5 metres above ground on tree trunks
(the type specimen at 7-8 m); only one male was
encountered on the ground (MALKMUS 1996c).
258
© M. Schroth
© M. Schroth
From top to base
Cyrtodactylus matsuii from Sg. Silau-Silau, 1450 m.
Fig. 268 Portrait. Fig. 269 Ventral view.
Distribution
Endemic to Borneo. Mt. Kinabalu: Headquarters
region (1450-1600 m a.s.l.).
Genus Gehyra GRAY, 1834
The distribution of the polytypic genus Gehyra
extends from Madagascar and islands of the Indian
Ocean, through Asia, the Indo-Australian Archi-
pelago, to Australia and islands of the Pacific
Ocean. Populations now also exist in Mexico and
the USA due to introduction.
Smaller geckos with vertical-oval pupils, and dor-
soventrally compressed body and tail. Skin finely
granular, sensitive to the touch, and tears easily.
Fingers and toes occasionally webbed, broad at

their bases, with well-developed, divided scansory
pads below. Outer four fingers and toes developed
normally, with thin, erect, clawed terminal
phalanges; inner fingers and toes also developed
normally, but without free terminal phalanges; fur-
nished with tiny, often concealed claws. Males with
preanofemoral pores (i.e. preanal and femoral pores
in a continuous series).
Species of the genus Gehyra live as commensals
of man in large cities as well as in forests. They
feed on a large variety of invertebrates. Females
produces clutches of two hard-shelled eggs each
several times per year.
Borneo is home to only one species which has also
been recorded from Mt. Kinabalu.
Gehyra mutilata (WIEGMANN, 1835)
Hemidactylus mutilatus WIEGMANN, 1835: 238 – Type local-
ity: Manila, Philippines
Gehyra mutilata, MOCQUARD 1890: 128; HANITSCH 1900a: 70;
MANTHEY 1983: 24; EBENHARD & SJÖGREN 1984: 24; MALKMUS
1992a: 126; TAN 1993: 4; MALKMUS 1994b: 245
Peropus mutilatus, SMITH 1931: 31
Diagnosis
A small gecko with an SVL of up to 60 mm. Body
dorsally with fine scales, without tubercles; skin
very easily torn; tail strongly compressed dorsov-
entrally, also without any conical tubercles; inner
fingers and toes with diminutive claws; fourth toe
with about 25 subdigital lamellae
of which 7-8 are divided; males
with 25-44 preanofemoral pores. Fig. 270 Male of Gehyra mutilata from Fraser’s Hill, West Malaysia.
Description
SVL up to 60 mm, TL up to 60 mm
or slightly less; body compressed
dorsoventrally; head oval, moder-
ately distinct from body; dorsal
surfaces of body and limbs with
small granular scales, no tubercles;
35-44 rows of large, imbricate,
oval-shaped ventrals between
weak ventrolateral folds; skin very
easily damaged; rostral quadran-
gular, medially grooved, in contact
with nostrils; 8-11 supralabials; 6-9
Gekkonidae
infralabials; large triangular to pentagonal mental,
followed by 2-3 pairs of large submaxillars; ear-
opening small, vertical-oval. Limbs short, hind
limbs with short fringes along the posterior edges;
fingers and toes with basal webbings; inner fingers
and toes with tiny claws; fourth toe with altogether
25 subdigital lamellae, 7-8 of which divided; wid-
ened, dorsoventrally compressed, segmented tail
without tubercles, but with finely serrated edges,
older specimens with a broad tail base; basal seg-
ments with 10-11, almost straight rows of small,
flat scales; median subcaudals greatly enlarged.
Male with a long, curved, very obtuse row of 25-44
preanofemoral pores; no postanal tubercles.
Entire upper surface light grey to light brown, oc-
casionally much darker, with or without small dark
spots and light dots; ventral surface yellowish to
dirty white.
Two hard-shelled eggs (8,5 × 10,5 mm) per clutch;
hatchlings 45-47 mm in total length.
Ecology/Ethology
The crepuscular and nocturnal geckos live as
commensals of man in villages and large cities, but
also in t rainforests and open country. They often
reside around and in houses, on poles, trees, or
fallen logs. They are common in the lowlands, but
also found in the Headquarters region of Mt.
Kinabalu.
Females produce several clutches per year in which
the eggs often stick to one another.
© W. Grossmann
259
Gekkonidae
Distribution
Sri Lanka, Andaman and Nicobar Islands east to
Taiwan, Malay Peninsula southeast to New Guinea,
Genus Gekko LAURENTI, 1768
The genus Gekko is distributed in south and south-
east Asia, north up to Korea and Japan, further-
more across the Indo-Australian Archipelago to the
Solomon and Santa Cruz Islands and on the Phil-
ippines.
Moderately large to large, relatively robust
geckos with dorsoventrally slightly compressed
bodies. Skin covered with granular scales, inter-
mixed with larger tubercles. Pupil vertical-oval,
with serrated edges. Fingers and toes well-de-
veloped, widened, occasionally with webbings
on bases, terminal phalanges not distinct from
others; claws limited to the outer fingers and toes.
Ventral surface of fingers and toes furnished with
strongly developed, undivided scansors. Preanal
Tab. 27 Distribution of Gekko-species on Borneo.
Key to the Bornean species of Gekko
1 Iris and back green .................................... smithi
• Iris brownish, back not green ........................... 2
2 Back predominantly brownish with small darker
spots, males with 32-40 preanofemoral pores;
small species, SVL of adults about 100 mm ......
.......................................................... monarchus
• Back predominantly bluish to greyish with reddish
spots, males with 10-24 preanal pores, no femoral
pores; large species, SVL of adults about 160 mm
................................................................... gecko
260
Philippines, Mascarenes, Mauritius, Seychelles,
Madagascar, Mexico, Cuba and Hawaii. Mt. Kina-
balu: Headquarters region.
or preanofemoral pores present, postanal tuber-
cles present.
The Sunda region, including Borneo, is home to
the largest species of this genus, the tokay gecko.
They are tree-dwellers and live in forests, but it is
not uncommon to find them living as commensals
of man. They are mainly active at dusk and during
the night, but sometimes their species-indicative
calls can be heard during the day. They feed on a
large variety of invertebrates although large spe-
cies of this genus can also overpower smaller ver-
tebrates. Their clutches consist typically of two eggs
which are “glued” to the surface. Females may use
the same deposition site several times . Males usu-
ally grow larger in total length than females.
Of the three species occurring on Borneo, only the
smallest, Gecko monarchus, has been recorded from
Mt. Kinabalu.
Gekko monarchus
(DUMÉRIL & BIBRON, 1836)
Platydactylus monarchus DUMÉRIL & BIBRON, 1836: 335 – Type
locality: “Amboine” [Ambon, Indonesia]
Gekko monarchus, SMITH 1931: 10, 31; MANTHEY 1983: 24;
E BENHARD & SJÖGREN 1984: 24; TAN 1993: 4; MALKMUS
1994b: 245
Diagnosis
Smallest species of Gekko on Borneo, reaching a
maximum SVL of about 100 mm. Dorsal surface
light brown to light grey ground colour with nu-
merous light and dark dots or small spots.

Fig. 271 Gekko monarchus from Cameron Highlands, Peninsular Malaysia.
Description
SVL males up to 102 mm, TL up to 125 mm, fe-
males smaller; body robust, slightly compressed
dorsoventrally; head oval, moderately distinct from
the body; dorsal surface with small scales, inter-
mixed with larger, roundish tubercles in 16-17 ir-
regular longitudinal rows; 30-38 roundish ventrals
at mid-body between ventrolateral folds; rostral
grooved medially, in contact with nostrils; 10-11
supralabials, 9-12 infralabials; mental small, tri-
angular; 1 or 2 pairs of large submaxillars; fingers
and toes moderately broadened, some with small
basal webbings; fourth toe with 16-17 subdigital
lamellae, 4-5 basal lamellae divided; tail segmented
by rings of tubercles; median, undivided subcaudals
broadened.
Both sexes with a curved row of enlarged preano-
femoral scales arranged at an obtuse angle, larger
and accompanied by 32-40 preanofemoral pores in
males; females without preanofemoral pores, some-
times possessing indented scales; both sexes with 2,
males occasionally with 3 postanal tubercles.
Gekkonidae
© U. Manthey
Dorsal surface cream-coloured to greyish brown,
speckled with numerous blackish brown dots; tu-
bercles yellowish or dark; often 6-7 paravertebral
blackish brown spots between neck and base of tail,
alternating with vertebral light markings; a dark,
almost W-shaped marking on the occiput; golden
to amber-coloured iris with a network of dark red
veins; ventral surface dirty white, occasionally
slightly translucent; hatchlings dark brown with
distinct yellow dots, dark markings indistinct.
Two hard-shelled eggs, 10-13 mm in diameter per
clutch; hatchlings 25-30 + 26-35 mm (SVL+TL).
Ecology/Ethology
Gekko monarchus inhabits lowlands and mountain-
ous regions up to altitudes of 1500 m a.s.l. This
species is common in and around human habita-
tions, in plantations, and in primary forests. It is
usually found living in pairs, more rarely in larger
groups. These geckos are mainly crepuscular and
nocturnal and present themselves as very shy. They
are territorial and do no tolerate other species of
261
Gekkonidae
housegeckos in their territories. The call of males
is not very loud and consists of a continuous series
of several “tock”-sounds.
Females produce clutches of eggs several times per
year; and are firmly attached to a suitable surface.
They often feed on empty shells once the hatchlings
Genus Hemidactylus OKEN, 1817
The distribution of the genus Hemidactylus extends
from southeast Europe through Africa, Asia, the Indo-
Australian Archipelago to the north of Australia (in-
troduced) and numerous islands of the Pacific Ocean
as well as northeastern south America. Hemidactylus
has been introduced to Mexico and the USA.
Small to moderately large geckos with vertical-oval
pupils and dorsoventrally compressed bodies.
Tab. 28 Distribution of Hemidactylus-species on Borneo.
Key to the Bornean species of
Hemidactylus
1 Tail roundish, with rows of enlarged tubercles ...
............................................................... frenatus
• Tail compressed, edges finely denticulated ........
............................................................... garnotii
Hemidactylus frenatus
DUMÉRIL & BIBRON, 1836
Hemidactylus frenatus DUMÉRIL & BIBRON, 1836: 366 – Re-
stricted type locality (LOVERIDGE 1947): Java
Hemidactylus frenatus, SMITH 1931: 10, 31; MANTHEY 1983:
24; EBENHARD & SJÖGREN 1984: 24, 25; MALKMUS 1988c: 174,
1989: 194, 1992a: 125; TAN 1993: 5; MALKMUS 1994b: 233,
245, 1996c: 288
262
have emerged in order not to waste a source of cal-
cium.
Distribution
From Thailand southeast to Irian Jaya including
the Philippines. Mt. Kinabalu: Kiau.
Granular dorsal scutellation intermixed with larger
tubercles. Fingers and toes dilated, with slender,
clawed phalanges and free, angled terminal
phalanges. Subdigital lamellae divided medially.
Males with preanal or femoral pores.
Usually living in and around houses, these geckos
feed largely on small insects. Several times per year,
females attach their clutches of two eggs to a suit-
able surface.
Both species occurring on Borneo are also present
on Mt. Kinabalu.
Diagnosis
A small housegecko of up to 65 mm SVL; dorsal
pattern absent or present and highly variable;
ground coloration light to dark brown; tail roundish
in cross-section, with distinct rings of backward-
pointing, conical tubercles.
Description
SVL up to 65 mm, TL up to 39 mm; body slender,
dorsoventrally compressed; dorsals small, with ir-
regularly scattered, weakly keeled tubercles; vent-
rolateral folds absent; ventrals smooth, roundish,
imbricate, in 28-36 rows; head oval, slightly distinct
from the body; rostral quadrangular, medially
grooved, touching nostrils; 10-12 supralabials; 7-10
infralabials; mental large, triangular or pentagonal
in shape; 2-3 pairs of submaxillary shields; fingers
and toes without basal webbings; 9-10 subdigital
lamellae beneath fourth toe, 7-9 of which V-shaped
and medially divided; tail roundish, only slightly
compressed dorsoventrally, with distinct rings of

© A. Nöllert
Fig. 272 Hemidactylus frenatus from Poring.
backward-pointing, conical tubercles; median row
of subcaudals broadened.
Males with a continuous row of 26-38 preanofemoral
pores, largest in the centre.
Dorsal surface in a large variety of shades of
brown ranging from greyish brown to yellowish
brown, highly variable as to lighter or darker; ei-
ther uniform or speckled with dark, often with
additional dark spots or streaks; dark lateral stripe
on the head; tail with or without faint bands; ven-
tral surface whitish to yellowish white, ventral
surface of tail sometimes reddish. Juveniles col-
oured like adults, but with more contrasting pat-
terns.
Usually two hard-shelled, almost round eggs
(9-10 mm in diameter) per clutch; hatchlings
18-22 + 18-23 mm (SVL+TL).
Ecology/Ethology
Hemidactylus frenatus is the most common house
gecko in Southeast Asia. Its altitudinal distribution
ranges up to approx. 1600 m a.s.l., but it is less
Gekkonidae
© P. Hoffmann
Fig. 273 Mating of Hemidactylus frenatus from Poring.
common at elevations above 1000 m a.s.l. It lives
on boulders, beneath rocks or rotting logs, on trees,
but is most commonly found on buildings of all
sorts and sizes. It inhabits villages just as well as
big cities and is usually encountered in the vicinity
of electric lights where it is drawn to at dusk. Al-
though in principal crepuscular and nocturnal, the
distinct cackling calls of males can be heard
throughout the day and have earned this gecko its
vernacular names in many regions, for example
“tingtock” and “tchikchak”.
Preceded by short matings, females produce eggs
4-6 times per year.
Distribution
Mexico, Madagascar; Mauritius, south and east
Africa, from south India and Sri Lanka east to Tai-
wan, from the Malay Peninsula southeast to New
Guinea, furthermore on many islands of the Pacific
Ocean, and the Philippines. Mt. Kinabalu: Poring
and Headquarters region (around buildings and their
immediate vicinity).
263
 Gekkonidae
Hemidactylus garnotii
DUMÉRIL & BIBRON, 1836
Hemidactylus garnotii DUMÉRIL & BIBRON, 1836: 368 – Type
locality: “L’île de Taiti” [Tahiti, Polynesia]
Hemidactylus garnotii, SMITH 1931: 10, 31; MANTHEY 1983:
24; EBENHARD & SJÖGREN 1984: 24, 25, fig. p. 27; MALKMUS
1989: 194; TAN 1993: 6; MALKMUS 1994b: 245
Hemidactylus cf. bowringii, MALKMUS 1992a: 125
Diagnosis
A small gekkonid species with a SVL of up to 65 mm
differing from the very similar H. frenatus by ser-
rated edges of its tail (vs. rings of tubercles).
Description
SVL up to 65 mm, TL up to 83 mm; body slender,
dorsoventrally compressed; dorsal scales small,
intermixed with a single tubercle or groups of tu-
bercles; ventrals small, smooth, rounded, in 29-34
rows between weakly developed ventrolateral folds;
head oval, slightly distinct from the body; rostral
quadrangular, medially grooved, in contact with
nostrils; 12-13 supralabials; 9-11 infralabials; men-
tal large, triangular in shape; 2 pairs of large
submaxillars; fingers and toes with small basal
webbings; 11-14 subdigital lamellae, 6-11 of which
divided; tail slender, dorsoventrally compressed,
with acute, finely serrated edges; dorsal surface
faintly segmented, with small scales; median row
of subcaudals broadened.
Fig. 274 Hemidactylus garnotii
© J.C. Murphy
264
Males with rows of 7-19 femoral pores on each
thigh; females with rows of 15-20 enlarged femo-
ral scales on each thigh, some slightly pitted; both
sexes with 1 postanal tubercle on each side.
Dorsal surfaces brownish in various shades, with
more or less distinct darker, often also with small
whitish spots; ventral surfaces whitish to cream-
coloured.
Ecology/Ethology
Hemidactylus garnotii lives in the lowlands and moun-
tainous regions up to approx. 1400 m a.s.l. whereby
it is more common at altitudes above 1000 m a.s.l.
The nocturnal gecko is found in and on houses as
well as in the forests.
Reproduction: Most populations consist of par-
thenogenetic females which produce clutches of
usually 2 eggs each several times per year.
Distribution
From India east to China, from Malay Peninsula
southeast to the Solomon Islands, furthermore on
several Pacific Ocean islands, Philippines, intro-
duced to Florida and New Zealand. Mt. Kinabalu:
Poring, Kiau.
Genus Hemiphyllodactylus BLEEKER, 1860
Hemiphyllodactylus is distributed from Sri Lanka
to south China and the Japanese Ryukyu Islands,
also inhabiting the Philippines, the Indo-Australian
Archipelago, and some islands of the Pacific Ocean.
Small, slender, dorsoventrally compressed geckos
with homogenous, granular scutellation. Tail usu-
ally somewhat shorter than SVL, at maximum of
equal length. Pupil vertical-oval. Outer four fin-
gers and toes occasionally webbed at bases, dilated,
with free, raised, clawed terminal phalanges; inner
fingers and toes rudimentary, diminutive, and lack
raised terminal phalanges, but sometimes with
claws; basal subdigital lamellae undivided, distal
ones medially divided. Males with preanal and
femoral pores.
Species of Hemiphyllodactylus feed on all sorts of
tiny insects. Females produce clutches of usually
two small, roundish, hard-shelled eggs.
 Gekkonidae

Borneo is home only to one species which has also Two hard-shelled eggs (5-6.5 × 6.5-8 mm) per
been recorded from Mt. Kinabalu. clutch; hatchlings 14-17 + 14-16 mm (SVL+TL).

Ecology/Ethology
Hemiphyllodactylus typus typus
This small nocturnal gecko usually inhabits moun-
BLEEKER, 1860 tainous regions above approx. 1000 m a.s.l. where
it is found in forests, agricultural areas, and in as
Hemiphyllodactylus typus BLEEKER, 1860: 327 – Type local- well as around buildings. Suitable day-shelters are
ity: “Agam” [West Sumatra] and “Goenong Parong (Java)” under wood, rocks, cushions of moss and other
(Gunung Parang, West Java)
objects.
Hemiphyllodactylus typus, SMITH 1931: 10, 21, 31; TAN 1993:
6; MALKMUS 1994b: 245 Females lay diminutive eggs attached to each other
Hemiphyllodactylus typus typus, MANTHEY 1983: 25 but not to the surface. Suitable oviposition sites
are often used by several females simultaneously.
Diagnosis
A small, reddish brown gecko, of up to 47 mm SVL Distribution
with a slim, roundish, almost patternless light India and Sri Lanka east to the Japanese Ryukyu
brown tail in great contrast to the body; body and Islands, Malay Peninsula, Greater and some of the
tail without tubercles. Lesser Sunda Islands, Philippines, New Guinea,
New Caledonia, and Polynesia. Mt. Kinabalu: Po-
Description ring, Kiau.
SVL up to 47 mm, TL up to 46 mm; body slim,
dorsoventrally compressed; dorsals small, smooth,
roundish or angular; no enlarged tubercles; no ven- Fig. 275 Hemiphyllodactylus typus typus from Sumatra.
trolateral folds; ventrals smooth, imbricate, about
40 rows; head oval, hardly distinct from the body;
rostral almost pentagonal, medially grooved, touch-
ing nostrils; 8-12 supralabials; 8-12 infralabials;
mental small, elongate, triangular in shape; no or
one pair of small submaxillars; fingers and toes with
or without basal webbings; inner fingers and toes
rudimentary, with or without tiny claws; 12-15
subdigital lamellae beneath fourth toe, distal 3-6
V-shaped; tail slender, round, with fine scales, no
tubercles; median three rows of subcaudals broad-
ened.
Males with an angular series of 6-15 preanal pores
(3-8 per side), separated from which are rows of 7-
13 femoral pores on either side; 3 round postanal
tubercles on either side;
Dorsal surface reddish brown, with slightly darker,
faint, V-shaped markings between sides of neck and
tail base; occasionally with an interrupted, indis-
tinct lateral stripe of the same colour extending from
rear margin of the eye to the leg insertion; some-
times with lighter dots arranged in longitudinal
© U. Manthey

series; dorsal surface of tail light to greenish brown,

ventral surface whitish at base, speckled with
brown, dark distally.

265
Gekkonidae
Genus Lepidodactylus FITZINGER, 1843
Lepidodactylus is a genus of southern and south-
eastern Asia, also inhabiting the Philippines, the
Indo-Australian Archipelago, New Zealand, and
islands of the Pacific Ocean.
Small, slender, dorsoventrally compressed geckos
with a fine, homogeneously granular scutellation.
Pupil vertical-oval. Fingers and toes occasionally
Tab. 29 Distribution of Lepidodactylus-species on Borneo.
Key to the Bornean species of
Lepidodactylus
1 Edge of compressed tail finely denticulated; 11-
14 upper labials ..................................... lugubris
• Edge of compressed tail nearly smooth; 9 upper
labials ............................................... ranauensis
Lepidodactylus ranauensis
OTA & HIKIDA, 1988
Lepidodactylus ranauensis OTA & HIKIDA, 1988: 616, fig. 1a-
d, 2 – Type locality: “Ranau (116°45’E, 5°50’N), Sabah,
Malaysia, Borneo”
Lepidodactylus ranauensis, MANTHEY & GROSSMANN 1997: 242,
fig. 175
Diagnosis
A small gecko of up to 45 mm SVL; body and
broad massive tail without tubercles.
Description
SVL up to 45 mm, TL slightly shorter; dorsals
small, no enlarged tubercles; no ventrolateral folds;
ventrals round, flat; head oval, slightly distinct from
body; rostral in contact with nostrils; 9 supralabials;
9-10 infralabials; mental triangular; 1 pair of large
266
webbed at bases, strongly dilated, usually with
undivided subdigital lamellae, latter V-shaped
distally and sometimes divided; inner fingers and
toes lack claws. Males with preanal and femoral
pores or preanofemoral pores.
These geckos feed on small invertebrates. Females
usually attach pairs of hard-shelled eggs to a suit-
able surface.
Borneo is home to two species of Lepidodactylus,
one of which lives on Mt. Kinabalu.
submaxillars; all fingers and toes dilated distally;
basal webbings only indicated between 3rd and fourth
toes; 15 subdigital lamellae beneath fourth toe, distal
ones V-shaped and occasionally divided; tail mus-
cular, dorsoventrally compressed, without lateral
fringes or tubercles; dorsal surface of tail finely
granular; median subcaudals slightly enlarged.
Males with an angular, continuous series of 37
preanofemoral pores extending up to the end of the
thighs; a large postanal tubercle on either side.
Dorsal surface a light shade of brown, with a dark
brown dorsolateral stripe from the occiput to the
tail; small light dots within this stripe; vertebral
region with an irregular, dark pattern; head speck-
led with blackish brown; limbs with dark bands;
Fig. 276 Cf. Lepidodactylus ranauensis from the Head-
quarters.
© I. Das
 Gekkonidae

original tail with a pattern

that corresponds with the
back;Entire dorsal surface
sometimes uniform grey
brown, locally lighter with
dark diffuse stripes; ven-
tral surfaces lighter.

Ecology/Ethology
These geckos have been
found on the walls of
buildings, in rain shelters,
and on shrubs and bushes.

© T. Ulber
Distribution
Sabah. Mt. Kinabalu:
Headquarters region. Fig. 277 Lepidodactylus ranauensis from a shelter near the Headquarters.

rus) was observed on the walls of buildings in the

Genus Ptychozoon
KUHL & VAN HASSELT, 1822
The distribution of the genus Ptychozoon is lim-
ited to the southeastern Asia, the western portion
of the Indo-Australian Archipelago and the south-
ern islands of the Philippines.
Moderately large geckos with dorsoventrally
strongly compressed bodies and tails and charac-
teristic lateral skin fringes along head, body, limbs,
and tail. Latter usually slightly shorter than head
and body together, rarely slightly longer. Skin finely
granular, dorsally mostly intermixed with larger
tubercles. Pupil vertical-oval. Fingers and toes di-
lated distally, with very large webbings. Outer four
fingers and toes with clawed, raised terminal
phalanges; subdigital lamellae broad and undi-
vided. Males with preanal and/or femoral pores.
All species are forest-dwellers mainly living on
trees, but occasionally one of them (P. rhacopho-
Headquarters region of Mt. Kinabalu. They are
equipped for short parachuting and feed on in-
sects. Clutches consist of two hard-shelled eggs
and are attached to a firm surface.
Three species live on Borneo, one, P. rhacopho-
rus, is endemic to Mt. Kinabalu.
Key to the Bornean species of
Ptychozoon
1 Sides of head with skin fringes, digits nearly fully
webbed .............................................................. 2
• Sides of head without skin fringes, digits 1/3 to ½
webbed .......................................... rhacophorus
2 Flanks with two rows of tubercles ............. kuhli
• Whole body without tubercles ........... horsfieldii

Tab. 30 Distribution of Ptychozoon-species on Borneo.

267
 Gekkonidae
Ptychozoon rhacophorus
(BOULENGER, 1899)
Gecko rhacophorus BOULENGER, 1899: 451 – Type locality:
“Kadamaian River, Kina Balu, 2100 feet, North Borneo”
Gecko rhacophorus, HANITSCH 1900a: 70, pl. 1: fig. 1, 1900c:
1; DE ROOIJ 1915: 55
Ptychozoon rhacophorus, SMITH 1931: 9; MANTHEY & DENZER
1992: 13, fig. 15; MANTHEY 1983: 25; EBENHARD & SJÖGREN
1984: 22, fig. 23; JACOBSON 1985: 42 fig. 42c; MALKMUS 1989:
195; T AN 1993: 8; MALKMUS 1994b: 245; M ANTHEY &
GROSSMANN 1997: 248, fig. 179
Diagnosis
Smallest species of Ptychozoon with an SVL of
max. 75 mm and a substantially shorter tail. Head
without lateral skin fringes, those along the body
and between toes less developed than in other spe-
cies of this genus.
Description
SVL of males up to 75 mm, TL much shorter, i.e.
up to about 50 mm; entire dorsal surface with fine
granular scales and small, round, smooth tubercles;
broad ventrolateral skin fringe between axilla and
groin with frayed edges; ventrals granular; head
oval, distinct from the head; sides of head with
conical tubercles; rostral broad, without median
groove, separated from nostrils; 9 supralabials; 10
infralabials; mental small, pentagonal; 6 pairs of
small submaxillars; limbs with ragged, partially
serrated lateral skin fringes on either side; fingers
Fig. 278 Ptychozoon rhacophorus from the Headquar-
ters.
© U. Manthey
268
and toes connected with webbings over nearly half
their lengths; 10-14, distally usually V-shaped
subdigital lamellae; skin fringes on base of tail
spike-like, then wavy with acute notches; tip of tail
tapering to a cone; dorsal surface of tail with small
scales, intermixed with numerous larger, round tu-
bercles; median subcaudals moderately broadened.
Males with about 17 preanal pores arranged in a
very obtuse row; no femoral pores; 2-3 roundish
postanal tubercles.
Dorsal surface greyish or greenish to brownish, with
often indistinct, darker, wavy bands, often with
lighter or green markings; ventral surfaces light
brown and speckled with dark, or brown with light
speckling; ventral surface of head often lighter with
dark dots.
Two hard-shelled eggs per clutch.
Ecology/Ethology
This predominantly nocturnal tree-dweller has to
date only been recorded from altitudes between 600
and 1600 m a.s.l. on Mt. Kinabalu. In the Head-
quarters region it is occasionally also found on the
outside walls of buildings. The “paragliding flight”,
for which the genus is renowned, has so far not
been observed in this species.
Females attach their eggs to a suitable surface.
Distribution
Endemic to Mt. Kinabalu: Headquarters region
and Sg. Kadamaian (600 m a.s.l.).
Fig. 279 Ventral pattern of Ptychozoon rhacophorus
from the Headquarters.
© U. Manthey
 Scincidae

Scincidae
Comprising some 1200 species, skinks or smooth lizards represent the most diverse family of lizards.
They are present in all temperate to tropical regions around the world. In most species the shiny scales
are homogenous, rounded and imbricate. Their pupils are round, and they have movable eyelids. Some
species are noted for their extremely reduced or absent limbs. The former case often goes along with a
reduction of the number of fingers and toes which otherwise number five. Like geckos, skinks have the
ability to autotomize and regenerate their tails. Juveniles of certain species differ extremely from adults
as far as their colorations and patterns are concerned (e.g. Dasia sp.).
Mt. Kinabalu is home to predominantly diurnal skinks. They are terrestrial, semiaquatic or fossorial with
a preference for levels of soil near the surface, others are often found among tree buttresses, yet others
inhabit the canopy, the banks of streams or even streams. On Mt. Kinabalu skinks may be found up to a
maximum altitude of approx. 2200 m (Sphenomorphus kinabaluensis). They feed on a large variety of
invertebrates; arboreal speciesare likely to also prey upon birds’ eggs, and larger species will occasion-
ally supplement their diet with smaller lizards. Most skinks lay two or more soft-shelled eggs, however
a few give birth to fully developed young.
On Borneo, this family is represented by 39 species in 10 genera with 8 genera and 15 species native to
Mt. Kinabalu.

Key to the Bornean genera of Scincidae (in part after MANTHEY & GROSSMANN 1997)

1 4 limbs present .................................................. 2

• Limbs absent ................................. Brachymeles A B
2 A light vertebral stripe and a dark dorsolateral
stripe from the tip of the snout to the leg (A) .....
................................................................. Lipinia
• Dorsal pattern different from that .................... 3
3 1 substantially enlarged preanal scale (B), dorsals
usually strongly keeled, humped .........................
.................................................... Tropidophorus
C D E
• No or several enlarged preanal scales, dorsals not
humped .............................................................. 4
4 1 frontoparietal (C) .................................. Emoia
• 2 frontoparietals (D) ......................................... 5
5 Supranasals present (E) .................................... 6
• Supranasals absent ................. Sphenomorphus
6 Body not elongate, adpressed limbs meet or F G H
overlap (F) ......................................................... 7
• Body elongate, adpressed limbs do not meet (G)
................................................................... Riopa
7 Dorsals keeled (H) ............................................ 8
• Dorsals smooth .............................. Lamprolepis

269
 Scincidae

8 4 supraoculars ................................................... 9
• 5 supraoculars, head and anterior portion of the I J
back with a distinct striped pattern (juveniles and
adults) (I) ....................................... Apterygodon
9 Dorsum with cross bands (more distinct in juve-
niles) (J) ..................................................... Dasia
• Dorsum without pattern, or small dots sometimes
arranged in longitudinal series, but not forming
distinct stripes; often with dark dorsolateral stripes
of various widths ................................... Mabuya

Genus Apterygodon EDELING, 1865 Apterygodon vittatum EDELING, 1865

Apterygodon is a monotypical genus endemic to
Borneo.
Moderately large, diurnal tree-dwellers with robust,
dorsoventrally compressed body, keeled dorsals and
long limbs (adpressed limbs overlap). Lower eye-
lid scaly; tympanum visible. Tail round, tail taper-
ing continuously to a point. Juveniles and adults
with a striped pattern on head and anterior part of
the back.
Apterygodon vittatum EDELING, 1865: 201 – Type locality:
“Bandjermasin, Bornéo” [Banjarmasin, South Kalimantan]
Apterygodon vittatum, MALKMUS 1987: 285; TAN 1993: 1
Apterygodon vittata, MALKMUS 1994b: 246
Dasia vittata, MANTHEY & DENZER 1982: 13, fig. 16; MANTHEY
1983: 26; MALKMUS 1985: 13
Lamprolepis vittatus, EBENHARD & SJÖGREN 1984: 22, fig. 24, 25
Diagnosis
A moderately large, robust tree skink of about 96 mm
max. SVL, distinguishable from tree-dwelling

Fig. 280 Adult specimen of Apterygodon vittatum from Santubong, Sarawak.

© U. Manthey

270
 Scincidae

skinks of other genera by its character-

istic light striped pattern on its other-
wise dark head and anterior body.

Description
SVL up to 96 mm, TL up to 108 mm;
lower eyelid covered with small scales;
tympanum visible; ear-opening small,
oval in shape; nostril in one nasal;
supranasals separated; frontonasal
wider than long, forming a suture with
rostral and frontal; 2 prefrontals, sepa-
rated from each other; frontal a little

© U. Manthey
larger than frontoparietals and inter-
parietals together; frontoparietals larger
than interparietals; nuchals distinct, oc-
casionally strongly keeled; 5 supra- Fig. 281 Juvenile specimen of Apterygodon vittatum from Poring.
oculars, the anterior 2 in contact with
frontal, second largest, fifth very small;
8-9 supraciliars; 7-8 supralabials; 6 infralabials; to find a pair occupying a tree together with their
dorsals and laterals with 3 or 5 strong keels; about offspring. They feed on insects.
30 scales around mid-body; preanals not enlarged; Several times per year, females deposit their eggs
males and females with 2 enlarged tarsal scales; in forks of branches, among roots of epiphytic
16-22 smooth subdigitals beneath fourth toe. plants, or beneath loose bark. Since a juvenile was
Head, anterior portion of the body, and back black; found near the ground (MANTHEY unpubl.) it may
a light, yellowish stripe from the tip of the snout to be assumed that eggs are also laid on the forest
the occiput; a yellowish dorsolateral stripe begin- floor.
ning above the eye and continuing bronze-coloured
on the body; another yellowish stripe from the tip Distribution
of the snout below the eye to above the insertion of Endemic to Borneo. Mt. Kinabalu: Poring and Sg.
the arm; limbs and posterior flanks brownish to Kadamaian at Kiau.
faintly olive, with small black spots and light dots;
ventral surface light green to whitish green. Juve-
niles with the same basic pattern on head and ante- Genus Brachymeles
rior part of the body as the adults, but more con-
trasted; the remainder of the upper body is bronze- DUMÉRIL & BIBRON, 1839
coloured and almost spotless; limbs and tail simi-
lar, but the former slightly darker and the latter Brachymeles is known only from the Philippines
slightly lighter than the posterior part of the body. and Borneo.
2-4 eggs per clutch. Skinks with short, degenerated limbs, or without.
Body long and round in cross-section; tail thick,
Ecology/Ethology round, ends in a blunt tip.
Apterygodon vittatum inhabits primary and second- Little is known about these skinks other than that
ary forests including Casuarina groves along sea they live in the soil near the surface. Some species
shores. This thermophile, diurnal skink shows a are oviparous, others give birth to live young.
preference for the medium and upper levels of trees Of the nine species known only one, Brachymeles
and are rarely encountered on the ground or even apus, is endemic to Sabah and occurs on Mt.
low on tree trunks. They are shy, and it is common Kinabalu.

271
Scincidae
Brachymeles apus HIKIDA, 1982
Brachymeles apus HIKIDA, 1982: 840, 6 fig. – Type locality:
“Bundu Tuhan (alt. about 1 300 m; 6°01’S; 116°32’E) near
the Headquarters of the Kinabalu National Park, Mt.
Kinabalu, Sabah, Malaysia”
Brachymeles apus, EBENHARD & SJÖGREN 1984: 24; TAN 1993:
1; MALKMUS 1994b: 245
Diagnosis
The only limbless species of skink on Borneo with
a cylindrical, blunt tail.
Description
SVL males 131 mm, females 119 mm; TL males 84
mm, females 81 mm. Lower eyelid with a single large
scale; no ear-opening; nostril between supranasal and
first supralabial; no postnasals; frontonasals wider than
long, forming a broad suture with rostral; prefrontals
separated from each other; frontal longer than wide;
parietals in broad contact behind interparietal;
frontoparietals separated; no or 1 pair of nuchals; 5
supraoculars, second largest; only 2 supraciliars in
the anterior corner of the eye; 6 supralabials, first very
large, third and fourth below eye; 5 infralabials; 1 large
postmental; dorsals and ventrals of equal size, smooth;
22-24 scales around mid-body; 108-113 vertebrals
between parietals and tail base; 5-8 enlarged preanals;
no limbs; tail cylindrical, ending in a blunt tip.
Dorsal surface of head and body dark reddish
brown, darker posteriorly; distal portion of the tail
dark brown to reddish black; tip of the snout lighter;
ocular scales darker; ventral surface light, spotless
cream-coloured; embryos reddish grey dorsally,
snout and ventral surface light grey.
Ecology/Ethology
The male holotype was collected on the road be-
tween Tenompok and Sosopodon (1300–1500 m
a.s.l.) on Mt. Kinabalu at around 2.00 a.m. while
the female paratype was dug out from a cultivated
Tab. 31 Distribution of Dasia-species on Borneo.
272
field not far from this point during the day. This
area was originally covered with primary rain for-
est. Their food is likely to consist of insects, their
larvae, and worms.
Brachymeles apus is a live-bearing species. The
paratype contained four embryos of 42-43 mm SVL
+ 20-21 mm TL.
Distribution
Sg. Purulon, Sabah. Mt. Kinabalu: Bundu Tuhan.
Genus Dasia GRAY, 1839
The distribution of the genus Dasia extends from
south India and Sri Lanka, east to the southern parts
of Vietnam, and south to Java. The genus is fur-
thermore present on southern Philippine islands.
Moderately large to large, diurnal tree-dwellers
with robust bodies, keeled dorsals, and long limbs
(adpressed limbs overlap). Juveniles with a char-
acteristic banded pattern which fades with increas-
ing age.
Only one (Dasia olivacea) of the three species inhab-
iting Borneo has been recorded from above 500 m
a.s.l. on Mt. Kinabalu.
Key to the Bornean species of Dasia
(in part after INGER & BROWN 1980)
1 Ventrals (counted in mid-line from mental to vent)
more than 56 ..................................................... 2
• Ventrals 56 or fewer .............................. olivacea
2 Adults with 5-8 dark rings between axilla and
groin; prefrontals usually separated in mid-line .
........................................................... semicincta
• Adults with 8-14 dark rings, prefrontals in broad
contact in mid-line ................................... grisea

Dasia olivacea GRAY, 1839
Dasia olivacea GRAY, 1839: 331 – Type locality: “Prince of
Wales’s Island” (= Pulau Pinang, West Malaysia)
Lygosoma olivaceum, B OULENGER 1887c: 251; MOCQUARD
1890a: 133; DE ROOIJ 1915: 203; SMITH 1931: 31
Dasia olivacea, MANTHEY 1983: 26; TAN 1993: 2; MALKMUS
1994b: 245
Diagnosis
A moderately large species reaching a maximum
SVL of up to 115 mm. Easily distinguishable from
all other Bornean species of Dasia by two ocelli on
the occipital region in adults and a not-banded tail
in juveniles.
Description
SVL up to 115 mm; lower eyelid with scales; tym-
panum deeply sunk in a very small ear-opening;
nostril in 1 nasal; supranasals separated; frontonasal
wider than long, usually forming a suture with the
rostral and the frontal; prefrontals usually separated
(rarely touching ); frontal of equal length or a little
Fig. 282 Dasia olivacea from South Thailand.
Scincidae
longer than the frontoparietals and interparietals
together; frontoparietals of equal length or a little
longer than the interparietal; parietals separated; 1
pair of nuchals; 4 supraoculars; 7-8 supraciliars; 7
supralabials, the fifth and sixth below the eye; 8
infralabials; dorsals with 3, 5, or 7, more or less
distinct keels; 28-30 scales around mid-body; 41-
46 vertebrals between the parietals and vent;
preanals not or only slightly enlarged; limbs long,
the adpressed hind limb reaches about the elbow;
17-21 smooth subdigitals beneath fourth toe; tail
muscular, long, tapering continuously to a fine
point; 85 medially broadened subcaudals.
Dorsal surface olive in various shades, with indis-
tinct cross bands of dark spots, some of them with
light centres; head often spotted with black; occiput
with 2 ocelli, more distinct in males; ventral surface
in variably light shades of green to whitish green.
Black-coloured bodies of hatchlings with narrow
reddish brown cross lines; limbs black, banded
light at their bases; dorsal surface of head light
brown with dark-margined scales; tail uniformly
© U. Manthey
273
Scincidae

yellowish to reddish behind the base; ventral sur-

face light greenish.
9-14 oval, 18-19 mm long eggs per clutch;
hatchlings 32-38 mm SVL + 40-50 mm TL.

Ecology/Ethology
Dasia olivacea inhabits lowlands and mountain-
ous regions up to approx. 1200 m a.s.l. where it
can be found in primary and secondary forests, in
parks, gardens, plantations, and Casuarina groves.
These shy skinks exhibit a natural curiosity and
claim territories which they vehemently defend
against conspecifics and other lizards. When dis-
turbed they always run up trees. It is almost im-
possible to observe these skinks for longer periods
as they are untiring foragers. Once a skink has
climbed up a tree, it will visit just about every
thicker branch, check its forks and epiphytic plants,
probably searching for food They descend to the
ground only to move to another tree. These lizards
feed mainly on arthropods.
Females produce eggs three to four times a year.

Distribution

© U. Manthey
South of 15°N, Myanmar to about Vietnam, Malay
Peninsula, Sumatra and some offshore islands,
Natuna Islands, Java, and Borneo. Mt. Kinabalu:
without precise locality data. Fig. 283 Juvenile specimen of Dasia olivacea.

Genus Lamprolepis FITZINGER, 1843
The vast distribution area of this genus is mainly
inhabited by only one species, i.e. Lamprolepis
smaragdinum, which is not present on Borneo. The
range of Lamprolepis covers Taiwan, some islands
of the Philippines, Borneo, and Java, several of the
Lesser Sunda Islands, Micronesia, New Guinea,
Australia (Cape York), and the Solomon and the
Santa Cruz Islands. Members of this genus are typi-
cally known only from a few specimens (except L.
smaragdinum).
Small to moderately large, slender skinks; usually
canopy dwellers. Body dorsoventrally compressed;
muscular limbs long (adpressed limbs overlap); tail
slender and relatively short. Lower eyelid covered
with scales like in Dasia and Apterygodon; ear-
opening small, but visible; Supranasals present or
absent; dorsals smooth.
Of the two species endemic to Borneo, one (Lamp-
rolepis nieuwenhuisi) has been recorded from Mt.
Kinabalu.
Key to the Bornean species of
Lamprolepis
1 More than 23 scale rows around mid-body .........
...................................................... nieuwenhuisi
• Less than 23 scale rows around mid-body ..........
.................................................................. vyneri

274

Lamprolepis nieuwenhuisi
(LIDTH DE JEUDE, 1905)
Lygosoma Nieuwenhuisi VAN LIDTH DE JEUDE, 1905: 195 – Type
locality: “Long Bloe” (= Long Blu, upper Mahakkam, East
Kalimantan)
Lygosoma nieuwenhuisi, SMITH 1931: 23, 31
Dasia nieuwenhuisi, MANTHEY 1983: 26; EBENHARD & SJÖGREN
1984: 24
Lamprolepis nieuwenhuisi, TAN 1993: 6
Lamprolepis nieuwenhuisi, MALKMUS 1994b: 246
Diagnosis
A moderately large species reaching an SVL of up
to 72 mm; 2-4 rows of enlarged vertebrals; 24-26
scales around mid-body; 18-20 subdigitals beneath
fourth toe.
Description
SVL up to 72 mm, TL up to 90 mm; lower eyelids
with scales; ear-opening very small, with small
lobules; nostril in 1 nasal; supranasals small, sepa-
rated; frontonasal as wide as long, forming a short
suture with rostral, in contact with frontal; prefron-
tals usually separated (rarely in contact); frontal
smaller than frontoparietals and parietals together;
frontoparietals smaller than interparietal; parietals
in contact behind interparietal; 1 pair of nuchals; 4
supraoculars; 9 supraciliars; fifth supralabial be-
low eye, twice as high as fourth; 24-26 scales
around mid-body; 2-4 rows of enlarged vertebrals;
preanals slightly enlarged; limbs long, adpressed
hind limb touching the elbow; 18-20 smooth sub-
digitals beneath fourth toe.
Colour in alcohol (after SMITH 1931): “Back light
brownish-green with distinct metallic gloss, and
numerous scattered black spots and black edgings
to most of the scales; base of tail above with black
transverse bands; top of head browner than back,
scales strongly outlined with black; below light
steel bluish or greenish.”
Tab. 32 Distribution of Mabuya-species on Borneo.
Scincidae
Ecology/Ethology
Lamprolepis nieuwenhuisi probably inhabits the
canopy and rarely descends close to the ground.
Distribution
Endemic to Borneo. Mt. Kinabalu: Kiau.
Genus Mabuya FITZINGER, 1826
Skinks of the genus Mabuya are distributed in south
America, Africa, southern and southeastern Asia.
Moderately large to large, diurnal ground-dwell-
ers with robust, dorsoventrally compressed bodies
and muscular, well-developed limbs and tails.
Lower eyelid covered with small scales in all spe-
cies inhabiting Borneo; tympanum deeply sunk in
a clearly visible ear-opening. Supranasals, prefron-
tals, and divided frontoparietals present. Dorsals
with several keels.
All of the four species known from Borneo occur
on Mt. Kinabalu.
Key to the Bornean species of Mabuya
1 More than 29 scale rows around mid-body ...... 2
• Less than 29 scale rows around mid-body ....... 3
2 Light narrow stripe from labials to forelimb or
beyond; small species, SVL of adults less than
70 mm .............................................. indeprensa
• Light stripe absent; large species, SVL of adults
more than 100 mm ........................ multifasciata
3 Less than 22 subdigitals; dorsals with 3 keels;
large species, SVL of adults more than 100 mm
.................................................................... rudis
• More than 21 subdigitals; dorsals with 5 or 7 keels;
small species, SVL of adults less than 70 mm ...
............................................................... rugifera
275
Scincidae
Mabuya indeprensa
BROWN & ALCALA, 1980
Mabuya indeprensa BROWN & ALCALA, 1980: 122 – Type
locality: “area northeast of San Jose, Mindoro Island”
Mabuia multicarinata, (not Tiliqua multicarinata GRAY, 1845)
MOCQUARD 1890a: 133, DE ROOIJ 1915: 161
Mabuya multicarinata, (not Tiliqua multicarinata GRAY, 1845)
SMITH 1931: 10, 31; MANTHEY 1983: 26; EBENHARD & SJÖGREN
1984: 24; TAN 1993: 7; MALKMUS 1994b: 246
Diagnosis
A small species of Mabuya, less than 70 mm SVL;
27-34 scales around mid-body; 40-48 scale rows
between parietals and base of tail; dorsals with 5-7
(9) keels; 18-24 subdigitals beneath fourth toe.
Description
SVL up to 67 mm; ear-opening very small, usually
without lobules; nostril in 1 nasal; 2 supranasals,
separated, long, narrow; frontonasal as wide as
long, forming a suture with rostral and frontal;
prefrontals close together or widely separated; fron-
tal about same length as frontoparietals and inter-
parietal together; parietals usually separated by a
large interparietal; 4 large supraoculars; 6-7 (6)
supralabials, fifth very large and below the eye;
usually 6 infralabials; dorsals with 5-7, rarely 9
keels (hatchlings and juveniles only 3); 27-34 (31)
scales around mid-body; 40-48 (45) vertebrals be-
tween parietals and base of tail; (52) ventrals be-
tween mental and vent; preanals not or only slightly
enlarged; limbs relatively short, adpressed hind
limb not reaching the elbow; 18-24 (21-22) subdig-
itals beneath fourth toe; tail abruptly tapering be-
hind its base.
Colour in alcohol (after BROWN & ALCALA 1980):
“Dorsal ground colour tannish or greenish olive to
light brown, usually with 2 to 4 longitudinal rows
of small darker brown to blackish blotches, some
specimens without pattern; vertebral area between
the inner rows occasionally forming a vaguely light
bluish vertebral stripe; narrow light dorsolateral
stripes sometimes present from posterior corner of
eye to tail, in some specimens present only anteriorly,
sometimes broken into scattered lighter blotches,
or very faint; lateral surfaces dark brown, marked
by a distinct, light narrow stripe extending from
labials posteriorly along side above fore limb for
276
varying distances as a line or row of spots; ventral
surface greyish-blue to slate with lighter fleshy ar-
eas on the chin as well as usually fore and hind
limb regions.”
Hatchlings 22.5 to 24.7 mm SVL in Philippine
populations.
Ecology/Ethology
This skink has been recorded from altitudes up to
1200 m on the Philippines where specimens were
found on the ground underneath leaves and in a
dry stream bed in secondary and primary forests.
Distribution
Philippines and Borneo (Sabah). Mt. Kinabalu:
without precise locality data.
Remark: The specimen classified by MOCQUARD
(1890a) as M. multicarinata was re-examined by
the author. According to its pholidosis (given in
brackets in the above description) the gravid fe-
male (50 mm SVL + 89 mm TL) could be allo-
cated to M. indeprensa without doubt. The number
of keels could not be verified due to the poor state
of the respective specimen. The only difference
worth mentioning was the presence of small lobules
which form an almost continuous ring within the
ear-opening. It can therefore be assumed that the
skinks determined as M. multicarinata by SMITH
(1931) also belong M. indeprensa, and that M.
multicarinata does not form part of the Bornean
herpetofauna (see MATSUI, HIKIDA & NAMBU 1985:
159).
Mabuya multifasciata (KUHL, 1820)
Scincus multifasciatus KUHL, 1820: 126 – Type locality: not
indicated – Locus typicus designata: Java (MERTENS 1930)
Mabuia multifasciata, MOCQUARD 1890a: 133; DE ROOIJ 1915:
162
Mabuya multifasciata, SMITH 1931: 10, 31; MANTHEY 1983: 26;
EBENHARD & SJÖGREN 1984: 24; MALKMUS 1992a: 130; TAN
1993: 7; MALKMUS 1994b: 236, 246, 1996c: 290
Diagnosis
SVL + TL up to 400 mm largest skink on Borneo,
usually with conspicuous white ocelli on the flanks.
30-25 scales around mid-body; dorsals with 3 (5)

Fig. 284 Mabuya multifasciata from Sumatra.
keels; 42-48 vertebrals between parietals and base
of tail; 16-20 subdigitals beneath fourth toe.
Description
SVL males up to 137 mm, females up to 129 mm,
TL more than 200 mm; ear-opening relatively large,
round, with several or no posterior lobules; nostril
in posterior portion of nasal; 1 postnasal; suprana-
sals separated, occasionally forming a suture;
frontonasal twice as wide as long; prefrontals usu-
ally in contact behind frontonasal, more rarely sepa-
rated; frontal as long as or slightly shorter than
frontoparietals and interparietal together; fronto-
parietals larger than interparietal; parietals sepa-
rated by a large interparietal; 1 pair of nuchals; 4
large supraoculars; 5-6 supraciliars; 6-7 suprala-
bials, fifth below the eye; 6-7 infralabials; 30-35
(usually 32-34) scales around mid-body; dorsals
with 3, very rarely 5 weak keels; 42-48 vertebrals
between parietals and base of tail; preanals not or
slightly enlarged. length of limbs correlating with
Scincidae
© U. Manthey
age, the adpressed hind limb reaching at least the
elbow. fourth toe with 16-20 smooth subdigitals.
Dorsal surface brownish to olive brown or olive
grey, almost unicoloured or with fine black lines;
thick, dark brown lateral stripe usually with black-
bordered, whitish spots; males and females often
yellowish, orange, deep red, or a combination of
these colours laterally, throat and chest often col-
oured correspondingly; ventral surface light beige,
yellowish, greenish, or light grey; hatchlings and
juveniles altogether darker; limbs and lateral stripe
almost dark brown to black, sometimes with light
dotting; dorsum golden brown, each scale bordered
with black.
Live-bearing species, 5-10 young; lengths vary
with the number of siblings 36-43 + 54-63 mm
(SVL+TL).
Ecology/Ethology
Mabuya multifasciata is by far the most common
skink in south-east Asia. It is not restricted to any
277
 Scincidae
© U. Manthey
Fig. 285 Just born Mabuya multifasciata from South Thailand.
particular type of habitat and can be found up to
approx. 1800 m a.s.l. (observation by the author
on Gunung Sibayak, Sumatra). It inhabits a wide
variety of forests, and can also be found in areas
with low vegetation. This skink is not very com-
mon on Mt. Kinabalu, probably due to a more com-
petitive situation caused by the presence of M. rudis
Both species prefer sun-exposed patches on wider
paths and clearings. The animals are attentive, love
to bask, and are excellent climbers. Besides a wide
variety of insects they prey also on smaller lizards.
Males in particular are highly territorial and de-
fend their territories vigorously against other
skinks.
Distribution
From the Andamans and Nicobars through Thai-
land to southern China, and from the Malay Penin-
sula to New Guinea, Palau Islands, and the Philip-
pines. Mt. Kinabalu: Poring, Kiau, Sg. Kadamai-
an, Bundu Tuhan.
278
Mabuya rudis BOULENGER, 1887
Mabuia rudis BOULENGER, 1887c: 188, pl. XI: fig. 3 – Type
locality: “Matang [Sarawak, Borneo] and Sumatra”
Mabuia rudis, MOCQUARD 1890a: 133; DE ROOIJ 1915: 161
Mabuya rudis, MANTHEY & DENZER 1982: 16; MANTHEY 1983:
26; EBENHARD & SJÖGREN 1984: 25, fig.; MALKMUS 1987: 285,
1988d: 9, 1992a: 130; TAN 1993: 7; MALKMUS 1994b: 246,
1996c: 290; MANTHEY & GROSSMANN 1997: 272, fig. 198
Diagnosis
A moderately large skink (up to 120 mm SVL)
varying considerably in coloration; 34-38 vertebrals
between parietals and base of tail; dorsals with 3
keels; 28-36 scales around mid-body; fourth toe
with 18-21 subdigitals.
Description
SVL up to 120 mm, TL up to 220 mm; ear-opening
round, occasionally with small anterior lobules;
nostril in posterior part of nasal; 2 supranasals;
frontonasal wider than long, forming a suture with
 Scincidae

rostral and occasionally

with frontal; prefrontals
usually separated by a nar-
row gap or, more rarely,
forming a short suture;
frontal little larger than
frontoparietals and inter-
parietal together; frontopa-
rietals larger than interpa-
rietal; parietals separated
by a large interparietal;
1 pair of nuchals, keeled
posteriorly; 4 supraocu-
lars, second largest; 6 su-
praciliars; 6-7 suprala-
bials, fifth largest and be-

© U. Manthey
low eye; 6-7 infralabials;
28-36 Scales around mid-
body; 34-38 vertebrals be-
tween parietals and base of Fig. 286 Mabuya rudis from Sg. Kipungit II, Poring.
tail; dorsals and laterals
with 3 strong keels; pre-
anals not or only slightly enlarged; adpressed hind
Mabuya rugifera (STOLICZKA, 1870)
limb reaching axilla or shoulder; fourth toe with
18-21 keeled subdigitals. Tiliqua rugifera STOLICZKA, 1870: 170, pl. X: fig. 3 – Type
Dorsal surface dark olive to reddish brown or dark locality: Camorta, Nicobars
brown, with several, black-spotted scales, some-
times arranged in longitudinal rows; a broad, light- Diagnosis
bordered, reddish brown to dark brown lateral stripe A small, sometimes very attractively coloured skink
from the tip of the snout to the tail base with or not exceeding 70 mm in SVL; 24-28 scales around
without lighter or black spots; chin and throat in mid-body; dorsals with 5 (6 or 3-4) keels; 40-48
various shades of blue, occasionally spotted with scale rows between parietals and base of tail; fourth
black or orange (males) or whitish to greenish toe with 18-24 subdigitals.
brown (females); ventral surface grass-green, yel-
lowish, brownish, brownish green, or greyish Description
brown. SVL up to 65 mm (rarely larger), TL 120 mm; ear-
2-4 eggs per clutch. opening very small, horizontal-oval, surrounded
completely by small lobules; nostril in posterior
Ecology/Ethology portion of nasal; usually no postnasal; supranasals
This is an inhabitant of lowlands and hilly areas narrow, widely separated; frontonasal wider than
preferring open situations such as clearings and long, in contact with rostral and frontal; prefrontals
paths through forests, up to approx. 1300 m. separated; frontal larger than frontoparietals and
interparietal together; frontoparietals larger than
Distribution interparietal; parietals in contact behind interpari-
Borneo, Sumatra and Mentawai Islands, Java, etal or separated by it; 1 pair of, or no, nuchals; 4
Sulawesi, Philippines (Sulu Archipelago east to supraoculars, second largest; 6 (rarely 5) supraci-
Jolo). Mt. Kinabalu: Poring: Sg. Kipungit II, liars; 9 supralabials, sixth, rarely fifth largest and
Bundu Tuhan. below eye; 7 infralabials; dorsals, laterals, and

279
 Scincidae
© C. Brühl
Fig. 287 Mabuya rugifera from Poring.
nuchals usually with 5 strong keels, laterals also
with 7 keels, some dorsals and laterals also with 6,
or only 3-4, keels; 24-28 scales around mid-body;
31 vertebrals between parietals and base of tail;
adpressed hind limb reaching the elbow; fourth toe
with 18-26 smooth subdigitals.
Colour variation of males (after BARTLETT 1895):
“Upper part of head, sides of neck, dark bronze-
green; rest of upper surface bright red-brown, finely
vermiculated with black; chin and throat pale co-
balt blue with scattered yellow spots; chest lemon
yellow; rest of ventral surface deep vermilion red.
Nearly black above; immaculate above and be-
low. Throat cobalt blue.
Throat pale cobalt blue, tinged with green on the
chest and fore-limbs; belly, ventral surface of hind
limbs and tail salmon pink.
Above blackish brown, immaculate; chin and
throat bright grass-green; belly and ventral surface
of tail salmon-red.
Above blackish brown, immaculate; eye-lids,
upper and lower lips, chin, throat and upper parts
of chest lemon yellow; belly and ventral surface of
tail salmon-red.”
Colour variation of females (after BARTLETT 1895):
“Above blackish brown with pale brown longi-
tudinal striations. Throat green, black spotted.
280
Above dark brown,
streaked with yellowish
brown from superciliary
line and corner of mouth to
base of tail; lips, chin and
throat pale blue, tinged
with green; nearly all
scales have a black termi-
nal band, these bands are
irregular and give the throat
a variegated appearance;
chest yellowish green;
belly and ventral surface of
tail pinkish salmon.
Chin and throat yellow-
ish green, spotted with
black; belly, ventral sur-
face of hindlimbs reddish
salmon tinged with green-
ish blue; of tail reddish salmon.”
Almost uniformly dark olive brown with weakly
indicated, lighter dorsolateral stripes and an equally
faint vertebral stripe in the posterior third of the
body (MANTHEY & GROSSMANN 1997 and MANTHEY
unpubl.). Lateral surface of head tinged with red-
dish; chin, throat, and chest yellowish green, re-
mainder of the ventral surface dirty white. Other-
wise, dorsal surface of body brown with numerous
black marks and 5 light brown stripes, dorsolateral
ones beginning at the posterior edge of the eye.
Head and neck rich reddish brown above, with
black spots, orange-red on lateral surface, with
black spots. Chin and throat faintly orange, remain-
der of ventral surface yellowish green.
Ecology/Ethology
Mabuya rugifera is a very shy, quite rarely encoun-
tered inhabitant of primary forests. It appears to be
more common in the lowlands although it has been
recorded from up to 1200 m a.s.l. in the Tengger
Mountains of Java. It is usually found along the
banks of smaller streams, on clearings, or on paths.
Distribution
Nicobars, Malay Peninsula, Borneo, Sumatra,
Pulau Nias, Mentawai Islands, Java. Mt. Kinabalu:
Poring.

Genus Riopa GRAY, 1839
The distribution of the genus Riopa extends from
continental Asia through the Malay Peninsula and
Greater Sunda Islands to Sulawesi and the Philip-
pines.
Slender, elongate ground-dwellers with short to
rudimentary limbs, adpressed limbs do not over-
lap or even meet. Supranasals present (vs absent in
similar-looking species of Sphenomorphus); lower
Tab. 33 Distribution of Riopa-species on Borneo.
Key to the Bornean species of Riopa
1 More than 37 scale rows around mid-body; a large
species, SVL of adults greater than 100 mm ......
........................................................... bampfyldei
• Less than 33 scale rows around mid-body; a small
species, SVL of adults smaller than 60 mm .......
............................................................. bowringii
Riopa bowringii (GÜNTHER, 1864)
Eumeces bowringii GÜNTHER, 1864: 91 – Type locality: Hong-
kong
Lygosoma whiteheadi MOCQUARD, 1890a: 144 – Type locality:
Kina Balu
(syn. after DE ROOIJ 1915: 264)
Lygosoma bowringi, DE ROOIJ 1915: 264; TAN 1993: 6
Lygosoma bowringii, SMITH 1931: 31
Riopa bowringii, MANTHEY 1983: 26; MALKMUS 1994b: 246
Mochlus bowringi, EBENHARD & SJÖGREN 1984: 24
Diagnosis
Small skink up to 58 mm SVL; flanks often col-
ourful (black, yellow, and reddish) with numerous
light dots; 4 supraoculars; 28-30 scales around mid-
body; fourth toe with 10-16 subdigitals; tail thick
at base, tapering gradually.
Scincidae
eyelid covered with fine scales or with an undi-
vided “window”. Tympanum visible, small.
A terrestrial to fossorial, diurnal and crepuscular
skinks feeding on smaller invertebrates; reproduc-
tion through oviposition.
Of the two species recorded from Borneo, one
(Riopa bowringii) occurs on Mt. Kinabalu.
Remark: The validity of this genus is subject to
debate with some authors assigning the respective
species to the genus Lygosoma (see GREER 1977).
Description
SVL up to 58 mm; TL greater than SVL; lower
eyelid with small scales; ear-opening small, with
or without 1-2 anterior lobules; nostril in a quad-
rangular nasal; supranasals large, in contact behind
rostral, rarely separated; prefrontals small, widely
separated; frontoparietals separated; parietals form-
ing a suture behind interparietal; usually 1 pair of
nuchals; 4 large supraoculars; 7 supraciliars, first and
seventh largest; 6-7 supralabials, fifth largest and
below eye; 6-7 infralabials; 26-32 (usually 28-30)
smooth scales around mid-body (dorsals rarely with
3 weakly developed keels); 52-62 vertebrals be-
tween parietals and base of tail; 4 preanals slightly
enlarged; adpressed limbs do no touch; fourth toe
with 10-16 subdigitals; tail thick at base, tapering
gradually.
Dorsal surface yellowish to bronze-brown; fine
black spots present on dorsals, forming a longitu-
dinal series; black dorsolateral stripe from the eye
to the anterior quarter of the tail, often broken by
reddish and yellow zones (sometimes bordered with
yellow above); a white line along supralabials
nearly reaching the ear-opening; lateral surface of
neck and body usually mottled with numerous light
and dark spots; tail often reddish ventrolaterally;
ventral surface orange to pink. Juveniles olive
brown above.
2-4 eggs per clutch.
281
 Scincidae
© I. Das
Fig. 288 Riopa bowringii from Kota Kinabalu, Sabah.
Ecology/Ethology
The skink is more commonly found at low eleva-
tions, rarely up to 1640 m a.s.l. and is not bound
Genus Sphenomorphus FITZINGER, 1843
Species of the genus Sphenomorphus are distrib-
uted in continental Asia, the Indo-Australian Ar-
chipelago, and the Philippines.
Often difficult to distinguish, these are small to
moderately large, usually slender ground- and tree-
dwellers. 4 fully developed limbs, usually overlap-
Tab. 34 Distribution of Sphenomorphus-species on Borneo.
282
to a particular type of habitat. In
fact, it inhabits towns just as well
as cultivated areas and forests
edges. It is often seen on banks
of streams. During the day, it is
rather secretive living beneath
leaf litter, rocks, fallen logs, and
similar cover. Main activity
peaks during early hours of dusk,
but it may emerge to forage also
in overcast weather conditions.
Distribution
India (Andamans), Myanmar,
Thailand, Cambodia, Laos, Viet-
nam, southern China, Hong Kong,
Malay Peninsula, Singapore, Bor-
neo, Java, Sulawesi and the Philippines (Sulu Ar-
chipelago). Mt. Kinabalu: without precise local-
ity data.
ping when adpressed. Lower eyelids covered with
small scales. No lobules around the ear-opening.
Dorsals and ventrals usually smooth. No supra-
nasals, 2 loreals, frontoparietals and interparietal
distinct and in broad contact. Reproduction in some
species ovoviviparous, in others oviparous.
Borneo is home to 14 species, of which 6 have been
recorded from Mt. Kinabalu. One, Sphenomorphus
tenuiculus, appears to be endemic to Mt. Kinabalu.

Key to the Bornean species of Sphenomorphus (in part after BACON 1967)
1 Limbs short, adpressed limbs do not meet, tail
thick at base; dark lateral stripe present; 26-30
scale rows around mid-body; 4 supraoculars
(count ending with the scale touching both
frontoparietal and parietal); 7-12 subdigitals
beneath fourth toe .................................... alfredi
• Limbs long, adpressed limbs (sometimes nearly)
meet or overlap ................................................. 2
2 Parietals in contact with supraoculars .............. 3
• Parietals not in contact with supraoculars; 36-40
scale rows; 12-14 subdigitals; tail thick at base .
................................................................. hallieri
3 Dark dorsolateral or lateral stripe present ........ 4
• Dark dorsolateral or lateral stripe absent ....... 10
4 4 supraoculars ................................................... 5
• 5 or more supraoculars ..................................... 8
5 Less than 28 scale rows .................................... 6
• More than 28 scale rows ................................... 7
6 4 longitudinal series of dark spots, 2 on the back
and 1 along each side; 24 scale rows; 21-23
subdigitals; tail thick at base ............. buttikoferi
• Dark spots on back not forming longitudinal
series, dark flanks whitish-spotted; 26 scale rows;
21-24 subdigitals; tail somewhat thick ...............
........................................................... tenuiculus
7 More than 26 subdigitals; 32-34 scale rows .......
.............................................................. shelfordi
• Less than 20 subdigitals; 32 scale rows ..............
.......................................................... modiglianii
8 Usually 5 supraoculars (rarely 6); black dorso-
lateral stripe with small scattered yellowish spots;
lower lateral parts of neck and throat neither
greyish to light blue nor deep blue; 32-38 scale
rows; 15-17 subdigitals; tail thick at base ...........
..................................................... kinabaluensis
• Usually 6 supraoculars ..................................... 9
9 37-42 scale rows; lower lateral parts of neck and
throat greyish to light blue (females), blue (males);
lower parts of flanks yellowish to orange
(females), dark with blue spots or blue (males);
16-19 subdigitals ........................... cyanolaemus
• Less than 36 scale rows; flanks whitish with
closely packed dark spots ................ murudensis
Scincidae
10 4 supraoculars; 22-24 scale rows; 18-23 subdigi-
tals ........................................................ stellatus
• 6 or more supraoculars; more than 30 scale rows
......................................................................... 11
11 A dark spot on each side of the neck; scales in 36
rows; 18-23 subdigitals ................ maculicollus
• No dark spot on neck; scales in more than 37
rows ................................................................ 12
12 Head and body black above, with small yellow
spots; 41-42 scale rows; 16-18 subdigitals ........
................................................................... haasi
• Head and body not black above .................... 13
13 Lips strongly barred, black bars as wide as or
wider than light ones, males without blue throats;
38-42 scale rows; 18-22 subdigitals .... sabanus
• Dark bars on lips absent, or if present narrower
than light bars, males with blue throat; 42-49 scale
rows; 17-23 subdigitals .......... multisquamatus
Sphenomorphus alfredi
(BOULENGER, 1898)
Lygosoma alfredi BOULENGER, 1898: 922, pl. LV: fig. 4 – Type
locality: “Savu, North Borneo” (= Sg. Sawun? = Sg. Sawan
4°46’N 115°31’E)
Sphenomorphus alfredi, TAN 1993: 8; MALKMUS 1994b: 246
Diagnosis
A very small, short-limbed, poorly known species
of Sphenomorphus with an SVL of about 33 mm
max., flanks with dark stripes; 4 supraoculars; 26-30
scales around mid-body; fourth toe with 7-12 sub-
digitals; tail thick at base, tapering gradually.
Description
SVL 33 mm, TL 35 mm; ear-opening small, round,
about diameter of eye; nostril in 1 nasal; frontonasal
wider than long, forming a broad suture with rostral,
a very narrow suture with frontal; prefrontals large,
in contact; frontal twice as long as wide, touching
first 2 supraoculars; parietals forming a suture be-
hind interparietal; no enlarged nuchals; 4 large su-
praoculars; 8 supraciliars; fourth to sixth supralabials
below eye; 26-30 smooth scales around mid-body;
283
 Scincidae
after BOULENGER 1898
Fig. CII Sphenomorphus alfredi
no enlarged preanals; adpressed limbs not touch-
ing; fourth toe with 7-12 smooth subdigitals; tail
thick at base, tapering gradually.
Colour in alcohol (after BOULENGER 1898): “Red-
dish brown above, with darker spots on the nape;
blackish spots forming a stripe on each side of head
and body, passing through the eye; whitish be-
neath.”
Ecology/Ethology: Unknown.
Distribution
Sphenomorphus alfredi is endemic to Borneo
(Sabah). Mt. Kinabalu: Sg. Mesilau.
Fig. 289 Female of Sphenomorphus cyanolaemus from Sumatra.
© U. Manthey
284
Sphenomorphus cyanolaemus
INGER & HOSMER, 1965
Sphenomorphus cyanolaemus INGER & HOSMER, 1965: 137 –
Type locality: “Sungei Seran, Labang, Bintulu District,
fourth Division, Sarawak”
Sphenomorphus cyanolaemus, TAN 1993: 8; MALKMUS ? 1994b:
236, 246;
Lygosoma variegatum, (not Lygosoma (Hinulia) variegatum
PETERS, 1867) SMITH 1931: 24 (part.)
Diagnosis
Due to its coloration a very attractive (throat of
males bright blue), long-limbed tree skink of the
lowlands, with an SVL of up to 60 mm; flanks with
dark stripes; 6 supraoculars; 37-42 scales around
mid-body; fourth toe with 16-19 subdigitals; tail
tapering abruptly behind its base.
Description
SVL males and females up to 60 mm; TL about
80 mm, males from Sumatra up to 105 mm (ZMB
58012); ear-opening vertical-oval; nostril in 1 large
nasal; rostral wider than long, forming a broad su-
ture with frontonasal, latter wider than long;
prefrontals in broad contact with each other, addi-
tionally with frontal and first supraciliar; frontal
longer than its distance from the tip of the snout, in
 Scincidae

Two eggs (6 × 11 mm),

hatchlings SVL
25-26 mm + TL 38-
39 mm.

Ecology/Ethology
This tree-dwelling
skinks live in lowland
primary rain forests.
A female was discov-
ered among the but-
tress of a large tree
immediately adjacent
to a river in Ketambe
(Aceh, Sumatra). It
© S. Manthey

used a hole in the

leaf-covered ground
immediately next to
Fig. 290 Male of Sphenomorphus cyanolaemus from Sumatra. the trunk to hide in

contact with first 3 supraoculars; parietals in little Fig. 291 Male of Sphenomorphus cyanolaemus from San-
contact behind interparietal; no nuchals; 6 supraocu- tubong, Sarawak.
lars; 12-15 supraciliars; 7 (rarely 6 or 8) supralabials,
fourth to sixth below eye, fifth or sixth largest; 6
infralabials; 37-42 (usually 38) smooth scales
around mid-body; 78-94 ventrals between mental
and vent; 2 enlarged preanals; limbs long, adpressed
hind limb reaching the axilla; fourth toe with 16-19
weakly keeled subdigitals; tail tapering abruptly
behind its base.
Dorsal surface bronze-coloured to olive green; back
with 2 longitudinal rows of dark spots; dark dorso-
lateral stripe of varying width between eye and in-
sertion of leg (occasionally extending on the tail),
interrupted by smaller and larger, yellowish or
whitish spots; ventrolateral surface in females
bright orange, yellowish, pink, or dirty yellow (fe-
males and males), each with lighter inclusions; lat-
eral and ventral surface of head and neck bluish
grey to dirty white (females) or cobalt-blue, some-
times chest and lateral surface of body coloured
likewise (males); ventral surface dirty yellow to
yellowish green with a bluish shine (females) or
pale blue (males); ventral surface of limbs and tail
© W. Denzer

dark orange to yellow. Juveniles with similar pat-

tern, but less intense colours, dorsal surface of tail
with a reddish shine, ventral surface pearlcoloured.

285
 Scincidae
(MANTHEY & GROSSMANN 1997). DENZER (1996)
observed this species usually at heights between
2 and 5 m on shaded trees on Santubong Penin-
sula (Sarawak). DAS (1995) also reported this spe-
cies as an inhabitant of buttresses in primary rain
forests. He found specimens sleeping 30-50 cm
above the ground on leaves of young trees whose
branches overhung mountain streams.
The soft-shelled eggs of this species were found
on Sumatra in an ant heap located between the
tree buttress beneath a dense thicket of fine
roots. From collecting the eggs until hatching a
period of 65 days passed (MANTHEY & GROSSMANN
1997).
Distribution
Sumatra and Borneo. Mt. Kinabalu: Poring.
Fig. 292 Sphenomorphus kinabaluensis from Mt. Kinabalu, 1825 m.
© S. Wilson
286
Sphenomorphus kinabaluensis
(BARTLETT, 1895)
Lygosoma kinabaluensis BARTLETT, 1895a: 94 – Type locality:
“Kina Balu, N. Borneo”
Lygosoma kinabaluensis, SMITH 1931: 11, 24, 31
Sphenomorphus kinabaluensis, MANTHEY 1983: 27 (part.);
EBENHARD & SJÖGREN 1984: 22, fig. 23, 24; MALKMUS 1985:
9, 1987a: 285; ? 1989a: 196, 1991a: 33, 1992a: 131; TAN
1993: 8; MALKMUS 1994b: 236, 246, 1996c: 290
Sphenomorphus cyanolaemus, MALKMUS 1996c: 290
Lygosoma varigatum, (not Lygosoma (Hinulia) variegatum
PETERS, 1867) DE ROOIJ 1915: 196 (part.)
Diagnosis
A fairly common, small to medium sized, long-
limbed skink of the more elevated Kinabalu region
with a SVL of up to 58 mm; flanks with or without

dark dorsolateral stripes; usually 5 supraoculars;
32-38 scales around mid-body; fourth toe with 15-
17 subdigitals; tail thick at base, tapering gradu-
ally.
Description
SVL males 52-58 mm, females 45-57.4 mm; ear-
opening vertical-oval; nostril in 1 large nasal;
rostral wider than long, forming a broad suture
with frontonasal, latter wider than long; prefron-
tals in broad contact with each other or separated
by a small shield; frontal longer than its distance
from the tip of the snout, touching first 3 supraocu-
lars (third just a little); parietals in contact for a
short distance behind interparietal; no nuchals; 5
(rarely 6) supraoculars; 7 supralabials, fourth to
sixth below eye; 7 infralabials; 32-38 smooth
scales around mid-body; 83-88 ventrals between
mental and vent; 2 enlarged preanals; limbs long,
adpressed hind limb reaching fingers or elbow;
fourth toe with 15-17 subdigitals; tail thick at base,
tapering gradually.
Dorsal surface light to medium brown or dark
brown to blackish; back with several longitudinal
rows of medium brown to yellowish spots, occa-
sionally with dark brown small dots; with or with-
out a dark dorsolateral stripe from the eye to the
insertion of the leg (occasionally continuing on the
tail), interrupted by smaller and larger yellowish
or whitish spots; ventrolateral surface reddish,
weakly orange, or greenish yellow, intermixed with
yellowish to whitish spots; ventral surface of head
and body whitish yellow, throat occasionally speck-
led with dark brown; ventral surface of tail flesh-
coloured to bright red or grey, lateral surface of
tail occasionally with the same colour and a grey
spotted pattern.
1-2 eggs (7.9 × 9.8 mm to 9.4 × 13.8 mm) per clutch,
hatchlings 20.1 + 27.5 mm to 21.5 + 28.3 mm
(SVL + TL).
Ecology/Ethology
This is a terrestrial skink which appears to pre-
fer higher elevations up to approx. 2100 m a.s.l.
It lives in the closer surroundings of the Head-
quarters on grassy margins of the roads and can
also be found in park-like structures and gardens.
The population density in the Headquarters re-
Scincidae
gion has gradually diminished over the past 15
years, a probable contributing factor being the
steep-banked water drains which parallel the
roads. In these canals we did not only find nu-
merous perished snakes but also many dead S.
kinabaluensis. Outside the Headquarters region,
the skink exhibits a preference for leaf litter
(sometimes in constantly shaded areas), but also
occurs in the vicinity of streams. However, it
avoids rocks of banks and does not flee through
the water.
OTA, HIKIDA, KON & HIDAKI (1989) found an ant
nest in a fallen log in which altogether 16 eggs (1-
2 each) were half buried.
Distribution
Sphenomorphus kinabaluensis is endemic to Bor-
neo (Sabah). Mt. Kinabalu: Marei Parei, Lumu
Lumu, Sg. Silau Silau, Sg. Liwago, Kamborangoh,
Sg. Mesilau, above Poring (800 m).
Sphenomorphus multisquamatus
INGER, 1958
Sphenomorphus multisquamatus (part.) INGER, 1958: 261, fig.
43, 44 – Type locality: “Kiau (3000 feet), Mount Kina Balu,
North Borneo”
Sphenomorphus multisquamatus, EBENHARD & SJÖGREN 1984:
24; MALKMUS 1985: 11, 1992a: 131; TAN 1993: 9; MALKMUS
1994b: 246; MANTHEY & GROSSMANN 1997: 279, fig. 203
Lygosoma variegatum, (not Lygosoma (Hinulia) variegatum
PETERS, 1867) BOULENGER 1887c: 246 (part.); MOCQUARD
1890a: 133; DE ROOIJ 1915: 196 (part.); SMITH 1931: 10, 24
(part.)
Sphenomorphus cf. variegatus, MANTHEY & DENZER 1982: 17;
Sphenomorphus variegatus, (not Lygosoma (Hinulia)
variegatum PETERS, 1867) MANTHEY 1983: 27
Sphenomorphus sabanus, (not I NGER , 1958) E BENHARD &
SJÖGREN 1984: 23, fig.
Sphenomorphus sp., MANTHEY & DENZER 1982: fig. p. 16
Diagnosis
A moderately large (SVL up to 69 mm) species
of Sphenomorphus, highly adapted to running wa-
ters; males with a blue, females with a light brown-
ish throat; no dark dorsolateral stripe; 6-7 su-
praoculars; 42-49 scales around mid-body; fourth
toe with 16-23 subdigitals; tail tapering rapidly
behind the base.
287
 © A. Nöllert © U. Manthey
288

Scincidae

Description
SVL males 64 mm, females 69 mm; TL greater than
SVL; ear-opening vertical-oval; nostril in one large
nasal; rostral very narrow, forming a broad suture
with hexagonal frontonasal; prefrontals pentagonal,
in broad contact; frontal longer than its distance from
the tip of the snout, touching first 3 supraoculars;
frontoparietals and interparietal of equal length; pa-
rietals in contact with each other behind interpari-
etal or separated by a small shield; no nuchals; 6-7
supraoculars, first largest, last smallest; 12-16 su-
praciliars; 6-7 (rarely 8) supralabials, fourth to sixth
below eye, fifth and sixth largest; 5 infralabials; 42-
49 smooth scales around mid-body; 83-101 scales
between mental and vent; 2 enlarged preanals;
adpressed hind limb reaching the shoulder; fourth
toe with 16-23 roundish, smooth subdigitals; tail
tapering rapidly behind the base.
Dorsal surface dark greyish brown, with 2-4 rows
of dark spots; laterally, a pattern of lighter and
darker spots of varying sizes; lateral surface of nape
and throat of males with a blue tinge; ventral sur-
face whitish without spots.
Remarks: It is worth noting that fig. 43 (right) of
the original description by INGER is inconsistent with
the description referring to the broad contact of the
prefrontals. It is not clear though whether this fault
has been made in the sketch or in the text, or whether
the suture between the prefrontals varies in length.
Ecology/Ethology
These are diurnal, very shy stream-dwellers that live
in pairs on and between rocks along the partly tor-
rential Sg. Kipungit I. Their flight distance is a stag-
gering 10-15 metres, and when fleeing they hide in
crevices or jump into the water. They feed on minute
insects on the moss-overgrown boulders.
Distribution
Sphenomorphus multisquamatus is endemic to
Borneo. Mt. Kinabalu: Poring: Sg. Kipungit I, Sg.
Langanan; Kiau, Bundu Tuhan.
Above: Fig. 293 Male of Sphenomorphus multisqua-
matus from Sg. Kipungit I.
Below: Fig. 294 Female of Sphenomorphus multisqua-
matus from Sg. Kipungit I.
Scincidae
Sphenomorphus sabanus
INGER, 1958
Sphenomorphus sabanus I NGER , 1958 – Type locality:
Sapagaya Forest Reserve, Sandakan District, North Bor-
neo
Sphenomorphus sabanus, EBENHARD & SJÖGREN 1984: 24;
MALKMUS 1987: 286; TAN 1993: 9; MALKMUS 1994b: 246
Diagnosis
A small species of Sphenomorphus reaching a
SVL of up to 58 mm; body almost without any
pattern, no dark dorsolateral stripe, labial shields
conspicuously banded black and white; 6 (7) su-
praoculars; 38-42 scales around mid-body; 79-91
rows of ventrals between mental and vent; fourth
toe with 18-22 subdigitals; tail tapering rapidly
behind its base.
Description
SVL males up to 51 mm, females up to 58 mm;
TL > SVL; ear-opening vertical-oval; nostril in 1
nasal; rostral much wider than long, forming a
broad suture with frontonasal, the latter wider than
frontal; prefrontals in broad contact or separated;
frontal longer than its distance from the tip of the
snout, touching first 3 supraoculars; parietals in
contact with each other behind interparietal; 6-7
(usually 6) supraoculars, first largest; 14-17 su-
praciliars; 7 supralabials, fourth to sixth below eye,
sixth largest; 5-7 (usually 6) infralabials; 38-42
smooth scales around mid-body; 79-91 rows of
ventrals between mental and vent; 2 enlarged pre-
anals; adpressed limbs overlap; fourth toe with
18-22 subdigitals; tail tapering rapidly behind its
base.
Dorsal surface almost uniform, dark brown to dark
grey, with indistinct lighter spots; in males lateral
surface of neck and body often with an orange
tinge; labials barred black and white, width of the
individual bars variable; ventral surface cream-
coloured.
2-3 eggs, 10-12 mm in length (INGER 1958).
Ecology/Ethology
One specimen was collected far from a water body
from under a fallen log in primary forest above
Poring Hot Springs.
289
 Scincidae
© C. Brühl

Fig. 295 Sphenomorphus sabanus from Poring.

Distribution Description
Sphenomorphus sabanus is endemic to Borneo SVL 46 mm, TL 60.5 mm; ear-opening vertical-oval;
(Sabah). Mt. Kinabalu: Poring. nostril in 1 nasal; frontonasal wider than long, form-
ing a broad suture with rostral and a narrower one
with frontal, the latter longer than its distance from
Sphenomorphus tenuiculus the tip of the snout and touching first 2 supraoculars;
parietals in contact with each other behind interpa-
(MOCQUARD, 1890) rietal; 4 supraoculars; 8 supraciliars; 7 supralabials,

Lygosoma teniuculum MOCQUARD, 1890a: 133, pl. VIII: fig. 2,

2a – Type locality: Kina Balu Fig. CIII Sphenomorphus tenuiculus
Lygosoma teniuculum, DE ROOIJ 1915: 180; SMITH 1931: 31
Sphenomorphus teniuculum, MANTHEY 1983: 27; TAN 1993: 9;
MALKMUS 1994b: 246
Sphenomorphus tenuiculus, EBENHARD & SJÖGREN 1984: 24

Diagnosis
A small, poorly known species of Sphenomorphus
of 46 mm SVL; colour in life unknown; 4 supra-
oculars; 26 scales around mid-body; 68 rows of
ventrals between mental and vent; fourth toe with
after MOCQUARD 1890
21-23 subdigitals; tail thick at base, tapering gradu-
ally.

290

fourth to sixth below eye, seventh largest; 6
infralabials; 26 smooth scales around mid-body; 57
vertebrals between parietals and tail base; 68 rows
of ventrals between submaxillars and vent; 2 en-
larged preanals; adpressed hind limb reaching the
wrist; fourth toe with 21-23 subdigitals; tail thick at
base, tapering gradually.
Colour in alcohol (after DE ROOIJ 1915): Brown
above, anteriorly spotted dark brown; a dark dor-
solateral band beginning at the nostril, spotted
Genus Tropidophorus
DUMÉRIL & BIBRON, 1839
The genus Tropidophorus is native to the Asian
mainland, the Philippines, Borneo, and Sulawesi.
Moderately large skinks with robust, slender bodies
and fully developed limbs. Dorsal surface covered
with characteristic, strongly keeled, humped scales
(exception T. beccarii). Lower eyelid with small
scales; ear-opening large and clearly visible; nostril
always in one nasal shield; prefrontals, frontopari-
Key to the Bornean species of
Tropidophorus
1 Dorsals smooth ..................................... beccarii
• Dorsals keeled ................................................... 2
2 Dorsals with 2 keels ................................ iniquus
• Dorsals with 1 keel ........................................... 3
3 30 scale rows around mid-body ......... perplexus
• More than 30 scale rows ................................... 4
4 32 scale rows, 5 supraoculars ................ brookei
• 34 scale rows, 4 supraoculars .............. micropus
Tab. 35 Distribution of Tropidophorus-species on Borneo.
Scincidae
whitish; flanks brown, white-spotted. Ventral sur-
face yellowish.
Remark: According to own measurements the tail
length given by DE ROOIJ (1915) is incorrect.
Ecology/Ethology: Unknown.
Distribution
Sphenomorphus tenuiculus is endemic to Mt. Kina-
balu. Mt. Kinabalu: Bundu Tuhan.
etals and interparietal present, supranasals absent.
In contrast to all other skinks inhabiting Borneo these
species possess a single large preanal plate.
All species included in this genus are more or less
strongly adapted to running waters in and at which
they live. Some species are known to be live-bearers.
Five species have been recorded from Borneo, but
only T. beccarii occurs at lower altitudes on Mt.
Kinabalu.
Tropidophorus beccarii (PETERS, 1871)
Amphixestus Beccarii PETERS, 1871: 574 – Type locality:
Sarawak
Tropidophorus Beccarii MOCQUARD 1890a: 135
Tropidophorus beccarii SMITH 1931: 10, 22, 31; MANTHEY 1983:
27; TAN 1993: 9; MALKMUS 1994b: 236, 246, 1996c: 290
Norbea beccarii EBENHARD & SJÖGREN 1984: 24
Tropidophorus mocquardi BOULENGER , 1894d: 735 – Type
locality: “Kina Baloo, N. Borneo” (syn. after SMITH 1923: 777)
Tropidophorus mocquardi DE ROOIJ 1915: 276; MANTHEY 1983:
27; MALKMUS 1994b: 236, 246
Norbea mocquardi EBENHARD & SJÖGREN 1984: 24
Sphenomorphus sp. MALKMUS 1987: 286, 1989: 197
Sphenomorphus sp. n. MALKMUS 1988d: 9, fig. 4
Sphenomorphus aquaticus MALKMUS , 1991b: 23 – Type
locality: “Kipungit I, hinter den Hot Springs/Poring, am
Südosthang des Mount Kinabalu (500 m NN)” (syn. after
HIKIDA & OTA 1994)
Sphenomorphus aquaticus MALKMUS 1992a: 131
291
 Scincidae
© R. Malkmus
Fig. 296 Tropidophorus beccarii from Sg. Luidan.
Diagnosis
A large species of Tropidophorus reaching 95-
98 mm SVL. The only species of this genus on
Borneo with smooth dorsals.
Description
SVL up to 98 mm, TL > 107 mm; ear-opening
vertical-oval; head shields and body scales
smooth; frontonasal wider than long; prefrontals
in broad contact or separated; frontal substantially
longer than its distance from the tip of the snout,
touching first 2 supraoculars; frontoparietals oc-
casionally separated by a small scale; parietals
forming a broad suture behind interparietal; 4-5
Fig. 297 Anale plate of Tropidophorus beccarii.
© R. Malkmus
292
supraoculars, first largest, last small-
est; 6-7 supraciliars; 7-8 supralabials,
fifth and sixth in contact with eye; 4-
5 infralabials; 28-36 smooth scales
around mid-body; ventrals scales larg-
est, 54-56 rows between submaxillars
and anal plate; one large preanal plate;
adpressed hind limb reaching at least
up to the wrist; fourth toe with 19-21
smooth subdigitals; tail slightly com-
pressed laterally.
Dorsal surface dark brown, with
slightly lighter bands; lateral surface
of head and body with light dots or
spots sometimes arranged in bands;
chin and throat whitish, the latter with
dark stripes and spots; remainder of ventral sur-
face yellowish.
Ecology/Ethology
The diurnal T. beccarii is the most common scincid
species along mountain streams of Mt. Kinabalu up
to elevations of approx. 1000 m. It is semi-aquatic
and lives on the banks of streams, but is often seen
perched on large boulders and logs scattered in the
streams. Its flight distance of 2-3 metres is relatively
short, but if disturbed, the skink disappears into a
crevice in a flash or jumps into cascades or pools
formed by the stream. With undulating movements
and limbs attached to the body it swims swiftly to
another place in the stream where it initially pushes
only the nostrils above the water surface. Rocky cliffs
overgrown with moss offer suitable resting places
for the night. Juveniles have also been located sleep-
ing on leaves overhanging water (DAS 1995). It ap-
pears that juveniles are exclusive to small rivulets
where they can be found between rocks and leaf lit-
ter, while the adults prefer larger streams. This may
be an indication that the latter also prey upon small
lizards including their own offspring.
A female examined by BOULENGER (1894d) con-
tained 4 fully developed embryos of 30 mm SVL.
Distribution
Tropidophorus beccarii is endemic to Borneo. Mt.
Kinabalu: Poring: Sg. Kipungit II, Sg. Langanan,
Kiau: Sg. Kadamaian; Sg. Luidan, Sayap: Sg.
Kemantis, Sg. Wariu.

Varanidae
The distribution of monitor lizards extends from
Africa through southern Asia and the Indo-Aus-
tralian Archipelago to Australia, Melanesia, and
Micronesia.
These are moderately large to very large lizards
with robust limbs, strong, clawed toes, and a long
powerful tail. The elongate skulls of monitor liz-
ards are heavily ossified withstanding the pressure
exerted when swallowing large food items. A strong
dentition enables them to tear apart large pieces of
prey. Distributed over a vast range monitor lizards
have adapted to a large variety of habitats, but are
generally rather a faunal element of the lowlands.
Representatives of this family may be found in
Genus Varanus MERREM, 1820
The range of the genus Varanus is identical to that
of the family Varanidae.
Moderately large to very large, muscular lizards.
Head elongate in shape, pupil round. Tongue very
long and slim with a deeply forked tip; retractable
into a skin sheath. Juveniles vividely coloured and
revealing contrasting pattern; markings in adults
less obvious, sometimes fading beyond recognition.
The diurnal monitor lizards of Borneo are excel-
lent climbers often residing high in the trees.
Two out of the three species inhabiting Borneo have
also been recorded from lower elevations on Mt.
Kinabalu.
Tab. 36 Distribution of Varanus-species on Borneo.
Varanidae
deserts as well as in rain forests, and species such
as Varanus bengalensis and V. salvator are occa-
sionally commensals of man in larger towns. De-
pending on the individual size monitor lizards prey
upon insects, amphibians, lizards, snakes, turtles,
fish, eggs of all sorts, and mammals of varying size.
They are also known to feed on carrion.
Sexes are difficult to distinguish, but males usu-
ally grow larger. Monitor lizards are oviparous;
clutch size varies with the species and individu-
ally.
Three species are known to occur on Borneo, two
of which have also been observed on Mt. Kina-
balu.
Key to the Bornean species of Varanus
1 Nostril distinctly closer to the eye than to the tip
of the snout ........................................................ 2
• Nostril distinctly closer to the tip of the snout than
to the eye ................................. salvator salvator
2 Neck very long, about as long as the distance from
the tip of the snout to the posterior margin of the
ear-opening .......................................... rudicollis
• Neck not elongated; distinctly shorter than the
distance from the tip of the snout to the posterior
margin of the ear-opening ................... dumerilii
293
 Varanidae
Varanus dumerilii (SCHLEGEL, 1839)
Monitor Dumerilii SCHLEGEL, 1839: 78 – Type locality: Borneo
Diagnosis
Moderately large monitor lizard, total length about
130 cm. Head relatively short, snout blunt; nos-
tril oval to slit-shaped, closable, situated near the
eye; neck shorter than head, nape with large,
roundish scales slightly larger than dorsals;
laterals relatively small; dorsum with 3-5 narrow,
lighter cross lines.
Description
Slender, monitor lizards; total lengths between 120
and 135 cm, 150 cm in exceptional cases; tail longer
than SVL; head short and broad, a blunt, flat snout;
neck shorter than head (i.e. the distance from the
tip of the snout to the posterior margin of the tym-
panum); nostrils closable, oval to slit-shaped, dis-
tance from the tip of the snout twice to three times
as long as that from the eye; tympanum large,
Fig. 298 Adult specimen of Varanus dumerilii.
© W. Grossmann
294
roundish, slightly sunk; legs and toes moderately
long, the latter with strongly curved claws; 37-45
scales between the angles of the mouth across the
occiput; nuchal scales large, larger than those on
the occiput and larger than dorsals; laterals sub-
stantially smaller than dorsals; 66-102 scales
around mid-body; ventrals elongate, quadrangular,
weakly keeled or smooth; 74-91 ventrals between
the gular fold and the insertion of the hindlegs; 2,
usually very distinct and large preanal pores; tail
compressed laterally with very small lateral scales;
subcaudals substantially larger; two keeled verte-
bral scale rows forming a double crest.
Sides of head and neck uniformly brown; 2 darker
dorsolateral stripes on the head between the poste-
rior margin of the eye and the nape, usually fusing
to form a cross band; upper side of the body black-
ish to dark brown, with 3-5 narrow, lighter cross
lines, sometimes broken into series of spots; upper
side of limbs spotted light or with light rings; tail
brown, indistinctly banded light and dark; venter
uniform faded yellowish brown, occasionally with
darker spots or bands.

© A. Götzke
Fig. 299 Juvenile specimen of Varanus dumerilii from
Poring.
Juveniles with more contrasting pattern; head and
nape often orange-red to red; cross lines of the body
correspondingly coloured or whitish yellow; up-
per side of the body, lateral stripes on the head and
neck often black; lower side of the neck with black
stripes or longitudinally elongated spots; lighter
markings dominating on the flanks; tail with alter-
nating light and dark bands; dark bands slightly
wider than their light counterparts.
4-9 eggs (30 × 45 to 25 × 65 mm, growing to 33 × 50
to 28 × 68 mm during incubation; hatchlings 90-
100 + 100-130 mm (SVL+TL) [HAUSCHILD 1998].
Ecology/Ethology
Varanus dumerilii is considered strictly arboreal
living in coastal forests, but also in the rain forests
of the lowlands and in hilly country more inland.
Despite its predilection for trees it is a good swim-
mer, hunting fish and aquatic skinks in shallow
water bodies. Crabs are a preferred food item; not
only the meat is taken but the whole body includ-
ing shell after removal of the legs. Even juveniles
will take on larger scorpions, beetle larvae, spiders,
and smaller birds (ZIEGLER & BÖHME 1996).
Distribution
Myanmar, Thailand, Peninsular Malaysia, Sumatra
and Batu Islands, Pulau Bangka, Pulau Belitung,
and Borneo. Mt. Kinabalu: Poring.
Varanidae
Varanus salvator salvator
(LAURENTI, 1768)
Stellio salvator LAURENTI, 1768: 56 – Type locality: “America”
(in error)
Varanus salvator salvator, MANTHEY 1983: 27; EBENHARD &
SJÖGREN 1984: 24
Varanus salvator, EBENHARD & SJÖGREN 1984: 24; MALKMUS
1992a: 132, 1994b: 246
Diagnosis
Very large monitor lizard of massive build; total
length up to 300 cm. Nostrils roundish-oval, usu-
ally situated twice as far or farther from the eye
than from the tip of the snout; nuchal scales not or
only little larger than the dorsals; tail muscular,
laterally compressed, oar-like; dorsum with round,
yellow, often ocellate spots in 5-6 cross rows.
Description
Very large, powerful monitor lizards easily exceed-
ing 200 cm, large specimens up to 300 cm in total
length; aged specimens dorsoventrally compressed;
tail longer than SVL; head long, snout slightly
pointed; length of the neck correlated to SVL: neck
shorter than head (distance between the tip of the
snout and the posterior edge of the tympanum) in
juveniles, longer than head in old specimens; nos-
trils large, roundish to oval, close to the tip of the
snout, twice the distance or more to the eye than to
the tip of the snout; tympanum large, vertical-oval;
limbs moderately long; claws of fingers and toes
slightly curved; 48-60 scales between the angles
of the mouth across the occiput; scales on the up-
per side of the head relatively large, flat, and
smooth, larger than nuchal scales, the latter usu-
ally slightly larger than the oval-shaped, keeled
dorsals; laterals usually slightly smaller than dor-
sals; 137-181 scales around mid-body; ventrals
keeled, elongate, rectangular with round posterior
limits, larger than dorsals, in 80-95 rows between
the gular fold and the insertion of the hind legs;
usually 1 distinct preanal pore on either side, occa-
sionally 2, very rarely 4; tail laterally compressed,
very powerful, oar-like, with a double vertebral
keel; subcaudals larger than lateral scales.
Entire upper side almost black in young specimens,
fading to grey with increasing age; juveniles with
295
 Varanidae
cross rows of bright yellow spots and/or ocelli on
the dorsal sides of the body and tail; nape, limbs
and toes with numerous small, yellow spots; head,
body and tail banded with yellow and black later-
ally. Yellow markings fading with increasing age
becoming more and more indistinct; old specimens
often plain dark grey dorsally and laterally.
Ventrally bright yellow (juveniles) to faded yellow-
ish (old specimens).
10-30 eggs (34-46 × 64-100 mm); hatchlings SVL
120-132 mm + TL 165-190 mm.
Ecology/Ethology
Varanus s. salvator is encountered predominantly
near or in water with no evident predilection for
specific types of water bodies, and has even been
found swimming in the open sea. It is not uncom-
mon to encounter sizeable populations inhabiting
smaller islands without freshwater foraging in the
tidal zones (MANTHEY unpublished observations
made in Sandakan Bay). Their powerful, oar-like
tail enables them to swim and dive even in rather
strong currents. Young and semiadult specimens are
often seen climbing, while larger adults prefer to
stay on the ground. Monitor lizards do not neces-
sarily take to flight when confronted. Larger speci-
mens in particular may decide to rather face the
opponent and are able to inflict severe injuries with
Fig. 300 Adult specimen of Varanus salvator salvator from Deramakot, Sabah.
© T. Eltz
296
their tails using it like a whip. If given the chance,
they will normally flee noisily into the undergrowth
or – if encountered in the immediate vicinity of a
water body – into the water.
Males are known for their combat rituals which
include head jerk, display, circling, clinch, wres-
tling phases and even mounting the defeated oppo-
nent. Every animal which can be overpowered is
considered prey. The diet of juveniles includes,
among others, also insects and frogs while adults
may not even shy away from dwarf deer.
Females bury their eggs or deposit them in cavities
in trees; clutch size depends on age and body size
of the respective female.
Varanus s. salvator appears to be fairly rare on Mt.
Kinabalu. One specimen was observed some 550 m
from the nearest water body between Kipungit II
and Langanan (700 m alt.) (MALKMUS 1992a), an-
other in the vicinity of Poring.
Distribution
From Sri Lanka through eastern India including the
Andamans and Nicobars, to southern China
(Kwangtung) and Hainan; from the Malay Penin-
sula southeast to the islands of Flores and Sulawesi
as well as on countless offshore islands through-
out the entire range and on Borneo. Mt. Kinabalu:
Poring and vicinity.
 Serpentes (snakes)
Snakes have colonized a multitude of ecological niches. Many species inhabit shrubbery and
trees, some are terrestrial, a great number of them are rather inconspicuous and have fossorial
habits, some live in freshwater throughout their lives, a number of others even inhabit the oceans.
Many snakes are excellent climbers and ascend, without much effort it appears, even the smoothest
tree trunks. Terrestrial species often are skilled diggers and burrowers. All snakes can swim. The
species of the genus Chrysopelea are even able to execute a controlled gliding flight.
Snakes comprise a large number of specialized feeders. While some species feed on mice and
other rodents, others may be equally specialized in snails or slugs, earthworms, frogs, lizards,
insects, even termites, amphibian or reptile eggs, or bats.
Observing these animals in nature often proves rather difficult. As a consequence of their
position high up in the food chain, they have to be much more infrequent than their natu-
ral prey. In addition to this they spend most of their time concealed and leave their hiding
places usually only for purposes such as foraging or mating. Most species are crep-
uscular and can therefore only be observed during the early hours of dawn and
dusk. Due to their noiseless type of locomotion and the inability to vocalize
they hardly give away their presence.
Most encounters are therefore
pure coincidences. This is par-
ticularly true in the case of
fossorial species which are
mostly found only by turning over
rocks and rotting logs. Some species hunt by
ambushing prey and often remain motionless for days.
This habit is usually aided by perfect camou- flage.
While most snakes are oviparous, some give birth to live young. In a few
© cases, an active incubation of the eggs can be observed, with the female
G.
Vo
ge coiling around her clutch and optimizing the incubation temperature by mus-
l
cular contractions.

As far as Borneo is concerned, the 135 species recorded from there are
grouped in 10 families and 61 genera. Of these, 7 families in 40 genera and
67 species occur on Mt. Kinabalu (comp. Fig. XIII).
Using the identification key, individual specimens can easily be assigned to
a family. Allocating a certain specimen of colubrid snake to the correct genus,
however, proves far more difficult as this is the family which accommodates the
greatest number of species. In this case it is usually necessary to evaluate a combi-
nation of colour and lepidotic traits. The sexes hardly differ with regard to their appear-
ances, but ontogenetic transformations are a common phenomenon meaning that juveniles
bear different colour patterns than the adults. This has been taken into consideration in the iden-
tification key.

Background: Sg. Kipungit I © P. Hoffmann

Serpentes

Distinguishing features in snakes

postocular Fig. CV Scales of the anal region.

preocular
loreal last ventral undivided
Fig. CIV Head scales postnasal anal
of a snake. prenasal
rostral

supralabial
infralabial
rostral
rostral
supranasal undivided
mental
prefrontal divided
subcaudal
frontal infralabial

supraocular mental groove

parietal submaxillary
gular
temporal
posttemporal first ventral

aglyph
dorsal snakes with no fangs
(keeled)

opistoglyph
dorsal rear-fangs snakes
ventral (smooth) vertebral

13
12
11 proteroglyph
10 snakes with fangs mounted on a immovable maxillary
9 bone in the front of the mouth
8
7
6 8
5 9
4 10
3 11 solenoglyph
2 12 13
1 snakes with fangs mounted on a rotatable maxillary
bone, so that the fang can be folded flat against the
2 4 6 10 12 roof of the mouth
1 8 13
3 5 7 9 11

Fig. CVI Methods of counting number of scale rows. Fig. CVII Different dentitions of snakes.

298
 Serpentes

Key to the families of Bornean snakes

A B

1 Ventrals different from dorsals (A) .................. 5

• Ventrals similar or indifferent from dorsals ..... 2
2 Eyes visible, not hidden beneath scales ........... 3
• Eyes hidden beneath scales (B); head not set off
from the body, distinctly rounded ... Typhlopidae
C
3 All scales very small (C), in 80 or more rows D
around the body .......................... Acrochordidae
• Scales in less than 30 rows around the body ... 4
4 5-6 Supralabials; mental groove present (D) ......
............................................... Cylindrophiidae
• 4 Supralabials; mental groove absent (E) ...........
.................................................... Anomochilidae E
5 Loreal shield grooved; anterior teeth of upper jaw
distinctly enlarged and hinged to fold back (F) ..
F
.......................................................... Crotalidae
• Loreal shield without groove; anterior teeth of the
upper jaw not distinctly enlarged ..................... 6
6 Supralabials with pits (G) ............... Pythonidae
• Supralabials without pits .................................. 7
7 Large, rhombic interparietal shield surrounded by
4 shields of more or less equal sizes (H) ............ G
...................................................... Xenopeltidae H
• Scales of the head not like that ......................... 8
8 Anterior teeth of upper jaw distinctly enlarged and
syringe-like (I); loreal shield absent (J) ..............
............................................................. Elapidae
• Anterior teeth of upper jaw not distinctly enlarged I
or syringe-like ................................................... 9 J
9 Prefrontal shields distinctly enlarged, considerably
larger than all other shields on the head; internasals
fused with nasals; no distinct frontal shield;
distinct parietals absent (K); small body size .....
.................................................... Xenophidiidae K
• Scalation of head not like that ........ Colubridae

299
Colubridae

Colubridae
This family constitutes the group of reptiles with the greatest number of genera and species of predomi-
nantly harmless snakes. They are rather heterogenous with regard to their appearances, ecologies, and
reproductive modes. Some species possess venom glands and grooved fangs, but few pose a real threat to
an adult, healthy human being. This is why the demarcation of this family of snakes against others is for
a large part based on dentition. As a rule, colubrids have aglyphous teeth (with no grooves or canals,
“true colubrids”) or are opisthoglyphous (with grooved fangs in the posterior upper jaw, “rear fanged
snakes”). Their ventral scales are always of a different size than the scales on the upper body. Pit organs
or other thermal sensors on the snout are absent, and neither are there rudiments of limbs found in the
skeleton. The division of this family into various subfamilies, their cladistic relationships, the
intersubfamiliar demarcation, and the systematic position of particular genera still require further re-
search.

Key to the Bornean genera of the family Colubridae

A B
1 Nostrils pointed upward (A) ............................. 2
• Nostrils pointed to the sides (B) ........................ 6
2 Pupil round ................................... Opisthotropis
• Pupil vertical-oval (C) ....................................... 3
C D
3 5 supralabials ........................................ Fordonia
• More than 5 supralabials ................................... 4
4 Keeled dorsals (D) in more than 40 rows ...........
.......................................................... Homalopsis
• Dorsals in less than 40 rows .............................. 5
5 Dorsals smooth .................................... Enhydris
E F
• Dorsals keeled (D) .............................. Cerberus
6 Pupil round ....................................................... 17
• Pupil vertical-oval (C) or a vertical slit (E) ...... 8 G H
• Pupil horizontal (F) ............................................ 7
7 Ventrals buckled (G); vertebrals not enlarged ....
......................................................... Dryophiops
• Ventrals rounded; vertebrals enlarged (H) .......... I J
............................................................. Ahaetulla
8 Asymmetrical scutellation of chin without men-
tal groove (I) ....................................................... 9
• Symmetrical scutellation of chin with mental
groove (J) ......................................................... 11
9 Subcaudals divided (K), 15 (17) dorsal scale rows
.......................................................................... 10
• Subcaudals entire (L), 13 dorsal scale rows ....... K L
......................................................... Aplopeltura

300
 Colubridae

10 One pre-chinshield anterior to 2 pairs of chin-

shields (M) ........................................ Internatus M N
• No pre-chinshield anterior 3 pairs of chinshields
(N) ........................................................... Pareas
1113-15 dorsal scale rows at mid-body ..................
...................................................... Dryocalamus
• More than 15 dorsal scale rows at mid-body .... 12
12 One row of small scales between prefrontals and fron-
tals and supraocular shields (O) ......... Stoliczkia O P
• No row of small scales between prefrontals and
frontals and supraocular shields .................... 13
13 Tail extremely long; dorsals strongly keeled (P)
....................................................... Lepturophis
• Tail of normal length; not keeled or weakly keeled
dorsals ............................................................. 14
14 Anterior maxillar teeth enlarged, separated from Q R
subsequent teeth by a gap (Q) ............. Lycodon
• Anterior maxillar teeth not enlarged .............. 15
15 Eye moderately large to small ....... Stegonotus
• Eye very large ................................................. 16
16 Vertebrals enlarged (R) ............................. Boiga
• Vertebrals not enlarged ........ . Psammodynastes
S T
17 Head distinctly set off from the neck (S) ...... 18
• Head not or hardly set off from the neck ....... 26
18 Lateral scales different from dorsals, vertebrals
larger than dorsals, in 3 rows (T) .... Xenodermus
• Body covered with uniform dorsals ............... 19
19 Ventrals and subcaudals round on their sides ....
........................................................................ 20
• Ventrals and subcaudals squarish on their sides (U) U
......................................................................... 24
20 2 or more loreal shields (V) ...................... Ptyas
• 1 loreal shield (W) .......................................... 21
21 More than 170 ventral scales .......................... 22
• Less than 170 ventral scales ........................... 36
22 Dorsals in 17 rows, more than 130 subcaudals .
......................................................... Xenelaphis V W
• More than 17 rows of dorsal scales; if in 17 rows,
then less than 120 subcaudals ....................... 23
23 Colour of tail in distinct contrast to colour of body
.......................................................... Gonyosoma
• Colour of tail not in distinct contrast to colour of
body ......................................................... Elaphe

301
Colubridae

24 Vertebrals larger than dorsals (X); 13–15 dorsal

scale rows around mid-body ....... Dendrelaphis X Y
• Vertebrals not larger than dorsals; 17–19 dorsal
scale rows at mid-body ................................... 25
25 17 dorsal scale rows at mid-body .... Chrysopelea
• 19 dorsal scale rows at mid-body ...... Gonyophis
26 Loreal shield present (Y) ................................ 30
• Loreal shield absent (Z) ................................. 27
27 More than 13 dorsal scale rows ...................... 28
• 13 dorsal scale rows; prefrontal and internasal
scales as one shield (AA) ................ Calamaria
28 7 or more supralabials .................................... 29 Z AA
• Less than 7 supralabials ........... Pseudorabdion
29 More than 7 supralabials ..................... Liopeltis
• 7 supralabials ............................ Oligodon (part)
30 Supralabials in contact with eye (BB) ........... 31
• Supralabials not in contact with eye (CC) .........
...................................................... Hydrablabes
3113 dorsal scale rows at mid-body .......................
..................................................... Gongylosoma
• More than 13 dorsal scale rows at mid-body 32
BB
32 5 or 6 supralabials ........................................... 33 CC
• More than 6 supralabials ................................ 34
33 Dorsals in 15 rows at mid-body .......... Elapoidis
• Dorsals in 17 rows at mid-body .. Oligodon (part)
34 Less than 135 ventrals ................. Oreocalamus
• More than 135 ventrals ................................. 35
35 Less than 75 subcaudals ........... Oligodon (part)
• More than 75 subcaudals ................. Sibynophis DD
36 Anterior dorsal scales smaller than those at mid-
body and arranged in oblique rows ....................
.................................................. Pseudoxenodon
• Dorsal scales uniform ..................................... 37
37 Last 2 teeth of upper jaw distinctly enlarged and
separated by a distinct interspace from the anterior EE
teeth (DD); the nape may be spread to form a hood
when molested (EE) ............... Macropisthodon
• Teeth of the upper jaw without or with small
interspace; the nape cannot be spread ............ 38
38 The current knowledge of the genera Amphiesma,
Rhabdophis, and Xenochrophis does not permit
to distinguish them on the basis of external traits
alone. Please refer to the species’ diagnoses.

302

Genus Ahaetulla LINK, 1807
The genus Ahaetulla is distributed throughout the
entire Oriental region, ranging from southern China
to India and in the south to the Indo-Australian
Archipelago. It is also present on some of the Phil-
ippine islands.
Moderately large, very slender tree snakes; body
slightly compressed laterally, head elongate and dis-
tinct from the body, with a sharply defined Canthus
rostralis, tapering to a point; eyes large, with hori-
zontal pupils which contract in bright light; nostrils
situated in the posterior part of a singular nasal; 15
rows of imbricate, unkeeled dorsal scales; vertebral
Tab. 37 Distribution of Ahaetulla-species on Borneo.
Key to the Bornean species of
Ahaetulla
1 Dorsal surface usually green, as an exception
brown or grey, but always uniform; dorsal side of
head without markings; anal scute usually divided
.................................................. prasina prasina
• Dorsal surface brown or grey, with ill-defined dark
crossbars; dorsal side of head stippled or spotted
with black, i.e. marked; anal scute entire ............
............................................................. fasciolata
Ahaetulla prasina prasina (BOIE, 1827)
Dryophis prasinus BOIE, 1827: 545 – Type locality: Java
Passerita prasina, SMITH 1931: 10, 32
Ahaetulla prasina, MANTHEY 1983: 27; MALKMUS 1994b: 246;
STUEBING 1991: 336
Dryophis prasinus, EBENHARD & SJÖGREN 1984: 24
Diagnosis
Extremely slim species (almost 2 metres in length,
but only between 120 and 150 cm on average);
Colubridae
scales slightly enlarged; ventrals rounded, sometimes
with a weak lateral keel; tail long, subcaudals ar-
ranged in pairs; hemipenis short and not forked.
The species of the genus Ahaetulla are diurnal and
live chiefly in bushes and shrubs through which
they can glide with great elegance and speed. It is
only occasionally that they descend to the ground
in search of prey. Their favourite food is lizards,
but they also take frogs, snakes, birds, small mam-
mals, and even fish. As far as is known, all species
give birth to live young.
These are “false colubrids” which do not pose a
threat to man.
Eight species are known, two of which occur on
Borneo, one on Mt. Kinabalu.
usually bright green without any pattern; noted for
its extremely pointed snout.
Description
Head distinct, with sharply edged Canthus rostralis,
1 large preocular (rarely followed by a second,
smaller one); 2 postoculars; usually 2, more rarely
3 loreals; 2+2 or 2+3 temporals; 8-10 (rarely 7)
supralabials, the 4th to 6th (rarely 4th and 5th or 5th to
7th) touching the eye; 9-10 (rarely 8) infralabials;
dorsal scales in 15 rows, smooth; 189-241 ventrals;
anal scute usually divided; 141-199 divided subcau-
dals.
Upper side normally leaf- to yellowish green, but
also dirty white, light grey, yellow, orange, light
brown, fawn-coloured, or bluish green. The vari-
ation in colour appears to be, at least partly, par-
ticular to certain populations. It is therefore likely
that not all of the mentioned colour varieties oc-
cur on Borneo. Always with black and whitish,
streaks which form bands on the interstitial skin
of the anterior third of the body; ventral sides
pale green with a yellowish white stripe along
the side.
303
 Colubridae
© U. Manthey
© U. Manthey
Juvenile Bornean specimens exhibit short, ob-
liquely arranged, bright blue streaks (markings) on
the sides of the neck and the anterior half of the
body.
This species is ovoviviparous, and a litter may con-
sist of 1 to 12 live young. The offspring measure
about 40 to 43 cm.
Ecology/Ethology
Ahaetulla prasina is found in lowland areas from
sea level to about 1370 m altitude, with a predilec-
304
Fig. 303-304 Ahaetulla prasina in different colour
varities from Thailand (left) and West Malaysia (above).
tion for the regions at low elevations. It inhabits
tropical wet lowland forests and tropical wet
montane forests, but is often found in secondary
tropical forests, in tropical dry, open forests, along
forests paths and tracks, in densely vegetated
shrublands, plantations, in bushy areas, on hedges
and trees around houses, and in gardens and other
vegetated areas as well. This snake is invariably
associated with dense vegetation, from tall grasses
to shrubs and high tree foliage. It is mainly arbo-
real, although juveniles may also be found forag-
ing on the ground. Strictly diurnal and very active,
A. prasina feeds primarily on lizards and frogs, also
taking small birds, small arboreal mammals and
other snakes.
This species is mild in disposition, it opens its
mouth when threatened, but it seldom attempts to
bite. It is harmless, but its bite has been reported to
produce slight local symptoms such as inflamma-
tion at the bite site, itching and mild pain.

The snakes have an interesting threat display in
which they extend the tongue halfway and leave it
extended for as long as they feel disturbed. The
longitudinal groove before the eye on the side of
the snout makes it possible for this species to see
straight ahead and focus on its prey in stereoscopic
vision.
There are different observations about the mating
of this species. HNIZDO (2000) observed mating in
the late afternoon, lasting 90 minutes. BULIAN &
Genus Amphiesma
DUMÉRIL, BIBRON & DUMÉRIL, 1854
The genus Amphiesma is widely distributed through-
out the southern, eastern and southeastern parts of
Asia ranging from Pakistan and India eastwards to
eastern China (inclusive of Taiwan), north into the
southernmost portions of Russia and Japan, to
Sumatra in the south and Sulawesi in the southeast.
Small to moderately large, terrestrial, semiaquatic,
harmless snakes; head distinct from the body; eyes
large with round pupils; internasals widened
anteriorly, nostrils positioned laterally; dorsal scales
keeled in 15-21 rows, with or without apical pits;
anal scute divided, subcaudals arranged in pairs;
hemipenis and Sulcus spermaticus undivided;
maxillar teeth arranged in a continuous series, in-
creasing in length posteriorly, the two posteriormost
teeth may be abruptly enlarged (MALNATE 1960).
The snakes inhabit lowland and/or montane situa-
tions up to about 2000 m altitude. They live in pri-
mary as well as in secondary forests, their fringes,
and bushlands. A more or less strict adaptation to
moist habitats is noticeable by their preferred oc-
currence along rivers, lakes, ponds, waterholes, or
small rapid mountain streams. They reproduce by
laying eggs.
Tab. 38 Distribution of Amphiesma-species on Borneo.
Colubridae
SIEBERT (2000) recorded that mating started in the
early morning, lasting for 5-6 hours. Gestation takes
about 5 months (BERGMAN 1956).
Distribution
The species ranges from southern China to India,
and south into the Indo-Australian Archipelago. It
also inhabits some of the Philippine islands. It is
also widely distributed on Borneo. Mt. Kinabalu:
Kiau, Bundu Tuhan.
Thirty-seven species are known in total (DAVID et
al. 1999), 4 occur on Borneo, 2 of which on Mt.
Kinabalu.
Taxonomic comment: STUEBING (1991, 1994) re-
corded Amphiesma stolatum from Borneo and from
Mt. Kinabalu, but this obviously constitutes a misi-
dentification of another species although it is as yet
unclear which. A. stolatum is a species distributed
considerably farther to the north. STUEBING & INGER
(1999) did not list this species any more, but also
did not offer an explanation.
Key to the Bornean species of
Amphiesma
1 19 dorsal scale rows at mid-body ..................... 2
• 17 dorsal scale rows at mid-body ..................... 3
2 Snout with a broad white stripe ......... flavifrons
• No white stripe on snout ........................ petersii
3 134-156 ventrals; 52-112 subcaudals ..................
....................................................... sarawacense
• 164-166 ventrals; 112-116 subcaudals ................
.............................................................. frenatum
305
 Colubridae

Amphiesma flavifrons (BOULENGER, 1887)

Tropidonotus flavifrons BOULENGER, 1887b: 96 – Type local-
ity: Mt. Kinabalu; North Borneo
Tropidonotus flavifrons, MOCQUARD 1890a: 140, pl. 9 fig. 2 a-c;
BOULENGER 1893b: 263; HANITSCH 1900a: 70; SHELFORD 1901a:
58; DE ROOIJ 1917: 87;
Natrix flavifrons, DUNN 1923: 2; SMITH 1931: 10, 32; DE HAAS
1949: 83; EBENHARD & SJÖGREN 1984: 24
Natrix (inc. sed.) flavifrons, MANTHEY 1983: 28

© M. Schroth
Amphiesma flavifrons, STUEBING 1991: 347; MALKMUS 1994b:
246, 1996c: 293; M ANTHEY & G ROSSMANN 1997: 313;
STUEBING & INGER 1999: 168

Diagnosis Fig. 306 Portrait of Amphiesma flavifrons from Poring,

Small, slender species, easily identified by the
white stripe across prefrontals and frontal. 19 dor-
sal scales in a row at mid-body; 75 cm max.
(STUEBING & INGER 1999).
Description
One loreal; 1-2 preoculars; 2-3 postoculars; 2+3
temporals; 7-9 supralabials, the 4th and 5th or the 5th
and 6th touching the eye; 19 dorsals at mid-body,
weakly keeled; 146-157 ventrals; anal scute entire;
87-103 subcaudals, divided.
Greenish dorsally, with a vertebral row of spots; two
rows of conspicuous white spots on either side of
Sg. Kipungit I.
the body, arranged parallel or alternating, the lower
one in contact with the ventral scales; upper side of
head greenish with black pattern; prefrontals entirely
white, frontal white anteriorly; labials white, bor-
dered with black; chin and throat white; ventral side
white, with large, irregularly scattered black
blotches, more densely and more finely spotted to-
wards the tail; underside of tail black, each subcaudal
scale with a grey spot.
This species is oviparous.

Fig. 305 Amphiesma flavifrons from Poring, Sg. Kipungit I.

© A. Nöllert

306

Ecology/Ethology
Found along small, clear, flowing waters in primary
or secondary forests. The snakes are often seen
swimming with the head held high. Amphiesma
flavifrons is both di- and nocturnal. It feeds on frogs,
their eggs, and tadpoles.
Two specimens were observed at an altitude of
about 600 m a.s.l. in a rapidly flowing, clear stream.
One of these snakes was basking on a rock, the
other was submerged clinging to a fallen tree trunk
(MANTHEY & GROSSMANN 1997).
MALKMUS (1996) discovered a specimen at around
10.00 a.m. on the confluence of the two Kipungit
streams, basking some 2,5 m high on a ginger plant.
Another one was found by him at around 1.00 p.m.
on the Sg. Kadamaian, and yet another at around 3.00
p.m. on the Sg. Liudan. Both the latter quickly took
to flight into the stream, crossed over to the opposite
bank and disappeared amongst a thicket of roots.
Distribution
Amphiesma flavifrons is a Bornean endemite. Mt.
Kinabalu: Kiau; 2100 feet (HANITSCH 1900a), Sg.
Kadamaian (1000 m), Sg. Liudan (1300 m), Sg.
Kipungit I (MANTHEY unpubl.), on the confluence
of the two Kipungit streams (500 m).
Fig. 307 Amphiesma sarawacense from Sg. Liwago.
Colubridae
Amphiesma sarawacense (GÜNTHER, 1872)
Tropidonotus sarawacensis GÜNTHER, 1872: 596 (new name
for Tropidonotus maculatus PETERS, 1871) – Type locality:
Sarawak and East Indies
Tropidonotus maculatus PETERS, 1871: 575 (not T. maculatus
EDELING, 1864) – Type locality: Sarawak and East Indies
Tropidonotus sarawacensis, BOULENGER 1887: 95; MOCQUARD
1890a: 140; DE ROOIJ 1917: 85
Tropidonotus maculatus, SHELFORD 1901a: 58
Tropidonotus saravacensis (sic!), HANITSCH 1900a: 70
Natrix saravacensis (sic!), SMITH 1931: 10, 11, 32
Natrix sarawacensis, DE H AAS 1949: 80, 92; E BENHARD &
SJÖGREN 1984: 24
Amphiesma sarawacensis, MANTHEY 1983: 27; STUEBING 1991:
348; MALKMUS 1994b: 239; STUEBING & INGER 1999: 170
Amphiesma sarawakensis, MALKMUS 1994b: 246
Diagnosis
Slender with a long tail; best identified by its col-
our pattern, especially the ventral pattern is char-
acteristic. 17 rows of dorsal scales at mid-body;
maximum length 78 cm.
Description
One loreal; 1 (rarely 2) preoculars; 3 (rarely 2) posto-
culars; 2+3 or 2+2 temporals; 8 (rarely 7 or 9) su-
pralabials, the 3rd to 5th or the 4th to 6th in contact
© A. Nöllert
307
Colubridae
with the eye; 17 rows of dorsal scales at mid-body,
all strongly keeled; 134-154 ventrals; anal scute di-
vided; 52-112 subcaudals, divided (STEJNEGER 1922).
Olive-brown above, with a series of dark transverse
bars, two series of yellow, red, or orange spots or
stripes; or blackish above, with orange spots ante-
riorly; a series of black spots on each side; head
vermiculated with black; labials yellow or red with
broad black sutures; lower surface checkered with
black and yellow, sometimes with the black pre-
dominating.
Juveniles brown; head above dark grey; labials
white with black outlines.
A clutch contains 4-5 eggs.
Genus Boiga FITZINGER, 1826
The genus Boiga is distributed over a vast geographi-
cal area ranging from Iran and parts of the former
USSR in the west right through to China and Tai-
wan in the east, covering the whole of Southeast Asia,
Indonesia, the Philippines in the south, to the north-
ern portion of Australia and the Solomon Islands.
Long to moderately long, slender snakes with long
tails; body compressed laterally, head large, very
distinct from body; eyes large, the pupil a vertical
slit; enlarged grooved rear fangs present; 17-31
dorsal scale rows at mid-body; dorsals smooth, anal
scute usually entire, subcaudals arranged in pairs.
The snakes of the genus Boiga are nocturnal and
arboreal. They feed on frogs, lizards, birds and their
eggs, small mammals, and even snakes. They re-
produce by laying eggs.
Altogether 30 species are known, five of which
occur on Borneo, and four of these have also been
recorded from Mt. Kinabalu.
Tab. 39 Distribution of Boiga-species on Borneo.
308
Ecology/Ethology
This diurnal to crepuscular snake has been re-
corded from along clear streams in montane as
well as lowland regions, but is also found far away
from water. Its diet is based on frogs, their larvae
and eggs, and fish. Its biology is altogether poorly
known.
Distribution
Amphiesma sarawacense is native to Borneo and
Peninsular Malaysia. Mt. Kinabalu: Kiau, Lobang,
Kenokok, Lumu Lumu, Bundu Tuhan, 2100 feet
(HANITSCH 1900a), Headquarters, Sg. Mesilau, Sg.
Liwago, Liwago trail (1500 m).
Remark: Some authors list Boiga multomaculata
for the fauna of Borneo, but there is no evidence
that this species would occur on the island.
Key to the Bornean species of Boiga
1 23-25 dorsal scale rows at mid-body .... cynodon
• 19-21 dorsal scale rows at mid-body ............... 2
2 19 dorsal scale rows at mid-body ........ drapiezii
• 21 dorsal scale rows at mid-body ..................... 3
3 Dorsum black with yellow rings on body and tail
...................................... dendrophila annectens
• Dorsum brown or grey, no rings on the body .. 4
4 Upper side of head without markings, brown or
olive ..................................................... nigriceps
• Upper side of head with dark, light-edged
markings ............................................... jaspidea
 Colubridae

Boiga cynodon (BOIE, 1827) 248-290 ventrals; anal scute entire; 114-165 subcau-
dals, in pairs.
Dipsas cynodon BOIE, 1827: 549 – Type locality: Sumatra Dorsal coloration usually yellowish brown with a
Boiga cynodon, MALKMUS 1992a: 136, 1994b: 246 variable dark (i.e. reddish brown, dark brown, or
blackish brown) banded pattern which becomes
Diagnosis
Very large, but slender with a long head very dis-
tinct from the neck; body laterally strongly com- Fig. 309 Juvenile Boiga cynodon from West Malaysia.
pressed; easily identified on the basis of build and
colour pattern (see below); 23-25 dorsal scale rows
at mid-body. Adult specimens of 200 to 240 cm
are average, but nearly 280 cm in length may be
reached.

Description
Body strongly compressed laterally; head very
elongate; anterior teeth distinctly enlarged; 1 lo-
real; 1 large preocular; 2-3 postoculars (STEJNEGER
1922); 2+2, 2+3 or 3+3 temporals; 8-10 supralabials,
usually the 4th to 6th or the 3rd to 5th (rarely 4th -7th,
4th -5th) in contact with the eye; 23-25 dorsal scale
rows at mid-body; vertebrals distinctly enlarged;

© G. Vogel
Fig. 308 Adult Boiga cynodon from West Malaysia.

© W. Grossmann

309
 Colubridae
more densely arranged posteriorly; tail with an al-
ternating pattern of very dark and yellow colours.
Exceptional specimens may be light brownish grey
with brown bands or dark grey with blackish grey
bands. The bands are partly bordered darker. Head
with the basic colour of the dorsum, lighter in the
labial region; a black stripe between the eye and
the angle of the mouth.
Juveniles show the same type of pattern, but are
substantially lighter; upper sides beige-coloured
with reddish brown bands and blotches. There are
a number of colour varieties, but the author is not
familiar with these as far as Borneo is concerned.
Clutches of 6-12 eggs several times per year.
Ecology/Ethology
Boiga cynodon inhabits regions of low altitude and
is usually found in thick vegetation along clear-
ings and on the edges of primary and secondary
tropical wet forests, open dry forests, tropical
montane forests, in rubber plantations, and close
to villages. This rather sluggish snake is predomi-
nantly arboreal and an excellent climber, but it
may be active also on the ground. During the day,
this nocturnal species retreats in tree holes and
becomes active at dusk. Boiga cynodon is often
discovered on branches overhanging small creeks,
or even seen swimming across fairly wide rivers.
It feeds mainly on birds (including small fowl)
and their eggs, but will also take mammals, liz-
ards, and snakes.
This species is widespread, but nowhere consid-
ered common. It is not really aggressive, although
rather variable in disposition, and may strike and
Fig. 310 An alarmed Boiga cynodon from West Malaysia.
© U. Manthey
310
bite. If alarmed, it flattens its neck vertically, dis-
playing the bright yellow colour of the throat. This
opisthoglyphous snake can inflict painful bites, and
envenomations of humans with marked clinical
effects have been reported.
One specimen was caught at night in a small mam-
mal trap set in the area of the park of Poring (MALK-
MUS 1992).
Distribution
Boiga cynodon is distributed from the Malay Pe-
ninsula to the Lesser Sunda Islands. It also occurs
on the Philippines. Mt. Kinabalu: Poring (500 m).
Boiga dendrophila annectens
(BOULENGER, 1896)
Dipsadonomorphus dendrophilus var. annectens BOULENGER,
1896c: 71 – Type locality: Borneo
Boiga dendrophila ssp., MANTHEY 1983: 27
Boiga dendrophila, MANTHEY & DENZER 1982: 19; EBENHARD
& SJÖGREN 1984: 24; MALKMUS 1994b: 246
Diagnosis
Very muscular and large; body laterally strongly
compressed; readily distinguished from other spe-
cies occurring in the same region by its conspicu-
ous yellow/black banded pattern; 21 dorsal scale
rows at mid-body. B. d. annectens differs from the
other subspecies of Boiga dendrophila with regard
to the number of bands and the coloration of the
head, throat, and belly. Maximum length 250 cm.
Description
One loreal; 1 preocular; 2 postoculars; 1-3 anterior
and 1-4 posterior temporals; 8, rarely 7 or 9, supra-
labials, usually the 3rd to 5th in contact with the eye;
10-12 sublabials; 21 dorsal scale rows at mid-body;
221-253 ventrals; anal scute entire; 89-118 subcau-
dals (STEJNEGER 1922), divided.
Dorsally and laterally iridescent black with (36)
42-69 bright yellow bands on the body and 15-26
bands on the tail; most bands closed around the
body, with some almost reaching mid-belly; one to
two scale rows in width; head black above,
unicoloured, or with small yellow speckles, of
which a small spot on the prefrontals, near the su-
 Colubridae

© U. Manthey
Top: Fig. 311 Juvenile Boiga dendrophila annectens
from Poring.
Right above: Fig. 312 Portrait of the specimen from
the top.
Right below: Fig. CVIII Drawing of the head scales
from a specimen from the HQ-Collection.

ture, is most constant. Upper labials yellow, with

© U. Manthey
fairly broad black margins; throat yellow, some
scales with black tips; belly black with lateral rows
of spots (VOGEL 2000).
BOETTGER (1893) reported about an aberrant speci-
men from Pontianak with only 26 bands on the body
and 9 on the tail, 223 ventrals, and 90 subcaudals.
In general, the various subspecies of Boiga
dendrophila produce clutches of 4-15 eggs (45.5-
51 × 24.5-25 mm) several times per year. A captive
female laid three clutches of altogether 25 eggs
within one year. Newly hatched juveniles measure
340-400 mm and may exhibit a pattern of reddish
bands.
No information is available for the subspecies dealt
with here.

311
 Colubridae
Ecology/Ethology
Boiga d. annectens inhabits lowland tropical wet
forests and mangroves. It is usually found in vari-
ous flooded and forested areas, closed swamps and
marshes, along the vegetated banks of rivers, and
other water-associated areas. On Mt. Kinabalu it
was found in dipterocarp primary forest near a small
stream. This arboreal snake is an excellent climber
and is especially found in thick vegetation and on
low branches overhanging water, but it may also
be active on the ground. It enters water readily
where it proves to be a skilful swimmer. During
the day, this nocturnal species retreats into tree
holes or hides in foliage and becomes really active
only after dusk. Boiga dendrophila feeds mainly
on mammals (including bats), birds and their eggs,
lizards, frogs, other snakes, and even fish. Accord-
ing to VOGEL (2000) the following food items were
recorded for B. d. annectens: birds, lizards (one
Bronchocela cristatella), bats, and snakes (prob-
ably one Boiga cynodon).
This species is widespread and common. It is an
aggressive snake which may strike and bite vi-
ciously. Larger specimens can inflict painful bites,
and envenomation of humans with marked clinical
effects have been reported.
Fig. 313 Boiga jaspidea from Thailand.
© G. Vogel
312
The biology of this particular subspecies has not been
described in particular. It is presumably similar to
that of the other subspecies.
MANTHEY & DENZER (1982) found a specimen in the
vicinity of a river which, at the time of the observa-
tion, was resting on the ground. It subsequently ac-
cepted a Hemidactylus frenatus as food.
Distribution
This subspecies is endemic to Borneo. It is reason-
ably common and has been recorded from all parts
of the island. Mt. Kinabalu: Poring (600 m).
Boiga jaspidea
(DUMÉRIL, BIBRON & DUMÉRIL, 1854)
Triglyphodon jaspideum DUMÉRIL, BIBRON & DUMÉRIL, 1854b:
1093 – Type locality: Java
Diagnosis
Comparatively small; slender; head short, distinct,
blunt; body laterally strongly compressed; most
easily identified on the basis of its colour pattern
(described below). 21 dorsal scale rows at mid-
body. Maximum length about 150 cm.

© G. Vogel
Fig. 314 Boiga jaspidea from Thailand.
Description
Eye very large; 1 loreal; 1-2 preoculars; 2 post-
oculars; 2+2 (rarely 2+3 [S TEJNEGER 1922])
temporals; 8-9 supralabials, the 3rd to 5th touching
the eye; dorsals in 21 rows, vertebrals distinctly
enlarged, wider than long; 243-267 ventrals; anal
scute entire; 140-166 subcaudals, divided.
Dorsally brown to reddish brown, with two rows
of dark spots or bars on the sides and a reddish
grey vertebral stripe, the latter spotted with dark
brown to its sides; laterally with a series of white
spots which may encroach on the vertebrals; head
of the same colour as the body with a dark spotted
pattern, the largest spots have light margins; a dark
stripe with a light outline on the nape; ventral side
bright yellow to yellowish in its anterior portion,
fading into white towards the tail, occasionally
black anteriorly and spotted with brown posteriorly.
One clutch contained 6 eggs (38-39 × 18-19 mm).
The hatchlings measured 390-400 mm.
Ecology/Ethology
This species is found from sea level to elevations
of about 800 m and inhabits primary forests and
Colubridae
the vicinity of villages. It is found in damp and
well-watered biotopes such as flooded forests,
marshes, swamps, and along river banks. Boiga
jaspidea is arboreal and nocturnal. Its diet consists
mainly of lizards, especially geckos, but it also takes
small mammals, snakes, and birds and their eggs.
Its anti-predator behaviour includes hissing and
flattening the neck laterally while simultaneously
expanding it vertically. When seized it will bite
readily.
A female has been reported as laying its eggs in
the nests of tree-dwelling termites, 3 metres above
the ground.
Distribution
Boiga jaspidea is distributed from the Malay Pe-
ninsula south and east to Java, Borneo and Sumatra.
Mt. Kinabalu: Poring (HQ-Collection).
Boiga nigriceps (GÜNTHER, 1863)
Dipsas nigriceps GÜNTHER, 1863: 359 – Type locality: “East
Indies”
Diagnosis
Large, but fairly slender; body laterally com-
pressed; most easily identified on the basis of its
colour pattern (see below). 21 dorsal scale rows at
mid-body. Maximum length 175 cm; males larger
than females.
Description
One loreal; 1 preocular; 2 postoculars; 1+2, 2+3,
or 3+3 temporals; 8 supralabials, the 3rd to 5th in
contact with the eye; 21 rows of dorsal scales at
mid-body; vertebrals distinctly enlarged; 240-265
ventrals; anal scute entire; 137-154 subcaudals,
divided.
Dorsally uniform brown in various shades, with fine
dark speckling; vertebrally two longitudinal rows
of black spots, usually covering the first scale row
next to the vertebrals; upper side of head olive;
supralabials white or orange red to red, with the
dark coloration of the head beginning right above;
tail above somewhat darker than the body; lower
parts of head and sublabials white, or yellowish to
deep yellow; venter from behind the throat faint
313
 Colubridae
© G. Vogel

Fig. 315 Boiga nigriceps from Java.

© G. Vogel

Fig. 316 Boiga nigriceps from Thailand.

314

© G. Vogel
Fig. 317 Juvenile Boiga nigriceps from Thailand.
pinkish, ventral scales weakly speckled on their
sides, both colorations intensify towards the tail
and continue on the lower side of the tail; under-
side of tail tip dark grey corresponding to the col-
our of the speckles. In juveniles, the colour of the
head is identical to that of the body (VOGEL in prep.).
3 to 5 eggs (41-45 × 17-21 mm, 46-56 g), hatchlings
measure 390-430 mm (BULIAN 2000).
Ecology/Ethology
Boiga nigriceps inhabits tropical rain forests in both
lowland and mountainous regions. It is often found
in hilly forested areas. This snake is arboreal and
nocturnal, feeding on frogs, small mammals, liz-
ards, birds, and other snakes. It is variable in dis-
position. Larger specimens can inflict painful bites,
and at least one case of envenomation of a human
with marked clinical effects has been reported.
BULIAN (2000) reported about the reproduction of
this species in Thailand: 1.5 to 2 hour long copula-
tions took place at early dawn over a period of 2-3
weeks. The female then laid 3 to 5 eggs. After 120
to 135 days, the babies hatched. After 2 years the
animals were mature.
Another captive female laid 3 eggs (48 × 17 mm)
which hatched after an incubation period of 120-
136 days. The hatchlings measured 390-430 mm
in length. Another female of the author’s also de-
posited two clutches of 3 eggs each.
Distribution
Boiga nigriceps is distributed from the Malay Pe-
ninsula southwards to Java, Sumatra, and Borneo.
Mt. Kinabalu: Poring (HQ-Collection).
Colubridae
Genus Calamaria BOIE in BOIE, 1827
The genus Calamaria is native to the region from
Myanmar in the northwest and China in the north,
through Vietnam, Thailand and Malay Peninsula,
south to Seram, Sulawesi, and the Philippines.
Small snakes; head weakly set off from the body; body
round in cross-section; tail short; eye distinct. Rostral
rounded; prefrontals paired, fused with the internasals
and in contact with the supralabials; parietal meets
with the last supralabial; enlarged paraparietal present;
4-5 supralabials, the 2nd and 3rd, 3rd, or 3rd and 4th touch-
ing the eye; nasal single, surrounding the nostril; no
loreal. Further pholidotic traits include 0-1 preocular,
1 postocular, no anterior temporal scale, and 5
sublabials. Dorsal scales in 13 rows throughout body
length; smooth; no apical pits; anal scute entire;
subcaudals paired.
Snakes of the genus Calamaria lead fossorial lives in
tropical rainforests. During the day, they may be dis-
covered beneath rocks, rotting wood, or in leaf litter,
where they forage for food at night. Although reli-
able information is limited, some specimens appear
to travel substantial distances at night, which may be
confirmed by the numbers of snakes found killed on
roads. At least one species, Calamaria griswoldi, was
repeatedly observed active during daylight hours.
If disturbed during the day, the snakes show a pecu-
liar way of freeing themselves or fleeing by sidewin-
ding movements. With the head on the ground, the
snake may raise a loop of the body above the ground
and throw it forward. Repeating this motion rapidly,
the snake appears to be rolling or tumbling forward
and moving with much greater speed than usual. An-
other defensive tactic of these snakes when held firmly
is to push the tip of the stubby tail, which is sharply
pointed in many species, against the hand holding it
(STUEBING & INGER 1999).
Certain species appear to be fairly common in some
regions, while of others only a few specimens have
ever been found. Their diets consist of worms, in-
sects and their larvae, possibly also small lizards
and their eggs. The snakes are oviparous, with the
eggs being deposited in leaf litter.
Fifty-one species have been described to belong to
this genus, 21 of which occur on Borneo including
the 11 species of Mt. Kinabalu.
315
Colubridae
Key to the Bornean species of Calamaria
1 3rd, or 2nd and 3rd, supralabials entering orbit .... 2
• 3rd and 4th supralabials entering orbit ............. 7
2 Preocular absent ............................................... 3
• Preocular present ............................................. 5
3 Supraocular and postocular fused into a single
shield ................................................. gracillima
• Supraocular distinct from postocular .............. 4
4 Mental touching anterior chin shields ........ lowi
• Mental not touching anterior chin shields ..........
............................................................ schmidti
5 2 infralabials touching anterior chin shields ......
.......................................................... borneensis
• 3 infralabials touching anterior chin shields .... 6
6 Narrow longitudinal stripes mid-dorsally; tail
short, tapering abruptly at end ......... battersbyi
• No longitudinal stripes mid-dorsally; tail longer,
tapering in posterior half ................... melanota
7 A light longitudinal stripe on scale rows 2 and 3
bordered below by a dark stripe on first row .....
............................................................ lumholtzi
• Stripes, if present, not as above ....................... 8
8 Preocular absent ............................................... 9
• Preocular present ........................................... 10
9 Most of the dorsal scales dark brown with a light
network or yellowish with a dark network ........
.......................................................... rebentischi
• Each dorsal scale above the 2nd row dark brown
without a light network ................... s. schlegeli
10 Mental not touching the anterior chinshields ... 11
• Mental touching the anterior chinshields ...... 18
11 Dorsum with wide dark transverse crossbands of
four or more scales in width .................. bicolor
• Dorsum lacking crossbands ........................... 12
12 Ventral and dorsal coloration dark brown with a
bold, continuous, white lateral stripe ... lateralis
• Ventral coloration different from dorsal coloration
....................................................................... 13
316
13 Almost all ventrals without dark pigmentation .
........................................................................ 14
• All ventrals behind the first quarter of body with
dark pigmentation at least on their lateral tips ..
........................................................................ 16
14 Eye larger than distance eye-mouth ...................
........................................................ leucogaster
• Eye smaller than distance eye-mouth ............ 15
15 Prefrontal not touching 3rd supralabial ...............
.......................................................... s. schlegeli
• Prefrontal usually touching 3rd supralabial ........
................................................................ bicolor
16 Reduction to 4 dorsal scale rows on tail at 14 or
more subcaudals from tip ...................... everetti
• Reduction to 4 dorsal scale rows on tail 13 or
less subcaudals from tip ................................ 17
17 Tail ending in a blunt tip ................... virgulata
• Tail tapering to a point ........................ modesta
18 Belly with dark crossbands, each more than one
ventral scale wide ....................... lumbricoidea
• Belly not cross-banded or with bands narrower
than one ventral .............................................. 19
19 Eye two-thirds of eye-mouth distance ...............
............................................................ hilleniusi
• Eye equal to or greater than eye-mouth distance
........................................................................ 20
20 Dorsum with longitudinal dark and light stripes
............................................................ griswoldi
• Mid-dorsal region not striped ........................ 21
21 Tail thick, tapering abruptly near the end to a blunt
tip ............................................................ prakkei
• Tail tapering gradually .................................. 22
22 Ventrals of males less than 145 in number; of fe-
males usually less than 164 ............... suluensis
• Ventrals of males 145 or more in number, of
females usually more than 164 ....... grabowskyi
 Colubridae

Tab. 40 Distribution of Calamaria-species on Borneo.

Calamaria bicolor
DUMÉRIL, BIBRON & DUMÉRIL, 1854
Calamaria bicolor DUMÉRIL, BIBRON & DUMÉRIL, 1854a: 78 –
Type locality: Borneo
Calamaria bicolor, MOCQUARD 1890a: 135; BARTLETT 1895c:
84; SHELFORD 1901a: 62; DE ROOIJ 1917: 165; INGER & MARX
1965: 149; MANTHEY 1983: 27; STUEBING 1991: 337; MALKMUS
1994b: 246
ing the anterior chin shields; 139-169 ventrals; 18-
28 subcaudals; tail thick, ending in a blunt point.
Colour of adults uniform dark brown above, or with
obscure black bands superimposed on ground col-
our; dark pigmentation usually ending abruptly on
scale rows 2 or 3; scales of 1st and 2nd rows some-
times with black anterior corners, forming a con-
tinuous black stripe in some specimens; head dark
brown above, with or without two oblique dark

Diagnosis
Third and fourth supralabials entering orbit; Fig. CIX Calamaria bicolor
preocular present; mental not touching anterior chin
shields; paraparietals surrounded by 5 scales and
shields; prefrontal touching 3rd supralabial; no dark
squares on ventrals; nasals orientated forward; max.
total length 45 cm.

Description
Five supralabials, the 3rd and 4th entering orbit, 5th after JAN & SORDELLI 1860-66

the largest, 1st the smallest; 5 sublabials, first 3 touch-

317
 Colubridae

bands across the yellow lip; ventrals yellow, usu- shields; paraparietal surrounded by 5 scales and
ally immaculate, rarely spotted with black; tail shields; reduction to four dorsal scale rows at a point
brown above, with dark bands; underside of tail 12 or more subcaudals from the terminal scute;
yellow, with or without a dark median line. Juve- max. total length 46.8 cm.
niles have a yellow ground colour with dark
crossbands. Description
Five supralabials, 3rd and 4th entering orbit, the 5th
Distribution the largest; 5 sublabials, first 3 touching the ante-
This species is distributed on the islands of Java rior chin shields; 150-190 ventrals; 20-29 sub-
and Borneo. Mt. Kinabalu: without more precise caudals; tail long, tapering gradually to a blunt tip.
localities. Colour dark brown above; each dorsal scale with a
dark network; scattered dark brown or black spots
forming dashes, or short lines above, usually the
Calamaria grabowskyi FISCHER, 1885 first three scale rows yellow just behind the head,
the yellow area bordered above by an irregular
Calamaria grabowskyi FISCHER, 1885: 50; pl. 4 figs. 1a-e – black stripe; scales of the 1st row with white cen-
Type locality: Telang and Tameanglaijang; Dusson Timor tres and dark edges forming a dark-edged white
District, southeast Borneo stripe running the length of the body; head brown
Calamaria grabowskyi, MOCQUARD 1890a: 136; DE ROOIJ 1917:
above with dark brown or black spots; a dark brown
155; SMITH 1931: 32; INGER & MARX 1965: 129; EBENHARD
& SJÖGREN 1984: 24; MANTHEY 1983: 27; STUEBING 1991: or black stripe on the side of the head occupying
338; MALKMUS 1994b: 246 the upper third of the supralabials and usually the
Calamaria baluensis BOULENGER, 1893a: 524 – Type locality: lower edge of the prefrontal; lower two thirds of
Mount Kinabalu, North Borneo (syn. after SMITH 1931: 32) supralabials yellow, with or without dark sutures;
Calamaria baluensis, BOULENGER 1894e: 335; BARTLETT 1895c:
83; SHELFORD 1901a: 62; DE ROOIJ 1917: 155; MANTHEY 1983:
underside of head yellow, usually with small dark
27; spots anteriorly; ventrals uniform yellow or, more
Calamaria grabowskii (sic!), SHELFORD 1901a: 62 often, with varying amounts of black pigment; the
black pigment, if present, it appears a short dis-
Diagnosis tance behind the head, taking the form of median
Third and fourth supralabials entering orbit; and lateral bands consisting of transversally elon-
preocular present; mental touching anterior chin gated dark spots; as the amount of dark pigment
increases, the rows of dark spots fuse
to form a band across the anterior edge
Fig. 318 Calamaria grabowskyi from Maliau Basin, Sabah. of each ventral; tail yellow below, with
a dark median band; subcaudals of
some specimens with dark anterior
edges.

Ecology/Ethology
The species is predominantly a forest
dweller. It inhabits the lowland as well
as montane regions up to altitudes of
about 1400 metres. Further details on
its ecology are unavailable.
© R. B. Stuebing

Distribution
This species is endemic to Borneo.
Mt. Kinabalu: Sayap, Sg. Kanda-
maian, Tenompok, Kenokok (1000 m).

318

Calamaria griswoldi LOVERIDGE, 1938
Calamaria lumbricoidea griswoldi LOVERIDGE, 1938: 43 – Type
locality: Luidan River; Bundu Tuhan; Mount Kinabalu; North
Borneo (3340 feet)
Calamaria lumbricoidea griswoldi, DE HAAS 1950: 570
Calamaria griswoldi, INGER & MARX 1965: 92; MANTHEY 1983:
27; MALKMUS 1985: 10, 1987: 287, 1989: 197, 1991a: 34,
1992a: 132, 1994b: 237, 246, 1996c: 291; STUEBING 1991:
339; INGER & TAN 1996b: 70; STUEBING & INGER 1999: 130
Diagnosis
Third and fourth supralabials entering orbit;
preocular present; mental touching anterior chin
shields; colour blackish brown above with a nar-
row white line between successive scale rows; im-
maculate yellowish white below; max. total length
50.7 cm. The only Bornean species of Calamaria
with light lines on the edges of the scale rows.
Description
Five or six supralabials, the 3rd and 4th entering the
orbit, 5th the largest; 5 sublabials, first 3 touching
the anterior chin shields; both pairs of chinshields
meeting at mid-line; 155-192 ventrals; 13-18
Fig. 319 Calamaria griswoldi from the Headquarters.
Colubridae
subcaudals; tail thick, tapering from base to a sharp
point.
Colour dark brown or black above; dark portion of
scales without network; blackish brown stripes on
the central two thirds of each scale row above the
first, yellowish stripes on the edges of adjacent scale
rows; scales of the first row yellow, immaculate in
the anterior part of the body; each scale usually
with a dark spot in the posterior half of the body;
head dark brown above; supralabials yellow in the
lower two thirds; head below immaculate yellow;
an oblique bar running forward and up from the
gular region onto the rear part of the parietals;
ventrals immaculate yellow, sometimes with irregu-
lar small black spots; subcaudals yellow, usually
with a faint zig-zag line mid-ventrally.
Ecology/Ethology
A species of the submontane and montane primary
forests living beneath leaf litter and in rotting tree
stumps. In selfdefence this snake will sting with
its tail tip. Calamaria griswoldi is a common vic-
tim of the road traffic in the Headquarters area of
Kinabalu Park.
© A. Nöllert
319
 Colubridae

Taxonomic comment: See

taxonomic comment under
Hydrablabes praefrontalis.

Diagnosis
Third and fourth supra-
labials entering orbit; preo-
cular present; mental not
touching anterior chin
shields; body dark brown
dorsally and ventrally with
a continuous lateral white
stripe on the 2 nd and 3 rd
scale rows for the entire
length of the body and tail;
max. total length 29 cm.
© M. Schroth

Description
Paraparietals surrounded
by 5 scales and shields; pre-
Fig. 320 Calamaria griswoldi from Sg. Tibabar. ocular present; 5 supra-
labials, the 3rd and 4th enter-

MALKMUS (1996c) found an adult male specimen

at 11.00 a.m. on a path; another specimen was ob-
served crossing a road during sunny and windy
weather with a low level of humidity at about noon
(MALKMUS 1992a). This may indicate that the spe-
cies is at least partly diurnal.
after MOCQUARD 1890
Distribution
This species is endemic to Borneo. Mt. Kinabalu: Fig. CX Calamaria lateralis
Tenompok, Bundu Tuhan (1370 m), Kambarangoh
Road (1550-1750 m), Liwago trail (1500-1750 m),
Sg. Tibabar, Headquarters (1500-1550 m), Lumu ing the orbit, 5th the largest; 5 sublabials, first 3
Lumu (1525 m), Sg. Liudan, Mesilau (1800 m). touching anterior chin shields; 146-151 ventrals;
16-23 subcaudals; tail tapering gradually to a point
from the end of the body.
Calamaria lateralis MOCQUARD, 1890 Colour above and below dark blackish brown
with a longitudinal white line from behind the
Calamaria lateralis MOCQUARD, 1890a: 136; Pl. 8; fig. 4 a-d –
eye over the entire length of body in scale rows
Type locality: Mount Kinabalu, Borneo 2 and 3.
Calamaria lateralis MOCQUARD, 1890b: 154 – Type locality:
Mount Kinabalu, Borneo Distribution
Calamaria lateralis,BOULENGER 1894e: 342; BARTLETT 1895c: This species has been recorded from Java and Bor-
84; SHELFORD 1901a: 62; DE ROOIJ 1917: 168; SMITH 1931:
32; INGER & MARX 1965: 146; MANTHEY 1983: 27; EBENHARD neo. On Borneo it has become known so far only
& SJÖGREN 1984: 24; STUEBING 1991: 339; MALKMUS 1994b: from Mt. Kinabalu, without more precise locali-
246 ties.

320

Calamaria leucogaster BLEEKER, 1860
Calamaria leucogaster BLEEKER, 1860b: 285, 288 & 293 – Type
locality: Ampat Lawang, Sumatra
Calamaria leucogaster, BOULENGER 1894e: 341; BARTLETT
1895c: 84; SHELFORD 1901a: 62; DE ROOIJ 1917: 164; SMITH
1931: 10, 11, 32; INGER & MARX 1965: 63; fig. 16; MANTHEY
1983: 28; EBENHARD & SJÖGREN 1984: 24; STUEBING 1991:
339; MALKMUS 1994b: 247
Diagnosis
Third and fourth supralabials enter orbit; mental
not touching anterior chin shields; paraparietal sur-
rounded by 5 shields and scales; no dark pigmen-
tation on the ventrals; max. total length 22.3 cm.
The only Bornean species of Calamaria with black
half-rings on the neck and on the tail.
Description
Preocular present; 5 supralabials, the 3rd and 4th
entering orbit, the 5th the largest, 2nd the next larg-
est; 5 sublabials, first 3 touching the anterior chin
shields; 126-157 ventrals; 12-26 subcaudals; tail
tapering to a sharp point near tip.
Ground colour dark brown, dorsal scales usually
with a light or dark network; body with or without
dark stripes, the striped pattern consisting of a nar-
row dark stripe along the edges of the vertebral and
paravertebral scale rows, a wider stripe on the ad-
jacent halves of the 4th and 5th scale rows, a stripe
along the adjacent halves of the 2nd and 3rd scale
rows, and a thin dark line along the upper edge of
the 1st scale row; lower two-thirds of the 1st scale
row cream-coloured; a black collar of 2 to 5 scales
in width, separated from parietals by 1 to 3 scales
and sometimes bordered posteriorly by a narrow
yellow ring; a dark ring around the body at the
base of the tail, usually a second black ring near
its tip; head dark brown mid-dorsally with a light
streak running through the centre of the prefron-
tal, the lateral edge of the supraocular, and the
lateral portion of the parietal; a dark longitudinal
stripe along the adjacent portions of the prefron-
tals and supralabials, continuing behind eye along
the adjacent edges of the parietal and supralabial
scales; remainder of upper lip and underside of
head cream-coloured; ventrals and subcaudals
cream-coloured; tail with a dark mid-ventral stripe
in some individuals.
Colubridae
Ecology/Ethology
This species is an inhabitant of lowland rain for-
ests. Further details are unknown; see generic di-
agnosis.
Distribution
This species occurs on the islands of Borneo and
Sumatra, with a doubtful record from Java (INGER
& MARX 1965). Mt. Kinabalu: Kiau (915 m),
Tenompok.
Calamaria lumbricoidea
BOIE in BOIE, 1827
Calamaria lumbricoidea H. BOIE in F. BOIE, 1827: 540 – Type
locality: Java
Calamaria temminckii, MOCQUARD 1890a: 136
Calamaria vermiformis, SHELFORD 1901a: 61; DE ROOIJ 1917:
154; SMITH 1931: 10, 11 27, 32
Calamaria lumbricoidea lumbricoidea, LOVERIDGE 1938: 43
Calamaria lumbricoidea, INGER & MARX 1965: 75; fig. 20;
E BENHARD & S JÖGREN 1984: 24; S TUEBING 1991: 340;
MALKMUS 1994b: 247
Calamaria vermiformis vermiformis, MANTHEY 1983: 28
Diagnosis
Third and fourth supralabials entering orbit;
preocular present; mental touching anterior chin
shields; paraparietal surrounded by 4 or 5 scales and
shields; belly yellow with black crossbars wider than
the width of one ventral; max. total length 64.2 cm.
Description
Paraparietal surrounded by 4 or 5 shields and scales;
5 supralabials, the 3rd and 4th entering orbit, the 5th
the largest; 5 sublabials, first 3 touching the anterior
chin shields; 137-229 ventrals; 13-27 subcaudals; tail
thick, tapering abruptly to a sharp point near the tip.
Colour dark brown or black above, with or without
narrow, light stripes; scales without network; first
one or two scale rows yellow; ventral surface yel-
low, with or without black bands; heads of juve-
niles yellow. Juveniles of most populations have
yellow bands of one-half to two scales in width
across the dorsum.
Ecology/Ethology
This snake is found in hilly areas up to altitudes of
about 1430 m in lowland tropical forests and tropical
321
 Colubridae
© U. Manthey
Fig. 321 Calamaria lumbricoidea from Sumatra.
and subtropical montane forests; it probably also
occurs in plantations and in the vicinity of rice pad-
dies. It is a terrestrial or semifossorial, secretive
snake, mainly active at night and often found among
litter or beneath rocks, fallen trees, and decaying
vegetation. Its diet is composed mainly of worms
and insect larvae, but it is reported to also take
skinks of the genus Lygosoma.
Distribution
This species inhabits the region between the Malay
Peninsula and the islands of Java, Borneo, and
Leyte. Mt. Kinabalu: Kiau (915 m), Sg. Kenokok
(1000 m), Lumu Lumu, Tenompok (1430 m), Bundu
Tuhan (1370 m).
322
Calamaria modesta
DUMÉRIL, BIBRON & DUMÉRIL, 1854
Calamaria modesta DUMÉRIL, BIBRON & DUMÉRIL, 1854a: 74 –
Type locality: Java
Calamaria modesta, I NGER & MARX 1965: 138, fig. 36;
E BENHARD & S JÖGREN 1984: 24; S TUEBING 1991: 340;
MALKMUS 1994b: 247
Calamaria lautensis, MANTHEY 1983: 28
Diagnosis
Third and fourth supralabials entering orbit;
preocular present; mental not touching anterior
chin shields; nasals orientated laterally; no light
stripe on scale rows 2 and 3; ventrals with dark
pigmentation, at least laterally; no oblique dark

stripe behind the eye; tail tapering to a point; max.
total length 45.5 cm.
Description
Paraparietal surrounded by 5 or 6 shields and scales;
5 supralabials, the 3rd and 4th entering orbit, the 5th
the largest; 5 sublabials, first 3 touching the ante-
rior chin shields; 140-154 ventrals; 12-19 sub-
caudals; tail tapering to a point; (ventral and
subcaudal counts for Bornean populations only).
Body and tail dark brown or black above; scales
with a light network and an occasional dark longi-
tudinal central streak, or scales solid brown except
for a light network at the edges, or scales solid black
with small yellow spots scattered along body; in
some specimens the dark central streaks unite to
form short lines; scales of 1st row with or without
light centres; head black above with yellow spots,
or yellowish brown with black spots; supralabials
spotted with dark; underside of head yellowish with
dark spots; ventral coloration highly variable;
amount of dark pigment on ventrals varies from
confined to the lateral edges to covering the entire
width of the ventrals; the light pigment is confined
to a narrow median stripe or to a series of small
semicircular spots.
Distribution
This species occurs on Java, Simalue, and Borneo.
Mt. Kinabalu: Bundu Tuhan (1370 m), Tenompok
(1430 m).
Calamaria schlegeli schlegeli
DUMÉRIL, BIBRON & DUMÉRIL, 1854
Calamaria schlegeli DUMÉRIL, B IBRON & DUMÉRIL, 1854a: 81 –
Type locality: Borneo
Calamaria bicolor (non Calamaria bicolor DUMÉRIL, B IBRON
& DUMÉRIL, 1854), MOCQUARD 1890a: 135; SMITH 1931: 32
Calamaria ? leucocephala, LOVERIDGE 1938: 43
Calamaria schlegeli schlegeli, INGER & MARX 1965: 162
Calamaria schlegeli, EBENHARD & SJÖGREN 1984: 24; STUEBING
1991: 341; MALKMUS 1994b: 247
Diagnosis
Third and fourth supralabials entering orbit; men-
tal not touching anterior chin shields; distinctly
bicoloured; dark dorsal pigmentation ending
abruptly above the 1st scale row; ventrals and 1st
Colubridae
scale row without dark pigment; nasals orientated
laterally; max. total length 46 cm.
Description
Paraparietal surrounded by 5 or 6 shields and scales;
preocular present or absent; 5 supralabials, the 3rd
and 4th entering orbit, the 5th the largest; 5 sub-
labials, first 3 touching the anterior chin shields;
129-180 ventrals; 19-44 subcaudals; tail long, ta-
pering throughout its length to a slightly blunt point.
Colour dark above, light below; scales above 1st or
2nd rows without network, dark brown or black;
head yellow above and below or black above and
yellow below, or various intermediate conditions;
ventrals and first one or two scale rows immacu-
late yellow.
Ecology/Ethology
Calamaria s. schlegeli is found in hilly areas up to
about 1000 m altitude, and inhabits chiefly tropi-
cal wet forests and tropical wet montane forests; it
probably also occurs in plantations and cultivated
areas. It is a terrestrial or semifossorial, secretive
snake, mainly active at night and often found among
forest litter or under rocks, fallen logs, and clumps
of decaying vegetation. Its diet is composed mainly
of worms, slugs, insect larvae, and it also takes
small frogs.
Distribution
The nominate subspecies ranges from the Malay
Peninsula to Borneo and Sumatra. Mt. Kinabalu:
without precise locality data.
Calamaria schmidti
MARX & INGER, 1955
Calamaria schmidti MARX & INGER, 1955: 197; fig. 27 – Type
locality: Bundu Tuhan, Mount Kinabalu, North Borneo
Calamaria schmidti, INGER & MARX 1965: 74; fig. 1; MANTHEY
1983: 28; EBENHARD & SJÖGREN 1984: 24; MALKMUS 1989:
197, 1994b: 238, 247, 1994d: 111, 1996c: 327; STUEBING
1991: 341; MANTHEY & GROSSMANN 1997: 313
Diagnosis
Four supralabials, the 2nd and 3rd entering orbit;
max. total length 28 cm.
323
 Colubridae
© M. Schroth
Fig. 322 Calamaria schmidti from the Headquarters.
Description
Paraparietal surrounded by 6 shields and scales;
preocular absent; 4 supralabials, the 2nd and 3rd en-
tering orbit, the 4th the longest, rarely the 4th enters
the orbit, too (MALKMUS 1994d); 5 sublabials, the
first 3 touching the anterior chin shields; 127-150
ventrals; 14-22 subcaudals; tail ending in a blunt
point. Only two specimens are known.
Upper side and flanks uniform black, with an in-
tense iridescence in blue, green, and shades of
golden. The latter colours are not created by pig-
mentation, but are spectral colours whose metallic
appearance is caused by the reflection of light. All
scales outlined with a fine white margin; lower parts
light grey, gradually darkening distally; mental and
anterior halves of the first pair of sublabials black-
ish giving the tip of the chin a sooty appearance.
The scale cover is silky, supple, and very smooth.
Ecology/Ethology
This species leaves its hiding place only at night or
after heavy rainfalls. Otherwise it can be found
beneath bark, in the humus of decaying logs, and
in leaf litter.
MALKMUS (1994d) came across a specimen at a tem-
perature of +19°C and relatively dry weather at
324
around 9.00 p.m. It was busy
crossing a road that cut through
a submontane Fagacea primary
forest with very structured ter-
rain. Another specimen was
found in the vicinity of the Head-
quarters of Kinabalu Park, and
yet another one was seen cross-
ing a road at 11.00 p.m. (MALK-
MUS 1996c).
One specimen had an earthworm
in its stomach, another fed on
earthworms in captivity.
Distribution
This species is an endemite of
Mt. Kinabalu. Mt. Kinabalu:
Bundu Tuhan (1370 m), Head-
quarters (1550 m), Kamborangoh
Road (1500-1600 m).
Calamaria suluensis TAYLOR, 1922
Calamaria suluensis T AYLOR, 1922: 189 – Type locality:
Cagayan Sulu, Philippine Islands
Calamaria pendlebury SMITH, 1931: 27 – Type locality: Kiau,
Mt. Kinabalu (syn. after INGER & MARX 1965:124)
Calamaria suluensis, I NGER & MARX 1965: 123, fig. 31;
E BENHARD & S JÖGREN 1984: 24; S TUEBING 1991: 341;
MALKMUS 1994b: 247
Calamaria pendleburyi, MANTHEY 1983: 28
Diagnosis
Third and fourth supralabials entering orbit;
preocular present; mental touching anterior chin
shields; paraparietal surrounded by 5 shields and
scales; each dorsal scale with a light network; a
continuous light stripe on the scales of the first row
only; no dark-edged saddles across the anterior part
of back; tail tapering gradually; max. total length
29,5 cm.
Description
Paraparietal surrounded by 5 shields and scales; 5
supralabials, the 3rd and 4th entering orbit, the 5th
the largest; 5 sublabials, first 3 touching the ante-
rior chin shields; 129-168 ventrals; 16-26 sub-
caudals; tail tapering gradually to a blunt tip.

Colour brown above, each dorsal scale above the
1st row with a fine dark network; scattered scales
with dark central spots forming dashes or short
lines; first 2 or 3 scale rows white in the anterior
portion; scales of 1st row with light centres, form-
ing a continuous white stripe right to the end of the
body; head dark brown above with scattered dark
spots; a dark stripe along the upper edges of the
supralabials, remainder of supralabials yellowish,
with or without dark sutures; head yellow, with
small dark spots on the labial sutures; ventrals usu-
ally uniform yellow with dark lateral corners, some
specimens with small dark spots in a broad series
down the centre of the belly; underside of tail yel-
low, with or without a dark median line.
Distribution
This species is endemic to Borneo and Cagayan
Sulu Island. Mt. Kinabalu: Sayap, Kiau (915 m),
Kenokok (1000 m), Bundu Tuhan (1370 m),
Tenompok (1430 m), Headquarters.
Calamaria virgulata BOIE in BOIE, 1827
Calamaria virgulata H. BOIE in F. BOIE, 1827: 540 – Type lo-
cality: Java
Calamaria virgulata, LIDTH DE JEUDE 1890c: 254; INGER & MARX
1965: 186, fig. 52; EBENHARD & SJÖGREN 1984: 24; STUEBING
1991: 341; MALKMUS 1994b: 247
See INGER & MARX (1965: 187) about the confusions between
Calamaria virgulata and C. modesta, and the synonymy with
C. modesta of most specimens referred to as C. virgulata in
the literature since BOULENGER (1894e).
Calamaria brachyura BOULENGER, 1895b: 481 – Type local-
ity: Mount Kinabalu, North Borneo (syn. after INGER & MARX
1965: 186)
Calamaria brachyura, SHELFORD 1901a: 62; DE ROOIJ 1917: 167;
SMITH 1931: 32; MARX & INGER 1955: 192; MANTHEY 1983: 27
Diagnosis
Third and fourth supralabials entering orbit;
preocular present; mental touching, or not
touching, the anterior chin shields; paraparietal
surrounded by 5 or 6 shields; max. total length
37,4 cm.
Description
Paraparietal surrounded by 5 or 6 shields and scales;
5 supralabials, the 3rd and 4th entering the orbit, the
Colubridae
5th the largest; 2nd the second largest; 5 sublabials,
first 3 touching the anterior chin shields; 160-260
ventrals; 8-30 subcaudals; tail thick, tapering
abruptly to a blunt tip at its end.
Colour dark brown above, each scale with a light
network, with or without longitudinal dark stripes;
a yellow ring of 1 to 3 scales in width on the oc-
ciput, or 4 to 8 scales behind the head, or absent;
dark longitudinal stripes between 1st and 2nd scale
rows, on 3rd scale row, between 4th and 5th scale
rows, and on 6th scale row; some specimens with
stripes on 3rd, 5th, and vertebral scale rows, some
without longitudinal stripes; an interrupted yellow
ring present or absent at the base of the tail; usu-
ally a yellowish spot at the tip of the tail; head dark
brown above and on the sides, upper lip usually
yellowish; underside of head yellowish, with or
without brown spots; ventrals whitish to brownish
black; dark pigment limited to the lateral corners
of the ventrals in some specimens, in others ex-
tending across the anterior margins of the ventrals;
in still others it covers all except the posterior edges
of the ventrals; underside of tail with the same pat-
tern as the belly except that a median dark stripe is
usually present.
Distribution
This species occurs on Sumatra, Nias, the Riau
Archipelago, Borneo, Java, Sulawesi, and the
southern Philippine islands. Mt. Kinabalu: Sayap,
Kenokok (1000 m), Bundu Tuhan, Sg. Luidan near
Bundu Tuhan, Mesilau.
Genus Chrysopelea SCHLEGEL, 1826
The distribution of the genus Chrysopelea extends
from India and Sri Lanka in the west, southern
China in the north, to the Philippines, Sulawesi,
Timor, Aru, and the Moluccan Islands in the east
and south.
Small to moderately large, elegant; body slender,
tail long, head slightly depressed and distinct from
body, with large eyes and round pupils; dorsal scales
with apical pits; ventrals keeled and notched, the
last one usually divided; anal scute and subcaudals
also divided.
325
Colubridae
The snakes of the genus Chrysopelea are “false
colubrids” (i.e. with an opistoglyphous dentition),
but their venom does usually not cause severe ef-
fects in humans.
The ventral and subcaudal scales are keeled later-
ally, thus enabling the snakes to climb with great
agility on vertical tree trunks. Some species are able
to “fly”, i.e. they can take to a descending (para-
)gliding flight with the ribs spread widely so that
the underside is shaped to work like a parachute.
The species of this genus are exclusively diurnal
and arboreal. They feed on lizards, frogs, small
mammals, birds, and snakes. They are oviparous.
Altogether 5 species are currently recognized, 2 of
which occur on Borneo, and both of the latter are
also present on Mt. Kinabalu.
Tab. 41 Distribution of Chrysopelea-species on Borneo.
Chrysopelea paradisi paradisi
BOIE in BOIE, 1827
Chrysopelea paradisi H. BOIE in F. BOIE, 1827: 547 – Type
locality: Java
Chrysopelea paradisi paradisi, MERTENS 1968: 202
Diagnosis
A slender, colourful tree snake; dorsal scales
weakly keeled,with apical pits; 198-239 ventrals,
the last of which divided; max. total length 120 cm,
but most specimens are only between 60 and 80 cm
long.
Description
One loreal; 1 preocular; 2 postoculars; occasion-
ally the loreal and the prefrontal shields are fused;
8-10 supralabials, the 4th to 6th, or the 5th and 6th, in
contact with the eye; 17 rows of dorsals at mid-
body, keeled, with apical pits; 198-239 ventrals,
the last one divided; anal scute divided; 106-149
subcaudals, divided.
Blackish green above; dorsals black with green
spots; usually with a flower-like pattern on the back,
326
Key to the Bornean species of
Chrysopelea
1 Belly smoky grey between the keels, sharply
contrasting with the yellow or white flanks, mid-
dorsal region red or orange with pairs of narrow
black bands which each frame a yellow one; 181-
201 ventrals ............................................... pelias
• Belly yellowish green; a longitudinal row of dor-
sal spots which cover 4 scales each may be present;
scales otherwise black with a light green or
yellowish green, pointed central spots; a banded
pattern only in juveniles; 198-239 ventrals .........
................................................ paradisi paradisi
each “flower” consisting of 4 deep red or orange-
coloured scales, but such pattern may also be ab-
sent; upper side of head blackish green with sev-
eral light or reddish bands; ventral side yellowish
green; ventrals typically with black margins
(MERTENS 1968).
A clutch containing up to 8 white eggs, 35-40 ×
10 mm (LIM & LEE 1989).
Ecology/Ethology
Chrysopelea paradisi paradisi is found from sea
level to elevations above 1300 m. It inhabits tropi-
cal wet forests, especially their clearings and edges,
tropical dry forests, wet and dry montane forests,
mangrove forests, open bushy areas, plantations,
and cultivated areas. It is also often encountered in
gardens and close to human habitations. This spe-
cies is arboreal, entirely diurnal, and very active. It
is an outstanding climber, living in tree foliage,
bushes, and basically any type of tall and thick veg-
etation, where it progresses or glides with incred-
ibly fast, elegant movements. This species feeds
chiefly on lizards, especially agamids and gekko-
nids, but also on frogs, small mammals including
bats, and birds.
 Colubridae

© W. Grossmann
Fig. 323 Chrysopelea paradisi paradisi from Thailand.

Chrysopelea p. paradisi is rather aggressive, strik- grey area along the mid-line of the ventrals being
ing and biting viciously when handled. At least one particularly diagnostic.
case of mild envenomation has been reported. This species is oviparous.

Distribution
This subspecies is native to the region between Fig. 324 Chrysopelea pelias from Sg. Kipungit I.
Myanmar and Thailand in the north and Java,
Sumatra, and Borneo in the south. Mt. Kinabalu:
Poring (HQ-Collection).

Chrysopelea pelias (LINNAEUS, 1758)

Coluber pelias LINNAEUS, 1758: 224 – Type locality: “In Indiis”
Chrysopelea chrysochlora, MALKMUS 1985: 10
Chrysopelea pelias, MALKMUS 1987: 287, 1994b: 247, 1996c:
291; STUEBING 1991: 342

Diagnosis
Small, slender, max. length about 75 cm, but usu-
© R. Malkmus

ally less than 65 cm;. 181-201 ventrals, the last one

usually divided; mostly easily identified on the
basis of its coloration with the sharply demarcated

327
 Colubridae

(STEJNEGER 1922); 17
rows of dorsals at
mid-body; 181-201
ventrals, the last one
often divided; anal
scute divided; 85-120
subcaudals, divided.
Upper side of head
with 2 or 3 red trans-
verse bars; dorsum
deep red to orange
© A. Nöllert

red with dark-edged,

light (i.e. white,
bluish white, dirty
white, or yellowish
brown) transverse lines which do not reach down
to touch the ventrals and which become more ob-
scure posteriorly; sides of body brownish to brown-
ish grey with small, whitish blue or yellowish spots;
chin and throat yellowish white to greyish white;
ventral side olive grey to smoke-grey between the
fine black ventral keels, sharply delimited against
the lemon-yellow stripe on the lateral edges of the
ventrals (MERTENS 1968).
© M. Schroth

Fig. 325 a. 326 Chrysopelea pelias from Poring.
Description
One loreal; 1 preocular; 2 postoculars; 9 supralabials,
the 4th and 5th, or the 4th to 6th, in contact with the
eye; 10-11 sublabials; 2+2 or 2+3 temporals
Fig. CXI Chrysopelea pelias from Sg. Kipungit I.
Ecology/Ethology
Chrysopelea pelias inhabits lowland tropical wet
forests and plantations up to 600 m altitude. This
snake is arboreal, diurnal, and active among foli-
age and bushes, but also on the ground in thick
vegetation. It feeds chiefly on lizards and frogs.
M ALKMUS (1987) found a specimen at around
11.00 a.m. which was just emerging for basking
from a cavity in an entanglement in a stream con-
sisting of branches, logs, rocks, and gravel. This
biotope was syntopically inhabited by Sphenomor-
phus sp. and Mabuya rudis. Lizards were accepted
as food in captivity which were seized in rapid
motion either at mid-body or behind the head.
Larger lizards were also secured with a body coil.
Frogs were left undisturbed.

Distribution
Chrysopelea pelias is distributed from Malay Pe-
ninsula to Borneo, Sumatra, and Java. Although
widespread it is considered rare throughout its
range. Mt. Kinabalu: above the Kipungit water-
fall (600 m); Sg. Mamut, Poring (500 m).

328

Genus Dendrelaphis BOULENGER, 1890
The genus Dendrelaphis inhabits large parts of Asia
and Australia.
Moderately large, very slender; tail long; head elon-
gated, distinct from the body; eyes large, pupils
round; 13-15 dorsal scale rows at mid-body, oblique,
imbricate; vertebrals enlarged; ventrals keeled lat-
erally, notched; anal scute divided; subcaudals in
pairs, keeled.
The diurnal snakes are arboreal and can be found
in primary forests as well as in cultivated areas.
They feed primarily on amphibians and lizards. All
species are oviparous,
About 20 species are known, of which 4 live on
Borneo including 2 on Mt. Kinabalu.
Taxonomic comment: STUEBING & INGER (1999)
present under the name of Dendrelaphis formosus
two photographs which undoubtedly show Dendre-
laphis cyanochloris. If the photographed specimen
Tab. 42 Distribution of Dendrelaphis-species on Borneo.
Dendrelaphis caudolineatus caudolineatus
(GRAY, 1834)
Ahaetulla caudolineata GRAY, 1834 (in 1832b-1835): pl. 81 –
Type locality: Not given; subsequently designated as
“Malayische Halbinsel” by MEISE & HENNIG (1932: 281)
Dendrelaphis caudolineatus, LOVERIDGE 1938: 43; STUEBING
1991: 342; MALKMUS 1994b: 247
Ahaetulla caudolineata, EBENHARD & SJÖGREN 1984: 24
Diagnosis
Readily identified by the combination of a striped
pattern and the imbricate arrangement of scales.
13 dorsal scale rows at mid-body; vertebrals en-
larged. A record length of 180 cm is known, but
140 cm is a normal adult size.
Colubridae
came in fact from Borneo, D. cyanochloris would
be confirmed as another species for the island.
Unfortunately no locality data was disclosed for
the photographs.
Key to the Bornean species of
Dendrelaphis
1 13 dorsal scale rows at mid-body ........................
............................ caudolineatus caudolineatus
• 15 dorsal scale rows at mid-body ...................... 2
2 A light lateral streak, bordered above and below
by black streaks .............................. pictus pictus
• No lateral streak ................................................. 3
3 More than 170 ventrals; eye very large; the anterior
part of the body red .............................. formosus
• Less than 170 ventrals; eye moderately large; the
anterior part of the body yellow ............ striatus
Description
Snout blunt; 1 preocular; 2 postoculars; 1 narrow
loreal; 2+2 temporals; 9 (rarely 10) supralabials,
the 5th and 6th, more rarely the 4th to 6th touching the
eye; dorsals large, obliquely arranged in only 13
rows; vertebrals enlarged; 163-190 ventrals, keeled
and notched; anal scute divided; 91-170 subcaudals,
divided.
Brownish or greyish yellow dorsally, with black
stripes composed of the black edges of the scales;
a yellow lateral stripe framed by two black stripes,
the lower one running along the outer edges of the
ventrals; upper side of head nearly uniform bronze
to brown; occasionally a black streak from the rear
edge of the eye over the nape onto the back where
it fades out; lips yellow; ventral side yellow; tail
329
 Colubridae
© U. Manthey
Fig. 327 Dendrelaphis caudolineatus caudolineatus from Pulau Langkawi, Peninsular Malaysia.
striped; a median dark line along the underside of
the tail.
Clutches of 5-8 eggs several times per year.
Hatchlings measured ca. 340 mm in length.
Ecology/Ethology
Dendrelaphis c. caudolineatus is found from sea
level to elevations of more than 1000 m with a
marked predilection for hilly areas. It inhabits low-
land tropical wet and dry forests, tropical open
montane forests, clearings and edges, bushy areas,
plantations, cultivated areas, gardens, and is also
found in the vicinity of houses. This agile active
species is arboreal and diurnal. It lives in tree foli-
age, in bushes, shrubs, and other dense vegetation.
It feeds chiefly on lizards, especially agamids and
gekkonids, and on frogs.
It is rather aggressive, biting readily, but is abso-
lutely harmless.
Distribution
Dendrelaphis c. caudolineatus is widespread and
common throughout its range. It occurs from the
330
Malay Peninsula south to Sumatra, Borneo, and Java,
and also inhabits the Philippines. Mt. Kinabalu: with-
out exact locality data.
Dendrelaphis pictus pictus
(GMELIN, 1789)
Coluber pictus GMELIN, 1789: 1116 – Type locality: Not given;
subsequently designated as Java by BOIE (1827: 530); later
corrected to Borneo by MERTENS (1930: 302) and LEVITON
(1968: 374)
Dendrelaphis pictus, LOVERIDGE 1938: 43; STUEBING 1991: 342;
MALKMUS 1994b: 247
Dendrelaphis cf. pictus, MALKMUS 1992a: 136
Diagnosis
Very slender; easily recognized on the basis of its
lateral stripe, bordered above and below with black,
and the radiant blue interstitial skin which becomes
well visible when the snake inflates its throat and
body. Dorsals imbricate, in 15 rows at mid-body;
vertebrals enlarged; max. total length 125 cm, but
 Colubridae

© U. Manthey
Fig. 328 Dendrelaphis pictus pictus from Sumatra.

usually only about 100 cm long; females substan- our traits as the adults. Incubation takes 85-126
tially larger than males. days.

Description Ecology/Ethology
Males with relatively larger eyes than females. 1 This snake is found from sea level to elevations of
preocular; 2 postoculars; 1 long loreal; 2+2, 1+1 more than 1350 m. It inhabits open lowland tropical
or 1+2 temporals; 7-10 supralabials, the 5th and 6th wet and dry forests, tropical and subtropical montane
(rarely the 4th ) in contact with the eye; dorsals in forests, especially clearings and edges, and also scrub-
15 rows at mid-body; vertebrals enlarged; 163-200
ventrals, keeled; anal scute divided; 112-169
subcaudals, divided and keeled. Fig. 329 Dendrelaphis pictus pictus from Thailand.
Bronze-coloured above; a yellowish lateral stripe
up to the tail, bordered with black above and be-
low; head brown above, laterally with a black stripe
from nasal to nape, behind which it disintegrates
into spots with blue, more rarely greenish blue,
spots in between; upper lips yellow; ventral side
whitish yellow to greenish yellow; tail uniform
bronze-coloured. In selfdefence the snakes inflate
to present the bright blue interstitial skin.
© U. Manthey

Several clutches per year of 3-8 eggs (22-38.5 ×

8.5-11 mm, 0.96-2.72 g); hatchlings measure
202-303 mm in length and possess the same col-

331
Colubridae
lands, open swamps, plantations, cultivated areas, the
vicinity of rice paddies, gardens, and the vicinity of
houses. Dendrelaphis p. pictus often lives close to
rivers, ponds, and in other watered environments. This
agile and active species is chiefly arboreal, being most
at home in tree foliage, in thick bushes and other
densely vegetated areas through which it glides
swiftly, although it often hunts its prey on the ground.
This snake is strictly diurnal and fond of basking. It
appears to feed nearly exclusively on terrestrial and
arboreal frogs, but lizards have also been reported.
When threatened these animals display a defence
behaviour which is primarily based on colour. They
Genus Elaphe
FITZINGER in WAGLER, 1833
The genus Elaphe is very widely distributed, rang-
ing through north and southcentral America and
large parts of Europe. It is furthermore present
throughout the tropical and temperate regions of
Asia. Various authors consider the neotropical spe-
cies as belonging to separate genera.
Large to moderately large species with slender,
muscular bodies. Head flat, elongate, slightly set
off from the body; eyes moderately large with round
pupils; 2, rarely 3, postoculars; loreal present; ven-
trals keeled; anal scute divided or entire; subcau-
dals arranged in two rows.
The largely crepuscular to nocturnal, occasionally
nervous rat snakes often display an aggressive de-
fence behaviour and are very agile. The various spe-
cies inhabit a variety of habitats, ranging from plan-
tations to primary rain forests, the vertical distribu-
tion extends from sea-level up to altitudes of above
2000 m a.s.l. Their diet consists of small mammals,
birds, reptiles, amphibians, fish, and even insects.
All species of Elaphe are oviparous.
Tab. 43 Distribution of Elaphe-species on Borneo.
332
enlarge the surface of the anterior body by later-
ally flattening the body and showing the blue in-
terstitial skin.
This beautiful species is widespread and one of the
most common snakes in tropical Asia. It is very
docile, but may bite readily and viciously when
handled; it is entirely harmless.
Distribution
The distribution ranges from India in the west to
Hong Kong and the Philippines in the east while
stretching to Sulawesi in the south. Mt. Kinabalu:
Sg. Langanan (700 m), Lumu Lumu.
Depending on which generic concept is applied, the
genus Elaphe comprises 24 to 40 species. Borneo is
home to three species, two of which occur on Mt.
Kinabalu.
Taxonomic comment: Certain authors list Elaphe
erythrura as native to Borneo, but this is evidently
based on a confusion with Elaphe flavolineata. Ac-
cording to SCHULZ (1996) Elaphe erythrura does not
occur on Borneo. In a recent review (HELFENBERGER
2001) the genus was split. Due to several reasons
we here keep the conservative taxonomie.
Key to the Bornean species of Elaphe
1 Anal scute divided; 2 supralabials entering orbit
.......................................... taeniura grabowskyi
• Anal scute single; 3 supralabials entering orbit ..
............................................................................ 2
2 Dorsal scales strongly keeled, lateral scales
weekly keeled, no black crossband on the nape;
supralabials 4-6 entering orbit ........ flavolineata
• Dorsal scales weekly keeled, lateral scales smooth,
a black crossband on the nape; supralabials 3-5
entering orbit ............................................ radiata

Elaphe flavolineata (SCHLEGEL, 1837)
Coluber flavolineatus SCHLEGEL , 1837: 141 (as young of
Coluber melanurus) – Type locality: Java and Sumatra, by
implication
Elaphe flavolineata, SMITH 1931: 10; MANTHEY 1983: 28;
E BENHARD & S JÖGREN 1984: 24; S TUEBING 1991: 343;
MALKMUS 1994b: 247; SCHULZ 1996: 125
Elaphe melanura, LOVERIDGE 1938: 43
Elaphe erythrura, S MITH 1931: 32; MANTHEY 1983: 28;
MALKMUS 1994b: 247
Elaphe erythrurus, STUEBING 1991: 343
Diagnosis
Moderately large to large, but relatively slender;
head only slightly set off from the body; eyes large;
ventrals keeled; the 4th to 6th supralabials in contact
with the eye; average total length about 150 cm, max.
total length 180 cm.
Description
One preocular; 2 postoculars; 2+2 or 2+3 temporals;
8-9 supraoculars, the 4th to 6th touching the eye;
10-11 sublabials; 19 dorsal scale rows, smooth or
Fig. 330 Elaphe flavolineata from Peninsular Malaysia.
Colubridae
slightly keeled on the sides, keeled along mid-dor-
sum; 198-242 ventrals, keeled; anal scute entire;
80-116 subcaudals.
Dorsum brownish olive in the anterior third of the
body; posterior third of the body uniform black
with a bluish gloss; a yellow to orange-coloured
vertebral stripe with slightly darker margins be-
gins on the nape fading posteriorly; anterior sides
of the body with usually paired, white/black or
black spots; interstitial skin with a reticulated
pattern; neck with a broad black stripe; a broad
black streak from behind the eye to the angle of
the mouth; a narrow black streak runs through the
supralabial shields; the pattern may occasionally
be rather faint. Ventral side yellowish white ante-
riorly, darker towards the tail; underside of the
tail dark grey to black. Juveniles possess the same
type of colour pattern, but display substantially
more intense colours.
Several clutches per year of up to 12 eggs each (51-
62 × 23-25.5 mm, about 21 g). Incubation takes
107-109 days.
© W. Grossmann
333
 Colubridae
Ecology/Ethology
Elaphe flavolineata is found between sea level and
elevations of at least 1370 m. It inhabits tropical wet
forests, especially their most open parts such as clear-
ings, edges, and track sides, open tropical dry for-
ests, savannas, scrub- and grasslands, plantations,
cultivated areas, rice paddies, and suburban areas. It
is often found close to human habitations, hidden
under piles of wood, and it frequently enters houses
in search of prey. This swift and rather elusive snake
is predominantly terrestrial, preferring a thick veg-
etation cover, and semi-arboreal, where it proves to
be a skilful climber. Elaphe flavolineata is diurnal,
mostly active in the early morning or late afternoon
hours. This species feeds on rodents, especially rats
and mice, and other small mammals, and on birds,
including small fowl; juveniles also take lizards and
amphibians. It is an aggressive snake, which strikes
fiercely when aroused. It bites readily and viciously,
and may inflict a relatively painful bite.
Fig. 331 Elaphe taeniura grabowskyi from the Kinabalu Park.
© R. B. Stuebing
334
Distribution
This species ranges from Vietnam through Thai-
land and the Malay Peninsula to the islands of
Borneo, Sumatra, and Java. Mt. Kinabalu: Kiau,
Bundu Tuhan (1370 m).
Elaphe taeniura grabowskyi
(FISCHER, 1885)
Elaphis grabowskyi FISCHER, 1885: 59, pl. IV, fig. 3 – Type
locality: Batu Hapu cave near Pengaron, Kalimantan, Selatan
(Southeast-Borneo), and Barabei (Southeast-Borneo), respec-
tively (3 syntypes)
Elaphe taeniura, EBENHARD & SJÖGREN 1984: 24; STUEBING
1991: 344, 1994: 933; MALKMUS 1994b: 247
Elaphe taenura (sic!) grabowskyi, MANTHEY & DENZER 1982:
14
Elaphe taeniura grabowskyi, MANTHEY 1983: 28; SCHULZ 1996:
261, pl. 35 C-D

Taxonomic comment: In the existing literature,
Batu Hapu cave is often given as the type locality
only (e.g. SCHULZ 1996) although FISCHER (1885),
in the original description, referred to several speci-
mens originating from two localities (the other one
being Barabei). As no singular holotype has been
fixed as yet, both of these have to be considered
type localities.
Diagnosis
Large, slender; head slightly set off from the body.
A characteristic trait is its dorsal stripe which is
particularly distinct in the posterior third of the
body. Anal scute divided; the 5th and 6th supralabials
in contact with the eye. This subspecies reaches
total lengths of between 170 and 190 cm.
Description
One preocular; 2, rarely 3 postoculars; 0-1 subocu-
lars; 9 supralabials; the 5th and 6th touching the eye;
10-13 sublabials; 25-27 dorsal scale rows at mid-
body, the central ones slightly keeled; 275-285
ventrals, keeled; anal scute divided; 102-114 sub-
caudals, divided.
Ground colour slate grey to greyish brown; head
coloured like the body, individual dorsal scales
with dark margins which together with the lighter
interstitial skin form a reticulated pattern; a light
vertebral stripe begins in the anterior portion of
the body and becomes more contrasting towards
the tail; the reticulated sides darken towards the
tail and eventually transform into a broad black
lateral stripe; a broad dark postocular stripe be-
hind the eye; supralabials white to yellow, occa-
sionally slightly orange; belly yellow to yellow-
ish white with a stripe running along the ventral
keels, this stripe being broken in the anterior part
of the body.
Juveniles are lighter with a more contrasting, lad-
der-like pattern.
About 15 eggs in a clutch.
Ecology/Ethology
Elaphe taeniura grabowskyi inhabits mainly re-
gions of middle to high elevations from 400 to at
least 2100 m, and is found chiefly in hilly and rug-
ged regions covered with tropical wet and dry for-
ests, tropical and subtropical montane forests, de-
Colubridae
ciduous and pine forests, and in rocky grasslands.
It lives in closed or open forests, on clearings, in
rocky and bushy areas, scrublands, plantations, on
the perimeters of cultivated areas, and also in the
vicinity of villages. It is also often found in the
vicinity and inside the numerous caves of karstic
and limestone geotypes, where it can be observed
living in absolute darkness up to several kilome-
tres from the entrance inhabiting crevices of the
wall and roof (e.g. at Gunung Api in Sarawak).
This unusual habit makes this species one of the
few true cave-dwelling snakes. This rather shy but
swift snake is basically terrestrial, occurring
mainly in moist, thick vegetation, but also has
arboreal habits, proving that it is a skilful climber.
It is both diurnal, avoiding direct sunlight, and
nocturnal. The diet is largely composed of mam-
mals, especially rodents such as rats and squir-
rels, and of birds and bats. Specimens living in
caves seem to feed essentially upon bats and spar-
rows and their eggs. It is an aggressive snake, strik-
ing at little provocation and biting fiercely when
given a chance to.
Distribution
This subspecies is only known from a few locali-
ties in the mountain ranges of Sumatra and Bor-
neo. Mt. Kinabalu: Kamborangoh Road (1600 m),
Headquarters, Lumu Lumu (STUEBING 1994), Bun-
du Tuhan.
Genus Gongylosoma FITZINGER, 1843
The genus Gongylosoma is distributed from Thai-
land, through Malay Peninsula to Java, Sumatra,
and Borneo.
Small, slender; head not distinct from the body;
tail long; eyes moderately large, pupil round; a
small loreal shield is present; nasal shields divided;
13 rows of dorsal scales without apical pits or keels;
anal scute divided; subcaudals in two rows.
Little is known about the biology of these terres-
trial snakes. They feed on invertebrates and lizards.
Their mode of reproduction is unknown.
Two species have been described for this genus,
and both occur on Borneo, one on Mt. Kinabalu.
335
Colubridae

Taxonomic comments: LEVITON (1964) has split the

Oriental genus Liopeltis auctorum into two groups
on the basis of meristic and morphological charac-
ters constant among species of the respective groups.
He referred the nominal species Coronella baliodeira
BOIE, 1827 and Ablabes longicaudus PETERS, 1871
to the genus Gongylosoma FITZINGER, 1843. LEVITON
(1964) was the last reviewer of the southeastern
members of this poorly studied group, and his ar-
rangement is followed here.
Key to the Bornean species of
Gongylosoma
1 Anterior part of body with 5 orange stripes; 77-
103 subcaudals .............................. longicaudum
• Anterior part of body with yellow spots or with
three yellow stripes; 58-79 subcaudals ...............
................................... baliodeirum baliodeirum

Tab. 44 Distribution of Gongylosoma-species on Borneo.

Gongylosoma baliodeirum baliodeirum stripes; supralabials light with black margins; belly
orange or yellowish white, with or without fine dark
BOIE, 1827 spots; the outer edges of the ventrals colored like
the dorsum.
Coronella baliodeira BOIE, 1827: 539 – Type locality: Java
Ablabes baliodeirus, MOCQUARD 1890a: 137; SHELFORD 1901a:
Presumably oviparous.
61
Ablabes baliodirus, DE ROOIJ 1917: 139 Ecology/Ethology
Gongylosoma baliodeira, SMITH 1931: 10, 32 This small species inhabits regions of low and
Gongylosoma baliodeirum, SMITH 1931: 32
moderate elevations from sea level up to at least
Liopeltis baliodeirus, MANTHEY 1983: 28; EBENHARD & SJÖGREN
1984: 24; MALKMUS 1994b: 247 1500 m and is thus found in both lowland tropical
Liopeltis baliodeira, MALKMUS 1992a: 134, 1994b: 239 wet forests and tropical and subtropical wet
montane forests. It is more common in hilly and
Diagnosis rugged areas. Gongylosoma b. baliodeirum is a ter-
This species is easily identified on the basis of restrial and rather secretive forest-dweller, living
its three rows of spots on the anterior body. 13
rows of dorsals; 1 loreal present. Total length ca.
40 cm. Fig. 332 Gongylosoma baliodeirum baliodeirum from Ulu
Katibas, Lanjak-Entimau Wildlife Sanctuary, Sarawak.
Description
Two preoculars; 2 postoculars; 1 loreal; 1+2 or 2+2
temporals; 7 supralabials, the 3rd and 4th touching
the eye; 8 sublabials; 13 dorsal scale rows through-
out the length of the body; dorsals smooth; 115-
145 ventrals (STEJNEGER 1922, KOPSTEIN 1941); anal
scute divided; 55-79 subcaudals (KOPSTEIN 1941,
MALKMUS 1992a, DAS 1995, DAVID & VOGEL 1996)
© R. B. Stuebing

in two rows.
Upper side dark brown, with some small, dark-bor-
dered, yellow spots on the nape and the anterior
portion of the body which may be fused to form

336

Fig. 333 Gongylosoma baliodeirum baliodeirum from Kelabit Highlands, Sarawak.
on the floor in among leaf litter, under rocks, and
beneath decaying vegetation. It is diurnal and re-
ported to feed on invertebrates and lizards. Accord-
ing to STUEBING & INGER (1999) its diet consists
mainly of spiders.
Genus Gonyophis BOULENGER, 1891
The genus Gonyophis is native to Borneo and Pe-
ninsular Malaysia.
Moderately large, slender; l body long, laterally com-
pressed; head elongate, distinct from the neck; eyes
moderately large, pupils round; dorsal scales keeled,
with apical pits, in 19 rows at mid-body; ventrals
with lateral keels and notches on their sides; tail long,
with keeled and notched subcaudals in two rows.
These are rare tree snakes of the lowlands and
montane regions whose biology is poorly known.
The genus Gonyophis is monotypic with the only
species living on Mt. Kinabalu.
Colubridae
© I. Das
Distribution
The distribution range extends from Peninsular Ma-
laysia to Borneo, Sumatra, and Java. Mt. Kinabalu:
Kiau, Poring: Hot Springs region (500 m) and be-
tween Sg. Kipungit I and Sg. Kipungit II (600 m).
Gonyophis margaritatus (PETERS, 1871)
Gonyosoma margaritatum, PETERS, 1871: 578 – Type locality:
Sarawak
Diagnosis
19 dorsal scale rows; keeled, notched ventrals; con-
spicuous by the bright yellow rings around the body
in its posterior part. Max. total length 170 cm.
Description
One loreal; 1 preocular; 2 postoculars; 8-9 suprala-
bials, the 4th to 6th or the 5th and 6th in contact with
the eye; 2+2 or 2+3 temporals; dorsals in 19 rows,
keeled, with apical pits; 230-249 ventrals, laterally
337
 Colubridae
© R.B. Stuebing

Fig. 334 Gonyophis margaritatus from Lanjak-Entimau Wildlife Sanctuary, Sarawak.

keeled and notched; anal scute divided; 108-130 Distribution

subcaudals, in pairs, laterally keeled. This species lives in Peninsular Malaysia and on
Dorsal ground colour bright green, each scale with Borneo. Mt. Kinabalu: Poring: Sg. Kipungit I.
black margins; posterior part of the body and tail
with several orange-coloured to deep red rings of
which the posterior most ones are the most bright Fig. 335 Gonyophis margaritatus from Sg. Kipungit I.
ones; head olive green with a black stripe between
eye and nape and between the parietal shields; un-
dersides yellow, ventrals often with black margins
and an orange-coloured dash.

Ecology/Ethology
These snakes are arboreal and inhabit forests of
the lowlands and montane regions up to altitudes
of 2000 m a.s.l. They have the ability to ascent even
© B. Leideritz

apparently perfectly smooth tree trunks. In captiv-

ity the only food accepted was fish. Its biology is
poorly known.

338

Genus Gonyosoma WAGLER, 1828
The genus Gonyosoma is distributed from Myanmar,
Laos and Vietnam, to Bali, Sulawesi and the Phil-
ippine Islands.
Large and slender, muscular; body slightly com-
pressed laterally; head slender; pupil round; 23 to
27 dorsal scale rows at mid-body; dorsals with elon-
gate apical pits; loreal shield at least twice as long
as high; anal scute and subcaudals divided.
The crepuscular and nocturnal snakes live on the
ground and on trees.
The genus comprises two species, one of which
lives on Borneo and on Mt. Kinabalu.
Gonyosoma oxycephalum (BOIE, 1827)
Coluber oxycephalus BOIE, 1827: 537 – Type locality: Java
Elaphe oxycephala, EBENHARD & SJÖGREN 1984: 24
Gonyosoma oxycephala, STUEBING 1991: 343; MALKMUS 1994b:
247
Diagnosis
Readily identified on the basis of its green body
which contrasts sharply with a grey tail; max. total
length of 240 cm, but usually only 160-180 cm.
Fig. 336 Gonyosoma oxycephalum from Poring, 600 m.
Colubridae
Description
One large preocular; 2 postoculars; 7-11 supra-
labials, the 5th and 6th or the 6th and 7th in contact
with the eye; 12-14 sublabials; dorsals in 23, 25,
rarely 27, rows; 229-263 ventrals, sharply bent on
their sides; anal scute divided; 120-157 paired sub-
caudals, divided.
Light green dorsally, some scales bordered with
black, others with yellow, the number of yellow-
bordered scales increasing posteriorly forming a
vague striped lateral pattern; upper side of head
greenish yellow; upper side of tail grey or brown;
ventral sides lighter; belly bright or pale yellow;
juveniles altogether somewhat darker (this colour
pattern refers to Bornean specimens only).
A clutch typically consists of 5-12 eggs (58-70 ×
25-27 mm). Hatchlings emerge after 87 to 125 days
and measure 40-55 cm in total length.
Ecology/Ethology
This species is found in lowland areas from sea level
to elevations up to about 1300 m where it inhabits
primary and secondary tropical forests, tropical wet
montane forests, mangrove forests, swamps and
marshes, wet scrublands, and plantations. It is rather
rarely encountered close to villages. Gonyosoma
oxycephalum is usually, but not exclusively, found
close to rivers, on river banks, near ponds and other
© P. Hoffmann
339
Colubridae
mesic areas covered with tangled vegetation; it is
most at home in trees and bushes overhanging wa-
ter. It is an agile, swift, and very active species which
is almost entirely arboreal, being partial to treetops,
tall bushes, and other thick vegetation; juveniles may
be more terrestrial. This strictly diurnal snake feeds
mainly on birds, but also readily hunts arboreal mam-
mals and bats; juveniles have been reported to prey
upon lizards. This species has an interesting threat
display, in which the neck is flattened top to bot-
Genus Hydrablabes BOULENGER, 1891
This genus is endemic to Borneo.
Head small, not distinct from neck; eyes small,
pupils round; an oblique suture between the two
nasal shields; 3 pairs of chin shields; body long
and round in cross-section; dorsals smooth, with-
out apical pits, in 15-17 rows; ventrals rounded;
tail moderately long; subcaudals in two rows.
These are terrestrial snakes with fossorial habits.
They are considered rare, and as a consequence
hardly anything is known about their biology. They
Tab. 45 Distribution of Hydrablabes-species on Borneo.
Hydrablabes praefrontalis
(MOCQUARD, 1890)
Ablabes periops var. praefrontalis MOCQUARD, 1890a: 137, pl.
IX; fig. 1a-c – Type locality: Mt. Kinabalu
Ablabes periops var. prefrontalis MOCQUARD, 1890b: 134 – Type
locality: Mt. Kinabalu
Hydrablabes praefrontalis, BOULENGER 1891b: 343; MOCQUARD
1892: 205; BOULENGER 1893b: 297; SHELFORD 1901a: 58; DE
ROOIJ 1917: 137; SMITH 1931: 32; M ANTHEY 1983: 28;
E BENHARD & S JÖGREN 1984: 24; S TUEBING 1991: 349;
MALKMUS 1994b: 247; STUEBING & INGER 1999: 172
Taxonomic comments: STUEBING & INGER (1999)
believed Hydrablabes praefrontalis to be a syno-
340
tom, and the bright blue-striped tongue extended and
retracted in an extremely slow, deliberate fashion.
Mating occurs usually at night on trees or shrubs
with a copulation taking 4-5 hours (SHIN 1994).
Distribution
Gonyosoma oxycephalum is distributed from
Myanmar, Laos and Vietnam in the north, to Bali,
Borneo, and the Philippines in the south. Mt.
Kinabalu: Poring.
appear to live almost exclusively near to water,
some specimens were found swimming in rain for-
est streams.
Two species are known, both live on Borneo, one
of which on Mt. Kinabalu.
Key to the Bornean species of
Hydrablabes
1 2 prefrontals ............................................ periops
• A single prefrontal ........................ praefrontalis
nym of Hydrablabes periops. They did, however,
not really synonymise the two species. As long as
there is no comprehensive review of the genus, we
consider the separation of the prefrontals to war-
rant species status for both taxa.
MOCQUARD published the description of this spe-
cies twice, i.e. once in Le Naturaliste, Paris: 144-
145, and again in Nouv. Arch. Mus. Hist. Nat.,
Paris, 3 (2): 115-168. MOCQUARD’s paper in the
“Naturaliste” was published on July 1st, 1890. In
the first part of this four-part series, MOCQUARD
wrote (in interpretation): “... The description of this
collection [Mr. Whitehead’s collection] will be soon
published in the Nouvelles Archives du Museum;
 Colubridae

but we believe that readers of the Naturaliste will

appreciate to find diagnoses of the new species in-
cluded in this collection in this interesting publi-
cation, which, at the same time, allows us to set a
date for these diagnoses.”
In the “Nouvelles Archives”, there is no date of pub-
lication. It is clear that the first part of the four-part
series in the “Naturaliste” was published before the
“Nouvelles Archives”, and most probably the sec-
ond part, too. It cannot, however, be clarified here
which publication serves as original description.
Unfortunately MOCQUARD used two different spell-
ings for this species in these two publications. If
the publication in the “Naturaliste” proves to have
been published first, the correct name should be
Hydrablabes prefrontalis.
after MOCQUARD 1890
Diagnosis
Prefrontals fused to form one shield; three pairs of Fig. CXII Hydrablabes praefrontalis
chinshields; max. total length 43.6 cm.

Description interrupted to form a series of points in the poste-

Prefrontals fused to form a single shield; 1 loreal;
2 preoculars; 2 postoculars; 2-3 suboculars; 1+2
temporals; 8-9 supralabials, none of which touch
the eye; 15 dorsal scale rows at mid-body; 178-
202 ventrals; 72 subcaudals, divided.
Olive brown dorsally; a dark vertebral stripe, some-
times hardly visible; two black dorsolateral stripes,
rior part of the body; lower sides yellow, the outer
margins of the ventrals dark brown; subcaudals with
brown margins; a dark median line along the tail.
Distribution
This species is endemic to Mt. Kinabalu, but no
precise locality data are known.

Being crepuscular and nocturnal colubrids they are

Genus Internatus YANG & RAO, 1992
The genus Internatus ranges from Malay Penin-
sula to Sumatra, Borneo, and Java. Isolated locali-
ties are known from central Thailand.
Small; head distinct; chin shields arranged asym-
metrically; 2nd or 3rd pairs of sublabials enlarged,
in contact with each other; 1 pre-chinshield an-
terior to the 2 pairs of chinshields; chinshields
wider than long; no preocular nor subocular
shields; 3rd, 4th, and 5th supralabials entering the
orbit; 2+2 temporals; supraoculars very small;
jaws with 5 to 6 teeth; dorsal scales in 15 rows,
very smooth, the vertebral scales enlarged; sub-
caudals paired.
specialized in a diet of slugs and snails. The latter
are extricated from their shells with the aid of the
anteriorly positioned fangs of the narrow, firm lower
jaw. All the species reproduce by laying eggs.
Four species have been described (DAVID & VOGEL
1996), two of which occur on Borneo, both on Mt.
Kinabalu.
Key to the Bornean species of
Internatus
1 5 or 6 upper labials ................................... laevis
• 7 upper labials ................................. malaccanus

341
 Colubridae

Tab. 46 Distribution of Internatus-species on Borneo.

Internatus laevis (BOIE, 1827)

Amblycephalus laevis BOIE, 1827: 520 – Type locality: Java

Amblycephalus laevis, BOULENGER 1896c: 441; SHELFORD 1901a:
68; DE ROOIJ 1917: 270; SMITH 1931: 11, 32
Amblycephalus vertebralis, LOVERIDGE 1938: 43; STUEBING
1991: 331
Pareas laevis, EBENHARD & SJÖGREN 1984: 24; STUEBING 1991:
330; MALKMUS 1992a: 136, 1994b: 247, 1996c: 293; STUEBING
& INGER 1999: 87
Internatus laevis, RAO & YANG 1992: 142

Diagnosis
Identified by the combination of asymmetrical

© A. Nöllert
chinshields and 5-6 supralabials. The long loreal
touches the eye. Internatus laevis can reach about
60 cm in total length.
Fig. 338 Underside of the specimen from below.

Fig. 337 Internatus laevis from the Kamborangoh Road.

© A. Nöllert

342
 Colubridae

Description
One loreal, in contact with the eye; no preocular;
1-2 postoculars; 2+2 temporals; 5-6 (7) suprala-
bials, the 3rd and 4th or the 3rd to 5th touching the
eye, the 6th very long; 15 dorsal scale rows at mid-
body; smooth; vertebrals enlarged; 148-176
ventrals; anal scute entire; 34-69 subcaudals, in
pairs. SMITH (1925b) reported about two specimens
from Mt. Perissen with unusually high ventral (196)
and subcaudal (73-78) counts. One of his speci-
mens had 7 supralabials.
Medium to dark brown dorsally, with irregular bark
bands which do not cross over mid-body; head usu-
ally slightly darker than body; lower sides brown
or yellow with exception of the dark spots which

© C. Brühl
mark the ends of the dorsal rings.

Ecology/Ethology Fig. 339 Internatus malaccanus from Poring.

This arboreal species inhabits tropical wet forests
and tropical wet montane forests. It is active dur-
ing the evening and night hours. In Peninsular
Malaysia, a specimen was seen in a small stream
(KIEW 1987). Nothing else is known of the biology
of this rare species. It is presumed to be similar to
other Internatus and Pareas species.
MALKMUS (1996c) found a specimen during rainy
weather on the Kamborangoh Road at 10.00 p.m.
Another one was discovered ca. 1 m up in a bush at
about 9.00 p.m. (MALKMUS 1992a).
Distribution
This species is distributed from Laos, through Thai-
land and Peninsular Malaysia, to Java, Sumatra,
and Borneo. On Borneo, it has been recorded from
Sabah as well as from Sarawak and Kalimantan.
No records are available for Brunei. Mt. Kinabalu:
Marei Parei, Kamborangoh Road (1600 m), Sg.
Langanan (850 m).
Internatus malaccanus (PETERS, 1864)
Astenodipsas Malaccana PETERS, 1864: 273, plate fig. 3 – Type
locality: “in der Nähe von Malacca” [Peninsular Malaysia]
Diagnosis
This species is identified by the combination of
asymmetrical chinshields with the second and third
infralabials in contact with one another and 7-8
supralabials. It attains a maximum length of of
about 600 mm.
Description
One loreal, in contact with the eye; no preocular; 2
postoculars; 2+2 temporals; 7-8 supralabials, the
3rd and 4th or the 3rd to 5th touching the eye, the last
large; 15 dorsal scale rows at mid-body, very faint
keeled; vertebrals enlarged; 165-181 ventrals; anal
scute entire; 35-58 subcaudals, in pairs.
Dark brown or black above with or without a white
vertebral line and with irregular light cross-bands
bordered with white; head white, with or without
black spots on each side of the nape; belly white
suffused with brown speckles on each side.
Oviparous.
Ecology/Ethology
BATCHELOR (1958) reported about a specimen found
in a rubber plantation. Nothing else is known about
the biology of this forest-dwelling species. It is
presumed to be similar to Pareas carinatus.
Distribution
Internatus malaccanus lives on Sumatra, Borneo,
and in the Malay Peninsula. Mt. Kinabalu: Por-
ing.

343
 Colubridae
Genus Lepturophis BOULENGER, 1900
The genus Lepturophis ranges from the Malay Pe-
ninsula south to Borneo and Sumatra.
Extraordinarily slim, elongate, cylindric; head dis-
tinct from the body, very depressed; eye moder-
ately large; pupil vertical-elliptical ; enlarged fangs
in the upper jaw; nostrils very large, each wedged
between two nasals and the first supralabial shield;
dorsals with distinct keels, finely serrated, lanceo-
late, slightly imbricate, with apical pits, in 17 rows
at mid-body; ventrals keeled laterally and notched;
tail extremely long and slim; subcaudals in two
rows.
These snakes are nocturnal and crepuscular, living
in secondary and primary forests They are prob-
ably largely arboreal, but have mostly been seen
active on the ground. The snakes are often found
along streams or in stream beds. Their favourite
Fig. 340 Adult Lepturophis albofuscus from Peninsular Malaysia.
© W. Grossmann
344
food is frogs and lizards and they reproduce by lay-
ing eggs.
This genus is monotypic. The only species has been
found also on Borneo and Mt. Kinabalu.
Lepturophis albofuscus
(DUMÉRIL, BIBRON & DUMÉRIL, 1854)
Sphecodes albo-fuscus DUMÉRIL, BIBRON & DUMÉRIL, 1854a:
394 – Type locality: Sumatra
Ophites albofuscus, MOCQUARD 1890a: 144
Lycodon albofuscus, SHELFORD 1901a: 59; DE ROOIJ 1917: 110;
SMITH 1931: 32; MANTHEY 1983: 28; EBENHARD & SJÖGREN
1984: 24; STUEBING 1991: 333
Lepturophis albofuscus, MALKMUS 1994b: 247
Taxonomic comment: For a discussion of the tax-
onomy of this species see DAVID & VOGEL (1996).
Lepturophis borneensis BOULENGER, 1900 is a syno-
nym of L. albofuscus.
 Colubridae

Diagnosis
Slim body with an ex-
tremely long tail; pupil
vertical-elliptical; ventrals
and subcaudals keeled;
max. length 207 cm, but
average specimens meas-
ure 120-160 cm.

Description
Snout very blunt; 1 preo-
cular; 2 postoculars; 1 lo-
real; 2+2 temporals; 8 (7)
supralabials, the 3rd to 5th
or the 4th and 5th in con-
tact with the eye; dorsal
scales in 17 rows at mid-
body, keeled; 220-259
ventrals; anal scute di-
vided; 113-206 subcau-

© W. Grossmann
dals, in pairs, keeled.
Dorsally uniform dark
grey, greyish brown, grey-
ish black, or black; ven-
tral sides yellowish white Fig. 341 Juvenile Lepturophis albofuscus from Peninsular Malaysia.
to white; juveniles dark
with 30-40 whitish bands
on the body and a broader band on the neck, the ently not occur on Java. Mt. Kinabalu: without
bands closer together posteriorly so that the snakes more precise locality data.
appear darker anteriorly.
Oviparous.

Ecology/Ethology Genus Lycodon FITZINGER, 1826

Lepturophis albofuscus inhabits lowland tropical
forests, where it is reported to live close to streams
in bushes and thick vegetation, on stream banks
and gravel beds. This snake is probably largely ar-
boreal, but it has mostly been seen active on the
ground in forests. Sometimes it was observed swim-
ming. It is nocturnal and feeds on frogs and liz-
ards, preying especially on the semiaquatic scincid
genus Tropidophorus.
This species is regarded as uncommon throughout
its range.
Distribution
This snake’s range extends from the Malay Penin-
sula to Borneo, Sumatra, and Nias. It does appar-
The genus Lycodon is distributed from the terri-
tory of the former USSR through large parts of In-
dia and China south to Sulawesi and the Philip-
pine Islands.
Small to medium-sized, slender snakes; head de-
pressed dorsoventrally, slightly set off from body;
eye relatively small, pupil vertical-oval; nostril
large; anterior maxillar teeth curved, with a gap
between the very large anterior and the subsequent
ones. Dorsal scales keeled or not, with apical pits,
in 17, 19, or 21 rows at mid-body; ventrals rounded,
bent, or with lateral keels; subcaudals arranged in
a single or two rows.

345
Colubridae
These crepuscular to nocturnal colubrids inhabit a
wide variety of habitats, ranging from cultivated
areas, through grass and sandy plains, to second-
ary and primary forests. All species of this genus
are terrestrial. They feed mainly on lizards, frogs,
and snakes, with the smaller species also preying
upon insects. Juveniles possess more intense col-
ours and more contrasting patterns. All species of
this genus are oviparous.
In total 26 species are known, three of which occur
on Borneo, one on Mt. Kinabalu.
Tab. 47 Distribution of Lycodon-species on Borneo.
Lycodon subcinctus sealei LEVITON, 1955
Lycodon subcinctus sealei LEVITON, 1955: 195 – Type local-
ity: Puerto Princessa, Palawan Island, Philippines
Lycodon subcinctus, STUEBING 1991: 333; MALKMUS 1994b: 247
Diagnosis
Adult Lycodon subcinctus are readily recognizable
on the basis of the usually three white rings on the
anterior portion of the body. L. s. sealei is distin-
guished from the other subspecies by a lower count
of subcaudals which, according to LEVITON (1955),
number less than 69. It differs from the other
Bornean species of Lycodon in that it lacks a
preocular shield and has a long loreal which touches
the eye. The longest specimen known measured
105 cm, with average specimens being between 80
and 100 cm long.
Description
One loreal; no preocular; 2-3 postoculars; 1+2 tem-
porals; 8 supralabials, the 3rd to 5th, 4th and 5th or 3rd
to 6th in contact with the eye; dorsals in 17 rows at
mid-body, keeled; 198-211 ventrals; anal scute
normally divided; 60-68 subcaudals, divided.
The pholidotic characteristics are adopted from
LEVITON (1955) and apply to this subspecies only.
Dark brown or black dorsally; nape region white;
white rings of variable width all along the body;
346
Key to the Bornean species of Lycodon
1 2 scales between the posterior nasal and orbit ....
............................................................ capucinus
• 1 scale between the posterior nasal and orbit .. 2
2 No preocular, an elongate loreal and the prefrontal
enter the orbit; one anterior temporal ..................
................................................ subcinctus sealei
• 1 preocular; loreal absent; prefrontal in contact with
the supralabials; 2-3 anterior temporals ...effraenis
fading from the tail end with increasing age;
adults greyish to brownish black with 2 white
bands or whitish specks in the anterior portion
of the body, the latter being remains of white
bands, very old specimens with hardly any such
markings at all; ventrum dirty to greyish white
or brown.
Several clutches per year of 5-11 eggs (31.5-36 ×
12.5-13.5 mm, average weight 3.65 g); hatchlings
measure about 240 mm in length.
Ecology/Ethology
This snake is found from sea level up to elevations
above 1500 m. It inhabits lowland tropical wet and
dry forests, tropical and subtropical wet montane
forests, shrublands, plantations, rice paddies, cul-
tivated areas, and the environs of villages. This swift
snake is terrestrial, retreating under rocks, stumps,
pieces of wood and vegetation, and arboreal. It is
nocturnal and forages actively in the darkness to
find its prey, that in essence consists of lizards, es-
pecially geckos and diurnal skinks, which are
started in their shelters.
Lycodon subcinctus is rather aggressive and bites
readily when molested.
Distribution
This subspecies is native to Borneo and Palawan
(Philippines). Mt. Kinabalu: Bundu Tuhan.
 Colubridae

rior portion of the body stretching it laterally and

Genus Macropisthodon BOULENGER, 1893
The genus Macropisthodon has a discontinuous
distribution range with records being known from
China, India, Sri Lanka, and Pakistan, as well as
from Thailand south to Sumatra, Borneo, and Java.
It comprises medium-sized species; with very stout
bodies. Head distinct from the body; eye moder-
ately large to large, pupil round; one pair of very
large fangs; dorsals strongly keeled, with apical
pits, in 19-27 rows; ventrals rounded; tail moder-
ately long; subcaudals in two rows.
The included species may be diurnal as well as
nocturnal and have a marked predilection for the
vicinity of water. Their defence behaviour resem-
bles that of the cobras in that they flatten the ante-
lifting it into a vertical position off the ground. Their
diet consists for a major part of frogs and toads,
their larvae, and fish. All species of this genus are
oviparous.
Four species are known, two of which occur on
Borneo, one on Mt. Kinabalu.
Key to the Bornean species of
Macropisthodon
1 Usually 8 upper labials; black or with black
transverse bands; neck partly red ........ flaviceps
• Usually 7 upper labials; a dark vertebral stripe;
neck partly blue ............................... rhodomelas

Tab. 48 Distribution of Macropisthodon-species on Borneo.

Macropisthodon rhodomelas lips, throat, and belly uniform pink to yellowish,

with a row of dots on the ventrals; juveniles deep
(BOIE in BOIE, 1827) reddish on their upper sides.
A single clutch comprised 25 eggs. No more infor-
Tropidonotus rhodomelas H. BOIE in F. BOIE, 1827: 535 – Type mation is available.
locality: Java

Diagnosis Fig. CXIII Macropisthodon rhodomelas

Fairly slender, body laterally compressed; easily
identified by its blue throat and the distinctly
marked dark vertebral stripe; max. length about
75 cm.

Description
One loreal; 1-2 preoculars; 3-4 postoculars; occa-
sionally 1 subocular; 2+2, 2+3 or 3+3 temporals;
7-8 supralabials, the 3rd and 4th, more rarely the 4th
and 5th touching the eye; 19 rows of dorsals at mid-
body, strongly keeled; 124-144 ventrals; anal scute
divided; 42-64 subcaudals, in two rows.
Dorsally reddish brown, with dark vertebral stripe
which splits into two branches on the nape to form a
“V”; nape bluish grey; upper side of head uniform after DE ROOIJ 1917
in the ground colour of the body; neck bluish; upper

347
Colubridae
Ecology/Ethology
Macropisthodon rhodomelas is found in lowland
and montane regions where it inhabits tropical wet
forests, tropical wet montane forests, swamps,
marshes, plantations, rice paddies, and gardens in
suburban areas. This diurnal and nocturnal snake
is terrestrial and semiaquatic, and lives close to
water in wet vegetated areas. It feeds chiefly on
frogs and toads; juveniles have been reported to
eat tadpoles.
Genus Oligodon FITZINGER, 1826
The genus Oligodon is very widely distributed with
populations in Pakistan, Iran and Turkmenistan in
the west, through large parts of India, China and
the Philippines in the east, south to Sulawesi.
Small to medium-sized; body muscular, round in
cross-section; head short, not distinct from body;
rostral shield large, separating internasals for two tirds
of their lengths; 2 temporals bordering parientals; a
loreal scale may be absent or present. Eye moderate
to small, pupil round. Dorsal scales smooth or feebly
keeled, in 13-23 rows, with or without apical pits;
ventrals rounded or with a weak keel; anal scute di-
vided or entire; tail short; subcaudals in two rows.
These terrestrial snakes are marked for their
strongly curved posterior teeth and a head which
ends bluntly in a large rostral shield. They can in-
flict profusely bleeding wounds if given an oppor-
tunity to bite. Some species are known to coil up
their tails to a spiral, lift it off the ground, and so
display its often brightly coloured underside in
selfdefence (WALLACH & BAUER 1996).
Depending on their sizes, these snakes prey upon
frogs, lizards, snakes, and insects and their larvae.
Some species appear to feed primarily on reptile eggs.
All species of this genus reproduce by laying eggs.
According to WELCH (1988) 63 species, according
to WALLACH and BAUER (1996) even 70 species and
135 nominal forms are valid, but this genus is in
urgent need of a comprehensive revision. Eight or
nine species are known from Borneo, respectively,
four of which from Mt. Kinabalu. The occurrence
of Oligodon meyerinkii requires confirmation, but
is included in the key below. Oligodon cinereus
348
This beautiful snake is common, especially in the
lowlands. When aroused, it raises the anterior part
of its body, flattens its neck like a cobra, and may
bite viciously.
Distribution
Macropisthodon rhodomelas is distributed from the
Malay Peninsula south to Borneo, Java, and Sumatra.
Mt. Kinabalu: Headquarters (1500-1600 m) [HQ-
Collection].
was also mentioned for Borneo by DE ROOIJ (1917)
under the name Oligodon violaceus. According to
STUEBING & INGER (1999) this is a misidentification.
Oligodon cinereus is a species living north of the
Malay Peninsula, for which reason we have ex-
cluded this species from the fauna of Borneo.
Key to the Bornean species of Oligodon
1 Body with stripes .............................................. 2
• Body without stripes ......................................... 3
2 Dark stripes without light spots ....................... 8
• Small red or yellow or white spots in a dark ver-
tebral stripe ....................................................... 7
3 Ventrals barred with dark pattern ........................
...................................................... purpurascens
• Ventrals unmarked, spotted on the outer edges or
mottled .............................................................. 4
4 19 dorsals at mid-body……………. semicinctus
• 15-17 dorsals at mid-body ................................ 5
5 17 dorsals at mid-body ................ subcarinatus
• 15 dorsals at mid-body ..................................... 6
6 Body with about 25 black rings, each with an
oval yellowish spot .............................. annulifer
• Narrow brownish or reddish crossbars on back ..
.......................................................... cf. signatus
7 Anal scute divided ......... vertebralis vertebralis
• Anal scute not divided ........................... everetti
8 7 stripes are clearly visible ............ octolineatus
• Only the dorsal stripe is clearly visible, the other
six stripes are wanting or very faintly visible .....
............................................................ meyerinkii

Tab. 49 Distribution of Oligodon-species on Borneo.
Oligodon everetti BOULENGER, 1893
Oligodon everetti BOULENGER, 1893a: 524 – Type locality: Mt.
Kinabalu
Oligodon everetti, B OULENGER 1894e: 339 pl. XI: fig. 1;
BARTLETT 1895c: 82; SHELFORD 1901a: 61; DE ROOIJ 1917:
130; SMITH 1931: 32; MANTHEY 1983: 28; E BENHARD &
SJÖGREN 1984: 24; STUEBING 1991: 334; MALKMUS 1994b: 247;
MANTHEY & GROSSMANN 1997: 369; STUEBING & INGER 1999:
104
Fig. 342 Oligodon everetti from Danum Valley, Sabah.
Colubridae
Diagnosis
Readily recognized on the basis of the colour pat-
tern described below. Ventral side uniform coral-
red; anal scute entire; 15 rows of dorsals at mid-
body. Total length less than 50 cm.
Description
One preocular; 2 postoculars; 1 small loreal; 7 su-
pralabials, the 3rd and 4th in contact with the eye;
© J.C. Murphy
349
 Colubridae

1+2 temporals; dorsals in 15 rows at mid-body; Oligodon octolineatus (SCHNEIDER, 1801)

132-154 ventrals; anal scute entire; 46-72 divided
subcaudals. Elaps octolineatus SCHNEIDER, 1801: 299 – Type locality: Not
Upper sides greyish brownish, with 3 broad black- given; subsequently designated by MERTENS (1969: 340) as
ish brown stripes, the middle one with yellowish probably being Java.
Simotes octolineatus, MOCQUARD 1890a: 137; DE ROOIJ 1917:
and reddish, black-bordered saddle blotches, the
123
lateral ones with interspersed light scales; head Oligodon octolineatus, SMITH 1931: 32; EBENHARD & SJÖGREN
brown, with a dark brown, V-shaped marking that 1984: 24; MALKMUS 1994b: 247
opens posteriorly; another dark brown band extend-
ing from the upper side of the head over the eyes; Diagnosis
lower side coral-red; ventrals laterally with dark Easily recognized by its distinct striped pattern.
margins. Dorsal scales in 17 rows at mid-body; 2 anterior
Like all species of this genus oviparous; otherwise temporals; ventrals slightly angulated; anal scute
unknown. entire. Max. length about 70 cm, average speci-
mens 50-60 cm.
Ecology/Ethology
Oligodon everetti is a ground-dwelling species of Description
the lowland forests. One specimen had a scincid One loreal; 1 preocular; 2 postoculars; 6 supra-
lizard of the genus Sphenomorphus in its stomach. oculars, the 3rd and 4th in contact with the eye; 2+2
This represents all the information available on its temporals; dorsals in 17 rows at mid-body; 161-
biology. 183 weakly angulated ventrals; anal scute entire;
43-61 subcaudals, divided. These values refer to
Distribution specimens from Borneo only.
This species is a Bornean endemite. Mt. Kinabalu: Dorsally pale brown with three pairs of dark stripes,
no precise locality data available. the median ones fusing on the head to form an
arrow-like pattern, the
interspace between them
Fig. 343 Oligodon octolineatus from Tawau Hills, Sabah. can be darker or lighter
than the ground colour.
Lateral stripes ending at
base of tail, dorsal ones
continue on the tail. A
dark ring on the head
passing through the eyes;
another band-like mark-
ing from the centre of the
head at the level of the
eyes onto the nape; lower
sides uniform pink. The
stripes may be bordered
with white in Bornean
specimens (M ERTENS
1969).
Clutches of up to 5 eggs.
© R.B. Stuebing

Ecology/Ethology
This snake is found in
lowland regions where it

350
 Colubridae

inhabits tropical wet and dry forests, but it is ob- Description

served mainly in clearings, open scrublands and
grasslands, along roads, in plantations, rice pad-
dies, cultivated areas, gardens, and in the vicin-
ity of villages. It is secretive, terrestrial and semi-
fossorial, retreating under rocks, logs, piles of
wood, and in vegetation. Oligodon octolineatus
is nocturnal and feeds on lizards, frogs, tadpoles,
and other snakes, but seems especially partial to
the eggs of lizards and birds as well as frog
spawn.
This species is mild-mannered and seldom attempts
to bite.
In Singapore, a clutch was discovered in a tree in
the borings of some longicorn beetle, as high as 7 m
above the ground (RIDLEY 1906).
Distribution
Oligodon octolineatus lives in Peninsular Malay-
sia as well as on the Indonesian islands of Bangka,
Belitung, Java, Nias, the Riau Archipelago,
Sulawesi, Sumatra and on Borneo. Mt. Kinabalu:
no precise locality data available.
Oligodon purpurascens (SCHLEGEL, 1837)
One to three preoculars; 2 or 3 postoculars; 1 or 2
suboculars; 1 loreal; 2+2 or 2+3 temporals; 8 su-
pralabials, the 4th and 5th or only the 5th in contact
with the eye; 19-21 rows of dorsal scales at mid-
body; 160-210 ventrals with slight angles; anal
scute entire; 40-60 subcaudals, divided.
Adults reddish brown, greyish brown, or dark grey
dorsally, or with a blend of the mentioned colours;
body with wide bands (saddle blotches), often with
undulated, dark (i.e. brown, grey, or blackish) bor-
ders, the interspaces with 2-3 darker lines; a dark
band from the angle of the mouth through the eye
to the forehead and back through the other eye to
the opposite angle of the mouth; a V-shaped dark
marking between the eyes opening towards the
nape; chin and throat whitish, ventral side usually
yellowish anteriorly, reddish more posteriorly; ven-
trals often with irregular, black, squarish blotches,
occasionally limited to the posterior portion or al-
together absent; Lower side of tail often reddish,
deep red, or whitish; older specimens altogether
more obscure; juveniles more contrasting, i.e. light
brown with dark brown bands and lines; laterally
with small white and reddish spots; ventral side
sparkling purple with irregular black speckling;
gular region often white.

Xenodon purpurascens SCHLEGEL,

1837: 90, pl. 3: fig. 13 & 14 – Fig. 344 Oligodon purpurascens from Bundu Tuhan, 1350 m.
Type locality: Mt. Parang, Java
Simotes purpurascens, DE ROOIJ
1917: 126
Oligodon purpurascens, SMITH
1931: 32; MANTHEY 1983: 28;
E BENHARD & SJÖGREN 1984:
24; S TUEBING 1991: 334;
MALKMUS 1994b: 241, 247,
1996c: 292

Diagnosis
19-21 rows of dorsals at
mid-body; anal scute en-
tire; two anterior tempo-
rals; the ventral scales may
or may not bear black
blotches of up to half the
© M. Schroth

individual scales’ sizes;

max. total length almost
100 cm.

351
 Colubridae
8-13 eggs ( 27-33 × 18-22 mm, 4.9-5.9 g); larger
clutches are not unlikely. Newly hatched specimens
measure about 210 mm.
Ecology/Ethology
This snake is found from sea level up to at least
1600 m altitude. It inhabits lowland tropical wet
and dry forests, tropical wet montane forests, and
rubber plantations. Oligodon purpurascens is
rarely found outside of forested areas (but was dis-
covered in demarcation hedges at Bundu Tuhan).
This slow, sluggish, terrestrial, and secretive spe-
cies spends much of its time beneath rocks, among
the bases of plants, and under decaying vegeta-
tion and logs. It is nocturnal and feeds mainly on
the eggs of frogs and lizards, but also preys upon
tadpoles and frogs. O. purpurascens has a gentle
disposition and is entirely harmless. Hatchlings
bite fairly readily.
MALKMUS (1996c) found a specimen on Mt. Kina-
balu in semi-open cultivated terrain. Another snake
was discovered on a path cutting through diptero-
carp montane forest at about 5.30 p.m. (MALKMUS
1994b).
Distribution
This species ranges from the Malay Peninsula to
the Sunda islands of Java, Sumatra, and Borneo.
Mt. Kinabalu: Sayap, Bundu Tuhan (1350 m),
Poring: Hot Springs region, Sg. Kipungit I (550 m).
Fig. 345 Oligodon purpurascens from West Malaysia.
© U. Manthey
352
Oligodon vertebralis vertebralis
(GÜNTHER, 1865)
Simotes vertebralis G ÜNTHER , 1865: 91 – Type locality:
Bandjermasin, south Borneo
Simotes vertebralis, MOCQUARD 1890a: 137
Oligodon vertebralis, BARTLETT 1895c: 82; SHELFORD 1901a:
61; DE ROOIJ 1917: 133; SMITH 1931: 32; MANTHEY 1983: 28;
E BENHARD & S JÖGREN 1984: 24; S TUEBING 1991: 335;
MALKMUS 1994b: 247
Taxonomic comment: STUEBING & INGER (1999)
indicate that this taxon might be conspecific with
Oligodon everetti, but in the absence of detailed
investigations, we continue to consider this to be
valid species at this point of time.
Diagnosis
Very similar to O. everetti differing from it only by
the divided anal scute and a spotted throat. 15 rows
of dorsal scales at mid-body. The largest of the few
known specimens measures 34.5 cm.
Description
One preocular; 2 postoculars; 1 small loreal; 1+2
temporals; 7 supralabials, the 3rd and 4th touching
the eye; 15 dorsal scale rows at mid-body; 154 ven-
trals; anal scute divided; 54 subcaudals, divided.
Brown above, with small yellow, black-edged spots,
largest along the vertebral line; head yellowish, with
two brown, black-edged bands, the anterior one
passing through the eyes; the posterior one across
the frontal; lower surfaces yellowish; throat spot-
ted with brown.
Like all its congeners this species is oviparous, but
no further details are known.
Ecology/Ethology
Owing to the very low number of specimens found
nothing is known about the ecology of this spe-
cies.
Distribution
The snake is a Bornean endemite which has to
date only been found on Mt. Kinabalu and in
Banjarmasin (Kalimantan). As far as Mt. Kina-
balu is concerned more precise locality data is
wanting.
 Colubridae

Genus Opisthotropis GÜNTHER, 1872
The genus Opisthotropis has a vast distribution
range from China through Vietnam, Kampuchea,
Laos, Thailand, Malaysia, Indonesia, to the Phil-
ippine Islands.
Small, slender, harmless colubrids. Body round in
cross-section; head small, weakly distinct from
body; nostrils directed upwards; eyes small, pupils
round; one, rarely two loreals; dorsals keeled, in
17 or 19 rows; tail long, tapering to a fine point;
subcaudals divided; ventrals rounded.
These snakes have adapted to a life in freshwater,
and are particular to rapidly flowing mountain
streams. They spend most of their time below the
water surface and are normally found below cas-
cades where they hide beneath rocks and boulders.
They feed on small fish, frogs, and tadpoles. As far
as is known, the snakes of the genus Opisthotropis
are live-bearing.
Altogether 14 species are known, of which only
one occurs on Borneo and also on Mt. Kinabalu.
Opisthotropis typicus (MOCQUARD, 1890)
Helicopsoides typicus MOCQUARD, 1890a: 141, Pl. IX, fig. 3a-c
– Type locality: Mt. Kinabalu
Helicopsoides typicus MOCQUARD, 1890b: 154
– Type locality: Mt. Kinabalu
Opisthotropis typica, BOULENGER 1891b: 343; Fig. 346 Opisthotropis typicus from Ulu Skrang, Lanjak-Entimau
BOULENGER 1893b: 285; SHELFORD 1901a: 58; Wildlife Sanctuary, Sarawak.
DE ROOIJ 1917: 52; SMITH 1931: 32; DE H AAS
1950: 531; MANTHEY 1983: 28; EBENHARD
& S JÖGREN 1984: 24; M ORI 1993: 67;
M ANTHEY & G ROSSMANN 1997: 374;
STUEBING & INGER 1998: 325, 1999: 176
Opisthotropis typicus, STUEBING 1991: 349,
1994: 934; MALKMUS 1994b: 247
Diagnosis
Moderately large, max. total length 50 cm; noted
for its depressed, “shovel-like” snout and very nar-
row internasals.
Description
Flat, “shovel-like” snout; fairly small eyes, sepa-
rated by small scales from the supralabials; dorsal
scales in 19 rows, strongly keeled with small
“knobs” on the keels; head scales slightly striated;
the first supralabials much longer than wide, the
last ones divided into larger upper and smaller lower
parts; internasals triangular and very narrow ante-
riorly, so that the nasals are sometimes in contact
with each other; prefrontals short; supraoculars
small, rarely divided; 1 loreal, occasionally divided;
2 preoculars; 2 postoculars; 2 suboculars; 1+2 tem-
porals; 11-12 supralabials, none of which touch-
ing the eye; 10 sublabials; 160-176 ventrals; anal
scute divided; 82-96 subcaudals, divided.
The entire upper side is olive brown to brown, the
ventral side yellowish.
Ecology/Ethology
Opisthotropis typicus live in the surrounds of wa-
ter bodies for which the keeled dorsal scales are an
indication for an aquatic biology. MORI (1993)
found a specimen in the evening which was creep-
ing on a boulder on a riverbank. It escaped into the

Taxonomic comments: The genus

name Opisthotropis is masculine with
the type species being O. ater (not O.
atra, as often quoted). The species
© R.B. Stuebing

name must therefore be given as O.

typicus since “typicus” is an adjective.
See also the taxonomic comment un-
der Hydrablabes praefrontalis.

353
Colubridae
water and hid beneath a loose rock. Other specimens
were discovered in primary rain forest and in sec-
ondary forest on the edges of ponds and a river,
respectively (STUEBING & INGER 1998). It is a rare
species which has been recorded from altitudes
between sea-level and 900 m a.s.l. (Kiau).
Distribution
Opisthotropis typicus is endemic to Borneo and for
a time was even considered endemic to Mt. Kina-
balu. It was only recently that it was also recorded
from other regions of Sabah, Sarawak, and from
Brunei (MORI 1993, DAS 1995, STUEBING & INGER
1998). Mt. Kinabalu: Kiau, Poring.
Genus Oreocalamus BOULENGER, 1899
The genus Oreocalamus is distributed in Peninsu-
lar Malaysia and on Borneo.
Small, terrestrial, harmless snakes closely resem-
bling those of the genera Calamaria and Macroca-
lamus. Head not distinct from body; eyes small,
pupils round. Like in Macrocalamus the first
supralabial touches the nostril; distinguished from
Calamaria and Macrocalamus by the presence of
two internasals and by its 17 rows of dorsals.
Fig. CIV Oreocalamus hanitschi (SMF 77913, Kaliman-
tan, Long Petah, 1172 m).
354
The reproductive biology and ecology are entirely
unknown.
Oreocalamus is a monotypical genus with its only
species also occurring on Mt. Kinabalu.
Oreocalamus hanitschi BOULENGER, 1899
Oreocalamus hanitschi BOULENGER, 1899: 453 – Type local-
ity: Mt. Kinabalu, 4200 feet
Oreocalamus hanitschi, HANITSCH 1900a: 70, 72; SHELFORD
1901a: 61; DE ROOIJ 1917: 141; LIM 1970; SMITH 1931: 11,
26, 32; DE HAAS 1950: 560; MANTHEY 1983: 30; EBENHARD &
SJÖGREN 1984: 24; STUEBING 1991: 349; MALKMUS 1994b: 247;
INGER & TAN 1996b: 69; MANTHEY & GROSSMANN 1997: 375;
STUEBING & INGER 1999: 178
Diagnosis
See generic diagnosis. A small fossorial snake with
a maximum length record of 57 cm.
Description
One loreal; 1 preocular; 1 postocular; 7-8 supra-
labials, the 4th and 5th touching the eye, the 7th is
the largest; 8 infralabials, of which the first 4 are
in contact with the anterior chinshields; 1+2
temporals; 17 dorsal scale rows along the entire
body, smooth; 125-132 ventrals; anal scute entire;
20-32 subcaudals, in pairs.
Sides and back blackish brown to blackish; dorsally
with darker scales which form some kind of a zigzag-
pattern and an almost continuous stripe on the low-
est dorsal scale rows; a thin line through the eye may
extend onto the body for a short distance; a V-shaped
marking on the occiput and nape; ventral side yel-
lowish white, sometimes with a few brown spots;
lateral margins of ventrals the colour of the dorsum;
lower side of tail brown with three darker stripes.
Ecology/Ethology
This species has been found at altitudes between
1000 and 1800 m. LIM (1970) collected a specimen
from under a piece of wood lying across a stream.
Distribution
Peninsular Malaysia (Pahang) and Borneo (Kali-
mantan, Long Petah, 1172 m; Sabah, Mt. Lumaku;
Sarawak, Kelabit Plateau). Mt. Kinabalu: Marei
Parei, Lumu Lumu (LIM 1970), Mesilau (1800 m),
at 4200 feet (HANITSCH 1900a).

Genus Pareas WAGLER, 1830
The genus Pareas ranges from eastern India,
through the south of China and Japan, south through
Indochina to Java and Borneo. Some islands of the
Philippines are also inhabited.
Small, slender, harmless snakes. Body distinctly
compressed laterally; head short and wide, dis-
tinctly set off from the body; eyes large, pupils
vertical; scutellation of chin asymmetrical lacking
a mental groove. In contrast to the genus Aplopel-
tura there are 15 (rarely 17) rows of dorsals at mid-
body, in contrast to Internatus two pairs of chin-
shields, but no pair of pre-chinshields, and the 2nd
and 3rd sublabials are enlarged. Subcaudals divided,
anal scute entire.
The crepuscular and nocturnal snakes are highly
specialised predators of slugs and snails. The lat-
ter are extricated from their shells by means of fangs
which are positioned in the front of a narrow, firm
lower jaw.
Tab. 50 Distribution of Pareas-species on Borneo.
Pareas nuchalis (BOULENGER, 1900)
Amblycephalus nuchalis BOULENGER, 1900a: 185, pl. XVII: figs.
1, 1b – Type locality: Matang, Kidi District, Sarawak
Pareas nuchalis, MALKMUS 1994b: 240, 247, 1996c: 55
Diagnosis
Chinshields asymmetrical; high counts of ventrals
(195-213) and subcaudals (105-118); supralabials 2
and 3 enlarged; longest specimen recorded 71.5 cm
(MALKMUS 1994b).
Description
Prefrontal shield in contact with the eye; 2 pre-
oculars; 2 postoculars; 3 narrow suboculars; 8-9
supralabials, none of which touch eye; 1-3 loreals;
3 anterior temporals; 15 dorsal scale rows at mid-
body; dorsals weakly keeled; vertebrals enlarged;
Colubridae
All species of this genus reproduce by laying eggs.
The taxonomy within this genus is hotly debated.
Following RAO & YANG (1992), 15 species are in-
cluded, one or two of which occur on Borneo, one
on Mt. Kinabalu.
Taxonomic comment: The existence of Pareas
carinatus on Borneo is doubted by STUEBING &
INGER (1999). Unfortunately no specimens were re-
examined by these authors. Although the present
author shares this opinion, a key to distinguish both
species is provided here.
Key to the Bornean species of Pareas
1 195-213 ventrals; 105-118 subcaudals ................
.............................................................. nuchalis
• 161-189 ventrals; 53-99 subcaudals ....................
............................................................. carinatus
195-213 ventrals; anal scute entire; 105-118 sub-
caudals, in two rows.
Dorsal side of body sand-coloured,, with mid-
dorsally connected, slightly backward pointing,
narrow bands spaced irregularly at 2-5 scales wide;
bands with diffuse contours, consisting of more
densely arranged clusters of tiny speckles, the lat-
ter are - more widely scattered - present on all dor-
sal scales; this pattern fades increasingly on the
sides of the tail. Pileus dark umber-coloured; sides
of head ochre; a black line, curving back behind
the upper edge of the eye, in a narrow V-shaped
fork to angle of mouth, then back to original level
of the horizontal stripe, widening to an X-shaped
marking in the nuchal region; the V- or U-shaped
branches of the fork meet 5 scales behind the
parietals and descend as large, deep black bands to
the margins of the ventrals; crossing the 6th and 7th
355
Colubridae
supralabials. An obliquely arranged, distally taper-
ing, i.e. thorn-shaped, dense cluster of speckles
from lower edge of eye to lip; lower sides of head
and ventrals whitish yellow, all over with tiny black
speckles; iris golden brown with darker gradations.
Like all species of Pareas, P. nuchalis is probably
oviparous.
Ecology/Ethology
A snake of the lowlands, with the highest record
stemming from 850 m above sea-level. It is noc-
turnal and arboreal, presumably feeding on slugs
and snails.
MALKMUS (1994b) found a specimen 2 metres up
in a bush in a bamboo forest at around 9.00 p.m.
sympatrically with Internatus laevis. He reported
about an interesting resting position of this particu-
lar specimen. The snake would lie curled up in its
shelter in 4-5 circular coils which are arranged
partly above each other, but partly also next to each
other with portions of the belly turned upwards.
Fig. CXV Pareas nuchalis from Sg. Langanan, 850 m.
356
© R. Malkmus
Fig. 347 Pareas nuchalis from Sg. Langanan, 850 m.
The sides of the body were strongly compressed.
The tail would form the lowermost coil, and the
head and neck were concealed below the upper or
mid-level body coils. The snake maintained this
peculiar posture for another minute even after it
had been taken out of its shelter. This behaviour
may indicate that internodes of bamboo serve as
primary day shelters.
Distribution
Endemic on Borneo. Mt. Kinabalu: ca. 300 m from
the Sg. Langanan (850 m a.s.l.).
Genus Psammodynastes GÜNTHER, 1858
This genus is distributed from Nepal and Assam
(India) through China in the north, to Sulawesi and
Flores in the east. It also inhabits many of the Phil-
ippine islands.
Small, slender, rear-fanged. Body elongate and dis-
tinct from body; snout with angular edges; eyes
large, pupils vertical; a single nasal shield; frontal
very narrow; dorsal scales smooth, without apical
pits, in 17-19 rows at mid-body; ventrals rounded,
subcaudals divided.
Nocturnal and diurnal, these are arboreal snakes.
They inhabit the lowlands and montane regions up
to about 2000 m altitude. The venom of these snakes
does normally not have serious effects on humans.

Tab. 51 Distribution of Psammodynastes-species on Borneo.
Their diet consists chiefly of lizards, but frogs are
also taken occasionally.
The females give birth to fully developed babies.
Two species are known, both live on Borneo, one
on Mt. Kinabalu.
Key to the Bornean species of
Psammodynastes
1 3rd infralabial not bordering the mental groove, 3
pairs of chin shields; dorsal side of head with dark
and light longitudinal streaks ...... pulverulentus
• 3rd infralabial bordering the mental groove; 2 pairs
of chin shields; dorsal side of head not streaked
................................................................... pictus
Psammodynastes pulverulentus
(BOIE in BOIE, 1827)
Psammophis pulverulenta H. BOIE in F. BOIE, 1827: 547 – Type
locality: Java
Psammodynastes pulverulentus M OCQUARD 1890a: 143;
SHELFORD 1901a: 65; DE ROOIJ 1917: 202; SMITH 1931: 10,
32; MANTHEY 1983: 30; E BENHARD & SJÖGREN 1984: 24;
STUEBING 1991: 335; MALKMUS 1994b: 247
Psammodynastes spec., MALKMUS 1996c: 292
Diagnosis
Small with a distinctive head pattern (see below);
pupils vertical slits; 3rd sublabial not touching the
mental groove; 3 pairs of chinshields. Max. length
63 cm, average about 50 cm.
Description (RASMUSSEN 1975):
One loreal (occasionally split into several shields);
1-3 preoculars; 1-4 postoculars; 1-3 anterior
temporals; 7-9 supralabials, normally the 3rd to 5th
in contact with the eye; 7-8 sublabials; 17 (rarely
19) rows of dorsals at mid-body; 139-178 ventrals;
anal scute normally entire; 38-74 subcaudals.
Colubridae
Dorsal coloration ochre, reddish brown, dark
brown, greyish brown, grey, or almost black. Fe-
males with a series of large oval rufous or brown
spots on each side of the spine, these spots some-
times confluent to form crossbars over the spine;
belly densely peppered with brown, and with dark
brown spots or longitudinal lines; flanks often
ornamented with white streaks, and ocelli anteriorly.
Males with a more or less distinct, dark, wide stripe
on the back, involving the median five rows and
half of the sixth row of the dorsals; similar stripes
on the flanks, separated from the former by a nar-
row white streak; markings in the median stripe
V-shaped with indistinct edges; belly not so densely
peppered as in females, the specks often forming
2-4 longitudinal lines; ocelli not as distinct as in
females. Head with symmetrical longitudinal mark-
ings in both sexes, on each side a dark streak
through the eye, sometimes with a white streak
below.
Psammodynastes pulverulentus is live-bearing with
several litters possible in a year. A litter may con-
tain up to 10 babies which measure 148-178 mm
in length at birth.
Ecology/Ethology
This snake is found from sea level up to about 2000 m,
with a marked predilection for hilly areas above
Fig. 348 Psammodynastes pulverulentus from Sg. Ki-
pungit I, 580 m.
© M. Schroth
357
Colubridae
700 m altitude. It inhabits lowland tropical wet and
dry forests, tropical and subtropical wet montane for-
ests, bamboo forests, moist scrublands, marshes and
swamps, rice paddies, hedges, and gardens in sub-
urban areas. This agile, swift snake is usually found
on low vegetation in the immediate vicinity of for-
est streams, ditches, ponds, and other water bodies.
It is both diurnal and nocturnal, sometimes terres-
trial, but mainly arboreal, found from fully exposed
areas to thick vegetation. It feeds chiefly on lizards,
but also hunts for frogs.
Genus Pseudorabdion JAN, 1862
The genus Pseudorabdion is distributed from Thai-
land in the north through the Malay Peninsula to
the Philippines, Sumatra, Borneo, and Sulawesi in
the south.
Small, harmless snakes; body slightly compressed
dorsoventrally, head not distinct from body; eyes
small, pupils round; characterized by the absence
of a preocular shield (present in Pseudorabdion
longiceps) so that the prefrontal borders the eye.
5-6 supralabials; anterior temporals wanting; 15
rows of smooth dorsal scales throughout the length
of the body; anal scute entire; subcaudals usually
arranged in pairs (LEVITON & BROWN 1959).
These are terrestrial, fossorial species which favour
the vicinity of water. They are often found in leaf
litter or under dead wood. Considering their small
size their diet is probably based upon insects,
worms, small lizards, frogs and tadpoles. Their bi-
ology is poorly known. The snakes of this genus
are oviparous.
Altogether 12 species are recognized (BROWN,
LEVITON & ROGELIO 1999), 4 of which live on Bor-
neo, one on Mt. Kinabalu.
Tab. 52 Distribution of Pseudorabdion-species on Borneo.
358
Psammodynastes pulverulentus is well-known for
its aggressive disposition, striking fiercely and bit-
ing readily and viciously when handled.
Distribution
Psammodynastes pulverulentus occurs throughout
the entire range of the genus, from Nepal and As-
sam (India) to China in the north, and from Thai-
land to Sulawesi and Flores and the Philippines
in the south. Mt. Kinabalu: Kiau, Sg. Kipungit I
(580 m).
Key to the Bornean species of
Pseudorabdion
1 Loreal shield absent; prefrontal in contact with
upper labials ...................................................... 2
• Loreal shield present; prefrontal not in contact
with upper labials .............................................. 3
2 Preocular present; internasals not in contact with
upper labials ........................................ longiceps
• Preocular absent; internasals in contact with upper
labials ..................................................... collaris
3 More than 125 ventrals; more than 40 subcaudals
........................................................ albonuchalis
• Less than 120 ventrals; less than 30 subcaudals .
........................................................ saravacensis
Pseudorabdion collaris
(MOCQUARD, 1892)
Idiopholis collaris MOCQUARD, 1892: 191 – Type locality:
Sebruang River, Borneo
Idiopholis everetti SHELFORD, 1901b: 517 – Type locality:
Sawah, North Borneo (syn. after INGER & LEVITON 1966: 310)
Pseudorabdion collaris, INGER & LEVITON 1966: 310, fig. 2
 Colubridae

Taxonomic comments: The

author is not quite convinced
that Pseudorabdion everetti
is indeed a synonym of Pseu-
dorabdion collaris. This prob-
lem can, however, only be
solved by the examination of
more material so that the
nomenclatural concept sug-
gested by INGER & LEVITON
(1966), which still represents
the latest review of the system-
atics of these species, shall be
followed here.

Diagnosis
A small, fossorial snake; max.
20.4 cm in length; distin-
guished from other species of
its genus by the following
combination: no loreal shield,
no preocular shield, frontal in
contact with eye.

© R.B. Stuebing
Description
Supraoculars, loreals, preocu-
lars, and anterior temporals
absent; eye surrounded by 1 Fig. 349 Pseudorabdion collaris from Malutut, Sabah.
or 2 postoculars, the frontal,
1 prefrontal, and the 3rd and
4th supralabials; 5-6 supralabials, the 5. the largest, Genus Ptyas FITZINGER, 1843
separating the sole posterior temporal from the posto-
culars; 15 dorsal scale rows along the entire length
of the body; 116-134 ventrals; anal scute entire; 25- The genus Ptyas inhabits large parts of China and
41 subcaudals (BRONGERSMA 1946), usually in pairs. India. To the south it reaches Sulawesi. In addition to
Dorsally blackish brown, blackish grey, or black, iri- this, it occurs on several islands of the Philippines.
descent; a yellowish, whitish, or reddish collar be- Large, slender snakes; body muscular, slightly com-
hind the parietals, may be wanting; ventral side lighter. pressed laterally; head long, distinct from body;
A female contained three eggs. eyes large, pupils round; dorsal scales smooth or
keeled with grooves, a distinct vertebral row may
Ecology/Ethology be present or absent. Ventrals rounded; tail long;
These small colubrids favour primary and second- subcaudals in two rows.
ary forests where they live in the layer of leaf litter The species of this genus are diurnal and truly ag-
or find shelter beneath rocks and fallen tree trunks. ile. They are encountered in a multitude of habi-
tats and do not necessarily avoid cultivated areas.
Distribution An interesting behaviour can be observed when the
This species is endemic to Borneo. Mt. Kinabalu: snakes cross through high grass. In order to obtain
Kamborangoh Road (HQ-Collection). a better overview they lift their heads and anterior

359
Colubridae
portions of the body high above the ground. At the
same time they hiss loudly by pressing out air.
The snakes of the genus Ptyas feed on frogs, snakes,
lizards, birds, and particularly on rodents. They
reproduce by laying eggs.
Eight species are known, two or three of which live
on Borneo, one on Mt. Kinabalu.
Taxonomic comment: The genus Zaocys COPE,
1861, was synonymised with Ptyas by WALL (1923b)
and TAYLOR (1965). This arrangement has not been
commonly accepted. LAZELL et al. (1991), based
upon a study of hemipenial morphology, placed both
genera in the synonymy of a greatly expanded ge-
nus Coluber. According to WALLACH (pers. comm.),
data from the internal anatomy support the merging
of Zaocys with Ptyas, but not with Coluber. We there-
Tab. 53 Distribution of Ptyas-species on Borneo.
Ptyas fuscus (GÜNTHER, 1858)
Coryphodon fuscus GÜNTHER , 1858: 112 – Type locality:
Borneo
Zaocys fuscus, SMITH 1931: 10; 32; EBENHARD & SJÖGREN 1984:
24; STUEBING 1991: 347; MALKMUS 1994b: 247
Taxonomic comment: It is obvious that STUEBING
& INGER (1999) confused Ptyas fuscus with P.
carinatus, so that the information provided by them
is left aside here.
Diagnosis
Most easily recognized by its lateral stripes on the
posterior portion of the body which are in stark
contrast with the light ventral side. Ptyas fuscus is
the only Bornean species with 16 rows of dorsals
at mid-body. Its maximum length is about 300 cm.
Description
One preocular; 1-2 small suboculars; 2 postoculars;
3 loreals; 2+2 temporals; 9 supralabials, the 5th and
360
fore recognize Ptyas as including the genus Zaocys
here (see also DAVID & VOGEL 1996).
STUEBING & INGER (1999) doubted that Ptyas korros
would occur on Borneo as this species had never
been confirmed for about 100 years. Notwithstand-
ing, Ptyas korros is considered in the following key.
Key to the Bornean species of Ptyas
1 An odd number of dorsal scale rows at mid-body
.................................................................. korros
• An even number of dorsal scale rows at mid-body
........................................................................... 2
2 More than 160 subcaudals, dorsals smooth ........
.................................................................. fuscus
• Less than 120 subcaudals, dorsals on the median
2-4 rows keeled ................................... carinatus
6th in contact with the eye; 16 rows of dorsal scales
at mid-body; dorsals smooth; 183-198 ventrals; anal
scute divided; 155-179 subcaudals in two rows.
Olive brown to brown dorsally, occasionally with
a red, black-bordered vertebral stripe and oblique
bars; a black lateral stripe from the mid-portion of
the body to the tail which touches the margins of
the ventrals and subcaudals and widens posteriorly
so that the tail may be entirely black; undersides
pale yellow.
Oviparous.
Ecology/Ethology
This snake is found in low and hilly areas where it
inhabits wet tropical forests, shrublands, and plan-
tations. It is usually found in dense, tangled vegeta-
tion in the immediate vicinity of water. Some indi-
viduals were found swimming and others up in trees
as high as 6 metres above the ground. Little is known
about its biology, except that it feeds mainly on
amphibians; BATCHELOR (1958) reported a specimen
feeding on a small chicken.

Fig. 350 Ptyas fuscus from Poring.
Distribution
This species lives in Peninsular Malaysia and on
Borneo, Sumatra, as well as on some of surround-
ing smaller islands. Mt. Kinabalu: Kiau, Poring.
Fig. 351 Ptyas fuscus from Niah Caves NP, Sarawak.
© S. Wilson
Colubridae
© A. Götzke
Genus Rhabdophis FITZINGER, 1843
The genus Rhabdophis is distributed from the
southwestern parts of Siberia and Japan, south
through China, to Borneo, Flores, Sulawesi, and
the Philippine Islands.
Moderately large, slender, aquatic colubrids; dorsals
keeled, in 17, 19, or 21 rows, without apical pits; head
slightly set off from body; eyes fairly large, pupils
round; two substantially enlarged fangs in the rear of
the upper jaw, which may transfer venomous saliva
into the blood stream when allowed to bite. Depend-
ing on the species the venom varies in toxicity and
may have a seriously dangerous effect on humans.
Internasals broad anteriorly, nostrils positioned later-
ally; tail long; anal scute and subcaudals divided.
The di- and nocturnal snakes occasionally exhibit
amphibious habits. All species are terrestrial and
feed for a major part on frogs, their tadpoles, and
fish. They reproduce by laying eggs.
361
 Colubridae
Tab. 54 Distribution of Rhabdophis-species on Borneo.
At least 15 species are currently known, four of
which live on Borneo, two on Mt. Kinabalu.
Taxonomic comment: Whether Rhabdophis sub-
miniatus occurs on Borneo is at present unclear. It
was listed by STUEBING (1991), but omitted by
STUEBING & INGER (1999) without any comment. It
is included in the key below.
Key to the Bornean species of
Rhabdophis
1 2 anterior temporals .......................................... 2
• 1 anterior temporal ............................................ 3
2 No black streak on the upper lips .... chrysargos
• A black streak from the eye across the upper lip
.................................... subminiatus subminiatus
3 Upper lips bright vermilion ............ murudensis
• A yellow, black-edged streak on the upper lip ...
..................................................... conspicillatus
Fig. 352 Rhabdophis chrysargos from Sumatra.
© S. Manthey
362
Rhabdophis chrysargos (SCHLEGEL, 1837)
Tropidonotus chrysargos SCHLEGEL, 1837: 312, pl. 12: fig. 6 &
7 – Type locality: “Forêts primitives à la cime de la montagne
Magmedon”, Java (= primary forests on the top of Mt.
Megamendung, near Bogor, Java)
Tropidonotus maculatus var. torquatus MOCQUARD, 1890a: 139
– Type locality: Kina Balu (syn. after BOULENGER 1893b: 258)
Tropidonotus chrysargus, DE ROOIJ 1917: 89
Natrix chrysarga, SMITH 1931: 10, 32; EBENHARD & SJÖGREN
1984: 24;
Natrix chrysarga chrysarga, DE HAAS 1949: 79, 88
Rhabdophis chrysarga, MANTHEY & DENZER 1982: 14; MANTHEY
1983: 30; MALKMUS 1985: 9, 1987: 287, 1989: 198, 1991a:
33, 1992a: 134, 1994b: 239; STUEBING 1991: 350; STUEBING
& INGER 1999: 180
Rhabdophis chrysargus, MALKMUS 1994b: 247, 1996c: 292
Diagnosis
Easily identified by its particular pattern on the neck
described below. 2 anterior temporals; max. length
98 cm.
Description
One loreal; 1 (rarely 2) preoculars; 3 (rarely 2 or 4)
postoculars; 8-9 supralabials, the 3rd to 5th or the 4th
to 6th in contact with the eye; 2+2 or 2+3 tempo-
rals; 19 dorsal scale rows at mid-body; dorsals
strongly keeled except for those in the first row;
133-184 ventrals (STEJNEGER 1922); anal scute di-
vided; 38-101 subcaudals, divided. It is apparent
that the subcaudal counts given in the literature
include data of at least one specimen with a muti-
lated tail. The variability range of this trait should
actually be substantially narrower.
Upper side of head medium, dark, or olive brown to
almost black; neck red with dark spots; a narrow
yellowish stripe on the supralabials continues more
“obscurely” up onto the nape; body olive grey with
dorsolateral light, dark-margined bars; ground col-
our of body changing from anteriorly to posteriorly,

i.e. yellow turns into brown, or red turns into brown,
or green becomes reddish brown; ventral side whit-
ish, margins of ventrals spotted with black. Juve-
nile specimens exhibit the same markings as the
adults, but more distinctly so. In addition, they
possess three light spots on the occiput; head dark,
bright yellow or white stripe from the labial shields
up onto the nape, where the stripe fuses with its
counterpart from the other side to form a wide V.
According to DE HAAS (1949), specimens from Mt.
Kinabalu have a tendency towards substantially
darker colours.
Clutches of 3-10 eggs (19.5-33.5 × 12-21 mm;
2.22-4.06 g) several times per year. Incubation
takes 51-61 days, hatchlings measure 148-220
mm in length. A case of Amphigonia retardata is
known.
Ecology/Ethology
Rhabdophis chrysargos is found from sea level up
to about 1600 m altitude, but is more common in
hilly, rugged areas and mountainous regions above
500 m. It inhabits lowland tropical wet forests,
tropical and subtropical wet montane forests, bam-
boo forests, forested mesic scrublands, marshes and
swamps, and occurs also near human habitations.
This snake is often found in the immediate vicin-
ity of streams and other water bodies. It is both
diurnal and nocturnal, mainly terrestrial, but can
sometimes also be seen basking on low vegetation.
It is usually found in thick, riparian vegetation close
to water, feeding on amphibians, small mammals,
lizards and birds.
This snake is widespread, generally common, and
is one of the most common snakes on Mt. Kina-
balu. It is considered harmless, but bites of at least
one other species of Rhabdophis (i.e. R. tigrinus)
have caused fatal envenomations in humans so that
the present species must be considered potentially
dangerous.
The snake was found on foggy days on hedge-like
thickets of Dipteris and Gleichenia ferns where it
proved to be a skilful climber (MALKMUS 1985).
Several specimens were discovered on roadsides
with a vegetation of grasses and ferns (Headquar-
ters), on grassy, more sun-exposed spots in the
rainforest not far from the Sg. Liwago, and on a
forested slope far from water east of the Sg. Liwago
Colubridae
© R. Malkmus
Fig. 353 Rhabdophis chrysargos from Poring.
(MALKMUS 1987). One snake was discovered in
primary rainforest (MALKMUS 1989), three others
along the Sg. Silau-Silau in the forenoon hours,
and another one was observed in dipterocarp for-
est between Sg. Kipungit II and Sg. Langanan in
the afternoon (MALKMUS 1992a).
Distribution
This species ranges from China in the north, south
to the Philippines and Sulawesi. Mt. Kinabalu:
Kiau, Sg. Langanan, Lobang (HAAS 1949), Head-
quarters, bank of the Sg. Liwago, east of the Sg.
Liwago, Kamborangoh Road (1600 m), Sg. Silau-
Silau (1450-1550 m), between Sg. Liwago and Sg.
Silau-Silau, Sg. Kipungit II,
Rhabdophis conspicillatus
(GÜNTHER, 1872)
Tropidonotus conspicillatus GÜNTHER, 1872: 596, fig. 4 – Type
locality: Matang, Sarawak, Borneo
Tropidonotus conspicillatus, HANITSCH 1900a: 70
Rhabdophis conspicillatus, STUEBING 1991: 350; MALKMUS
1994b: 247
Diagnosis
Easily identified by its particular head pattern de-
scribed below. Only one anterior temporal. Max.
about 40 cm in length.
Description
One loreal; 1 (rarely 2) preoculars; 2-3 postocu-
lars; 1+2 or 1+3 temporals; 7-8 supralabials, the
363
 Colubridae
3rd through 5th, or the 3rd and 4th, in contact with the
eye; 19 rows of keeled dorsals at mid-body, lowest
row may be smooth; 136-152 ventrals; anal scute
divided; 36-60 subcaudals, in pairs (DE ROOIJ 1917,
DE HAAS 1949, STEJNEGER 1922).
Brown or reddish brown above, with a dark network
and two longitudinal series of yellow spots or trans-
verse lines; head with small yellow, dark-edged
spots; a yellow, black-edged streak on the upper lip,
another from the eye to corner of the mouth; usually
a pale transverse mark on the neck, narrow dorsally,
widening on the sides, where it connects with the
coloration of the lower side. Lower surface yellow,
red or pink, uniform, or dotted with brown. Young
animals more vividly coloured.
Probably oviparous just like the other species of
the genus
Ecology/Ethology
This snake is found at low elevations where it in-
habits tropical wet forests. Little is known of its
biology. One specimen was collected from the dry
leaf litter in the forest floor in Sarawak (DAVID &
VOGEL 1996) so it is probably a terrestrial forest-
dweller. Rhabdophis conspicillatus is active at
night. There is no information about its diet.
Fig. 354 Rhabdophis murudensis from near Kamborangoh Road.
© R. Gerhardt
364
This snake is uncommon in some areas, and com-
mon in others. It is regarded as harmless, but its
bite is actually potentially dangerous.
Distribution
This species lives in Peninsular Malaysia, on
Sumatra, Borneo, and some smaller Indonesian
islands. Mt. Kinabalu: near Tampassuk River
(HANITSCH 1900a), Sg. Mamut/Poring.
Rhabdophis murudensis (SMITH, 1925)
Natrix murudensis SMITH, 1925: 5 – Type locality: Mt. Murud,
Borneo, 5500-6000 feet
Natrix murudensis, SMITH 1931: 10, 12, 26, 32; DE HAAS 1949:
82, 95; EBENHARD & SJÖGREN 1984: 24
Rhabdophis chrysarga, MANTHEY & DENZER 1982: 12 fig., 14
Natrix (inc. sed.) murudensis, MANTHEY 1983: 28
Rhabdophis murudensis, MALKMUS 1994b: 247; INGER & TAN
1996b: 65; STUEBING & INGER 1999: 184
Diagnosis
Easily recognized by its bright red upper labial
shields. One anterior temporal. Max. length about
100 cm.
Description
One loreal; 1-2 preoculars; 3 posto-
culars; 1+2 temporals; 9 supralabi-
als, the 4th to 6th touching the eye; 19
dorsal scale rows at mid-body, dor-
sals strongly keeled except in the low-
est row; 176-187 ventrals; anal scute
divided; 63-97 subcaudals in two
rows.
Dorsum grey with short, indistinct,
black crossbars; a row of light spots
associated with the ends of these dark
markings appears at mid-body and
runs down the back, converging to a
single row anterior to the vent and
extending to the end of the tail; head
and nape brown, upper lip bright ver-
milion; eye brown, with a light green
segment in the upper portion; part of
the neck bright reddish orange with
regularly arranged black spots; belly

greyish yellow, with longitudinal series of small
black spots, best marked towards the outer margins
of the ventrals; tail dark grey below.
Ecology/Ethology
Rhabdophis murudensis has been found at eleva-
tions between 1000 and 2500 m above sea level. It
thus appears to be strictly submontane. One speci-
men was kept alive on a diet of frogs.
This is the only species of snake which ascends up
into the fog forest region of Mt. Kinabalu. No fur-
ther details are available on its biology.
Genus Sibynophis FITZINGER, 1843
The genus Sibynophis has a wide-ranging distribu-
tion, extending from Sri Lanka and the Indian sub-
continent, north to Nepal, central, eastern and south-
ern China, Japan, through Southeast Asia, and the
Philippines.
Small, slender; head moderately distinct from body;
pupils round; 17 rows of dorsals at mid-body,
smooth, with tubercles, but without apical pits. 8-
10 supralabials, the 4th to 6th touching the eye. Anal
scute and subcaudals divided; hemipenis entire.
Ground-dwelling, diurnal snakes, their diet consists
primarily of lizards, but also includes frogs and
insects. The species of this genus are oviparous.
Altogether six species are known, one occurs on
Borneo and on Mt. Kinabalu.
Sibynophis melanocephalus
melanocephalus (GRAY, 1835)
Lycodon melanocephalus GRAY, 1835 (in 1832b-1835): pl. 83:
fig. 2 – Type locality: Not given; subsequently designated
as “India“ by GÜNTHER (1858a: 28, in error), and “Singapore?”
by BOULENGER (1893a: 186)
Sibynophis geminatus, DE ROOIJ 1917: 54, fig. 32; SMITH 1931:
10, 25, 32; MANTHEY 1983: 30
Sibynophis melanocephalus, MORGAN 1973; E BENHARD &
SJÖGREN 1984: 24; STUEBING 1991: 347; MALKMUS 1994b: 247
Diagnosis
Small slender; head round, hardly distinct; best
identified by the following combination of traits:
Colubridae
Presumably oviparous like the other species of this
genus, nothing precise has as yet become known
about the species’ reproductive biology.
Distribution
Rhabdophis murudensis is an endemite of Borneo.
Here, it was discovered only on Mts. Murud and
Kinabalu. Mt. Kinabalu: Kiau, Headquarters
(MANTHEY pers. comm.) Kamborangoh Road, Sum-
mit Trail (2500 m) (KNOWLES, pers. comm., at the
same time representing the maximum altitudinal
record for reptiles on Mt. Kinabalu).
pupil round, 1 loreal shield, and 17 rows of dorsals
at mid-body. Its colour pattern, described below, is
also characteristic. Max. length 90 cm.
Description (MORGAN 1973)
Nasal double, nostril in-between; 1 preocular; 2
postoculars; 1 loreal; 8-10 supralabials, normally
the 4th to 6th touching the orbit; 2+2 temporals; 8-
10 sublabials; 17 rows of dorsals throughout length
of body, smooth; 128-177 ventrals; anal scute di-
vided; 87-149 subcaudals.
Head reddish brown, olive brown, or yellowish
brown, with well-defined, subcircular spots of light
pigment randomly distributed over the cephalic
scales; labials with or without darker mottling;
nuchal area reddish brown, or yellowish brown,
with light, lateral blotches or a streak; dorsal as-
pect of body and tail reddish brown or olive brown,
with light stripes on the upper half of dorsal scale
row 7; usually with a series of alternating dark bars,
tapering to spots on the tail, extending into the lat-
eral light stripes. Chin, throat, ventrals, and subcau-
dals yellow; a series of dark spots at the edges of
the ventrals and subcaudals; a row of mid-ventral
flecks present or not.
Oviparous.
Ecology/Ethology
The snake is found in lowland areas, but it is princi-
pally an inhabitant of hilly and mountainous regions,
at elevations up to 1000 m. It occurs in lowland tropi-
cal wet forests, tropical and subtropical wet montane
forests, bamboo forests, plantations, and in cultivated
areas. Sibynophis m. melanocephalus is a secretive,
365
 Colubridae
terrestrial snake, preferring tangled vegetation along
rivers, moist bushes and clumps of bamboo. It is
diurnal and seems to feed on forest floor skinks,
frogs and insects, especially grasshoppers.
Genus Stegonotus
DUMÉRIL & BIBRON, 1853
The genus Stegonotus is native to the islands of the
Indo-Australian Archipelago between Borneo and
New Guinea. It also inhabits the Philippines.
Small to medium-sized; body long, round or
slightly compressed in cross-section; head distinct
from body; eyes moderately large or small, pupils
vertical-elliptical; dorsals usually with apical pits,
in 15, 17, or 19 rows, lack keels; ventrals obtusely
angled laterally; tail fairly long; subcaudals in 1-2
rows.
Nocturnal terrestrial snakes, they are partly semi-
fossorial. They are found in a multitude of habitats
Fig. 355 Stegonotus borneensis from the Headquarters.
© A. Nöllert
366
Distribution
Sibynophis m. melanocephalus is distributed from
the Malay Peninsula in the north to Sumatra and
Borneo. Mt. Kinabalu: Kiau.
ranging from rain forests to cultivated land. The
snakes of the genus Stegonotus feed on reptile eggs,
small frogs, lizards, snakes, and small mammals.
They are oviparous, producing clutches of 6 to 12
eggs.
Ten species have been described, one species oc-
curs on Borneo and on Mt. Kinabalu.
Stegonotus borneensis INGER, 1967
Stegonotus borneensis INGER, 1967: 82 – Type locality: Nanga
Tekalit, Kapit District, Third Division, Sarawak
Stegonotus borneensis, E BENHARD & S JÖGREN 1984: 24;
MALKMUS 1985: 10, 1987: 288, 1989: 198, 1991a: 34, 1992a:
134, 1994b: 239, 247, 1996c: 292; STUEBING 1991: 347; INGER
& TAN 1996b: 70; MANTHEY & GROSSMANN 1997: 393

Diagnosis
Unicoloured, almost dark brown to black; pupil ver-
tical-elliptical; Each ventral scale with an obtuse,
lateral angle; vertebrals enlarged. Max. total length
130 cm.
Description
Two preoculars; 2 postoculars; 1 loreal; 9 supra-
labials, the 4th and 5th in contact with the eye; 9-10
sublabials; 2+3 temporals; 17 dorsals at mid-body;
vertebrals enlarged; 194-233 ventrals; anal scute
entire, 60-79 subcaudals in pairs.
Head and body greyish black dorsally and later-
ally; supralabials grey with a pinkish tinge; chin
grey with a hint of brown; ventrals light grey, each
with a blackish band anteriorly.
Oviparous.
Ecology/Ethology
This species lives in the rain forests and is predomi-
nantly terrestrial.
MALKMUS (1994b) found a specimen which was
climbing confidently in branchwork. When it felt
threatened it struck out quickly, but did not attempt
to bite. During its transport it strangled an 8,5 cm
large Cyrtodactylus baluensis which caused the
expulsion of two eggs in the latter.
Another, semi-adult specimen was discovered at
around 8.00 p.m. as it crept with very slow mo-
tions over the forest floor (MALKMUS 1989). When
it was picked up, it contracted firmly around the
hand and produced a scent reminiscent of a billy
goat.
One specimen was located beneath a sun-exposed
slab of rock on a grassy roadside above a trickle of
spring water (MALKMUS 1991a, 1992a). When un-
covered, it hid its head under its body coils. It was
due to shed very soon.
Another specimen was found very active at about
9.00 p.m. during a heavy downpour (MALKMUS
1996c).
Distribution
This species has to date only been found in Sabah
and Sarawak. Mt. Kinabalu: Bundu Tuhan (1370
m), Sg. Silau-Silau (1450-1500 m), Headquarters
exit (1500 m), Kamborangoh Road (1500-1750 m),
golf course/Mesilau (1750 m).
Colubridae
Genus Stoliczkia JERDON, 1870
The genus Stoliczkia is native to Sabah on Borneo
and the Khasi Hills in India.
Moderately large, harmless colubrids. Head distinct
from body; body slender, compressed laterally; eyes
small, sometimes prominent, pupil round or verti-
cal-oval; nostrils pointed forward; tail long;
Dorsalls strongly keeled, elliptical, juxtaposed, in-
creasing in size towards the venter. A single nasal
scale. Ventrals well-developed; subcaudals entire.
These are terrestrial snakes whose biology is en-
tirely unknown.
Two species have been described, of which one lives
on Borneo and on Mt. Kinabalu.
Taxonomic comment: The generic name Stoliczkia,
described by JERDON in 1870 and named in honour
of the famous naturalist Dr. Stoliczka, was emendated
by BOULENGER (1899) to read Stoliczkaia. He rea-
soned that the spelling Stoliczkia would contravene
grammar rules. Notwithstanding this, the original
spelling used by JERDON must be considered proper
according to the rules laid down by the Code
(I.C.Z.N. 1985) as it was used twice in this spelling
in the respective volume (once in the description and
again in the index) rendering a spelling error un-
likely [Art. 32 (c) (ii)]. For reasons unknown, JERDON
used the ending –ia to latinize the name Stoliczka,
perhaps following the classic Latin where the names
of persons often end in -ia or -ium. Furthermore the
name Stoliczkaia is preoccupied by Stoliczkaia
NEUMAYR, 1875 (Mollusca) and is therefore unavail-
able anyway (WILLIAMS & WALLACH 1989). The
proper spelling is thus Stoliczkia JERDON, 1870
(DAVID pers. comm.).
Stoliczkia borneensis (BOULENGER, 1899)
Stoliczkaia borneensis BOULENGER, 1899: 452 – Type locality:
Mt. Kinabalu, Borneo, 4200 feet
Stoliczkaia borneensis, HANITSCH 1900a: 70, 71; SHELFORD
1901a: 57; DE ROOIJ 1917: 45, Fig. 26; SMITH 1931: 32; DE
HAAS 1950: 530; MANTHEY 1983: 30; EBENHARD & SJÖGREN
1984: 24; STUEBING 1991: 329; MALKMUS 1994b: 247;
STUEBING & INGER 1999: 77
Stoliczkia borneensis, MANTHEY & GROSSMANN 1997: 394
367
 Colubridae
© I. Das
© P. Hoffmann
Diagnosis
Readily recognized by its exposed skin between
some dorsal scales. A row of small scales between
the prefrontals and the frontals and supraoculars.
Max. total length 75 cm.
Description
Internasals very narrow; prefrontals large, separated
by smaller scales from the frontal and the supra-
oculars; 2 or 3 preoculars; another 7-8 small scales
may occasionally surround the eye; nostrils large; 1
loreal; 10-11 supralabials, none of which, or the 6th
368
Stoliczkia borneensis
Fig. 356 (above) from Crocker Range NP, Sabah.
Fig. 357 (left) from the Headquarters, 1600 m.
and 7th, in contact with the eye; temporals very small;
dorsals in 30-35 rows, partly separated from each
other by uncovered skin; 205-210 ventrals; anal scute
entire; 124 undivided subcaudals.
Reddish to reddish brown, or grey to brownish black
above, with 3 longitudinal rows of large bluish,
blackish blue, or black spots; ventral side brown,
margins of ventrals yellowish.
Ecology/Ethology
Nothing is known about the biology of this spe-
cies, other than that it is a terrestrial mountain
dweller.
Distribution
This species is endemic to Sabah and has only been
found on Mts. Trus Madi and Kinabalu and the
Crocker Range. Mt. Kinabalu: 4200 feet (HANITSCH
1900a), View trail (1600 m).

Genus Xenelaphis GÜNTHER, 1864
The distribution of the genus Xenelaphis extends
from Myanmar and the Malay Peninsula to Sumatra,
Borneo, and Java.
Large, slender, harmless; body muscular, round;
head distinct from nape; eye fairly large, pupil
round; dorsals smooth, in 17 rows at mid-body;
vertebrals slightly enlarged, hexagonal; ventrals
rounded; tail long; subcaudals in 2 rows.
The rat snakes of the genus Xenelaphis inhabit low
and medium elevations up to about 1100 m a.s.l.
Fully grown specimens show a predilection for ro-
dents, such as mice, rats, and squirrels, but also
feed on fish, while juveniles prefer frogs and liz-
ards. Their biology is generally poorly known.
Tab. 55 Distribution of Xenelaphis-species on Borneo.
Xenelaphis ellipsifer BOULENGER, 1900
Xenelaphis ellipsifer BOULENGER, 1900a: 184, pl. XVI – Type
locality: Sarawak river
Xenelaphis ellipsifer, DE ROOIJ 1917: 95; EBENHARD & SJÖGREN
1984: 24; S TUEBING 1991: 329; MALKMUS 1994b: 247;
STUEBING & INGER 1999: 79
Taxonomic comment: STUEBING & INGER (1999)
doubted the occurrence of this species on Mt.
Kinabalu, but failed to provide a reason for their
doubts.
Diagnosis
Dorsal scales smooth, in 17 rows at mid-body. A
large species most easily recognized by the colour
pattern described below. Max. length recorded so
far 232 cm, but it may even grow larger according
to TWEEDIE (1983).
Description
One preocular; 1 large subocular; 2 postoculars; 1
loreal; 8-9 supralabials, either separated from the
eye by the subocular, or the 4th or 5th in contact
Colubridae
As far as is known to date, the species of this ge-
nus are oviparous.
Two species have been described, both inhabit
Borneo, and one occurs on Mt. Kinabalu.
Key to the Bornean species of
Xenelaphis
1 140-179 subcaudals, brown above with more or
less distinct black transverse bands anteriorly ....
........................................................ hexagonotus
• 134 subcaudals, body with 18 black-edged brown
streaks, separated by cream-coloured interspaces
.............................................................. ellipsifer
with the eye; 2+2 temporals; 17 rows of dorsal
scales; 186-203 ventrals; anal scute divided; 129-
134 subcaudals.
Cream-coloured above, with 18 to 20 large, ellipti-
cal or round, black-bordered, brown markings; sides
cream-coloured, slightly lighter than the dorsum,
spotted or marbled with black; head and nape pale
brown, sides of neck with interrupted, black, elon-
gate markings; upper labials yellow; posterior por-
tion brown with a black lateral stripe; lower side
uniformly yellow or cream-white.
Probably oviparous.
Ecology/Ethology
TWEEDIE (1983) stated that only six specimens were
known. Most of them seem to have been caught in
mountainous forest areas between 800 and 1100 m
above sea level. Nothing else is known about the
biology of this apparently very rare species.
Distribution
This species lives on Sumatra, Borneo, and in Pe-
ninsular Malaysia. Mt. Kinabalu: without precise
localities.
369
Colubridae
Genus Xenochrophis GÜNTHER, 1864
The genus Xenochrophis is native to the region
between Pakistan in the west, China in the north
and east, south to Sulawesi.
Moderately large, harmless water snakes. Body
muscular, round in cross-section; head long, nar-
row, distinct from body; eyes large, pupils round;
no substantially enlarged teeth; dorsal scales in 19-
17 rows at mid-body, 17-15 a single head-length
anterior to the anal scute. Dorsals keeled except
for the lowest rows; ventrals rounded; tail long; anal
scute and subcaudals divided, the latter at least
partially with pits; hemipenis forked.
The diurnal, highly aquatic snakes are renowned
for biting readily. Their diet consists of fish and
frogs and their tadpoles. They reproduce by laying
eggs, partly in large numbers.
Tab. 55 Distribution of Xenochrophis-species on Borneo.
Xenochrophis maculatus (EDELING, 1864)
Tropidonotus maculatus EDELING, 1864a: 203 – Type locality:
Martapura, near Bandjermasin, Borneo.
Tropidonotus maculatus, MOCQUARD 1890a: 138; SHELFORD
1901a: 58; DE ROOIJ 1917: 90
Natrix maculatus, SMITH 1931: 32
Natrix (inc. sed.) maculata, MANTHEY 1983: 28
Natrix maculata, EBENHARD & SJÖGREN 1984: 24;
Xenochrophis maculatus, STUEBING 1991: 351; MALKMUS 1994b:
247
Diagnosis
Moderately large; muscular; dorsal scales keeled.
Eye very large; 2 anterior temporals present; no
substantially enlarged teeth. Most easily identified
by the colour pattern described below. Max. length
about 100 cm.
Description
Eye very large; 1 loreal; 1 preocular; 3 (rarely 2
or 4) postoculars; 2+3 temporals (rarely 1+3); 9
370
Ten species are known, two of which live on Bor-
neo, both also on Mt. Kinabalu.
Taxonomic comment: Xenochrophis piscator was
listed by STUEBING (1991) as a third species of this
genus for Borneo, but was then again removed from
the list of native snakes by STUEBING & INGER (1999).
The latter conforms with the author’s opinion.
Key to the Bornean species of
Xenochrophis
1 All ventrals with dark margins; a dark lateral point
on the ventrals; upper side with no black triangles
and no red laterally ........................... maculatus
• Ventrals laterally with alternating black margins,
formed by the black lateral triangles; upper side
with lateral black triangles interspaced by red ...
..................................................... trianguligerus
(rarely 8 or 10) supralabials, the 4th to 6th (rarely
the 3rd to 5th or the 5th to 7th ) bordering the eye; 19
dorsal scale rows at mid-body, all dorsals
strongly keeled; 138-156 ventrals; anal scute di-
vided; 84-117 subcaudals (FUKADA 1964), in two
rows.
Olive to brown dorsally; 4 rows of small, square,
black spots, the interspaces filled with two longi-
tudinal rows of yellow spots; supralabials yellow
with black margins; head dark above, with black
spots; lower sides yellow, the ventrals bordered with
black and a black dot laterally.
Oviparous.
Ecology/Ethology
Xenochrophis maculatus is a snake of the plains,
inhabiting lowland tropical wet forests, marshes,
swamps, plantations, and rice paddies. It is invari-
ably found in the vicinity of water or in wet areas.
It is diurnal, terrestrial, and semiaquatic. Little is

Fig. 358 Xenochrophis maculatus from Sumatra.
known about its biology, other than that it feeds on
frogs, toads, and fish.
This snake is mild-mannered and docile, very sel-
dom attempting to bite.
Distribution
Xenochrophis maculatus is native to the Peninsu-
lar Malaysia, Borneo, Sumatra, and some smaller
Indonesian islands. Mt. Kinabalu: without precise
localities.
Colubridae
© M. Kunkel
Xenochrophis trianguligerus (BOIE, 1827)
Tropidonotus trianguligerus BOIE, 1827: 535 – Type locality:
Java
Tropidonotus trianguligerus, DE ROOIJ 1917: 84
Natrix trianguligera, EBENHARD & SJÖGREN 1984: 24
Natrix trianguligerus, SMITH 1931: 32
Xenochrophis trianguligera, STUEBING 1991: 351
Xenochrophis trianguligerus, MALKMUS 1994b: 247
Sinonatrix trianguligera, MALKMUS 1992a: 136
Sinonatrix trianguligerus, MANTHEY 1983: 30
371
 Colubridae

9 supralabials, the 4th to 6th

touching the eye; dorsals in
19 rows at mid-body (17
were quoted by M ALKMUS
1992a), keeled except for the
lowest row; 131-150 ventrals
(FISCHER 1885); anal scute di-
vided; 57-105 subcaudals, in
pairs.
Dark olive above, with small
black spots; a lateral series of
large, triangular, black spots,
the points reaching the ven-
© U. Manthey

trals, and sometimes forming

bands across the belly, sepa-
rated by red interspaces;
Fig. 359 Xenochrophis trianguligerus from Peninsular Malaysia. sometimes red spots inside
the black triangles; on the
posterior part of the body the
Diagnosis spots are indistinct; head dark; upper labials yel-
Moderately large; muscular; dorsal scales keeled. low or white, with black sutures; lower surface
Eye of normal size; 2 anterior temporals present; yellow, uniform, or the ventrals edged with black.
no substantially enlarged teeth. Most easily recog- Young animals are more vividly coloured.
nized by the lateral, dark-coloured triangles and 5-8 eggs (29-34 × 15-17 mm; 4.39-5.29 g). Incu-
the red interspaces. Record length 135 cm, but av- bation takes 59-60 days.
erage specimens measure only 80-90 cm in length.
Ecology/Ethology
Description Xenochrophis trianguligerus is found from sea level
Eye moderately large; 1 loreal; 1 (rarely 2) preo- to altitudes above 1350 m. It inhabits regions cov-
culars; 3-4 postoculars; usually 2+3 temporals; ered by lowland tropical wet forests, tropical wet
montane forests, and plantations, where it occurs
mostly in rivers, streams, marshes, swamps, flooded
Fig. 360 Xenochrophis trianguligerus from Peninsular rice paddies, and ponds. Xenochrophis triangu-
Malaysia.
ligerus is invariably found in the thick vegetation
immediately adjacent to water, or submerged in the
water among aquatic plants. It is diurnal, and largely
aquatic, although it is also at home on land in the
riparian vegetation. This snake feeds on amphib-
ians and fish.
This species is sometimes rather aggressive, strik-
ing and biting viciously when molested, but it is
harmless.

Distribution
Xenochrophis trianguligerus is distributed from
© U. Manthey

Laos, Cambodia, Myanmar and Vietnam in the

north, to Sulawesi in the south. Mt. Kinabalu:
Bundu Tuhan (1370 m).

372
 Crotalidae

Crotalidae
As far as Borneo is concerned, this family comprises small to medium-sized snakes, usually with short or
moderately elongated bodies and tails. The almost triangular-shaped head is distinctly set off from the
body and has small eyes with vertical pupils. The upper jaw is furnished with solenoglyphous fangs.
These are long, hinged, hollow teeth which are folded back parallel to the skull roof into protective skin
pockets when the mouth is shut. On opening it, they are erected up to a right angle by individual muscles.
The venom is injected through the ducts of the teeth. The potency of the venoms varies with the species;
some require immediate medical attention to avert fatalities, but the venom of several species remains
unknown.
The family is marked for the presence of a sensory organ whose presence is recognizable externally by
loreal pits. This organ is able to distinguish minute differences in temperatures and so enables the snake
to locate prey, predators, and sexual partners.
Pit vipers may be oviparous or ovoviviparous.

Key to the Bornean genera of the family Crotalidae

1 Gular and intersupraocular scales smooth ....... 2 A

• Gular and intersupraocular scales keeled (A) .....
................................................... Tropidolaemus
2 More than 40 subcaudals .............. Trimeresurus
• Less than 35 subcaudals ........................ Ovophis

Snakes of the genus Ovophis may lay eggs or give

Genus Ovophis
BURGER in HOGE & ROMANO HOGE, 1981
The genus Ovophis is distributed from central Ne-
pal and Bangladesh in the west, eastwards to central
China and Japan in the east, and China in the north,
southwards, through the Indochinese Peninsula and
the Malay Peninsula to Sumatra and Borneo.
Small to medium-sized; muscular; body stout; tail
short; head triangular, distinct from neck, thick,
often swollen, above with small, asymmetrically
arranged scales. Two to four rows of small scales
between suboculars and supralabials; subcaudals
divided or entire.
These are ground-dwellers which prefer higher el-
evations. Despite their apparent plumpness they can
strike in a flash. Their venoms may be fatal for
humans.
birth to fully developed young. It is known of some
species that the females guard their clutches (of eggs).
Five species are currently recognized, one of which
inhabits Borneo and Mt. Kinabalu.
Ovophis chaseni (SMITH, 1931)
Trimeresurus chaseni SMITH, 1931: 29, pl II: fig. 1 – Type
locality: Kiau, Mt. Kinabalu
Trimeresurus chaseni, LOVERIDGE 1938: 45; MANTHEY 1983: 30;
EBENHARD & SJÖGREN 1984: 24
Ovophis chaseni, STUEBING 1991: 357; MALKMUS 1994b: 248;
STUEBING & INGER 1999: 223
Diagnosis
Distinguished from the closely related O. monti-
cola, by lower number of dorsal scale rows and
sublabials. Max. total length nearly 65 cm.

373
 Crotalidae
© I. Das
Fig. 361 Ovophis chaseni, paratype from Kiau (ZRC 2.2173).
Description
Body stout; head flat, very distinct from body; eye
small; shields on upper side of head relatively large,
subimbricate, weakly keeled; supraoculars very
large; 6 supralabials, the 2nd below, but not bor-
dering, the loreal pit, the 3rd the highest one, sepa-
rated from eye by 2 small scales; 10-11 sublabials;
19, 17-19, or 15 rows of dorsals, the median ones
on the anterior body strongly keeled, those on the
posterior body weakly keeled; 130-143 ventrals;
anal scute entire; 20-30 subcaudals, in pairs.
Brownish above, with irregular blackish, light-
margined spots which fuse to form bands in the
posterior portion of the body. Lower side yellow-
ish, with a multitude of small grey spots. An ob-
lique, black stripe with a white lower border be-
hind the eye.
Ecology/Ethology
A submontane species, recorded from altitudes
between 915 and about 1550 metres (HQ-Collec-
tion).
374
Distribution
This species is endemic to Borneo, it has only been
found in Sabah (Kg. Tagodon, Maliau Basin,
Sinsuron, Crocker Range, and Mt. Kinabalu). Mt.
Kinabalu: Kiau, Headquarters.
Genus Trimeresurus LACEPÈDE, 1804
The distribution range of the genus Trimeresurus
extends from northern India in the west, eastwards
through large parts of China to Taiwan, southwards
through the Malay Peninsula, to the Philippine Is-
lands and to the Indonesian islands of Wetar and
Timor.
Small to medium-sized venomous snakes. Head
usually very distinct from body, its upper side cov-
ered with supraoculars, internasals and many small,
irregularly arranged scales. In contrast to the ge-
nus Tropidolaemus the chinshields are smooth.
One subocular shield; eye with vertically slit pupil;
 Crotalidae

sensory pit with one nasal pore; body round or oval

in cross-section; tail short to moderately long, pre-
hensile; anal scute entire; subcaudals most often
divided.
The crepuscular and nocturnal, bush- and tree-
dwellers species live in forests as well as in open,
bushy areas, grasslands, plantations and gardens.
They feed on smaller mammals, birds, lizards, and
frogs. Most species are ovoviviparous, but a few
are oviparous.
Thirty species are known at present, of which four
or five live on Borneo. The occurrence on this is-
land of Trimeresurus albolabris, based on an old
record, has not been confirmed. Three species are
found on Mt. Kinabalu.
Key to the Bornean species of
Trimeresurus
1 Snout strongly projecting, bilobed or spatulate;
1-3 supraocular plates, flat or slightly elevated ..
........................................................... borneensis
• Snout not strongly projecting, bilobed or spatulate;
1 flat supraocular scute ...................................... 2
2 19 dorsal scale rows at mid-body ....... malcolmi
• 21 dorsal scale rows at mid- body .................... 3
3 148-157 ventrals .................. popeiorum sabahi
• 182-191 ventrals ............................... sumatranus

Tab. 57 Distribution of Trimeresurus-species on Borneo.

Trimeresurus borneensis (PETERS, 1872)
Atropophis borneensis PETERS , 1872: 41 – Type locality:
Sarawak, Borneo
Trimeresurus puniceus, STUEBING 1991: 357 (part.)
Diagnosis
Easily recognizable by its projected, spatulate, up-
turned tip of the snout. Max. length 83 cm; aver-
age-size 45-55 cm.
brown spots and greenish brown to reddish brown
markings; upper side of the head dark with a light
brown line from the posterior margin of eye to be-
hind angle of the mouth; numerous light brown dots
on labials; venter light to greyish brown, finely
dotted with dark; lower side of tail dark brown.
Juveniles more contrasting with yellow tail.
7-14 eggs in a clutch.

Description Fig. 362 Trimeresurus borneensis from Poring.

Slender; head distinctly set off from body; tip of
snout spatular; 1-3 supraoculars, flat or slightly
button-shaped, separated by 9-11 scales; 3 preocu-
lars; 1 subocular; 9-11 supralabials, the 2nd high,
bordering anterior margin of loreal pit entirely or
almost entirely, the 1st separated from the nasal
shield; 19-21 rows of smooth dorsals at mid-body;
152-180 ventrals; anal scute entire; 45-58 subcau-
dals, divided.
© C. Brühl

Dorsally light, medium, to reddish brown, more

rarely greenish, with an irregular pattern of dark

375
 Crotalidae
Ecology/Ethology
This snake is found between sea level and altitudes of
at least 1200 m, with a predilection for hilly regions
at elevations between 500 and 1000 m. It inhabits low-
land tropical wet forests, tropical wet montane for-
ests, swamps, wet shrublands, and plantations. It of-
ten lives in damp places or close to water. This noc-
turnal species is both arboreal, frequenting logs, wood
piles, thick bushes and tree foliage up to 20 m above
the ground, and terrestrial, feeling at home among
leaves and plants on the forest floor. Juveniles some-
times climb into low vegetation holding on to twigs
with their prehensile tails. In this position they often
pull the body into a flat coil. As a rule, the snake is
mostly encountered in tangled wet vegetation.
T. borneensis feeds mainly on small mammals, liz-
ards and frogs. It is an aggressive snake, striking
promptly at little provocation and biting readily.
Nothing is known about its venom. By analogy with
T. puniceus, this snake should be regarded as poten-
tially dangerous. HAILE (1963) reported on a bite of
a Trimeresurus chaseni in Brunei, but it is almost
certain that T. borneensis was involved. This bite
nearly killed a middle-aged man in excellent health.
It has been reported that the females would coil up
around their clutches and keep this position until
their young eventually hatch.
Distribution
Trimeresurus borneensis inhabits the Peninsular parallely arranged, red spots between dark green cau-
Malaysia, Borneo and Sumatra. It is distributed in dal scales; subcaudals light green, sometimes framed
all parts of Borneo. Mt. Kinabalu: Poring.
Fig. 363 Trimeresurus malcolmi, juvenile from Sg. Silau-Silau, 1600 m.
© R. Malkmus
376
Trimeresurus malcolmi LOVERIDGE, 1938
Trimeresurus sumatranus malcolmi LOVERIDGE, 1938: 45 - Type
locality: Sungii River, near Kiau, Mt. Kinabalu, ca. 3300 feet
Trimeresurus sumatranus malcolmi, M ANTHEY 1983: 30;
MALKMUS 1985: 9, 1987: 290, 1994b: 242, 248
Trimeresurus malcomi (sic!), STUEBING & INGER 1998: 327 (error
typographicus)
Trimeresurus malcolmi, STUEBING & INGER 1999: 227
Trimeresurus sumatranus, SMITH 1931: 11, 29 (part.); EBENHARD
& SJÖGREN 1984: 24 (part.); STUEBING 1991: 357 (part.)
Diagnosis
Max. total length 133 cm; closely related to Trime-
resurus sumatranus, but differing by fewer rows
of dorsal scales, and lower ventral and subcaudal
counts.
Description
Three preoculars; 3 postoculars (inclusive one
subocular); 2+2 temporals; 8-9 supralabials, the 1st
separate from nasal; 1 postsubocular; 19 rows of
keeled dorsals at mid-body; 163-174 ventrals; anal
scute entire; 61-81 subcaudals.
Upper sides of head and body black with bright
green spots consisting of triangular green areas on
individual scales; a tendency to form indistinct
transverse lines; ventrals dark laterally and on their
anterior margins, dark or light green on their pos-
terior edges; tail patterned with series of uniform,
with black.
Juveniles grass-green above,
often with black frames be-
tween the scales; tail in stark
contrast, i.e. light brown with
indistinct, dark, transverse
bars, posterior third of tail
dark brown. Lateral surfaces
yellowish with reddish
stripe; venter greenish yel-
low in the anterior quarter,
otherwise cream-yellow;
supra- and sublabials bright
yellowish green; lateral
stripe white above and red-
dish below, on bottom rows
of dorsals.
 Crotalidae

© K.E. Linsenmair
Fig. 364 Recently we received this photo of a snake from Poring. This seems to be Trimeresurus sumatranus. Due
to the fact, that Trimeresurus sumatranus is very similar to T. malcolmi superficially, we hesitate to include T.
sumatranus to the herpetofauna of Mt. Kinabalu until specimens are at hand, to check the pholidosis.

Ecology/Ethology Trimeresurus popeiorum sabahi

Trimeresurus malcolmi has been recorded from
altitudes between 915 and 1600 m. Most of the
specimens were collected on the forest floor or as
victims of road traffic. MALKMUS (1987) found a
specimen on a roadside vegetated with ferns and
grass at around 9.00 p.m. during a rainshower. This
species is nocturnal. Its diet is unknown, but adult
specimens fed on rats and ground squirrels in cap-
tivity (STUEBING & INGER 1998). A juvenile ac-
cepted frogs and tadpoles from forceps (MALK-
MUS 1987).
Distribution
This species is endemic to Mt. Kinabalu: Sg.
Kenokok (at Kiau), Kiau, Bundu Tuhan, Lumu
Lumu, Headquarters, Kamborangoh Road
(1600 m), Kundasang (1525 m), Sg. Silau-Silau
(1500 m).
REGENASS & KRAMER, 1981
Lachesis gramineus (not Coluber gramineus SHAW, 1802),
HANITSCH 1900a: 70; SHELFORD 1901a: 68 (part?); DE ROOIJ
1917: 285 (part.)
Bothrops gramineus, MOCQUARD 1890a: 144
Trimeresurus gramineus (not Coluber gramineus SHAW, 1802), SMITH
1931: 10, 30, 32; LOVERIDGE 1938: 45; MANTHEY 1983: 30
Trimeresurus popeorum sabahi REGENASS & KRAMER, 1981: 190
– Type locality: Mount Kinabalu, Kulapis River, British
North Borneo
Trimeresurus popeorum, E BENHARD & SJÖGREN 1984: 24;
STUEBING 1991: 357
Trimeresurus sumatranus malcolmi (not Trimeresurus s.
malcolmi LOVERIDGE, 1938), MALKMUS 1992a: 136
Trimeresurus popeorum sabahi, MALKMUS 1994b: 248
Taxonomic comment: STUEBING & INGER (1999)
again used the old name Trimeresurus popeorum
and referred to the correction by the original author.

377
Crotalidae
It therefore appears appropriate to quote the com-
ment given by DAVID & VOGEL (1996), reading: “This
species was named in honour to Clifford H. Pope
and Sarah H. Pope.” The original spelling of the spe-
cific epithet, popeiorum, was corrected into
popeorum by SMITH (1943: 518) on the basis that it
was indeed a clerical error. Unfortunately, accord-
ing to the Art. 32 (c, ii) of the Code (I.C.Z.N. 1985),
such a change does not fall into the category of a
“correction of an incorrect original spelling.” Ac-
cording to the Art. 33 (d), the use of a termination -
orum in a subsequent spelling of a species-group
name that is a genitive based upon a personal name
in which the correct original spelling terminates with
-iorum, is an incorrect subsequent spelling, even if
the change is deliberate. The original spelling,
popeiorum, must therefore be conserved.”
Diagnosis
Differing from the other subspecies by the combi-
nation of the following traits: 21 dorsal scale rows
at mid-body, males with fewer than 158 ventrals,
females with fewer than 156 ventrals. Max. total
length 78 cm.
Description
Supralabials 9-11, the 1st separate from the nasal
shield; internasals enlarged, separate from each other
by 1-2 scales; 9-14 scales between the narrow
supraoculars; temporals weakly or not keeled in adult
specimens; 21 dorsal scale rows at mid-body; dor-
sals keeled except for those in the first row; 10-13
sublabials; 148-159 ventrals; 59-74 subcaudals.
Upper sides of head and body uniformly green; tip
of tail brown; sides of head and ventral sides
slightly lighter than body above; adult specimens
without postocular stripe. Males with distinct lat-
eral stripe on first row of dorsals from neck to
subcaudals; such stripe is missing or hardly recog-
nizable in females. Hemipenis bilobate, smooth,
without hooks or spikes.
Trimeresurus popeiorum sabahi is live-bearing. The
mainland form produces about 10 young in a litter
which measure 18-20 cm in length.
Ecology/Ethology
Trimeresurus popeiorum sabahi is found between
sea level and elevations of at least 1475 m. It is
378
common along streams, but nothing is known of its
biology. The nominate form inhabits lowland tropi-
cal wet forests, but is mostly found in tropical and
subtropical wet montane forests, bamboo forests, wet
montane shrublands, swamps, and plantations. It
often occurs along densely vegetated banks of riv-
ers and streams or close to water. This nocturnal
snake is largely arboreal and lives in bushes, tea and
coffee plants, shrubs, and low tree foliage. It feeds
on frogs, small mammals and lizards.
Trimeresurus popeiorum sabahi is an aggressive
snake, striking and biting fiercely when disturbed.
Many cases of envenomations involving humans
have been reported of the nominate form, especially
in tea estates, resulting mostly in only moderate
local symptoms. However, fatalities are known, and
this species should be regarded as potentially dan-
gerous.
Distribution
This subspecies is a Bornean endemite with records
being as yet limited to Sabah and Sarawak. Mt.
Kinabalu: Sayap, Kiau, Kenokok, without local-
ity 640 m (HANITSCH 1900a), Sg. Takobang at Kiau,
Sg. Kadamaian at Kiau, Bundu Tuhan, Sg. Luidan
at Bundu Tuhan, Headquarters. R EGENASS &
KRAMER 1981 mentioned a specimen of Mt. Kina-
balu at an elevation of 4175 m. This is obviously a
mistake, as Mt. Kinabalu is only 4095 m high. Per-
haps the high was 4175 feet (1270 m) or 1475 m
due to an error.
Genus Tropidolaemus WAGLER, 1830
The genus Tropidolaemus shows a disjunct distribu-
tion pattern. It occurs from the Malay Peninsula and
Indonesia to the Philippines, but there is also a spe-
cies known from southern India.
Characterized by the absence of nasal pores, strongly
keeled, small scales on head, and keeled gular scales.
Second supralabial does not border the anterior mar-
gin of the loreal pit, and is bordered above by a
prefoveal shield (DAVID & VOGEL 1998).
The species of the genus Tropidolaemus are crep-
uscular and nocturnal and live predominantly on
bushes and shrubs. They are very lethargic

ambushers preying upon small mammals, lizards,
frogs, and birds.
Tropidolaemus are live-bearing.
Three species are known, one of which occurs on
Borneo.
Tropidolaemus wagleri WAGLER, 1830
[Tropidolaemus] wagleri WAGLER, 1830: 175 – Type locality:
Asia
Taxonomic comment: The taxonomy of this spe-
cies or species complex is highly unsatisfactory and
currently under investigation (KUCH pers. comm.).
This form might have the name Tropidolaemus
wagleri schlegelii B LEEKER , 1857 in future
(ISKANDAR & COLIJN in prep.). For the authorship
of this species see DAVID & VOGEL (1996).
Diagnosis
Large, stout, max. length about 130 cm, average
80 cm in females, males are smaller and thinner.
Tail length/total length ratio is below 20%.
Description (valid for Bornean specimens only
[DAVID & VOGEL 1998])
10-13 strongly keeled scales between the supraocu-
lars; 3 preoculars; 1 enlarged supraocular; 1-3 posto-
culars; 1 subocular; 0-6 small loreals; 7-11 su-
pralabials (FUKADA 1964), none in contact with eye;
10-12 sublabials; dorsal scales keeled, in 21-27
rows, outer row smooth; 130-150 ventrals; anal
scute entire; subcaudals in pairs, 46-54.
Coloration variable, a mixture of various shades of
green and yellow, forming a banded pattern dorsally
which is often not clearly defined and extends onto
the tail; upper side of head green with yellowish
spots; labial shields yellow or green; ventrals green
with yellow margins or yellow with dark green
margins.
Juveniles green with reddish white dorsolateral
spots; a whitish stripe, bordered with reddish be-
low, from the loreal pit to the occiput; tail with a
reddish or black tip.
Ovoviviparous, giving birth to about 15 young, with
a maximum record of 41 juveniles in captivity (LIM
1990).
Crotalidae
Ecology/Ethology
T. wagleri is found in the lowlands up to elevations of
at least 1300 m, but is most frequent in the well-wa-
tered lowlands. It inhabits lowland tropical wet for-
ests, tropical wet montane forests, closed or open
swamps and marshes, mangroves, and coastal
swamps. It often occurs close to water, along banks
of forest rivers and streams, in forested swamps, where
it lives on structures overhanging water, in bushes
close to rice paddies, and in ditches in villages. This
very sluggish, crepuscular and nocturnal species re-
treats during the day in hollow tree trunks or in tan-
gled vegetation, where it can remain coiled motion-
less for a long time, and becomes active after dusk.
Adults are strictly arboreal, preferring low elevations
offered by bushes and shrubs, but juveniles often for-
age on the ground and in tall grass. This snake feeds
mainly on mammals, lizards and frogs, but also takes
birds; juveniles prey upon frogs and lizards.
Tropidolaemus wagleri is a very docile snake, rarely
striking or attempting to bite unless handled roughly.
However, some specimens are more irritable than
others, and the venom is potent. Severe cases of en-
venomation have been reported, which lead to both
local and systemic symptoms. This species should
be regarded as potentially dangerous.
Distribution
This snake ranges from the Malay Peninsula and
the Greater Sunda Islands to the Philippines. On
Borneo, this species is wide-ranging. Mt. Kina-
balu: Headquarters (HQ- Collection).
Fig. 365 Tropidolaemus wagleri from Deramakot, Sabah.
© T. Eltz
379
Cylindrophiidae

Cylindrophiidae
This family comprises small to moderately large snakes. The body is cylindrical in cross section, the
head not distinct. A mental groove is present. Of the 6 supralabials, the 3rd to 4th are in contact with the
eye. A loreal is absent. There are 19-23 rows of dorsal scale at mid-body; they are smooth.
Teeth are present on the palatine and pterygoid; the maxillary teeth count 9-13. The ectopterygoid is
robust and overlaps maxilla and pterygoid extensively. The premaxillary nasal process is long and ex-
tends between the nasals (CUNDALL et al. 1993).

eels and other fish, lizards, and invertebrates such

Genus Cylindrophis WAGLER, 1828
The genus Cylindrophis is distributed from Laos,
Myanmar, and Vietnam south to the Aru Islands; it
also occurs on Sri Lanka.
Small species; body cylindrical; head covered with
large, symmetrical shields, very small and not dis-
tinct from body; mental groove present; eyes very
small, pupil round or vertical-elliptic; no loreal; no
preocular. 5-6 supralabials; nostril situated in a sin-
gle nasal; 17-21 smooth dorsal scale rows at mid-
body; ventrals hardly or not at all widened; tail very
short, with a blunt tip.
These small snakes of semifossorial habits live be-
neath leaf litter, loose bark, or other materials cov-
ering the ground, but they are more rarely found in
the soil itself. They prey upon truly fossorial snakes,
as earthworms.
The snakes give birth to fully developed young.
Ten species are known (SMITH & SIDIK 1998), three
of which live on Borneo, one on Mt. Kinabalu.
Key to the Bornean species of
Cylindrophis
1 17 dorsal scale rows at mid-body ... engkariensis
• 19 or more dorsal scale rows at mid-body ....... 2
2 Two longitudinal whitish or black stripes on the
back; head and tail yellowish or red, lower part of
tail white ................................................. lineatus
• No longitudinal stripes on the back, head and tail
above coloured like the body, lower part of tail
vermilion ....................................... ruffus ruffus

Tab. 58 Distribution of Cylindrophis-species on Borneo.

Cylindrophis ruffus ruffus Description

(LAURENTI, 1768)
Anguis ruffa LAURENTI, 1768: 71 – Type locality: “Suriname”
[in error]
Diagnosis
Small; subfossorial; readily identified by red un-
derside of tail tip. Loreal and preocular shields
absent; max. total length nearly 90 cm.
No loreal; no preocular; 1 postocular; 6 suprala-
bials, the 3rd and 4th in contact with eye; dorsals
in 19 or 21 rows at mid-body, smooth; 185-245
ventrals; anal scute divided; 5-10 entire subcau-
dals.
Dorsally iridescent black, dark brown, or blackish
brown; irregularly spaced whitish, reddish, or
beige-coloured bands, not encompassing the body
entirely, the last, or the two posteriormost ones, on

380

Above: Fig. 366 Dorsal view of Cylindrophus r. ruffus
from Peninsular Malaysia.
Right: Fig. 367 Ventral view of Cylindrophus r. ruffus
from Thailand.
the tail reddish or bright red, the first one on the
neck likewise; these bands may, however, be ab-
sent; throat with almost entirely black scales with
white tips; ventral side alternating blackish brown
and whitish; underside of tail tip red.
Juveniles are coloured more vividly with the col-
lar being particularly more distinct.
Females give birth to litters of 5-13 babies of about
20 cm in length.
Ecology/Ethology
Cylindrophis ruffus ruffus is found at low eleva-
tions and inhabits non-flooded lowland tropical wet
forests, plantations, cultivated areas and gardens,
but is more frequently found in swamps, in the vi-
cinity of rice paddies, and in suburban and urban
ditches, canals, and gardens. It is subfossorial, liv-
ing close to the surface in loose, wet soil or in
muddy areas where it can burrow with ease. It is
also found in rodent burrows or other holes, under
decaying wood or vegetation, piles of wood, boards,
rocks, stumps, or in forest litter. Nocturnal, it is
Cylindrophiidae
© U. Manthey
© I. Das
often seen foraging at night on the ground during
the rainy season, and it takes to shallow water where
it swims effortlessly. This species feeds chiefly on
other terrestrial snakes, but also takes slow-mov-
ing burrowing lizards, eels and other fish; nestling
mammals and invertebrates (including worms and
myriapods) have also been reported. When it is
surprised or threatened, it raises its tail and curls it
over its back, so that the red ventral colour flashes
a warning, possibly as a mimicry of cobras.
Distribution
Cylindrophis r. ruffus inhabits a region ranging from
Myanmar and China to the islands of Borneo,
Sumatra, Java, and Sulawesi. Mt. Kinabalu: Head-
quarters (HQ-Collection).
381
Elapidae

Elapidae
Elapid snakes are characterized by having proteroglyphous teeth. They possess two grooved, largely
syringe-like teeth in the anterior portion of the upper jaw which are connected to venom-producing
glands. These serve to inject venom through the ducts and grooves. The family is furthermore typified by
its members having round pupils and no loreal shields. It comprises small species of some 50 cm
(Calliophis), as well as the largest venomous snake, the king cobra (Ophiophagus hannah), which may
grow up to 585 cm in total length. Their venoms can be very potent so that a bite may be hazardous, often
even fatal, for humans. These snakes reproduce by laying eggs.

Key to the Bornean genera of the family Elapidae

1 Tail flattened; inhabiting the ocean .. Seasnakes
• Tail more or less round .................................... 2 A B
2 Vertebrals much larger than dorsals (A) .............
............................................................ Bungarus
• Vertebrals not much larger than dorsals .......... 3
3 13 dorsal scale rows at mid-body .... Calliophis
• More than 13 dorsal scale rows at mid-body ... 4
4 2 large occipital shields (B) .......... Ophiophagus
• No large occipital shields ......................... Naja

Genus Bungarus DAUDIN, 1803 vertebrals distinctly enlarged; subcaudals entire, or

The distribution of the genus Bungarus extends
from Afghanistan in the west, through large parts
of China in the east, to the south through the Malay
Peninsula, the Greater Sunda Islands and Sulawesi,
to Java and Bali.
Moderately large to large; muscular; body round
or triangular in cross-section; tail short; head little
distinct from body; eyes small, pupils round; loreal
shield absent; dorsals smooth, in 13-17 rows;
intermixed with single divided subcaudals.
These nocturnal ground-dwellers often present
themselves as inoffensive and even too lazy to bite
during the day. When disturbed they attempt to
conceal the head beneath their own body coils. They
feed primarily on snakes and, to a lesser extent, on
lizards and fish, but also take rodents and birds in
captivity. They reproduce by laying eggs.
The genus Bungarus comprises altogether 12 or
13 species, two of which occur on Borneo, and both
on Mt. Kinabalu.

Tab. 59 Distribution of Bungarus-species on Borneo.

382

Key to the Bornean species of
Bungarus
1 15 (rarely 17) dorsal scale rows at mid-body; all
subcaudals entire, with the exception of the last
pair. Body normally encircled with complete,
alternatingly black and yellow-coloured rings ...
.............................................................. fasciatus
• 13 dorsal scale rows at mid-body; posterior
subcaudals paired; dorsal surface not encircled
with black and yellow rings .............................. 2
2 Dorsal surface with dark, white and red bands in
the posterior part of the body; tail banded ..........
............................................. flaviceps baluensis
• Dorsal surface uniformly dark, tail red without
bands ..................................... flaviceps flaviceps
Fig. 368 Bungarus fasciatus from Peninsular Malaysia.
Elapidae
Bungarus fasciatus (SCHNEIDER, 1801)
Pseudoboa fasciata SCHNEIDER, 1801: 283 (description based
on RUSSELL 1796: 3, pl. 3) – Type locality: Mansoor Cottah,
Bengal
Bungarus fasciatus, EBENHARD & SJÖGREN 1984: 24; STUEBING
1991: 351; MALKMUS 1992a: 136, 1994b: 247
Diagnosis
Easily identified on the basis of its yellow and black
banded pattern, and by having a usually blunt tail
tip; head hardly distinct from body; max. length
225 cm, average specimens about 150 cm. Females
shorter than males.
Description
One preocular; 2 postoculars; 7 supralabials, the
3rd and 4th border the eye, the 6th largest; 1+2 tempo-
rals; 15 dorsal scale rows at mid-body; dorsals
© W. Grossmann
383
Elapidae
smooth; 200-234 ventrals; anal scute entire; 23-41
subcaudals, entire. Body and tail distinctly trian-
gular in cross-section; tail with very blunt tip.
Body and tail with alternating black and yellowish
to yellow bands of equal widths, continued on ven-
tral side. A black stripe from the eye over head and
neck; labial shields and lower side of neck yellow;
underside generally lighter than upper side render-
ing the bands light yellow and greyish black.
Bungarus fasciatus is oviparous, with a clutch con-
sisting of 5-15 eggs. Hatchlings measure 250-349 mm
in length.
Ecology/Ethology
This snake is found in lowland and hilly regions
up to altitudes of at least 2300 m, but with a pre-
dilection for the lowlands. It inhabits the margins
of lowland tropical wet and dry forests, tropical
and subtropical montane forests and scrublands,
and especially swamp and marshes, plantations,
and cultivated areas like rice paddies. Bungarus
fasciatus is found in dry, open areas, in scrublands
and fields or on roads, as well as in thickly veg-
etated damp places in the vicinity of water, where
it is quite frequent. This snake is a secretive, ter-
restrial species, which is predominantly noctur-
nal, emerging after dusk from its day-time retreat
in the ground or under vegetation, logs and rocks.
It often occurs in the vicinity of human habita-
tions, and is found under piles of wood, on paths
and roads; it may enter water. This species feeds
chiefly on other snakes, but also takes lizards and
amphibians; fish and eggs have also been reported
in the diet.
Bungarus fasciatus is a dangerously venomous
snake, with a highly effective venom; however, few
human fatal accidents have been reported.
MALKMUS (1992a) found a specimen in the maze
of adventitious roots of a tree.
Distribution
The distribution of this species covers the region
between India in the west and Hong Kong in the
east, southwards through Indochina, Thailand, and
the Malay Peninsula to Sumatra, Java and Borneo.
On Borneo, the snake is wide-ranging and has been
recorded from all parts of the island. Mt. Kina-
balu: Poring (500 m).
384
Bungarus flaviceps baluensis
LOVERIDGE, 1938
Bungarus flaviceps baluensis LOVERIDGE , 1938: 44 – Type
locality: Kenokok River, near Kiau, Mt. Kinabalu, ca. 3300
feet
Bungarus flaviceps, BOULENGER 1896: 372 (forma D); SHELFORD
1901a: 67 (part.); DE ROOIJ 1917: 245; SMITH 1931: 32;
E BENHARD & S JÖGREN 1984: 24; S TUEBING 1991: 352;
MALKMUS 1994b: 247
Bungarus flaviceps baluensis, BRONGERSMA 1948: 27; MANTHEY
1983: 30; VOGEL & HOFFMANN 1997: 13; STUEBING & INGER
1999: 191; KUCH & GÖTZKE 2000: 19
Taxonomic comment:
The author is convinced that this taxon is much
rather a good species than a subspecies. For a de-
tailed discussion see VOGEL & HOFFMANN (1997).
Diagnosis
13 rows of dorsals at mid-body; readily distin-
guished from all other snakes of the region by its
colour pattern described below; max. total length
133 cm.
Description
One preocular; 2 postoculars; 1+2 temporals; 7 su-
pralabials, 3rd and 4th touching eye; 13 dorsal scale
rows at mid-body; 206-225 ventrals; anal scute
entire; 42-52 subcaudals, entire anteriorly, divided
more posteriorly.
Head and nape red, more rarely yellowish brown
to yellow; anterior portion of upper body black with
a light vertebral line; dorsals of first two rows white
with black centres; posterior half of body banded
black and coral-red, the eight black bands split by
white bands, gradually increasing from one white
band on the tail to four white bands at mid-body.
Another group of white bands on the anterior half
of the body is limited to an indication. The belly is
white anteriorly, with black margins at the ventrals
and coral red posteriorly with remainders of the
bands (VOGEL & HOFFMANN 1997, KUCH & GÖTZKE
2000).
Like all Elapids, this taxon should be oviparous.
Ecology/Ethology
This krait is found in lowland and hilly regions,
with a predilection for the latter, where it inhabits

Fig. 369 Bungarus flaviceps baluensis from Kundasang, ca. 1400 m.
tropical wet forests. The snake is terrestrial, per-
haps semiaquatic, and nocturnal. It lives on the for-
est floor and hides under vegetation and decaying
logs. One female was found on stones in a small,
clear small stream, in a high-canopy dipterocarp
dominated primary forest in the afternoon. It is
known to feed on other snakes and sometimes liz-
ards, but other aspects of its biology are unknown.
This subspecies is very rare. Only a few specimens
have as yet been found.
K UCH & G ÖTZKE (2000) found two specimens
copulating at 7.30 to 8.00 p.m. on a path through
a high-canopy dipterocarp dominated primary for-
est at Poring Hot Springs (550 m). The air tem-
perature was about 25°C, and the air was satu-
rated with water due to regular showers late in
the afternoon. Both snakes were about 150 cm,
but the male was a little bit longer than the fe-
male. The two snakes were twisted around each
other. Disturbed by the light of a tourch the speci-
mens tried to flee, but did not interrupt the copu-
lation. For at least 20 minutes, the left hemipenis
Elapidae
© P. Hoffmann
of the male could be traced in the females cloaca
by the swelling.
Distribution
This subspecies was found at Mt. Kinabalu and Mt.
Trus Madi (1550 m) in Sabah. Mt. Kinabalu:
Kundasang (ca. 1400 m), Kenokok River near Kiau
(1005 m), Poring: Poring Hot Springs (550 m).
Genus Calliophis GRAY, 1835
Small to medium-sized; Very slender; head small,
not distinct from body; eyes small, pupil round;
body round in cross-section; scales small 13 rows
of dorsals at mid-body; ventrals rounded; tail
short; subcaudals divided. A single, elongate tem-
poral scale, Haderian gland and parietal bone ori-
gin for the AES muscle, and a posterior extension
of the Haderian gland beyond the orbit (SLOWINSKI,
BOUNDY & LAWSON 2001).
385
Elapidae
When they feel threatened, the ground-dwelling
and nocturnal Coral snakes exhibit interesting
defence behaviour in that they curl up the tail lat-
erally to form a flat spiral, which is then presented,
to the potential attacker with its colourful under-
side up.
All species of this genus are oviparous.
Eight species are known, two of which occur on
Borneo, and both also on Mt. Kinabalu.
Tab. 60 Distribution of Calliophis-species on Borneo.
Calliophis bivirgata tetrataenia
(BLEEKER, 1859)
Elaps tetrataenia BLEEKER 1859: 201 – Type locality: Sintang,
West Borneo
Calliophis bivirgata (tetrataenia), S LOWINSKI , B OUNDY &
LAWSON 2001: 239
Diagnosis
Easily identified by its red head and tail; body with
striped pattern (in contrast to the other subspecies);
Max. total length 150 cm, average 130-140 cm;
females substantially smaller than males, reaching
a little more than 80 cm (BRONGERSMA 1948).
Description (BRONGERSMA 1948):
One large preocular; 2 small postoculars; 6 (7) supra-
labials, 3 rd and 4th touching eye; 1+2 temporals
(rarely 1+1 or 1+1+1); 13 dorsal scale rows at mid-
body, dorsals smooth; 239-306 ventrals; anal scute
entire; 36-49 subcaudals, divided.
Body dark or blackish blue; a pair of light longitu-
dinal lines both dorsolaterally and laterally, the lat-
eral one wider and situated on the bottom two rows
of dorsals; head, tail, and all lower sides of body
and tail uniformly bright red.
This snake is oviparous, but no details of its repro-
ductive biology are known.
Ecology/Ethology
This snake is found from sea level up to elevations
of at least 1750 m. It inhabits lowland tropical wet
386
Key to the Bornean species of Calliophis
1 Ventral surface of body and tail uniformly bright
red or coral-red; more than 34 subcaudals ..........
.......................................... bivirgata tetrataenia
• Ventral surface of body and tail with black cross-
bars; less than 35 subcaudals ...............................
............................................ intestinalis thepassi
forests, tropical wet montane forests, and plantations.
This nocturnal snake is a secretive, terrestrial and
semifossorial animal, living in forested, closed ar-
eas along streams under decaying logs, bark, and
fallen vegetation, among roots and beneath vegeta-
tion clumps. Calliophis bivirgata tetrataenia seems
to feed mainly on snakes, but also preys upon liz-
ards and frogs. It is rarely encountered, but might
nevertheless be common in some areas. This snake
is not aggressive, but will bite if molested. It is dan-
gerously venomous, with a highly effective venom.
Distribution
This subspecies is endemic to Borneo and has been
recorded from all parts of the island. Mt. Kinabalu:
Headquarters (1750 m) [HQ-Collection].
Calliophis intestinalis thepassi
(BLEEKER, 1859)
Elaps thepassi BLEEKER, 1859: 201 – Type locality: Sintang,
Borneo
Doliophis intestinalis var. everetti, BOULENGER 1896c: 404
Doliophis intestinalis, SHELFORD 1901a: 67; DE ROOIJ 1917: 253
Maticora intestinalis nigrotaeniatus, SMITH 1931: 10, 28
Maticora intestinalis everetti, MANTHEY 1983: 30; MALKMUS
1985: 10, 1987: 289, 1988: 174; MALKMUS & SAUER 1993:
153; MALKMUS 1994b: 241, 247, 1996c: 293
Maticora intestinalis, SMITH 1931: 32; EBENHARD & SJÖGREN
1984: 24; STUEBING 1991: 352
Calliophis intestinalis (thepassi), SLOWINSKI, BOUNDY & LAWSON
2001: 239

Diagnosis
Small, slender, max. length about 72 cm. Distin-
guished from related forms by the colour pattern
described below.
Description
One preocular; 2 postoculars; 1+2 temporals; 6 su-
pralabials, 3rd and 4th touching eye; 13-15 rows of
smooth dorsal scales at mid-body; 203-263 ventrals;
anal scute entire; 16-28 subcaudals, divided.
Upper surface and flanks iridescent black; two par-
allel, pale grey stripes from the eyes over the pari-
etals and posttemporals right to the tail tip; a red
area on the supralabial shields, interrupted only by
a black bar below eye; sublabials and preinfra-
maxillars with some black spots, more of those on
temporal, parietal and frontal shields; a reddish
brown medial stripe on about 5 small scales poste-
rior to the parietal notch which may extend up to
the tail in some individuals; prefrontal shield with
a light band; bottom row of dorsal scales whitish,
each scale with a halfmoon-shaped marking open
below; lower side red with black bars, the latter 2-
2.5 scales in width and spaced by 3 to 4 scales;
occasionally the underside may be uniformly red;
upper side of tail with reddish brown, doubly in-
terrupted medial stripe; lower side of tail red with
black bands.
This snake is oviparous, laying up to 4 eggs at a time.
Ecology/Ethology
Calliophis intestinalis thepassi is found from sea
level up to altitudes of at least 1750 m, with a pre-
dilection for forested hilly areas. It inhabits low-
land tropical wet forests, tropical and subtropical
wet montane forests, and plantations. It is a noc-
turnal, secretive, and semifossorial snake which
lives beneath or in the forest litter, under rocks,
roots and fallen logs and other debris, and under
vegetation clumps. This snake feeds only on other
small, fossorial snakes. It has an distinct defensive
display, when molested, it flattens the body and
raises its tail to show the red underside of it. Some-
times it may flip over exposing the bright pattern
of the belly.
One specimen was discovered crossing a road at
an elevation of 1600 m at around 8.00 p.m. (MALK-
MUS 1994b). MALKMUS & SAUER (1993) reported
Elapidae
© R. Malkmus
Fig. 370 Calliophis intestinalis thepassi from Sg. Silau-
Silau.
on the effects of a bite inflicted by this subspecies.
It resulted in severe local swelling which, however,
had disappeared after two days.
Distribution
This subspecies is endemic to Borneo and has been
found in all parts of the island. Mt. Kinabalu: Kiau
(900 m), Bundu Tuhan, between Headquarters and
Kundasang (1400-1550 m), Kamborangoh Road
(1600-1750 m). STUEBING (1991) mentions Lumu
Lumu with regard to MALKMUS (1988). This is an
error, as MALKMUS does not mention a localitiy for
his specimen.
Genus Naja LAURENTI, 1768
The genus Naja inhabits a vast area including large
parts of Africa and Asia.
Moderately large; slender; max. length about 2 m;
body muscular, round in cross-section; tail moder-
ately long; head little distinct from body; eyes large;
pupils round. Slim rips in the neck enable a spread-
ing of the surrounding skin to form the famous
“hood”. Nostril between two nasals and an inter-
nasal shield; no loreal shield. dorsals in 15-25 rows
at mid-body, without apical pits; All, or most, sub-
caudals divided.
The display of the hood, while the anterior third of
the body is simultaneously lifted vertically off the
ground and the head is held horizontally and directed
towards the source of the disturbance, serves as a
distinct warning to a potential antagonist. The two
hollow teeth in the upper jaw facilitate the injection
of large amounts of venom with a single bite.
387
 Elapidae
The food spectrum of the cobras comprises smaller
rodents, birds, fish, amphibians, lizards, and snakes.
All the included species are oviparous.
Altogether 18 species of Naja are known, only one
of which occurs on Borneo and on Mt. Kinabalu.
Naja sumatrana MÜLLER, 1887
Naja tripudians var. sumatrana MÜLLER, 1887: 277 – Type
locality: Solok, Sumatra
Naja tripudians, SHELFORD 1901a: 67
Naja tripudians var. miolepis, BOULENGER, 1896c: 385 – Type
locality: Sarawak (syn. after WÜSTER & THORPE 1989: 403)
Naja tripudians var. miolepis, DE ROOIJ 1917: 248
Naja naja miolepis, SMITH 1931: 10, 28, 32; MANTHEY 1983:
30
Naja naja, EBENHARD & SJÖGREN 1984: 24
Naja sumatrana, WÜSTER & THORPE 1989: 403; STUEBING 1991:
353; MALKMUS 1994b: 247
Diagnosis
Distinguished from all other snakes occurring on
Mt. Kinabalu, except the colubrid Macropisthodon
rhodomelas, by its ability to form a “hood”. No
large shields behind the parietals. Max. total length
150 cm, average 100-120 cm.
Fig. 371 Juvenile Naja sumatrana from South Thailand.
© U. Manthey
388
Description
One preocular; 3 postoculars; 7 supralabials, 3rd and
4th touching eye; 8 sublabials; 15-25 dorsal scale
rows at mid-body, dorsals smooth; 162-206
ventrals; anal scute entire; 40-55 subcaudals, the
anterior ones entire, the posterior ones divided.
Ground coloration yellow to yellowish green, on
Borneo also black; upper side of the head brown-
ish yellow or olive; lips and throat yellowish; hood
without pattern; eyes dark; tongue pink; venter usu-
ally light yellow, pale yellow, or black; individual
specimens with irregularly shaped brown spots on
the underside of the hood. Juveniles occasionally
with light transverse lines.
A clutch of this species may comprise up to 23 eggs.
Ecology/Ethology
Naja sumatrana is found in lowland and hilly re-
gions up to at least 1500 m altitude. It inhabits the
margins of lowland tropical wet and dry forests, es-
pecially along clearings, tropical and subtropical
montane forests, swamps and marshes, scrublands,
plantations (where it feels much at home), cultivated
areas, rice paddies, gardens, and urban areas. It is
frequently found in densely vegetated wet places in
the vicinity of water. Naja sumatrana is terrestrial,
although it enters water readily, and mainly noctur-
nal, emerging after dusk from its retreat made in any
hole of appropriate size and becoming an active for-
ager. It is often encountered in the vicinity of hu-
man habitations. It feeds on mammals, especially
rats, and amphibians; lizards, birds and other snakes
are also taken. This species is very common every-
where within its range. It is not aggressive, although
it displays its defensive posture at the least provoca-
tion and defends itself vigorously when cornered.
This snake can spit its venom accurately towards an
aggressor. It is a dangerously venomous animal, with
a very potent venom. Numerous human fatalities
have been reported from its bites.
Distribution
The species ranges from the Malay Peninsula and
Borneo to Palawan Island in the Philippines. It is
furthermore native to Sumatra and the surround-
ing islands.
It is present in all parts of Borneo. Mt. Kinabalu:
Kiau. Bundu Tuhan (1370 m).
 Pythonidae

Pythonidae
These are medium-sized to large snakes found in the tropics and subtropics. They inhabit nearly all types
of biotopes, there are desert dwellers, arboreal forest dwellers, and species which are highly aquatic.
Their skeletons include rudiments of a pelvic girdle, and their jaws carry large numbers of teeth. Their
labial shields are pitted serving as highly developed sensory organs which are able to perceive minute
differences in temperature. Pythons reproduce by laying eggs which, in some species, are actively incu-
bated by the female snake.

Genus Python DAUDIN, 1803
The Genus Python is widely distributed in large
parts of Africa and Asia and, with Python reticu-
latus, contains one of the two largest species of
snakes on earth.
Some species of this genus are very slender, others
are rather plump. Head always distinct from body;
distinguished from related genera by a very deep
first labial pit and the absence of apical pits
(UNDERWOOD & STIMSON 1990).
Most of the species of Python are primarily terres-
trial snakes and only a few could be termed semi-
arboreal. They inhabit a wide range of biotopes,
and can occasionally be found in the vicinity of
human settlements. They are often encountered near
water bodies. Their diet consists primarily of mam-
mals and birds, more rarely of reptiles. All species
of this genus are oviparous with the females ac-
tively incubating their eggs by coiling around the
clutch and raising their body temperature.
The genus comprises ten species, two of which live
on Borneo, both on Mt. Kinabalu.
Key to the Bornean species of Python
1 Rostral and first 2 supralabials with a pit each;
less than 280 ventrals ..................... breitensteini
• Rostral and first 4 supralabials with pits; more than
295 ventrals ....................................... reticulatus

Tab. 61 Distribution of Python-species on Borneo.

Python breitensteini STEINDACHNER, 1880
Python breitensteini STEINDACHNER, 1880: 276 – Type local-
ity: Teweh, Borneo
Python breitensteini, KEOGH, BARKER & SHINE 2001: 122
Diagnosis
A snake of the lowlands. Body very stock; tail short;
distinguished from Python brongersmai by a sin-
gle supraocular shield above the eye and a row of
small or granular scales covering the area from
below the eye to the nasal; differing from Python
curtus by the fact that the anterior pair of parietals
are in broad contact at the medial suture, whereas
they are not or only weakly in contact at Python
curtus (KEOGH, BARKER & SHINE 2001). Max. total
length 170 cm.
Description
Very muscular, stocky body; head distinct; tail short,
2 preoculars, 1-2 postoculars; 1 supraocular; a se-
ries of small, granular subocular scales extending

389
 Pythonidae
© B. Leideritz
Fig. 372 Python breitensteini from Poring.
up to the nasal; 9-13 supralabials; 48-61 dorsals at
mid-body; 150-169 ventrals; anal scute entire; 22-
32 subcaudals (AULIYA & ABEL 1999).
Basic coloration brown or yellowish brown, later-
ally with light, dark-bordered spots; a row of light
spots the spine, broadening towards tail and fusing
to form an uninterrupted line framed with dark.
Head light above, with an indication of a line which
begins on the rostral and splits up into 4 lines
posteriorly; a dark streak from in front of the eye
to onto the body; an oblique postocular stripe from
beneath the eye backwards.
Up to 32 eggs (50-70 × 35-50 mm on average [ERDE-
LEN et al. 1997]) may be laid in a clutch. Neonates
measure 33-40 cm and weigh around 61 g.
Ecology/Ethology
This species is particularly common in cultivated
areas. Here it lives in rice paddies, rubber planta-
tions, coconut groves, and other types of second-
ary vegetation. It spends most of its time in rodents’
burrows, piles of grass, or earthworks.
Rodents form the basis of its diet (ERDELEN et al.
1997), but larger mammals, chickens and ducks
have also been recorded as prey.
Altogether, the biology of this subspecies in the
wild is still poorly known.
390
Combat dances have occasionally been observed
(EVEN 1997, KIRSCHNER & WICHMANN 1998). After
a period of gravidity of about 60 to 90 days, up to
32 eggs may be laid in a clutch, in nature usually at
some time between July and November. The eggs
are deposited in heaps of grass, accumulations of
plant matter, stumps of dead coconut palms, or old
rotting, horizontal tree trunks. Python breitensteini
females incubate their eggs actively by coiling,
more tightly or more loosely, as the situation may
require, around the clutch and raising the body tem-
perature through muscular contractions if and when
necessary. It is possible that the female even col-
lects water on her body to supply moisture to her
eggs. Spoilt or infertile eggs are sorted out and
pushed aside with the head. At a temperature of
29-31°C the neonates hatch after 59-61 days. One
to two days may pass between the cutting open of
the egg shell and the actual hatching of a baby.
Males mature at an age of three years, females at
four years of age.
Distribution
This species is a Bornean endemite ranging over
large parts of the island. Mt. Kinabalu: Poring
(500-600 m a.s.l.).
Python reticulatus (SCHNEIDER, 1801)
Boa reticulata SCHNEIDER, 1801: 264 - Type locality: Not given;
based on specimens cited by SEBA from “Orient” (1734: pl.
62), “mountains of Japan” (1735: pl. 79; in error), and “Nova
Hispania” (1735: pl. 80; in error), respectively.
Python reticulatus, MALKMUS 1992a: 132, 1994b: 246
Diagnosis
Very large, max. length 980 cm, but adult speci-
mens from Kalimantan measure around 300 cm
(RIQUIER 1998). The reticulated pattern which cov-
ers the entire body is a characteristic trait of this
snake.
Description (only animals from Borneo, AULIYA
& ABEL 2000a)
Of muscular, although slender build; tail distinct
from body; head very distinct; eye with a vertical
pupil; rostral and first four supralabials with deep
 Pythonidae

pits; 12-16 supralabials, 6th to 8th in contact with

eye; 3-5 loreals; 2 preoculars; 3 postoculars; 61-82
dorsals at mid-body; 271-333 ventrals; anal scute
entire; 55-98 predominantly divided subcaudals.
Both sexes possess anal spurs, but those of the
males are larger.
Basic coloration of body varying from pale yellow
to dark or olive brown; back covered with roundish,
oval, or rhombic markings, forming a reticulated
pattern through fusion; these spots are yellowish
to brown and bordered with black and yellow; a
black longitudinal line from the tip of the snout to
the nape crossing through centre of head; sides of
head with a black temporal stripe between eye and
neck; belly yellowish, sparsely speckled with black.

© C. Brühl
When exposed to bright light, the body shows a
metallic gleam.
12 to 103 eggs (10.3-11.6 × 6.3-7.5 cm); hatch- Fig. 373 Python reticulatus from Deramakot, Sabah.
lings measure 60-80 cm and weigh 120-170 g.

Ecology/Ethology convulsions known from other species of Python

Python reticulatus is found from sea level to up to are rarely observed. The young emerge after a pe-
above 1300 m altitude and inhabits lowland pri- riod of about three months. Males mature after 2-3
mary and secondary tropical wet forests, tropical years, females about one year later (after VAN DER
open dry forests, tropical wet montane forests, MEER MOHR 1926, TRUTNAU 1980, WERNER 1981,
rocky shrublands, swamps, marshes, plantations, ROSS & MARZEC 1994, ERDELEN et al. 1997).
and cultivated areas, as well as suburban or even
urban areas. It is most often found close to water, Distribution
along river banks and canals, and is frequently dis- The distribution of Python reticulatus ranges from
covered in or around human settlements, near or India in the west to Laos and Vietnam in the east,
inside houses. Active at dusk and mainly at night, south through Myanmar, Thailand, and the Malay
this Python is terrestrial and semiarboreal, often Peninsula to Sulawesi and the Philippines. This spe-
concealing itself in dense foliage of low trees over- cies is widely distributed on Borneo. Mt. Kina-
hanging water or in large hollow tree trunks. It is balu: Sg. Kipungit II (640 m a.s.l.).
also a very good swimmer. Its diet consists chiefly
of mammals such as large rats, rabbits, small deer
Fig. 374 Python reticulatus from Poring, Sg. Kipungit II.
and domestic animals (cats, dogs and pigs), but it
also takes birds, especially fowl, large lizards, frogs
and fish (AULIYA & ABEL 2000b).
Although variable in disposition, it is often rather
aggressive, and may bite viciously. Larger speci-
mens are able to inflict severe wounds and must be
considered potentially dangerous animals.
Gravidity in Python reticulatus is said to take 65 to
© K.E. Linsenmair

105 days. Oviposition takes place mainly during

the months of February and March, with clutches
being deposited in cavities in tree trunks. The fe-
male coils around her clutch, but the muscular

391
 Typhlopidae

Typhlopidae
This family comprises small, worm-like, fossorial snakes whose bodies maintain about the same diam-
eter throughout the entire length. The head is not distinct from the body, the tail is short with a blunt tip.
The eyes are hidden beneath scales and invisible in some species. The scales of the body are fairly
uniform with no modified ventrals or subcaudals. They feed on ants and termites, their larvae and eggs,
other small insects, and worms.
Most of the representatives of this family reproduce by laying eggs, but a few give birth to fully devel-
oped young. The two genera present in the Sunda region, Ramphotyphlops and Typhlops, cannot be
distinguished by external traits.

Typhlops sp.
A species of the genus Typhlops was recently pho-
tographed on Mt. Kinabalu. Its wide rostral shield
and the high count of dorsal scale rows suggest that
this might be Typhlops koekkoeki BRONGERSMA,
1934 (WALLACH pers. comm.). Typhlops koekkoeki
has so far been known only from a single speci-
men collected on Bunju Island off the east coast of
Borneo. The photograph does unfortunately not
permit an unquestionable identification. If it is not
Typhlops koekkoeki, it is most certainly an as yet
undescribed species (WALLACH pers. comm.).
Locality: Mt. Kinabalu: Poring.

Fig. 375 Typhlops sp. from Poring.

© K.E. Linsenmair

392
 Xenopeltidae

Xenopeltidae
These are snakes of medium lengths with cylindrical bodies. The bones of the skull are well-united. The
premaxillary bone touches the maxillary, bearing well-developed teeth in line with maxillary teeth. The
ectoptergoid is loosely attached to the maxilla. The prefrontal is sutured to the nasal. The dentary is very
loosely attached anteriorly to the anterior end of the articular, but free behind and movable. There is no
coronoid and no postfrontal. A supratemporal supports a short quadrate. Hypapophyses are absent in the
posterior part of vertebral column. – This family contains only one genus.

scales, tapering to a very flat, round, broad snout;

Genus Xenopeltis BOIE, 1827
The genus Xenopeltis inhabits a large portion of
Asia, ranging from the south of China to Myanmar,
and south to the Indo-Australian Archipelago. It
also inhabits some of the Philippine islands.
Max. length nearly 1 metre; head large, not set off
from body, covered above with large, symmetrical
eye small, pupil vertical-elliptic; body round, cov-
ered with large, very smooth scales with an irides-
cent sheen in bright sunlight. 15 dorsal scale rows;
tail very short, about 10% of total length; anal scute
divided; subcaudals entire.
The genus Xenopeltis comprises only two species,
one of which occurs on Borneo and on Mt. Kina-
balu.

Fig. 376 Xenopeltis unicolor from South Thailand.

© U. Manthey

393
Xenopeltidae
Xenopeltis unicolor BOIE, 1827
Xenopeltis unicolor BOIE, 1827: 564 – Type locality: Java.
Diagnosis
Max. total length 90 cm, tail length < 10 cm. Defi-
nition see generic diagnosis.
Description
Head flattened; snout rounded; nostrils between two
nasals; frontal as long as its distance from the rostral;
preoculars large; 2 postoculars, the anterior one
larger; 1 small supraocular present, smaller than the
upper postocular; a large, septagonal interparietal
in the centre of 4 parietals; 8 supralabials, the first
in contact with internasals, 4th and 5th in contact with
eye; 1 pair of small chinshields, in contact with the
three anterior sublabials; 15 dorsal scale rows
throughout; dorsals smooth; 164-193 ventrals; anal
scute entire; 26-32 divided subcaudals.
Upper side brown or blackish, iridescent in bright
sunlight; scales with light margins; supralabial
scales pale yellow; lower side white; young with a
light collar.
This species is oviparous with a clutch comprising
between 6 and 17 eggs.
Fig. CVI Xenopeltis unicolor, juvenile.
394
Ecology/Ethology
Xenopeltis unicolor inhabits lowland and hilly re-
gions covered with tropical wet and dry forests,
but is chiefly found close to water in open vegeta-
tion such as marshes, scrublands, hedges, planta-
tions, cultivated areas, rice paddies, in gardens and
other suburban areas, and along ditches and
streams. It is semifossorial, living for most of the
time under the surface in wet, loose or sandy soil.
It may dig its own burrows, but more often lives in
holes in the ground and occupies burrows origi-
nally excavated by rodents, is found under rocks,
tree stumps, clumps of grass, logs, and among roots
and decaying vegetation. Active at night and at day,
it may often be seen foraging among vegetation on
the ground. This species feeds on lizards, other
snakes, small mammals and amphibians; it may also
prey upon ground-nesting birds.
Distribution
The distribution of this species ranges from Cam-
bodia in the east and China in the north, south
through Myanmar, Thailand, and the Peninsular
Malaysia, to large parts of Indonesia to the Philip-
pine Islands. It is also wide-ranging on Borneo. Mt.
Kinabalu: Poring (HQ-Collection).
after dE R OOIJ 1917
 Testudines

Testudines (turtles, tortoises, and terrapins)

There are about 300 species of turtles* distributed worldwide in the entire warm and most of the temperate
zones. They comprise terrestrial as well as aquatic reptiles marked for their bony shells which are in most
cases rigid, but furnished with hinges and therefore movable in some genera. Their vertebral columns are
largely firmly fused to their shells and, as a consequence, are freely movable only in the neck and tail
portions. The head, limbs, and the tail can be retracted more or less deeply into the openings of the shell.
All species inhabiting the Sunda region belong to the suborder Cryptodira which retract the head and
neck into the shell by vertically bending the vertebral column into an “S”. Their jaws are furnished with
horn-covered shear-like structures and lack teeth. Depending on their ways of life, these chelonians have
distinctly flattened to roundish limbs. Freely movable toes are not found in all families. These animals
can live to very old ages with some species reaching 150 years. They reproduce by laying eggs with
clutch sizes varying between 1-2 and up to 100 eggs. Many species are known to possess the ability for
a delayed fertilization (Amphigonia retardata). Juvenile specimens often exhibit shell shapes and colora-
tions different from those of adults and usually require several years to mature.
Borneo is home to 11 species which represent 3 families and 10 genera. Observations on Mt. Kinabalu
are limited to 2 species representing 2 families. *(here used also for tortoises and terrapins)

Distinguishing features in turtles

nuchal gular
keel
humeral
marginal
length of carapace (CL)

axillar
length of plastron (PL)

costal
pectoral
Carapace inframareginal
abdominal
central inguinal
femoral
Plastron
anal
caudal
width of carapace (CW)
Fig. CXVII Scutes of the shell of a turtle.

Key to the families of Bornean turtles (after MANTHEY & GROSSMANN 1997)
1 Limbs elongate, substantially flattened, flipper- A B
like (A) ............................................ Cheloniidae
• Limbs short, more or less round ....................... 2
2 Shell soft, leather-like, flat, without horny scutes
(B) ................................................. Trionychidae C D E
• Shell hard, with horny scutes (C) ...................... 3
3 Feet with distinct toes (D) ............. Bataguridae
• Feet without free toes, “elephant foot-like” (E) .
......................................................... Testudinidae

395
Bataguridae

Bataguridae
The family Bataguridae comprises about 70 species which are distributed in West and Southeast Europe,
North Africa, West, Central, South, and Southeast Asia. The genus Rhinoclemmys is the only representa-
tive of this family to occur in Central and South America. Save for a few exceptions, Batagurids are
highly adapted to a life at and in water. Their bony shells are fairly variable in shape, but usually rather
flat, and always covered with horny scutes. Both halves of the shell are usually rigid, but in some species
they are united by connective tissue that allows for flexibility. Due to these hinges parts of the ventral
shell can be moved (e.g. Cuora species). The limbs bear free toes which are interlinked by membranes,
usually with 5 claws on the front and 4 claws on the hind feet.
Asian tortoises are in general omnivores, which consume animal as well as vegetarian food, but there are
also some specialized feeders. They reproduce by laying eggs with soft, parch-like shells in varying
numbers.
On Borneo, this family is represented by 9 species in 8 genera with only one genus and one species native
to Mt. Kinabalu.

Key to the Bornean genera of Bataguridae (in part after MANTHEY & GROSSMANN 1997)

1 6 or 7 centrals (A) .............................. Notochelys A B

• 5 centrals ............................................................ 2
2 Front feet with 4 claws (B) .................... Batagur
• Front feet with 5 claws ...................................... 3
3 Axillar and inguinal scutes present (C) ............. 4
• Axillar and inguinal scutes tiny or absent, ante-
rior and posterior sections of plastron joined by a
hinge (D) and movable ............................. Cuora C D
4 Anterior section of plastron with hinge (D) and
movable ................................................ Cyclemys
• Plastron without hinges ..................................... 5
5 Plastron with brown radial pattern (E) ................
............................................................ Heosemys
• Plastron without radial pattern .......................... 6
6 Carapace olive to brownish, males with three black
stripes and short nasal trunk ................. Callagur E
• Carapace dark brown to blackish, males without
nasal trunk and no striped pattern ..................... 7 F
7 Whitish yellow spots on eyes, angles of mouth,
and occiput (F), plastron blackish .......................
..................................................... Siebenrockiella
• No whitish yellow spots on the head, plastron
uniformly light .......................................... Orlitia

396
 Bataguridae

Genus Heosemys STEJNEGER, 1902

Heosemys inhabit the southwest of India, South-

east Asia, the Sunda region, and the Philippine Is-
lands.
Semiaquatic terrapins with a distinct vertebral keel
on a flattened carapace; anterior marginals usually
smooth, posterior ones serrated; plastron joined rig-
idly with carapace; toes of the front feet with 5,
those of the hind feet with 4 claws.
Only one species of this genus is represented on

© W. Grossmann
Borneo and also occurs on Mt. Kinabalu.

Heosemys spinosa (GRAY, 1831)

Fig. 378 Ventral view of a juvenile to semiadult speci-
Emys spinosa GRAY, J.E. (1831): 20 – Type locality: Penang men of Heosemys spinosa.
Forest Hill Tortoise, JACOBSON 1979: 33, fig. 32a
Geoemyda spengleri spengleri, (not Testudo spengleri GMELIN,
1789) MANTHEY 1982: 31
Geoemyda spengleri, (not Testudo spengleri GMELIN, 1789) distinctly serrated marginal scutes (limited to the
MALKMUS 1994: 246 anterior and posterior ones in adult specimens).

Diagnosis Description
A moderately large (CL up to 22 cm), brown to CL up to 22 cm, PL up to 18 cm; shell flat, very
reddish brown terrapin with a vertebral keel and wide; 5 strongly keeled centrals, all wider than long;

Fig. 377 Dorsal view of a juvenile to semiadult specimen of Heosemys spinosa.

© W. Grossmann

397
 Bataguridae

4 costals, each with a spike in young, smooth in
old specimens; 11 marginals, terminating in a point
in juveniles, tips of the anterior and posterior
marginals slightly raised, lateral marginals of old
specimens almost smooth; 1 nuchal longer than
wide, 2 supracaudals; 1 axillar, 1 inguinal, no
inframarginals; plastron large with wide bridge.
Carapace of juveniles almost round and very spiky,
more elongate with increasing age and progres-
sively smoother. Limbs with large horny scales;
front toes webbed over half their lengths, hind toes
only with basal membranes; inguinal region and base
of tail with pointed tubercles; tail very short.
Carapace reddish to reddish light or dark brown,
vertebral keel light brown throughout in young
specimens; plastron yellowish with a brown ra-
dial pattern on each scute in juveniles, in various
shades of brown without radial pattern in old
specimens; head and limbs brown to greyish
brown, side of neck with a small yellowish, red-
dish, or light brown spot.

Ecology/Ethology
H. spinosa inhabits small,
shallow, fast-flowing,
clear streams in the forests
covering the hills and
mountains, but is a rather
poor swimmer. It often
leaves the water for longer
periods of time to frequent
shaded moist locations or
to bury itself in the layer
of leaf litter. It feeds pri-
marily on plants and fruits,
but snails and worms are
© W. Grossmann

also consumed. Activity is

limited to the cooler hours
of the mornings and eve-
nings.
1-3 large, elongate eggs,
CL of hatchlings 51-
63 mm.

Distribution
Burma, Thailand, Penin-
sular Malaysia, Singa-
pore, Sumatra incl. Pulau
Bangka, Batu Islands, and
Pulau Natuna Besar, Bor-
neo. Mt. Kinabalu: with-
out precise locality.
© W. Grossmann

From top to base: Heosemys

spinosa (adult, CL 20 cm)
Fig. 379 Dorsal view.
Fig. 380 Ventral view.

398
 Trionychidae

Trionychidae
This family of cryptodirans includes about 30 species. Their distribution range extends from North America,
through Africa and Asia east to New Guinea.
They are characterized by soft shells of roundish to elongate-oval shapes. The carapace and plastron are
not covered by horny scutes but a leathery skin. With marginal bones being absent, the halves of the
shells are joined by ligaments. The surface of the plastral bones is distinctly structured, usually leaving
their depressions (callosities) well visible through the skin. The elongate heads of soft-shelled turtles
terminate in trunk-like noses of various lengths. Their horny beaks are covered with fleshy lips. The
individual genera are mainly distinguished by anatomical traits, such as the absence of presence of bony
plates, their numbers and situations. Both the front and hind feet are furnished with three free claws each.
Soft-shelled turtles have a reputation for biting more or less readily. They feed mainly on other animals
and reproduce by laying eggs in larger numbers.
On Borneo, this family is represented by 3 species in 3 genera with only one genus and one species native
to Mt. Kinabalu.

Key to the Bornean genera of Trionychidae (after MANTHEY & GROSSMANN 1997)

1 Long nasal trunk present (A) ............................ 2 B

A
• Long nasal trunk absent (B) ............ Pelochelys
2 Carapace with vertebral stripes (C), 4 callosities
............................................................. Dogania
• Carapace without vertebral stripes, 5 callosities
.................................................................. Amyda C

Genus Dogania GRAY, 1844

Dogania is distributed from Myanmar and the

Malay Peninsula to Java, Borneo and the Philip-
pines.
Relatively small soft-shelled turtles (CL up to 35 cm)
with an oval carapace and a dark, often indistinct,
vertebral stripe. Large elongate head with long na-
sal trunk.
The genus is monotypical.

Dogania subplana (GEOFFROY, 1809)

© W. Grossmann

Trionyx subplanus GEOFFROY, 1809: 11, pl. 5, fig. 2 – Type

locality: unknown
Dogania subplana, DE ROOIJ 1915: 326; MANTHEY 1983: 30;
MALKMUS 1994a: 246
Pelochelys bibroni (not Trionyx bibronii OWEN, 1853), JACOBSON
Fig. 381 Portrait of Dogania subplana from Peninsular
1986: pl. 43. Malaysia.

399
Trionychidae

Description
CL up to 35 cm, carapace very
flat, oval; plastron with 4 weakly
developed callosities; head
large, elongate, with long snout,
terminating in a long nasal trunk;
feet flipper-like, each carrying 3
claws, all toes webbed over their
entire lengths; tail very short,
longer in males than in females.
Carapace olive greyish brown or

© W. Grossmann
yellowish brown, with a median
black stripe and 2-3 pairs of
black spots, without any addi-
tional pattern, or with irregularly
shaped and scattered yellowish,
more rarely reddish brown or
brown, spots; plastron white,
yellowish, cream-coloured, or
grey; head grey to olive, spotted
and striped with black; limbs
olive grey, with small yellow
dots; sides of head and lateral
part of nape red in juveniles,
ventral side bluish, fading with
increasing age.
3-7 perfectly round eggs (22-

© W. Grossmann
31 mm in diameter).

Ecology/Ethology
D. subplana shows a preference
for clean, cool, fast-flowing
streams in lowland and hilly
country. It climbs very well and
often seeks shelter among rocks
outside the water. D. subplana
is crepuscular to nocturnal,
spending most of the day in hid-
ing. It feeds predominantly on
other animals.

Distribution
© W. Grossmann

Myanmar incl. Mergui Islands,

Thailand, Peninsular Malaysia,
Singapore, Borneo, Sumatra
incl. Pulau Singkep and Pulau
Natuna Besar, Java, Philippine Fig. 382-384 Different colour pattern of the carapace in Dogania subplana
Islands. Mt. Kinabalu: Sayap. from Peninsular Malaysia.

400

Glossary
abiotic: inanimate; physical and chemical aspects
of the environment of an organism (contrary
to biotic)
adaptive radiation: the evolutionary development
of new species from an existing species as a
result of adaptations to various ecological
niches within a common geographical distri-
bution range
adult: sexually mature, grown-up
allopatric: living in different geographical regions
(contrary to sympatric)
amphigonia retardata: 1: delayed fertilization,
2: the storage of viable sperm by a female
amplexus: the mating position of anuran amphib-
ians, the embracing of the female‘s body by
the male‘s forelimbs
annuli: rings or ring-like markings
apical: at the tip of a structure
aquatic: living in the water
arboreal: living in a forested environment
arboricol: living on trees
autotomy: the capability to break off the tail, or
parts of it, at predesignated places as a defence
mechanism
benthic community: plants and animals which
inhabit the bottom of aquatic ecosystems
biodiversity (biological diversity): all aspects of
the diverse animated world (e.g. species diver-
sity, genetic diversity, diversity of ecosystems)
canthus rostralis: skin-covered, bony ridge be-
tween eye and nostril
carnivore: meat-consuming species
caudal: referring to the tail
conspecific: belonging to the same species
convergence: evolutionary process through which
unrelated, geographically often widely sepa-
rated species develop similar structures (habi-
tus, behaviour) due to similar ecological chal-
lenges
crepuscular: active at twilight hours (dawn and/
or dusk)
distal: more distant from the centre of the body or
a defined point of reference than other parts of
the body or an organ
Glossary
diurnal: active during the day
dorsal: referring to the back
dorsolateral: referring to the side of the back
dorsoventral: in a vertical direction
endemic: being exclusive to a certain geographi-
cal region; if a species has developed within
that region it is called autochthonous
endotrophic: feeding on internal resources; here:
tadpoles which develop without a free-rang-
ing stage and are therefore independent from
an external food supply; they develop directly
from the yolk mass.
fossorial: living subterraneanly (underground)
genotype: all genetic traits which determine the
structure and function of an organism
gravid: carrying eggs, being “pregnant”
habitat: the natural environment an animal lives
in
habitus: general appearance, body build
hemipenis: paired mating organ of male reptiles
inc. sed.: incertae sedis, preliminary taxonomic
assignment
incubation: the development (“brooding”) of eggs
infra-: (prefix) lower
inguinal: being situated in or near the groin
isolation mechanism: any genetically induced dif-
ference between species which prevents these
from mating under natural circumstances
juvenile: not yet sexually mature, not fully grown
lateral: referring to the side
mandible: the lower jaw
maxilla: the upper jaw
median: referring to the middle or centre
microhabitat: a portion of the macrohabitat which
offers the environmental circumstances an or-
ganism requires for its survival
mimesis: the adaptation of coloration, body shape,
and/or behaviour to the immediate environment
by mimicking respective structures of the lat-
ter in order to escape the attention of potential
predators (e.g. the imitation of plant structures
º phytomimesis)
monotypic: containing only one unit per superior
systematic rank
401
Glossary

natural selection: the contribution of various geno- syntopic: occurring together, inhabiting the same
types to the offspring of the subsequent gen- biotope
eration; over time this results in changes to the taxon (pI. taxa): any scientifically named category
genetic composition of populations of plants or animals (e.g. species, genus, fam-
niche partitioning: the tendency of co-existing ily)
species to occupy different ecological niches taxonomy: the science of classifying organisms
within their commonly inhabited habitat terminal: “far from the centre”, belonging to the
niche: the specific habitat within an ecosystem typi- most distant end
cally inhabited by a species type locality: locus typicus: the locality at which
nocturnal: active at night the holotype of a species has been collected
omnivore: a species that consumes both plant and type specimen: most holotype; reference specimen
animal material upon which a species has been described sci-
oviposition: the lying of eggs entifically
papilla: a small, fleshy projection ventral: referring to the belly
paravertebral: being situated next to the vertebral ventrolateral: being situated on the side of the
column belly
phenotype: the obvious traits of an organism which vertebral: referring to the centre or middle line of
are the result of the interaction between geno- the back, along the vertebral column
type and environment
phylogenesis: the evolutionary history of a taxo-
nomic group
population: group of individuals of a species which
Conversion table
inhabit a certain geographical region at a cer-
tain point of time 1 cm (centimetre) = 0.39 in. (inch)
reproductive isolation: spatial and temporal sepa- 1 m (metre) = 3.2808 ft. (feet)
ration of populations of the same species; an 1 g (gram) = 0.04 oz. (ounces)
important precondition for the development of 1 l (litre) = 0.264 gal. (American gallons)
new species 1°C (degree Celsius) = 2.63°F (degrees Fahrenheit)
resource: everything an organism requires for its
existence (e.g. food, spawning sites, etc.)
sexual dimorphism: the differences in body shape
and/or coloration between males and females
of the same species
sic!: really so! (really so written)
sit-and-wait: foraging strategy in which a preda-
tor remains immobile until a prey comes within
striking range
speciation: the process of the development of spe-
cies
stratification: the tiering or storey-like, vertically
organised structures of a habitat (e.g. the vari-
ous storeys of a rainforest)
sub- (prefix): below, beneath, under
supra- (prefix): upper, above
symphysis: the junction between two pieces of
bone
synonym: in systematics, a taxonomic name re-
ferring to an already named species

402

Bibliography
Literature used in the introductory chapters and
further readings are marked by an asterisk (*).
AHL, E. (1927): Zur Systematik der asiatischen Arten der
Froschgattung Rhacophorus. – Sitz. Ber. naturf. Freunde
Berlin: 35-47.
– (1931): Liste der rezenten Amphibien und Reptilien.
Anura III Polypedatidae. – Das Tierreich, Berlin, 55: xvi
+ 477 pp.
ALCALA, A.C. & W.C. BROWN (1982): Reproductive biol-
ogy of some species of Philautus (Rhacophoridae) and
other Philippine anurans. – Philipp. J. Biol., 11: 203-
226.
ALTIG, R. (1970): A key to the tadpoles of the continental
Unites States and Canada. – Herpetologica, Johnson City,
26: 180-207.
ALTIG, R. & G. F. JOHNSTON (1989): Guilds of anuran lar-
vae: relationships among developmental modes, mor-
phologies, and habitats. – Herpetological Monogr., 3:
81-109.
ANDERSSON, L.G. (1923): Some reptiles and batrachians
from Central-Borneo. – Medd. Zool. Mus., Kristiana,
61(7): 119-125.
AULIYA, M & F. ABEL, (1999): Neue Erkenntnisse zu Py-
thon curtus (Taxonomie, Autökologie, Biologie und
Schutz). – herpetofauna, Weinstadt, 21 (120): 25-34.
– (2000a): Zur Taxonomie, geographischen Verbreitung
und Nahrungsökologie des Netzpythons (Python
reticulatus). Teil 1. – herpetofauna, Weinstadt 22 (127):
5-18.
– (2000b): Zur Taxonomie, geographischen Verbreitung
und Nahrungsökologie des Netzpythons (Python
reticulatus). Teil 2. – herpetofauna, Weinstadt, 22 (128):
19-28.
BACON, J.P.Jr. (1967): Systematic Status of Three Scincid
Lizards (Genus Sphenomorphus) from Borneo. –
Fieldiana, Zool., Chicago, 51(4): 63-76.
BARBOUR, T. (1904): A new batrachian from Sarawak,
Borneo. – Proc. Biol. Soc. Washington, 17: 51.
– *(1912): A contribution to the zoogeography of East In-
dian Islands. – Mem. Mus. Comp. Zool., Cambridge,
44(1): 1-203.
– (1921): A new Bornean Lizard. – Proc. New Engl. Zool.
Club, Boston, 7: 87-89.
– (1938): Notes on “Nectophryne”. – Proc. Biol. Soc.
Washington, 51: 191-195.
BARBOUR, T. & G.K. NOBLE (1916): New amphibians and a
new reptile from Sarawak. – Proc. New Engl. Zool. Club,
Boston, 6: 19-22.
Bibliography
BARKER, D. & T. BARKER (1994): The Blood-Python and
other subspecies of the Short-Tailed Python (Python
curtus). – The Vivarium, 6 (3): 30-33, 35.
BARTLETT, E. (1895a): The crocodiles and lizards of Borneo
in the Sarawak Museum, with descriptions of supposed
new species, and the variation of colours in the several
species during life. – J. Str. Br. Roy. As. Soc., Singapore,
28: 73-96.
– (1895b): Notes on the batrachians or frogs and toads of
Borneo and adjacent islands. – Sarawak Note Book, no.
1: 7-14.
– (1895c): Notes on the snakes of Borneo and adjacent
islands. Part II. – Sarawak Note Book, no. 1: 78-85.
BATCHELOR, D.M. (1958): Some notes on the snakes of
Asahan, Malacca. – Malay. Nat. J., Kuala Lumpur, 12:
103-111.
*BEAMAN, J.H. & R.S. BEAMAN (1990): Diversity and dis-
tribution patterns in the flora of Mount Kinabalu. In:
BAAS, P., K. KALMAN & R. GEESNIK (Eds.): The plant
diversity of Malesia – Dordrecht (Kluwer Academic
Publ.): 147-160.
*BEAMAN, R.S., J.M. BEAMAN, C. MARSH & P. WOODS
(1985): Drought and forest fires in Sabah in 1983 – Sabah
Soc. J., 1985: 10-30.
*BECCARI, O. (1904): Wanderings in the great forests of
Borneo – London
BERGMAN, R.A.M. (1956): The anatomy of Dryophis prasinus.
I. – Proc. Kon. Ned. Akad. Wet. Ser. C, 59: 263-271.
BERRY, P. Y. (1972): Undescribed and little-known tadpoles
from West-Malaysia. – Herpetologica, Johnson City,
28(4): 338-346.
– (1975): The amphibian fauna of Peninsular Malaysia. –
Kuala Lumpur (Tropical Press), 130 pp.
BERRY, P.Y. & J. HENDRICKSON (1963): Leptobrachium
nigrops, a new pelobatid frog from the Malay Peninsula,
with Remarks on the Genus Leptobrachium in South-
east Asia. – Copeia, Washington, (4): 643-648.
BLEEKER, P. (1858/59): Reptiliën en visschen van Sinkawang
(westkust van Borneo). – Nat. Tijdschr. Ned. Ind.,
Batavia, 16: 188-189.
– (1859): Reptiliën van Sintang, verzameld door A. H.
THEPASS. – Nat. Tijdschr. Ned. Indië, (4, part II), 20: 200-
202.
– (1860a): Reptiliën van Agam. – Nat. Tijdschr. Ned. Ind.,
Batavia, 20: 325-329.
– (1860b): Over de reptiliën-fauna van Sumatra. – Nat.
Tijdschr. Ned. Ind., Batavia, 21: 284-298.
BOETTGER, O. (1892): Katalog der Batrachier-Sammlung
im Museum der Senckenbergischen naturforschenden
Gesellschaft in Frankfurt/M. – Frankfurt/M (Gebr.
403
Bibliography
Knauer), 73 pp.
– (1893): Übersicht der von Kapt. Storm auf Borneo
gesammelten Reptilien und Batrachier. – Mitt. geogr.
Ges. Lübeck, 2(5): 85-89.
– (1901): Die Reptilien und Batrachier. In: Ergebnisse einer
zoologischen Forschungsreise in den Molukken und
Borneo im Auftrage der Senckenbergischen naturfor-
schenden Gesellschaft, ausgeführt von Dr. Willy
Kükenthal – 2. Teil Wissenschaftliche Reiseergebnisse,
Band III. – Abh. Senck. naturf. Ges., Frankfurt/ M, 25:
321-402.
BOIE, F. (1827): Bemerkungen über Merrem’s Versuch eines
Systems der Amphibien, 1. Lieferung: Ophidier. – Isis
van Oken, Jena, 20: 508-566.
BOSCHMA, H. (1922): Über den Trichterapparat der Larven
von Megalophrys montana KUHL. – Bijdr. Dierk.,
Amsterdam, 22: 9-12.
BOSSUYT, F. & M.C. MILINKOVITCH (2000): Convergent adap-
tive radiations in Madagascan and Asian ranid frogs re-
veal covariation between larval and adult traits. – PNAS,
Athens, 97(12): 6585-6590.
BOULENGER, G.A. (1882): Catalogue of the batrachia
salientia s. ecaudata in the collection of the British Mu-
seum 2. Ed. – London (Trustees Brit. Mus.), 495 pp.
– (1885a): Catalogue of the lizards in the British Museum
(Natural History). I. Geckonidae, Eublepharidae,
Uroplatidae, Pygopodidae, Agamidae. – London (Trust-
ees Brit. Mus.), 575 pp.
– (1885b): A list of reptiles and batrachians from the Is-
land of Nias. – Ann. Mag. Nat. Hist., London, 5. Ser. 16:
388-389.
– (1887a): On new batrachians from Malacca. – Ann. Mag.
Nat. Hist., London, 5. Ser. 19: 345-348.
– (1887b): On new reptiles and batrachians from North
Borneo. – Ann. Mag. Nat. Hist., London, 5. Ser. 20: 95-
97.
– (1887c): Catalogue of the lizards in the British Museum
(Natural History). III. Lacertidae, Gerrhosauridae,
Scincidae, Anelytropsidae, Dibamidae, Chameleontidae.
– London (Trustees Brit.Mus.), 575 pp.
– (1891a): Description of new Oriental reptiles & batra-
chians. – Ann. Mag. Nat. Hist., London, 6. Ser. 7: 279-
283.
– (1891b): Remarks on the herpetological fauna of Mount
Kina Baloo, North Borneo. – Ann. Mag.Nat. Hist., Lon-
don, 6. Ser. 7: 341-345.
– (1891c): On new or little known Indian and Malayan
reptiles and batrachians. – Ann. Mag. Nat. Hist., Lon-
don, 6. Ser. 8: 288-292.
– (1892): An account of reptiles and batrachians collected
by Mr. C. Hose on Mt. Dulit, Borneo. – Proc. Zool. Soc.
London,: 505-508.
– (1893a): Description of new reptiles and batrachians
obtained in Borneo by Mr. A. Everett and Mr. C. Hose. –
Proc. Zool. Soc. London,: 522-528.
404
– (1893b): Catalogue of the snakes in the British Museum
(Nat.Hist.). Vol. I. – London (Taylor & Francis): 1-448.
– (1894a): On the herpetological fauna of Palawan and
Balabac. – Ann. Mag. Nat. Hist., London, 6. Ser. 14: 1-90.
– (1894b): A list of reptiles and batrachians collected by
Dr .E. Modigliani on Sereinu (Sipora), Mentawei Islands.
– Ann. Mus. civ. stor. nat. Genova: 613-618.
– (1894c): Third report on additions to the batrachian col-
lection in the Natural-History Museum. – Proc. Zool.
Soc. London: 640-646.
– (1894d): Second report on additions to the lizard collec-
tion in the Natural History Museum. – Proc. Zool. Soc.
London: 722-736.
– (1894e): Catalogue of the snakes in the British Museum
(Natural History). Vol. II. Containing the conclusion of
the Colubridae Aglyphae. – London, (Taylor & Francis):
xi + 382 pp., pl. 1-20.
– (1895a): Description of 4 new batrachians discovered
by Mr. Charles Hose in Borneo. – Ann. Mag. Nat. Hist.,
London, 6. Ser. 16: 169-171.
– (1895b): Description of two new snakes of the genus
Calamaria. – Ann. Mag. Nat. Hist., London, 6. Ser. 16:
481.
– (1896a): Description of new batrachians in the British
Museum. – Ann. Mag. Nat. Hist., London, 6. Ser. 17:
401-403.
– (1896b): Description of two new batrachians obtained
by Mr. A. Everett on Mount Kinabalu, North Borneo. –
Ann. Mag. Nat. Hist., London, 6. Ser. 17: 449-450.
– (1896c): Catalogue of the snakes in the British Museum
(Nat. Hist.). Vol. III. Containing the Colubridae
(Opisthoglyphae and Proteroglyphae), Amblycephalidae
and Viperidae. – London (Taylor & Francis): xiv + 727
pp., 25 pls.
– (1898): Third report on additions to the lizard collection
in the Natural History Museum. – Proc. Zool. Soc. Lon-
don: 912-923, 1 pl.
– (1899): Description of three new reptiles and a new batra-
chian from Mt. Kina Balu, North Borneo. – Ann. Mag.
Nat. Hist., London, 7. Ser. 4: 451-453.
– (1900a): Description of new reptiles and batrachians from
Borneo. – Proc. Zool. Soc. London: 182-187.
– (1900b): Description of new reptiles from Perak, Malay
Peninsula. – Ann. Mag. Nat. Hist., London, 7. Ser. 5:
306-308.
– (1908): A revision of the Oriental pelobatid batrachians
(genus Megalophrys). – Proc. Zool. Soc. London: 407-430.
– (1918): Remarks on the batrachian genera Cornufer,
TSCHUDI, Platymantis, GÜNTHER, g.n. and Staurois, COPE. –
Ann. Mag. Nat. Hist., London, 9. Ser. 1: 372-375.
– (1920): A monograph of the South Asian, Papuan,
Melanesian and Australian frogs of the genus Rana. –
Rec. Ind. Mus., Calcutta, 20: 1-226.
BOURRET, R. (1942): Les Batraciens de l’Indochine. – Hanoi
(Institut océanographique de l’Indochine), x + 547 pp.

*BRANDON-JONES, D. (1996): The Asian Colobinae as indi-
cators of Quaternary climatic change – Biol. Jour. Lim.
Soc., 59: 327-350.
BRONGERSMA, L.D. (1937): On a small collection of am-
phibia from Central East Borneo. – Zool .Meded. Leiden,
20: 6-9.
– (1946): Note on Idiopholis collaris MOcQUARD. – Zool.
Meded., Leiden, 26: 249-250.
– (1948): Notes on Maticora bivirgata (BOIE) and on
Bungarus flaviceps REINH. – Zool. Meded. Leiden, 30(1):
1-29.
BROWN, A.E. (1902): A collection of reptiles & batrachians
from Borneo and the Loo Choo Islands. – Proc. Acad.
Nat. Sci. Philadelphia: 175-186.
BROWN, W.C. & A.C. ALCALA (1980): Philippine Lizards of
the family Scincidae. – Silliman Univ. Nat. Sci.,
Dumaguete City, Mon., Ser. 2: xi + 246 pp.
BROWN, R. M., A. E. LEVITON & V. S. ROGELIO (1999): De-
scription of a new species of Pseudorabdion (Serpentes:
Colubridae) fom Panay Island, Philippines with a revised
key to the genus. – Asiatic Herp. Res., 8: 7-12.
BULIAN, J. (2000): Zur Kenntnis von Boiga nigriceps
(GÜNTHER, 1863). – Reptilia, Münster 5 (5): 56-60.
BULIAN, J. & B. SIEBERT (2000): Keine Probleme mit
Ahaetulla prasina (BOIE, 1827) - Freilandbeobachtungen,
Haltung, Zucht und Aufzucht. – Sauria, Berlin, 22 (3):
33-39.
CHANNING, A. (1989): A re-evaluation of the phylogeny of
Old World treefrogs. – S.-Afr. Tydskr. Dierk., 24(2): 116-
131.
COCHRAN, D.M. (1926): A new Pelobatid batrachian from
Borneo. – J. Wash. Acad. Sci., 16(16): 446-447.
COPE, E.D. (1860): Catalogue of the Colubridae in the
Museum of the Academy of Natural Sciences of Philadel-
phia. Part 3. – Proc. Acad. Nat. Sci. Philadelphia, 12:
553-566.
– (1865): Sketch of the primary groups of Batrachia
Salienta. – Nat. Hist. Rev.: 97-120.
*CORNER, E.J.H. (1978): Plant Life. In: Kinabalu, Summit
of Borneo – Kota Kinabalu (Sabah Soc. Monogr.): 113-
178.
CUNDALL, D., V. WALLACH & D. A. ROSSMAN (1993): The
systematic relationships of the snake genus Anomochilus.
– Zool. J. Linn. Soc., 109: 275-299.
DAS, I. (1995): Amphibians and reptiles recorded at Batu
Apoi, a lowland dipterocarp forest in Brunei Darussalam. –
Raffles Bull. Zool., Singapore, 43(1): 157-180.
– (1996): Spatio-temporal resource utilization by a Bornean
rainforest herpetofauna: Preliminary results. – Tropical
Rainforest Research: 315-323.
DAS, I. & A.M. BAUER (1998): Systematics and biogeogra-
phy of Bornean geckos of the genus Cnemaspis Strauch,
1887 (Sauria: Gekkonidae), with the description of a new
species. – Raffles Bull. Zool. 46(1): 11-28.
DAS, I. & G. ISMAIL (2001): A Guide to the Lizards of
Bibliography
Borneo. – www.arbec.com.my/lizards/
DAVID, P. & G. VOGEL (1996): The snakes of Sumatra. An
annotated checklist and key with natural history notes. –
Frankfurt am Main (Edition Chimaira), [second un-
changed edition 1997]: 1-261.
– (1998): Redescription of Trimeresurus huttoni Smith,
1949 (Serpentes, Crotalinae), with a discussion of its
relationships. – Hamadryad, 22 (2): 73-87.
DAVID, P., G. VOGEL & O. PAUWELS (1998): Amphiesma
optatum (HU & DJAO, 1966) (Serpentes, Colubridae): an
addition to the snake fauna of Vietnam, with a list of the
species of the genus Amphiesma and a note on its type
species. – J. Taiwan Mus., 51(2): 83-92.
*DAVIS, D.D. (1962): Mammals of the lowland rainforest of
North Borneo – Bull. Nat. Mus. Singapore, 31: 1-129.
DAVISON, G.W.H. (1984): Foot-flagging display in Bornean
frogs. – Sarawak Mus. J., Kuching, 33(54): 177-178.
DENZER, W. (1994a): Tree hole breeding in the toad
Pelophryne brevipes (PETERS, 1867). – Amph. Rept.,
Leiden 15: 224-226.
– (1994b): Zur Biologie von Pelophryne brevipes (PETERS,
1867). – herpetofauna, Weinstadt, 16(93): 17-20.
– (1996): Reptilien und Amphibien der Santubong Halbinsel,
Sarawak, Borneo. – Sauria, Berlin, 18(4): 35-42.
DESPAX, M.R. (1912): Sur trois collections de reptiles et de
batraciens provenant de l’Archipel Malais. – Bull. Mus.
Hist. Nat. Paris, 18: 198-205.
DRING, J. (1983a): Frogs of the genus Leptobrachella
(Pelobatidae). – Amphibia-Reptilia, Leiden, 4: 89-102.
– (1983b): Some new frogs from Sarawak. – Amphibia-
Reptilia, Leiden, 4: 103-115.
– (1987): Bornean treefrogs of the genus Philautus (Rhaco-
phoridae). – Amphibia-Reptilia, Leiden, 8: 19-47.
DUBOIS, A. (1980): Notes sur la systématique et la répartition
des Amphibiens Anoures de Chine et des régions avois-
inantes. IV. Classification générique et subgénérique des
Pelobatidae Megophryinae. – Bull. Soc. Linn. Lyon, 49:
469-482.
– (1981): Liste des genres et sous-genres nominaux de
Ranoidea (Amphibiens Anoures) du monde, avec iden-
tification de leurs espèces-types: conséquences
nomenclaturales. – Monit. zool. ital., (n.s.), suppl. 15:
225-284.
– (1982): Leptophryne FITZINGER, 1843, a senior synonym
of Cacophryne DAVIS, 1935 (Bufonidae). – J. Herp., 16:
173-174.
– (1983): Note préliminaire sur le genre Leptolalax DUBOIS,
1980 (Amphibiens, Anures), avec diagnose d’une espèce
nouvelle du Vietnam. – Alytes, Paris, 2(4): 147-153.
– (1986): Miscellanea taxinomica batrachologica (1). –
Alytes, Paris, 5(1-2): 7-95.
– (1992): Notes sur la classification des Ranidae
(Amphibiens Anoures). – Bull. mens. Soc. linn. Lyon,
61(10): 305-352.
D UBOIS , A. & A. O HLER (1998): A new species of
405
Bibliography
Leptobrachium (Vibrissaphora) from northern Vietnam,
with a review of the taxonomy of the genus Leptobrachium
(Pelabatidae, Megophryinae). – Dumerilia, Paris, 4(1):
1-32.
– (2000): Sytematics of Fejervarya limnocharis (GRAVEN-
HORST, 1829) (Amphibia, Anura, Ranidae) and related
species. 1. Nomenclatural status and type-specimens of
the nominal species Rana limnocharis GRAVENHORST,
1829. – Alytes, Paris, 18(1-2): 15-50.
DUELLMANN, W.E. (1993): Amphibian Species of the World:
Additions and Corrections. – Univ. Kansas Mus. Nat.
Hist., Spec. Publ. No. 21: iii + 372 pp.
DUELLMANN, W.E. & L. TRUEB (1986): Biology of Amphib-
ians – New York ( McGraw Hill Book Comp.), 670 pp.
DUMÉRIL, A.H.A. & G. BIBRON (1836): Erpétologie générale
ou Histoire Naturelle complète des Reptiles. – Paris, Vol.
III, iv + 518 pp.
– (1839): Erpétologie générale ou Histoire Naturelle
complète des Reptiles. – Paris, Vol. V, viij + 855 pp.
DUMÉRIL, A.M.C., G. BIBRON & A.H.A. DUMÉRIL (1854a):
Erpétologie générale ou Histoire Naturelle complète des
Reptiles. – Paris, Vol. VII (1), xvi + 780 pp.
– (1854b): Erpétologie ou Histoire Naturelle complète des
Reptiles. – Paris, Vol. VII (2), xii + 784-1556.
DUNN, E.R. (1923): On a collection of reptiles from
Sarawak. – J. Malay. Br. Royal Asiat. Soc. 1: 1-4.
EBENHARD, T. & P. SJÖGREN (1984): Life on a tropical moun-
tain. 2. The reptiles of Mt. Kinabalu. – Nature
Malaysiana, 9(3): 16-27.
EDELING, A.C.J. (1865): Recherches sur la faune erpéto-
logique de Bornéo. – Ned. Tijdschr. Dierk., Leiden, 2:
200-204.
EMERSON, S.B. (1991): The ecomorphology of Bornean tree
frogs (family Rhacophoridae). – Zool. J. Linn. Soc.
101(4): 337-357.
– (1992): Courtship and nest-building behavior of a
Bornean frog, Rana blythi. – Copeia, Washington, (4):
1123-1127.
EMERSON, S.B. & D. BERRIGAN (1993): Systematics of South-
east Asian Ranids: Multiple origins of voicelessness in
the subgenus Limnonectes (FITZINGER). – Herpetologica,
Johnson City, 49(1): 22-31.
EMERSON, S.B. & R.F. INGER (1992): The comparative ecol-
ogy of voiced and voiceless Bornean frogs. – J. Herp.
26(4): 482-490.
EMERSON, S.B., R.F. INGER & D. ISKANDAR (2000): Molecu-
lar systematics and biogeography of the Fanged Frogs
of Southeast Asia. – Molecular Phylogenetics and Evo-
lution, 16(1): 131-142.
EMERSON, S.B. & M.A.R. KOEHL (1990): The ineractions
of behavioral and morphological change in the evolu-
tion of a novel locomotor type: “Flying” frogs. – Evolu-
tion 44(8): 1931-1946.
ERDELEN, W., F. ABEL & M. RIQUIER (1997): Status, Popu-
lationsbiologie und Schutz von Bindenwaran (Varanus
406
salvator), Netzpython (Python reticulatus) und
Blutpython (Python curtus) in Sumatra und Kalimantan,
Indonesien. Abschlußbericht zum Forschungsprojekt im
Auftrag des Bundesministeriums für Umwelt, Natur-
schutz und Reaktorsicherheit. (Unveröffentlicht) 157 pp.
EVEN, E. (1997): Blood Pythons, Python curtus; subspe-
cies, care and breeding. – Litt. Serpent., 17(6): 116-124.
FEI, L, YE, C. & Y. HUANG (1991) Key to Chinese Am-
phibia. – Chongqing, Editions of Sciences and Tech-
niques, i-iv + 1-2 + 1-364. [In Chinese]
FISCHER, J.G. (1885): Über eine Kollektion von Amphibien
und Reptilien aus Südost-Borneo. – Arch. Naturgesch.,
Berlin: 41-72.
FITZINGER, L.J. (1826): Neue Classification der Reptilien nach
ihren natürlichen Verwandtschaften. Nebst einer
Verwandtschafts-Tafel und einem Verzeichnisse der
Reptiliensammlung des k. k. zoologischen Museums zu
Wien. – Wien (J. G. Huebner), viii + 66 pp.
– (1843): Systema Reptilium. Fasc. 1. Amblyglossae –
Vindobonae. – Wien (Braunüller & Seidel), 106 pp.
FRAHM, J.B., W. FREY, H. KÜRSCHNER & M. MENZEL (1990):
Mosses and liverworts of Mt. Kinabalu – Kota Kinabalu
(Sabah Parks Publ. 12)
FROST, D.R. (ED.) (1985): Amphibian species of the world.
A taxonomic and geographical reference. – Lawrence
(Allen Press & The Association of Systematic Collec-
tions), 732 pp.
– (2000): Amphibian species of the world: An online ref-
erence. V2.20 (1. September 2000)
FUKADA, H. (1964): A small collection of snakes of the
Kyoto University Borneo Expedition. – Bull. Kyoto
Gakugei Univ., Ser. B, 25: 71-77.
GEOFFROY, S. H. (1809): Sur les tortues molles, nouveau
genre sous le nom de Trionyx, et sur la formation des
carapaces. – Ann. Mus. Hist. nat., Paris, 14: 1-20.
*GHAZALLY, I. & L. DIN (1995): Sayap - Kinabalu Park
Sabah – Petaling Jaya (Pelanduk Publ.)
GMELIN, J. F. (1789): Caroli a Linné Systema per Regna
tria Naturae. – G.E. Beer, Leipzig, 1(3): 1033-1516.
GLAW, F. & M. VENCES (1994): A Fieldguide to the Am-
phibians and Reptiles of Madagascar. – Leverkusen (M.
Vences & F. Glaw Verlags GbR), 480 pp.
*GOOSE, J.P. (1972): Fishes of Kinabalu National Park –
Bull. Inst. Sci. Nat. Belg., 48 (3).
GOSSNER, U. L. (1960): A simplified table for staging anu-
ran embryos and larvae with notes on identification. –
Herpetologica, Johnson City, 16: 183-190.
GRAVENHORST, J.L.C. (1829): Deliciae Musei zoologici
Vratislaviensis continens Chelonias et Batrachia, Fasc.I:
xiii + 104 pp.
GRAY, J.E. (1827): A synopsis of the genera of saurian rep-
tiles in which some new genera are indicated, and the
others reviewed by actual examination. – Philos. Mag.,
London, 2(2): 54-58.
– (1831a): Zoological miscellany. – London, 1-86.

– (1831b): Synopsis Reptilium or short description of the
species of reptiles. Part I. – Cataphracta. Tortoises, Croco-
diles, and Enaliosaurians. – London (Treuttel, Wurtz &
Co) Part 1: vii + 85 pp.
– (1839): Catalogue of the slender-tongued saurians, with
descriptions of many new genera and species. – Ann.
Mag. Nat. Hist., London, 2: 331-337.
– (1842): Description of some new species of reptiles,
chiefly from the British Museum collection. – The Zoo-
logical Miscellany: 57-59.
– (1845): Catalogue of the specimens of Lizards in the
collection of the British Museum. – London (British
Museum Natural History), xxvii + 289 pp.
GRAY, J. E. & T. HARDWICKE (1830-1832): Illustrations of
Indian zoology; chiefly selected from the collection of
Major-General Hardwicke, F.R.S., L.S., M.R.A.S., M.R.I.A.,
&c., Volume I (parts I-X). – London (Treuttel, Wurtz,
Treuttel, Jun. & Richter): (1+2), pl. 1-100.
– (1832b-1835): Illustrations of Indian zoology; chiefly
selected from the collection of Major-General Hardwicke,
F.R.S., L.S., M.R.A.S., M.R.I.A., & C. Volume II (parts XI-
XX). – London (Adolphus Richter and Co & Parbury,
Allen, and Co): (1+2), pl. 1-102.
GREER, A.E. (1970): The relationship of the skinks referred
to the genus Dasia. – Breviora, Cambridge, 348: 1-30.
– (1974): The generic relationships of the scincid lizard
genus Leiolopisma and its relatives. – Austr. J. Zool.,
Suppl. Ser., No. 31: 1-67.
– (1977): The systematics and evolutionary relationships
of the scincid lizard genus Lygosoma – J. nat. Hist., 11:
515-540.
GRIJS, P. DE (1937): Eine neue Eidechse aus Nord-Borneo:
Calotes kinabaluensis. – Zool. Anz. Leipzig, 117: 136-
138.
*GRISWOLD, J.A. (1939): Up Mt. Kinabalu. – Sci. Mon. N.Y.,
116: 401-414, 504- 518.
GÜNTHER, A. (1858a): Catalogue of Colubrine snakes of
the British Museum. – London, xvi + 281 pp.
– (1858b): Catalogue of the Batrachia Salientia in the col-
lection of the British Museum. – London, 160 pp.
– (1863): Third account of new species of snakes in the
collection of the British Museum. – Ann. Mag. Nat. Hist.,
London, 3. Ser. 12: 348-365.
– (1864): The reptiles of British India. – London, xxvii +
452 pp., 26 pls.
– (1865): Fourth account of new species of snakes in the
collection of the British Museum. – Ann. Mag. Nat. Hist.,
London, 3. Ser. 15: 89-98.
– (1872): On the reptiles and amphibians of Borneo. – Proc.
Zool. Soc. London: 586-600.
HAAS, C.P.J. DE (1949): The genus Natrix in the collection
of the Raffles Museum and its distrubution in the Malay
Peninsula. – Bull. Raffles Mus., Singapore, 19: 78-97.
– (1950): Checklist of the snakes of the Indo-Australian
Archipelago (Reptilia, Ophidia). – Treubia, Bogor, 20:
Bibliography
511-625.
HaILE, N.S. (1963): Snake bites man: two recent Borneo
cases. – Sarawak Mus. J., Kuching, 11: 291-298.
*HALL, R. & J.D. HOLLOWAY (Eds.) (1998): Biogeography
and geological evolution of SE Asia – Leiden (Backb.
Publ.).
HANITSCH, R. (1900a): An expedition to Mt.Kinabalu, Brit-
ish North Borneo. – J. Str. Br. Roy. As. Soc., Singapore,
34: 49-88.
– (1900b): Notes on the flying frog Rhacophorus
nigropalmatus. – J. Str. Br. Roy. As. Soc. Singapore, 34:
96-97.
– (1900c): Check List of the Reptiles and Amphibians in
the Raffles Museum, Singapore. – Singapore, 6 pp.
HARDING, K.A. (1982): Courtship display in a Bornean frog.
– Proc. Biol. Soc. Washington, 95(3): 621-624.
HARDWICKE, F.R. & J.E. GRAY (1827): A synopsis of the
species of saurian reptiles, collected in India by Major-
General Hardwicke. – Zool. J. London, 3: 214-229.
HAUSCHILD, A. (1998): Haltung und Nachzucht des Duméril-
Warans, Varanus dumerilii (S CHLEGEL , 1839). –
herpetofauna, 20(115): 27-34.
HENNIG, W. (1936): Revision der Gattung Draco (Agami-
dae). – Temminckia, Leiden, 1: 153-220.
HERRMANN, H.J. & T. ULBER (1992): Über Haltung sowie
Verhalten von Ansonia hanitschi INGER, 1960 (Anura:
Bufonidae) in Biotop und Terrarium. – Sauria, Berlin,
14(2): 15-18.
HEYER, W. R. (1971): Mating calls of some frogs from Thai-
land. – Fieldiana Zool., Chicago, 58(6): 61-82.
HIKIDA, T. (1982): A new limbless Brachymeles (Sauria:
Scincidae) from Mt.Kinabalu, N.-Borneo. Copeia, Wa-
shington, (4): 840-844.
– (1990): Bornean Gekkonid lizards of the genus
Cyrtodactylus (Lacertilia: Gekkonidae) with descriptions
of three new species. – Jpn. J. Herpetol. 13(3): 91-107.
HIKIDA, T. & H. OTA (1994): Sphenomorphus aquaticus
MALKMUS, 1991, a junior synonym of Tropidophorus
beccarii (PETERS, 1871) (Reptilia: Squamata: Scincidae).
– Bonn. zool. Beitr. 45(1): 57-60.
HNIZDO, J. (2000): Neue Erkenntnisse zur Fortpflan-
zungsbiologie von Ahaetulla prasina (BOIE, 1827). –
Sauria, Berlin, 22 (2): 41-44.
HOFFMANN, P. (1994): Fund der seltenen Rana palavanensis
am Mt. Kinabalu auf Borneo. – Salamandra 30(3): 223-
224.
– (1995a): Untersuchungen zur Anurenfauna des Mt.
Kinabalu (Borneo) – Teil I: Einführung, Familien
Pelobatidae, Bufonidae, Microhylidae. – Sauria, Berlin,
17(1): 3-10.
– (1995b): Untersuchungen zur Anurenfauna des Mt.
Kinabalu (Borneo) – Teil II: Familien Ranidae,
Rhacophoridae. – Sauria, Berlin, 17(3): 9-17.
– (1995c): Zur Kenntnis des Engmaulfrosches Chaperina
fusca MOCQUARD, 1892 vom Mt. Kinabalu. – herpeto-
407
Bibliography
fauna, Weinstadt, 17(96): 27-29.
– (1998a): Brutpflegende Wasserfrösche aus Borneo. –
DATZ, Gelsenkirchen, 51(6): 384-385.
– (1998b): Mount Kinabalu – Skizze eines Lebensraumes.
– Elaphe, 6(2): 83-90.
– (1998c): Anmerkungen zur Fortpflanzungsbiologie und
zum Ruf von Philautus petersi (BOULENGER, 1900). –
herpetofauna, Weinstadt, 20(116): 15-18.
– (2000): Metaphrynella sundana (PETERS, 1867) – ein
Frosch ruft im Amplexus. – Sauria, Berlin, 22(2): 37-40.
HONDA. M., M. KOBAYASHI, H.-S. YONG, H. OTA & T. HIKIDA
(1999): Taxonomic re-evulation of the status of Draco
cornutus GÜNTHER, 1864 (Reptilia: Agamidae). – Am-
phibia-Reptilia 20: 195-210.
HORST, R. (1883): On new and little known frogs from the
Malayan Archipelago. – Notes Leyden Mus., Leiden, 5:
235-244.
HUBRECHT, A.A.W. (1881): On a new genus and species of
Agamidae from Sumatra. – Notes Leyden Mus., Leiden,
3: 51-52.
I.C.Z.N. (International Commission on Zoological Nomen-
clature) (1958): Opinion 524. Interpretation of the nomi-
nal species Coluber ahaetulla LINNAEUS, 1758, and ad-
dition to the “Official list of generic names in zoology”
of the generic name Ahaetulla LINK, 1807 with Ahaetulla
mycterizans LINK, 1807 as type species (Class reptilia).
– Opin. Decl. Int. Comm. Zool. Nom., 19 (11): 261-276.
– (1985): International code of zoological nomenclature.
Third edition. – London, International Trust for zoologi-
cal Nomenclature, xx + 338pp.
INGER, R.F. (1954a): On a collection of amphibians from
Mt. Kinabalu, North Borneo. – J. Wash. Acad. Sci., 44:
250-251.
– (1954b): Systematics and zoogeography of Philippine
amphibia. – Fieldiana Zool., Chicago, 33(4): 181-531.
– (1956): Some amphibians from the lowlands of North
Borneo. – Fieldiana Zool., Chicago, 34(36): 389-424.
– (1958a): Three new skinks related to Sphenomorphus
variegatus (PETERS). – Fieldiana Zool., Chicago, 39(24):
257-268.
– (1958b): A new Gecko of the genus Cyrtodactylus with
a key to the species from Borneo and the Philippine Is-
lands. – Sarawak Mus. J., Kuching, 8: 261-264.
– (1958c): A new toad from Sarawak. – Sarawak Mus. J.,
Kuching, 8(11): 476-478.
– (1958d): Notes on the Bornean Glass Snake. – Sarawak
Mus. J., Kuching, 8(11): 479-481.
– (1960a): Notes on the toads of the genus Pelophryne. –
Fieldiana Zool., Chicago, 39(39): 415-418.
– (1960b): A review of the oriental toads of the genus
Ansonia STOLICZKA. – Fieldiana Zool., Chicago, 39(43):
473-503.
– (1960c): A review of the agamid lizards of the genus
Phoxophrys HUBRECHT. – Copeia, Washington, (3): 221-
225.
408
– (1962-1963): Royal society expedition to North Borneo
1961: Report 8, Amphibia. – Proc. Linn. Soc. London,
175: 45-46.
– (1964): Two new species of frogs from Borneo. –
Fieldiana Zool., Chicago, 44(20): 151-159.
– (1966): The systematics and zoogeography of the am-
phibia of Borneo. – Fieldiana Zool., Chicago, 52: 1-402.
– (1967): A new colubrid snake of the genus Stegonotus
from Borneo. – Fieldiana Zool., Chicago, 51: 77-83.
– (1978): The frogs and toads. Chapt. 13 in: Kinabalu, sum-
mit of Borneo. – Kota Kinabalu (Sabah Society), 311-
320.
– (1980): Relative abundance of frogs and lizards in for-
ests of Southeast Asia. – Biotropica, 12(1): 14-22.
– (1983a): Morphological and ecological variation in the
Flying Lizards (genus Draco). – Fieldiana Zool., Chi-
cago, neue Ser. 18: 1-35.
– (1983b): Larvae of Southeast Asian species of
Leptobrachium and Leptobrachella (Anura: Pelobatidae)
In: RHODIN, A. & MIYATA, K.(eds), Advances in herpetol-
ogy and evolutionary biology. – Cambridge (Museum
of Comparative Zoology), 13-32.
– (1985): Tadpoles of the forested regions in Borneo. –
Fieldiana Zool., Chicago, new Ser. 26: iv + 89 pp.
– (1986): Diets of tadpoles living in a Bornean rain forest.
– Alytes, Paris, 5(4): 153-164.
– (1989): Four new species of frogs from Borneo. – Malay.
Nat. J., Kuala Lumpur, 42: 229-243.
– (1992a): A bimodal feeding system in a stream-dwell-
ing larva of Rhacophorus from Borneo. – Copeia, Wash-
ington, (3): 887-890.
– (1992b): Variation of apomorphic characters in
streamdwelling tadpoles of the bufonid genus Ansonia
(Amphibia: Anura). – Zool. J. Lin. Soc., 105: 225-237.
– (1996): Commentary on a proposed classification of the
family Ranidae. – Herpetologica, Johnson City, 52(2):
241-246.
INGER, R.F. & J.P. BACON (1968): Annual reproduction and
clutch size in rain forest frogs from Sarawak. – Copeia,
Washington: 602-606.
INGER, R.F. & W.C. BROWN (1980): Species of the scincid
genus Dasia GRAY. – Fieldiana Zool., Chicago, new Ser.
3: 1-11.
*INGER, R.F. & P.K. CHIN (1962): The fresh-water fishes of
North Borneo – Fieldiana, Zool., 45.
INGER, R.F. & J. DRING (1988): Taxonomic and ecological
relations of Bornean stream toads allied to Ansonia
leptopus (GÜNTHER) (Anura: Bufonidae). – Malay. Nat.
J., Kuala Lumpur, 41: 461-471.
INGER, R.F. & K.J. FROGNER (1980): New species of Nar-
row-Mouth Frogs (genus Microhyla) from Borneo. –
Sarawak Mus. J., Kuching, 27(48): 311-322.
INGER, R.F. & B. GREENBERG (1966): Ecological and com-
petitive relations among three species of frogs (genus
Rana). – Ecology, 47(5): 746-759.

INGER, R.F. & P.A. GRITIS (1983): Variation in Bornean frogs
of the Amolops jerboa species group, with description
of two new species. – Fieldiana Zool., Chicago, new
Ser.19: 1-12.
INGER, R.F. & N.S. HAILE (1959): Two new frogs from
Sarawak. – Sarawak Mus. J., Kuching, 9:270-276.
INGER, R.F. & W. HOSMER (1965): New species of scincid
lizard of the genus Sphenomorphus from Sarawak. – Is-
rael J. Zool., 14: 134-140.
INGER, R.F. & W. KING (1961): A new cave-dwelling lizard
of the genus Cyrtodactylus from Niah. – Sarawak Mus.
J., Kuching, 11: 274-276.
INGER, R.F., M. LAKIM, A. BUIN & P. YAMBUN (1997): A new
species of Leptolalax (Anura: Megophryidae) from
Borneo. – Asiat. Herp. Research 7: 48-50.
INGER, R.F. & A.E. LEVITON (1966): The taxonomic status
of Bonean snakes of the genus Pseudorabdion JAN and
of the nominal genus Idiopholis MOCQUARD. – Proc. Ca-
lif. Acad. Sci., 34: 307-314.
INGER, R.F. & H. MARX (1965): The systematics and evolu-
tion of the Oriental colubrid snakes of the genus
Calamaria. – Fieldiana Zool., Chicago, 49: 1-304.
INGER, R.F. & R.B. STUEBING (1989): Frogs of Sabah. –
Sabah Parks Publication No. 10- Kota Kinabalu (Sabah
Parks Trustees), 129 pp.
– (1991): A new species of frog of the genus Leptobra-
chella SMITH (Anura: Pelobatidae), with a key to the spe-
cies from Borneo. – Raffl. Bull. Zool., Singapore, 39(1):
99-103.
– (1992): The montane amphibian fauna of northwestern
Borneo. – Malay. Nat. J., Kuala Lumpur, 46: 41-51.
– (1994): First Record of the lizard genus Pseudocalotes
(Lacertilia: Agamidae) in Borneo, with description of a
new species. – Raffl. Bull. Zool., Singapore, 42(4): 961-
965.
– (1996): Two new species of frogs from Southeastern
Sarawak. – Raffl. Bull. Zool., Singapore 44(2): 543-549.
– (1997): A field guide to the frogs of Borneo. – Kota
Kinabalu (Natural History Publications in assoc. with
Science and Technology Unit), 205 pp.
INGER, R.F., R.B. STUEBING & F.L. TAN (1995): New spe-
cies and new records of Anurans from Borneo. – Raffl.
Bull. Zool., Singapore 43(1): 115-131.
– (1996): Frogs and toads. – In: WONG, K.M. & A. PHILLIPPS
(Eds.): Kinabalu - Summit of Borneo. – Kota Kinabalu
(Sabah Soc.; Sabah Parks): 352-367.
INGER, R.F. & F.L. TAN (1990): Recently discovered and
newly assigned frog larvae (Ranidae and Rhacophoridae)
from Borneo. – Raffl. Bull. Zool., Singapore, 38(1): 3-9.
– (1996a): Checklist of the frogs of Borneo. – Raffl. Bull.
Zool., Singapore, 44(2): 551-574.
– (1996b): The natural history of amphibians and reptiles
in Sabah. – Kota Kinabalu (Natural History Publications
(Borneo) Sdn. Bhd.), v + 101 pp
INGER, R,F., F.L. TAN & P. YAMBUN (2000): The frog fauna
Bibliography
of three Parks in Sabah, Malaysia – Kinabalu Park,
Crocker Range Park and Tawau Hills Park. – Sabah Parks
Nat. Journ., 3: 7-28.
INGER, R.F., B. & A. TAUFIK (1996): New species of ranid
frogs (Amphibia : Anura) from Central Kalimantan,
Borneo. – Raffl. Bull. Zool., Singapore, 44(2): 363-369.
INGER, R.F. & H.K. VORIS (1988): Taxonomic status and
reproductive biology of bornean tadpole - carrying frogs.
– Copeia, Washington, (4): 1060-1062.
– (1993): A comparison of amphibian communities
through time and from place in Bornean forests. – J. Trop.
Ecol., 9: 409-433.
INGER, R.F., H.K. VORIS & K.J. FROGNER (1986): Orga-
nization of a community of tadpoles in rain forest streams
in Borneo. – J. Trop. Ecol., 2: 193-205.
INGER, R.F., H.K. VORIS & P. WALKER (1986): Larval tranport
in a Bornean ranid frog. – Copeia, Washington, (2): 523
-525.
INGER, R.F. & R.J. WASSERSUG (1990): A centrolenid-like
Anuran larva from Southeast Asia. – Zool. Scien. 7: 557-
561.
ISKANDAR, D.T. (1978): A new species of Barbourula: First
record of a discoglossid anuran in Borneo. – Copeia,
Washington, (4): 564-566.
JACOBSON, S.K. (1979): A guide to Mount Kinabalu Na-
tional Park. Sabah National Parks Publication No.3 –
Kota Kinabalu (Sabah National Park Trustees), 55 pp.
– (1986): Kinabalu Park. Sabah Parks Publication No.7 –
Kota Kinabalu (Sabah National Park Trustees), 65 pp.
*JENKINS, D.V. (1978): The first hundred years. In: LUPING,
D.M., C. WEN & E.R. DINGLEY (Eds.): Kinabalu - Sum-
mit of Borneo – Kota Kinabalu (Sabah Soc. Monogr.):
45-74.
JONG, J. K.DE (1930): Notes on some reptiles from Dutch -
East Indies. – Treubia, Bogor, 12(4): 115-119.
KABISCH, K., H.-J. HERRMANN, P. KLOSSEK, H. LUPPA & K.
BRAUER (1994): Die Schaumdrüse von Polypedates
leucomystax und die chemische Zusammensetzung des
von ihr produzierten Schaumes. – Beitr. z. Biol. d.
Anuren, tetra-Verl., Melle: 146:155.
KAMPEN , P.N. VAN (1907): Amphibien des indischen
Archipels. In: M. WEBER: Zoologische Ergebnisse einer
Reise in Niederländisch Ostindien. – Leiden (E. J. Brill),
383-418.
– (1910): Eine neue Nectophryne - Art und andere
Amphibien von Deli (Sumatra). – Nat. Tijdschr. Ned.
Ind., Batavia, 69: 18-24.
– (1923): The amphibia of the Indo-Australian Archi-
pelago. – Leiden (E. J. Brill), xii + 304 pp.
K AUP , F. (1825): Einige Bemerkungen zu Merrems
Handbuch. – Isis van Oken, 18: 589-592, pl. III.
– (1827): Zoologische Monographier. – Isis van Oken, 20:
508-566 + 610-625.
KEOGH, J.S., D.G. BARKER & R. SHINE (2001): Heavily ex-
ploited but poorly known: systematics and biogeogra-
409
Bibliography
phy of commercially harvested pythons (Python curtus
group) in Southeast Asia. – Biol. J. Linn. Soc., London,
73: 113-129.
KIEW, B.H. (1984): A new species of frog (Kalophrynus
baluensis n. sp.) from Mt. Kinabalu, Sabah, Malaysia. –
Malay. Nat. J., 38: 151-156.
– (1987): An annotated checklist of the herpetofauna of Ulu
Endau, Johore, Malaysia. – Malayan Nat. J., 41: 413-423.
KIRSCHNER, A. & M. WICHMANN (1998): Bemerkungen zur
Fortpflanzung und Pflege des Buntpythons (Python
curtus) insbesondere von P. c. breitensteini STEINDACHNER,
1880 und P. c. brongersmai STULL, 1938. – Herpetofauna,
Weinstadt, 20 (114): 5-12.
*KITAYAMA, K. (1991): Vegetation of Mount Kinabalu Park,
Sabah, Malaysia, Map of physiognomically classified
vegetation – Honolulu (East-West Center)
– *(1992): An altitudinal transect study of the vegetation
on Mount Kinabalu, Borneo – Vegetatio, 102: 149-171.
KOPSTEIN, F. (1938): Ein Beitrag zur Eierkunde und zur
Fortplanzung der malaiischen Reptilien. – Bull. Raffles
Mus., Singapore, 14: 81-167, pl. 27.
– (1941): Über Sexualdimorphismus bei malaiischen
Schlangen. – Temminckia, Leiden, 6: 109-185.
KOSUCH, J. (1994): Genetische Untersuchungen zur
systematischen Stellung indonesischer Vertreter der
Gattung Rana (Amphibia, Anura, Ranidae). – Unpub-
lished diploma thesis, Mainz University.
KUCH, U. & A. GÖTZKE (2000): Eine Freilandbeobachtung
des Kinabalu-Kraits, Bungarus flaviceps baluensis
LOVERIDGE, 1938 (Serpentes: Elapidae). – Sauria, Berlin
22(2): 19-22.
KUHL, H. (1820): Beiträge zur Kenntnis der Amphibien In:
KUHL, H.: Beiträge zur Zoologie und vergleichenden
Anatomie. – Frankfurt a.M. (Hermannsche Buchhand-
lung), 75-132.
*KURATA, S. (1976): Nepenthes of Mount Kinabalu – Kota
Kinabalu (Sabah Parks Publ. 2).
*KÜRSCHNER, H. (1990: Die epiphytischen Moosgesell-
schaften am Mount Kinabalu (Nord-Borneo, Sabah,
Malaysia) – Nova Hedwigia, 51 (1/2): 1-75.
LAKIM, M.B., YAMBUN, P. & S. AIRAME (1999): A compara-
tive study of the amphibian population in reference to
habitat disturbance across an elevational gradient in
Kinabalu Park, Sabah, Malaysia. – Sabah Parks Nat.
Journ., 2: 27-44.
LAURENTI, J.N. (1768): Specimen medicum, exhibens
synopsin reptilium. – Vienna, 214 pp.
LAZELL, J.D., KEIRANS, J.E. & G.A. SAMUELSON (1991): The
Sulawesi Black Racer, Coluber (Ptyas) dipsas, and a
remarkable ectoparasitic aggregation. – Pacific Sci., 45:
355-361.
LESSON, R.P. (1830): Observations générales sur les rep-
tiles recueillis dans le voyage. In: DUPERREY, L. I.: Voy-
age autor du monde, exécuté par ordre du Roi, sur la
corvette de sa Majesté, la Coquille, pendant les annes
410
1822, 1824 et 1825. – Zoologie 2(1): 1-65.
LEVITON, A.E., (1955): Systematic notes on the Asian snake
Lycodon subcinctus. – Phil. J. Sci., Manila, 84: 195-203.
– (1963): Contribution to a review of Philippine snakes.
III. - The genera Maticora and. – Phil. J. Sci., Manila,
92: 523-550.
– (1964): Contributions to a review of Philippine snakes,
II. - The genera Liopeltis and Sibynophis. Phil. J. Sci.,
Manila, 92 [1963] (3): 367-381.
– (1968): Contribution to a review of Philippine snakes.
XII. - The snakes of the genus Dendrelaphis (Serpentes:
Colubridae). – Phil. J. Sci., Manila, 97: 371-396.
LEVITON A. E. & W. C. BROWN (1959): A review of the snakes
of the genus Pseudorabdion with remarks on the status
of the genera Agrophis and Typhlogeophis (Serpentes:
Colubridae). – Proc. Calif. Acad. Sci., 19 (14): 475-508.
LIDTH DE JEUDE, T.W. VAN (1893): On reptiles from North-
Borneo. – Notes Leyden Mus.,Leiden, 15(3): 250-257.
– (1905): Zoological results of the Dutch Scientific Expe-
dition to Central-Borneo. The reptiles. – Notes Leyden
Mus., Leiden, 25(4): 187-202.
LIEM, D.S.S. (1970): The morphology, systematics and evo-
lution of the Old World treefrogs (Rhacophoridae and
Hyperoliidae). – Fieldiana Zool., Chicago, 57: 1-145.
LIEM, K.H. (1959): The breeding habits and development
of Rana (SCHLEG.) (Amphibia). – Treubia, Bogor, 25(1):
89-111.
LIM, B.L. (1970): A genus of snakes (Oreocalamus) new to
Malaya. – Sarawak Mus. J., Kuching, 18(36/37): 410-411.
– (1990): Venomous land snakes of Malaysia. In: P.
GOPALAKRISHNAKONE & L. M. CHOU (eds.): Snakes of
medical importance (Asia-Pacific region). – Singapore
(National University of Singapore): 387-417.
LIM, B.L. & I. DAS (1999): Turtles of Borneo and Peninsu-
lar Malaysia. – Kota Kinabula (Natural History
Paublications), 151 pp.
LIM, F.L.K. & M.T.M. LEE (1989): Fascinating snakes of
Southeast Asia - An introduction. – Kuala Lumpur (Tropi-
cal Press), 124 pp.
LIM, K.K.P. & P.K.L. NG (1991): Nepenthiphilous larvae
and breeding habits of the sticky frog, Kalophrynus pleu-
rostigma TSCHUDI (Amphibia: Microhylidae). – Raffl.
Bull. Zool., Singapore, 39 No 1: 209-214.
LINK, H.F. (1807): Beschreibung der Naturalien-Sammlung
der Universität zu Rostock. Zweite Abtheilung. – Rostock
(Adlers Erben): 51-100
LINNAEUS, C. (1758): Systema naturae per regna tria natu-
rale, secundum classes, ordines, genera, species, cum
characteribus, differentiis, synonymis, locis. – Tomus I.,
edit. 10: 824 pp.
LLOYD, M., R.F. INGER & F.W. KING (1968): On the diver-
sity of reptile and amphibian species in a Bornean rain
forest. – Amer. Naturalist, 202: 497-515.
LOVERIDGE, A. (1938): New snakes of the genera Calamaria,
Bungarus and Trimeresurus from Mt. Kinabalu, North

Borneo. – Proc. Biol. Soc. Washington, 51: 43-46.
*LOW, H. (1852): Notes of an ascent of the mountain Kina
Balow – J. Ind. Arch., VI: 1-17.
*LOWRY, J.B., D.W. LEE & B.C. STONE (1973): Effect of
drought on Mount Kinabalu – Malay. Nat. J., 26: 178-
179.
*LUPING, D.M., C. WEN & E.R. DINGLEY (1978): Kinabalu
- Summit of Borneo – Kota Kinabalu (Sabah Soc.
Monogr.).
*MACKINNON, K., G. HATTA, H. HALIM & A. MANGALIK
(1997): The ecology of Kalimantan – London (Oxford
Univ. Press).
MALKMUS, R. (1985): Amphibien und Reptilien vom Mount
Kinabalu (4101m), Nordborneo. – herpetofauna,
Weinstadt, 7(35): 6-13.
– (1986): Ein seltener Ruderfrosch vom Mt.Kinabalu Nord-
Borneo - Rhacophorus everetti (BOULENGER). – Sauria,
Berlin, 8(1): 3-4.
– (1987): Herpetologische Beobachtungen am Mt.Kina-
balu Borneo. – Mitt. Zool. Mus. Berlin, 63(2): 269-292.
– (1988a): Cyrtodactylus baluensis (MOQUARD,1890). –
Sauria, Berlin, 10(1): 29.
– (1988b): Bufo juxtasper als Bewohner der Kulturland-
schaft Tambunans (Nord-Borneo). – Sauria, Berlin, 10(4):
9-11.
– (1988c): Wanderungen im Mount Kinabalu - National-
park/ Nordborneo. – Nat. Mus., Frankfurt/M, 118(6):
161-181.
– (1988d): Herpetologische Studien an einem Waldbach
am Mt.Kinabalu, Borneo. – herpetofauna, Weinstadt,
10(53): 6-11.
– (1989): Herpetologische Beobachtungen am Mount
Kinabalu, Borneo II. – Mitt. Zool. Mus. Berlin, 65(2):
179-200.
– (1991a): Zur Herpetofauna des oberen Liwago, Mt.
Kinabalu/ Nordborneo. – herpetofauna, Weinstadt,
13(72): 26-34.
– (1991b): Sphenomorphus sp.n. (Sauria: Scincidae) vom
Mt. Kinabalu/ Nord- Borneo. – Sauria, Berlin 13(3): 23-
28.
– (1991c): Die Bogenfingergeckos der Gattung Cyrtodac-
tylus am Mt. Kinabalu/ Nord Borneo. – herpetofauna,
Weinstadt, 13(74): 6-10.
– (1992a): Herpetologische Beobachtungen am Mt.
Kinabalu, Nord-Borneo. III . – Mitt. Zool. Mus. Berlin,
68(1): 101-138.
– (1992b): Leptolalax pictus sp.n. (Anura: Pelobatidae)
vom Mount Kinabalu/Nord-Borneo. – Sauria, Berlin
14(3): 3-6.
– (1993): Zur Zeichnungsvariabilität und zum Totstell-
verhalten des Ruderfrosches Rhacophorus angulirostris
MOCQUARD, 1890. – Sauria, Berlin 15(4): 35-38.
– (1994a): Einige Bemerkungen zu Kalophrynus baluensis
KIEW, 1984. – Sauria, Berlin 16(2): 9-13.
– (1994b): Herpetologische Beobachtungen am Mt.
Bibliography
Kinabalu, Nord-Borneo. IV . – Mitt. Zool. Mus. Berlin,
70: 217-249.
– (1994c): Vijf kikkers bij de Mesilau-grot op Mt. Kinabalu
(noordelijk Borneo). – Lacerta 52(4): 86-90.
– (1994d): Een zeldzame slang van Mt. Kinabalu: Schmidts
Grondslang (Calamaria schmidti). – Lacerta 52(5): 111-
113.
– (1995a): Over het voortplantingsbiotoop van Philautus
op Mount Kinabalu (noordelijk Borneo). – Lacerta 53(3):
68-74.
– (1995b): Wer rief hier: Frosch oder Heuschrecke? –
Sauria, Berlin, 17(2): 35-38.
– (1995c): Die Krötenfrösche der Gattung Megophrys
KUHL & VAN HASSELT, 1822 am Mount Kinabalu, Nord
Borneo. – herpetofauna, Weinstadt, 17(94): 6-10.
– (1995d): Kopfseitenflecken bei beiden Geschlechtern von
Rhacophorus angulirostris AHL, 1927 (Amphibia:
Rhacophoridae). – Salamandra, Rheinbach, 31(4): 245-
246.
– (1995e): Borneose Bergagamen (Phoxophrys) van Mount
Kinabalu. – Lacerta, Leiden, 54(2): 65-69.
– (1996a): Voortplanting van de Dwergpad Pelophryne
misera. – Lacerta, Leiden, 54(4): 129-133.
– (1996b): Bemerkungen zur Rufaktivität einiger
Froscharten am Mount Kinabalu, Nord-Borneo. –
herpetofauna, 18(101): 20-26.
– (1996c): Herpetologische Beobachtungen am Mount
Kinabalu, Nord-Borneo. V. – Mitt. Zool. Mus. Berl.
72(2): 277-295.
– (1996d): Leptobrachium gunungensis sp. n. (Anura:
Pelobatidae) vom Mount Kinabalu, Nord-Borneo. – Mitt.
Zool. Mus. Berl. 72(2): 297-301.
– (1996e): Untersuchungen zum Aktivitätsrhythmus von
Staurois BOULENGER, 1918 am Mt. Kinabalu. – Sauria,
Berlin 18(4): 1-16.
– (1998): Zur Fortpflanzungsbiologie der Kaskadenfrösche
der Gattung Meristogenys YANG, 1991. – herpetofauna,
Weinstadt, 20(115): 21-26.
– (1999): Die Kaulquappengemeinschaft eines Bergbaches
in Borneo. – herpetofauna, Weinstadt, 21(120): 13-19.
– (2000): Die Schlankfrösche der Gattung Leptolalax
DUBOIS, 1980 am Mount Kinabalu, Nord-Borneo. –
herpetofauna, Weinstadt, 22(124): 11-15.
MALKMUS, R, C. BRÜHL & T. ELTZ (1999): Amfibiën en
reptilen van Deramakot (Sabah, Maleisië). - Lacerta,
Leiden, 57(6): 191-198.
MALKMUS, R & J. KOSUCH (1999): Beschrijving van het
kikkervisje van Leptolalax arayai MATSUI, 1997, een
kikker uit de familie Megophryidae. – Lacerta, Leiden,
57(5): 161-165.
– (2000): Beschreibung einer neuen Ansonia-Larve
(Ansonia guibei) von Borneo. – Salamandra, Rheinbach,
36(2): 121-124.
MALKMUS, R, J. KOSUCH & J. KREUZ (1999): Die Larve von
Staurois tuberilinguis BOULENGER, 1918. Eine neue
411
Bibliography
centrolenidenähnliche Kaulquappe aus Borneo (Anura:
Ranidae). – Herpetozoa 12 (1/2): 17-22.
MALKMUS, R. & M. MATSUI (1997): Megophrys kobaya-
shii, ein neuer pelobatider Frosch vom Mount Kinabalu.
– Sauria, Berlin 19(1): 31-37.
MALKMUS, R. & K. RIEDE (1993): Nachtrag zu Leptolalax
pictus MALKMUS, 1992. (Anura: Pelobatidae). – Sauria,
Berlin 15(3): 7-9.
– (1996a): Die Baumfrösche der Gattung Philautus vom
Mount Kinabalu – Teil I: Überblick und die aurifasciatus-
Gruppe mit Beschreibung einer neuen Art (Philautus
saueri n. sp.). – Sauria, Berlin, 18(1): 27-37
– (1996b): Die Baumfrösche der Gattung Philautus vom
Mount Kinabalu – Teil II: Die vermiculatus-Gruppe mit
Beschreibung einer neuen Unterartrt (Philautus
aurantium gunungensis n. ssp.) und die hosei-Gruppe. –
Sauria, Berlin, 18(2): 21-28.
– (1996c): Zur Bioakustik von Megophrys (BOULENGER,
1899) und Kalophrynus KIEW, 1984, zweier seltener
Froscharten aus Borneo. – herpetozoa, Wien, 9(1/2): 151-
155.
MALKMUS, R. & H. SAUER (1993): Die Folgen eines Bisses
von Maticora intestinalis everetti. – Salamandra, 29(2):
153-154.
– (1996): Ruhestellung von Pareas nuchalis und Erst-
nachweis dieser Art im Nationalpark Mount Kinabalu/
Malaysia. – Salamandra, 32(1): 55-58.
MALNATE, E.V. (1960): Systematic division and evolution
of the colubrid snake genus Natrix, with comments on
the subfamily Natricinae. – Proc. Acad. Nat. Sci. Phila-
delphia, 112(3): 41-71.
MANTHEY, U. (1982): Die Gattung Ptychozoon KUHL, 1822
Teil 2. – Sauria, Berlin, 4(3): 5-12.
– (1983): Exkursion am Mt. Kinabalu 4100 m, Nord -
Borneo. Teil 3: Checkliste der Herpetofauna oberhalb
600m üNN. – herpetofauna, Weinstadt, 5(23): 20-31.
MANTHEY, U. & W. DENZER (1982): Exkursion am Mt.
Kinabalu 4100 m, Nord - Borneo. Teil 2: Herpetologi-
sche Eindrücke. – herpetofauna, Weinstadt, 4(21): 11-
19.
– (1991): Die Echten Winkelkopfagamen der Gattung
Gonocephalus KAUP (Sauria: Agamidae) III. Gonoce-
phalus grandis (GRAY, 1845). – Sauria, Berlin 13(3): 3-
10, 5 Abb.
– (1992): Die Echten Winkelkopfagamen der Gattung
Gonocephalus (Sauria: Agamidae) V. Die bellii-Gruppe.
– Sauria, Berlin 14(3): 7-20.
– (2000): Description of a new genus, Hypsicalotes gen.
nov. (Sauria: Agamidae) from Mount Kinabalu, North
Borneo, with remarks on the generic identity of
Gonocephalus schultzewestrumi URBAN, 1999. – Ha-
madryad, Madras 25(1): 13-20.
MANTHEY, U. & W. GROSSMANN (1994): Gleitfliegen bei
Bronchocela cristatella (KUHL, 1820) (Sauria: Agami-
dae). – Sauria, Berlin 16(3): 37-38.
412
– (1997): Amphibien & Reptilien Südostasiens. – Münster
(NTV), 512 pp.
MANTHEY, U. & C. STEIOF (1998): Rhacophorus cyano-
punctatus sp. n. (Anura: Rhacophoridae), ein neuer
Flugfrosch von der Malaiischen Halbinsel, Sumatra und
Borneo. – Sauria, Berlin, 20(3): 37-42.
MARMAYOU, J., A. DUBOIS, A. OHLER, E. PASQUET & A. TILLIER
(2000): Phylogenetic relationships in the Ranidae. Inde-
pendent origin of direct development in the genera Philautus
and Taylorana. – C.r.Acad. Sci. Vie., 323: 287-297.
MARX, H. & R.F. INGER (1955): Notes on the snakes of the
genus Calamaria. – Fieldiana Zool., Chicago, 37: 167-
209.
MATSUI, M. (1979): Amphibians of Sabah. 1. Systematic
and natural history notes. – Contr. biol. Lab. Kyoto Univ.,
25(4): 303-346.
– (1982a): Amphibians of Sabah. 1I. Acoustic characte-
ristics of three common Anuran species. – Contr. biol.
Lab. Kyoto Univ., (26): 123-129.
– (1982b): Call Characteristics of several Anuran species
from East Kalimantan. – Contr. biol. Lab. Kyoto Univ.,
(26): 131-139.
– (1995): Call produced by a “voiceless” frog, Rana blythi
BOULENGER, 1920, from Peninsular Malaysia (Amphibia,
Anura). – Tropical Zoology, 8: 325-331.
– (1986): Three new species of Amolops from Borneo.
(Amphibia, Anura, Ranidae). – Copeia, Washington, (3):
623-630.
– (1996): Call characteristics and systematic relationships
of a Malayan treefrog Nyctixalus pictus (Anura:
Rhacophoridae). – Herp. J., 6: 62-64.
– (1997): Call characteristics of Malaysian Leptolalax with
a description of two new species (Anura: Pelobatidae). –
Copeia 1997(1): 158-165.
MATSUI, M., T. CHANARD & J. NABHITABHATA (1996): Dis-
tinct specific status of Kalophryne pleurostigma
interlineatus (Anura: Microhylidae). – Copeia, Wash-
ington, (2): 440-445.
MATSUI, M., T. HIKIDA & H. NAMBU (1985): The amphib-
ians and reptiles collected from Borneo, Malaysia. – Bull.
Toyama Sci.Mus. no.8: 151-159.
MATSUI, M., T. SETO & T. UTSUNOMIYA (1986): Acoustic
and karyotypic evidence for specific separation of
Polypedates megacephalus from P. leucomystax. – J.
Herp., 20(4): 483-489.
MATSUI, M., G.-F. WU & H.-S. YONG (1993): Acoustic char-
acteristics of three species of the genus Amolops (Am-
phibia, Anura, Ranidae). – Zool. Soc. Kapan, 10: 691-
695.
MATSUI, M., H.-S. YONG, K. ARAYA & A.A. HAMID (1996):
Acoustic characteristics and systematic relationships of
aboreal microhylid frogs of the genus Metaphrynella
from Malaysia. – J. Herp., 30(3): 424-427.
MCGUIRE, J.A. & B.H. KIEW (2001): Phylogenetic system-
atics of Southeast Asian flying lizards (Iguania:

Agamidae: Draco) as inferred from mitochondrial DNA
sequence data. – Biol. J. Linnean Society, London, 72:
203–229.
MEER MOHR, J. C. VAN DER (1926): Aanteekeningen over
Python reticulatus. – Trop. Natuur, 15: 9-14.
MEISE, W. & W. HENNIG (1932): Die Schlangengattung
Dendrophis. – Zool. Anz., Leipzig, 93(11/12): 273-297.
– (1935): Zur Kenntnis von Dendrophis und Chrysopelea
. – Zool. Anz., Leipzig, 109: 138-150.
MERTENS, R. (1927): Neue Amphibien und Reptilien aus
dem Indo-Australischen Archipel. – Senckenbergiana,
Frankfurt/M, 9(6): 234-242.
– (1934): Die Schlangengattung Dendrelaphis BOULENGER
in systematischer und zoogeographischer Beziehung. –
Arch. Nat., Leipzig, 3: 187-204.
– (1968): Die Arten und Unterarten der Schmuckbaum-
schlangen (Chrysopelea). – Senck. biol., Frankfurt/M,
49(3/4): 191-217.
– (1969): Über die Variabilität der Achtstreifennatter,
Oligodon octolineatus. – Senck. biol., Frankfurt/M, 50(5/
6): 339-345.
MITTLEMAN, M.B. (1952): A generic synopsis of the lizards
of the subfamily Lygosominae. – Smithson. Misc. Coll.,
117(17): 1-35.
*MJÖBERG, E. (1928): I Tropikernas Villarde Urskogar –
Stockholm.
– *(1930): Forest life and adventures in the Malay Archi-
pelago – London (George Allen & Unwin Ltd.)
MOCQUARD, F. (1890a): Recherches sur la faune herpéto-
logique des îles de Bornéo et de Palawan. – Nouv. Arch.
Mus. Hist. Nat., Paris, 3(2): 115-168.
– (1890b): Diagnoses d’espèces nouvelles de reptiles et
de batraciens des îles Bornéo et Palawan. – Le Naturaliste,
Paris (Sér. 2), 12: 144-145.
– (1892): Nouvelle contribution à la faune herpétologique
de Bornéo. – Mem. Soc. Zool. France, 5: 190-206.
MORGAN, E.C. (1973): Snakes of the subfamily Sibynophii-
nae. Lafayette, Louisiana, University of Southwestern
Louisiana, Ph.D. dissertation: 1-260.
MORI, A. (1993): A note on the sidewinding locomotion in
two colubrid snakes, Ophisthotropis typica and Pseu-
doxenodon macrops. – The Snake, 25(1): 67-70.
*MOTLEY, J. & L. L. DILLWYN (1855): Contributions to the
natural history of Labuan and the adjacent coasts of
Borneo - Reptilia. – London (John VAN Voorst), 39-49.
*MOULTON, J.C. (1915): An account of the various expedi-
tions to Mt. Kinabalu. – Sarawak Mus. J., 6: 137-176
MÜLLER, F. (1890): Fünfter Nachtrag zum Katalog der
herpetologischen Sammlung des Basler Museums. –
Verh. naturf. Ges. Basel, 8(2): 249-296, pl. 1-3.
MÜLLER, S. (1836): Bombinator baleatus. – Verh. Batavia
Genootsch., 16: 96-97.
MUSTERS, C.J.M. (1983): Taxonomy of the genus Draco L.
(Agamidae, Lacertilia, Reptilia). – Zool. Verh., Leiden,
199: 1-120.
Bibliography
NARINS, P.M., A.S. FENG, H.-D. YONG & J. CHRISTENSEN-
DALSGAARD (1998): Morphological, behavioral, and ge-
netic divergence of sympatric morphotypes of the tree
frog Polypedates leucomystax in Peninsular Malaysia. –
Herpetologica, 54(2): 129-142.
NOBLE, G.K (1926): An analysis of the remarkable cases of
distribution among the amphibia, with descriptions of new
genera. – Am. Mus. Nov., New York, (212): 1-24.
OHLER, A. & A. DUBOIS (1999): The identy of Elachyglossa
gyldenstolpei ANDERSON, 1916 (Amphibia, Ranidae), with
comments on some aspects of statistical support to tax-
onomy. – Zoologica Scipta, 28: 269-279.
OTA, H. & T. HIKIDA (1988): A new species of Lepidodac-
tylus (Sauria Gekkonidae) from Sabah, Malysia. –
Copeia, Washington, (3): 616-621.
– (1991): Taxonomic review of the lizards of the genus
Calotes CUVIER 1817 (Agamidae Squamata) from Sabah,
Malaysia. – Tropical Zool., 4: 179-192.
– (1996): The second specimen of Calotes kinabaluensis
de GRIJS (Squamata: Agamidae) from Sabah, Malaysia,
with comments on the taxonomic status of the species. –
J. Herp., 30(2): 288-291.
– (2000) Aphaniotis nasuta (DE JONG, 1930), a junior syn-
onym of A. ornata (van LIDTH DE JEUDE, 1893) (Squamata:
Agamidae). Curr. Herpetol. 19(1): 11-14.
OTA, H., T. HIKIDA, M. KON & T. HIDAKI (1989): Unusual
nest site of a scincid lizard Sphenomorphus kinabalu-
ensis from Sabah, Malaysia. – Herp. Rev. 20(2): 38-39.
OTA H., S. SENGOKU & T. HIKIDA (1996): Two New Species
of Luperosaurus (Reptilia: Gekkonidae) from Borneo. –
Copeia, Washington, (2): 433-439.
PARKER, H. W. (1928): The brevicipitid frogs of the genus
Microhyla. – Ann. Mag. Nat. Hist., London, 10. Ser. 2:
473-499.
– (1934): A monograph of the frogs of the family
Microhylidae. – London (Brit. Museum Trustees), viii +
208 pp.
*PARRIS, B.S., R.S. BEAMAN & J.M. BEAMAN (1992): The
plants of Mount Kinabalu. 1. Ferns and Fern Allies –
Kew (Royal Botanic Gardens).
*PAYNE, J., C.M. FRANCIS & K. PHILLIPPS (1985): A field
guide to the mammals of Borneo – Kuala Lumpur (Sabah
Soc. & World Wildlife Fund Malaysia).
*PENDLEBURY, H.M. & B.E. CHASEN (1932): A zoological
expedition to Mt. Kinabalu. – J. Fed. Malay. St. Mus.,
Singapore, 17: 1-38.
PETERS, U. (1981): Drei bemerkenswerte Nachzuchten:
Netzpython, Nashorn-Leguan und Gopher-Schildkröte.
– Das Aquarium, 15 (139): 33-37.
PETERS, W. (1864): Über neue Amphibien (Typhloscincus,
Typhlops, Asthenodipsas, Ogmodon ). - Mber. k. preuss.
Akad. Wiss., Berlin: 271-276, 1 Pl.
– (1867): Herpetologische Notizen. – Mber. Königl. Akad.
Wiss. Berlin: 13-37.
– (1871): Über neue Reptilien aus Ostafrica und Sarawak
413
Bibliography
(Borneo), vorzüglich aus der Sammlung des Hrn. Mar-
quis J. Doria zu Genua. – Mber. Königl. Akad. Wiss.
Berlin,: 566-581.
– (1872): Übersicht der von den Herren. Mse. G. Doria
und Dr. O. Beccari in Sarawak auf Borneo von 1865 bis
1868 gesammelten Amphibien. – Ann. Mus. civ. stor.
nat. Genova, 3: 27-45, pls. 2-6.
*PHILLIPS, A. & A. LAMB (1988): Pitcher-plants of East Ma-
laysia and Brunei. – Nat. Malaysiana, Kuala Lumpur,
13(4): 8-27.
PINOU, T. & H.G. DOWLING (2000): “Coluber” versus
“Ptyas”: A regrettable tangle of names. – Herp. Rev., 31
(3): 136-138.
POPE, C.H. & S.H. POPE (1933): A study of the green pit-
vipers of southeastern Asia and Malaysia, commonly
identified as Trimeresurus gramineus (Shaw), with de-
scription of a new species from Peninsular India. – Amer.
Mus. Novitates, 620: 1-12.
RAO, D.Q. & D.T. YANG (1992): Phylogenetic systematics
of Pareinae (Serpents) of southeastern Asia and adjacent
islands with relationship between it and the geology
changes. [In Chinese, English summary]. – Acta zool.
Sinica, 38(2): 139-150.
RASHID, S.M.A. & C.H. DIONG (1999): Observartions on
Varanus salvator feeding on Oligodon octolineatus. –
Hamadryad, Madras, 24(1): 48-49.
RASMUSSEN, J.B., (1975): Geographical variation, includ-
ing an evolutionary trend, in Psammodynastes pulveru-
lentus (BOIE, 1827) (Boiginae, Homalopsidae, Serpentes).
– Vidensk. Meddr. dansk. naturh. Foren., 138: 39-64.
REGENASS, U. & E. KRAMER (1981): Zur Systematik der
grünen Grubenottern der Gattung Trimeresurus
(Serpentes, Crotalidae). – Rev. Suisse Zool., Genève,
88(1): 163-205.
RIDLEY, H.N. (1906): Curious nesting place of Simotes
octolineatus. – J. Str. Br. Roy. As. Soc.,Singapore, 46: 263.
RIEDE, K. (1993): Bioakustik der Rufgemeinschaften eines
malaysischen Tieflandregenwaldes. – Verh. Deutsch.
Zool. Ges., Stuttgart, 86(1): 269.
RIQUIER, M.A. (1998): Status, population biology and con-
servation of the Water Monitor (Varanus salvator), the
Reticulated Python (Python reticulatus), and the Blood
Python (Python curtus) in Sumatra and Kalimantan, In-
donesia – Project Report Kalimantan – In: ERDELEN, W.
(Ed.): Conservation, trade and sustainible use of lizards
and snakes in Indonesia. – Mertensiella, Rheinbach 7:
119-129.
ROOIJ, N. DE (1915): The reptiles of the Indo-Australian
Archipelago. I. Lacertilia, Chelonia, Emydosauria. –
Leiden (E.J.Brill), xiv + 384 pp.
– (1917): The Reptiles of the Indo-Australian Archipelago.
II. Ophidia. – Leiden (E.J.Brill), xiv + 334 pp.
ROUX, J. (1906): Synopsis of the toads of the genus
Nectophryne B. & P., with special remarks on some
known species and description of a new species from
414
German East Afrika. – Proc. Zool. Soc. London: 58-65.
ROZKOŠNÝ, R. & D. KOVAC (1994): A new species of
Odontomyia meigen (Iinsecta: Diptera: Startiomyidae)
from Sabah, Borneo. – Raffl. Bull. Zool., Singapore,
42(4): 859-867.
SANCHEZ-HERRAIZ, M.J., MARQUEZ, R., BARBADILLO, L.J. &
J. BOSCH (1995): Mating calls of three species of anurans
from Borneo. – Herp. J. 5(4): 293-297.
SCHLEGEL, H. (1826): Notice sur l’erpétologie de l’île de
Java; par M. Boié. – Bull. sci. nat. géol., Paris, 9 (2):
233-240.
– (1827): Erpetologische Nachrichten. – Isis van Oken:
281-294.
– (1837): Essai sur la physionomie des serpents. – La Haye
(J. Kips, J. HZ. et W. P. van Stockum), Partie Générale xxviii
+ 251 pp. + Partie Descripive 606 pp. + xvi.
– (1837-1844): Abbildungen neuer oder unvollständig
bekannter Amphibien, nach der Natur oder dem Leben
entworfen. – Düsseldorf (Arnz & Comp.), xiv + 141 pp.
– (1848): Descriptions de plusieurs espèces nouvelles du
genre Lophyrus. – Bijdr. Dierk., Amsterdam, 1: 4-6.
– (1858): Handleiding tot de beoefening der dierkunde. –
Breda, Koninklijke Militaire Academie, xx + 630 pp.
SCHMIDT, A. A. (1976): Erst-Nachzucht des Zipfelfrosches
Megophrys nasuta. – Salamandra, Frankfurt/M, 12(2):
55-68.
SCHNEIDER, J.G. (1801): Historiae Amphibiorum naturalis
et literariae. Fasciculus secundus continens Crocodilos,
Scincos, Chamaesauras, Boas, Pseudoboas, Elapes,
Angues, Amphisbaenas et Caecilias. – Jena, 364 pp.
SCHULZ, K.-D. (1996): A monograph to the colubrid snakes
of the genus Elaphe FITZINGER. – Havlickuv Brod (Koeltz
Scientific Books), i-iii + 1-439, 58 pls.
SEBA, A. (1734): Locupletissimi rerum naturalium thesauri
accurata descriptio, et iconibus artificiosissimis expressio,
per universam physices historiam. Opus, cui, in hoc re-
rum genere, nullum par existit. Ex toto terrarum orbe
collegit, digessit, descripsit, et depingendum curavit.
Tomus I. Amstelaedami [= Amsterdam], Janssonio-
Waesbergios, J. Wetstenium & Gul. Smith: (32), 1-178,
pl. 1-111.
– (1735): Locupletissimi rerum naturalium thesauri
accurata descriptio, et iconibus artificiosissimis expressio,
per universam physices historiam. Opus, cui, in hoc re-
rum genere, nullum par existit. Ex toto terrarum orbe
collegit, digessit, descripsit, et depingendum curavit.
Tomus II. Amstelaedami [= Amsterdam], Janssonio-
Waesbergios, J. Wetstenium & Gul. Smith: (32), 1-154,
pl. 1-114.
SHELFORD, R. (1901a): A list of the reptiles of Borneo. – J.
Str. Br. Roy. As. Soc., Singapore, 35: 43-68.
– (1901b): On two new snakes from Borneo. – Ann. Mag.
Nat. Hist., London, 7. Ser. 8: 516-517.
– (1905): A new lizard and a new frog from Borneo. –
Ann. Mag. Nat. Hist., London, 7. Ser. 15: 208-210.

– *(1916): A naturalist in Borneo – London (T. Fisher
Unwin Ltd.).
S HIN, C. K. (1994): Observations in the copulatory
behaviour of Gonyosoma oxycephalum (BOIE). – Ma-
layan Naturalist, 48 (2/3):4-5.
SLOWINSKI, J.B., J. BOUNDY & R. LAWSON (2001): The phy-
logenetic relationships of Asian Coral Snakes (Elapidae:
Calliophis and Maticora) based on morphological and
molecular characters. – Herpetologica 57(2): 233-345.
SMEDLEY, N. (1931): Notes on the Giant Frog, Rana
macrodon. – Bull. Raffles Mus., Singapore, 5: 59-62.
*SMITH, J.M.B. (1980): The vegetation of the summit zone
of Mount Kinabalu – New Phytol., 84: 547-573.
SMITH, L.A. & I. SIDIK (1998): Description of a new species
of Cylindrophis (Serpentes: Cylindrophiidae) from
Yamdena Island, Tanimbar Archipelago, Indonesia. –
Raffles Bull. Zool., Singapore, 46(2): 419-424.
SMITH, M.A. (1923): A review of the lizards of the genus
Tropidophorus on the Asiatic Mainland. – Proc. Zool.
Soc. London: 775-781.
– (1924): New tree-frogs from Indo-China and the Malay
Peninsula. – Proc. Zool. Soc. London: 225-233.
– (1925a): On a collection of reptiles and amphibians from
Mt. Murud, Borneo. – Sarawak Mus. J., Kuching, 8: 5-
14.
– (1925b): Contributions to the herpetology of Borneo. –
Sarawak Mus. J., Kuching, 8: 15-34.
– (1927): Contributions to the herpetology of the Indo-
Australian region. – Proc. Zool. Soc. London: 199-225.
– (1931): The herpetology of Mt. Kinabalu, North Borneo,
13455ft. – Bull. Raffles Mus., Singapore, 5: 3-32.
– (1937): The names of two Indian vipers. – J. Bombay
nat. Hist. Soc., 39 (4): 730-731.
– (1943): The fauna of British India, Ceylon and Burma,
including the whole of the Indo-Chinese Subregion. Rep-
tilia and Amphibia. Vol. III, Serpentes. – London (Tay-
lor & Francis) xii + 583 pp.
*SMYTHIES, B.E. (1968): The birds of Borneo – Edinburgh
(Oliver & Boyd).
*STAENIS, C.G.G.J. VAN (1964): Plant geography of the
mountain flora of Mt. Kinabalu – Proc. Roy. Soc., Ser.
B, 161: 7-38.
*STAPF, O. (1894): On the flora of Mt. Kinabalu, North
Borneo – Trans. Linn. Soc. London, Bot., 4: 69-263.
*STEIN, N. (1978): Coniferen im westlichen Malaiischen
Archipel – The Hague, Boston, London (Junk Publ.)
STEINDACHNER, F. (1881): Über eine neue Pythonart (Py-
thon Breitensteini) aus Borneo. – Sitz. Ber. Kais. Akad.
Wiss., Wien, Abt.1, 82: 267-268.
STEJNEGER, L. (1902): Ptychozoon kuhli, a new name for P.
homalocephalum. – Proc. Biol. Soc., Washington, 15:
37.
– (1907): Herpetology of Japan and adjacent territory. –
Bull. U. S. Nat. Mus., 58: xx + 577 S.,
– (1922): List of snakes collected in Bulungan, northeast
Bibliography
Borneo by Carl Lumholtz, 1914. – Nyt. Mag. Naturv.
Kristiana, 60: 77-84.
*St. JOHN, R.W. (1862): Life in the forests in the Far East.
Vol. 1 -London (Smith Elder & Co.).
STOLICZKA, F. (1870): Observations on some Indian and
Malayan amphibia and reptiles. – J. Asiat. Soc. Bengal,
Calcutta, 39: 134-228.
STUEBING, R. (1991): A checklist of the snakes of Borneo. –
Raffles Bull. Zool., Singapore 39(2): 323-362.
– (1994): A checklist of the snakes of Borneo: Addena
and corrigenda. – Raffles Bull. Zool., Singapore, 42(4):
931-936.
STUEBING, R. & R.F. INGER (1998): Additional records on two
rare snakes from Borneo, with the confirmation of
Trimeresurus malcolmi LOVERIDGE as a distinct species. –
Raffles Bull. Zool., Singapore 46(2): 325-328.
– (1999): A fieldguide to the snakes of Borneo. – Kota
Kinabalu (Nat. Hist. Publ.), 254 pp.
TAN, F.L. (1993): Checklist of lizards of Sabah. – Kota
Kinabalu (Sabah Parks Trustees), 18 pp.
TAYLOR, E.H. (1920): Philippine amphibia. – Phil. J. Sci.,
Manila, 16(3): 215-357.
– (1922a): Additions to the herpetological fauna of the
Philippine Islands. II. – Phil. J. Sci., Manila, 21: 257-
303.
– (1922b): The snakes of the Philippine Islands. – Manila
(Bureau of Printing), 312 pp.
– (1960): On the caecilian species Ichthyophis monochrous
and Ichthyophis glutinosus with descriptions of related
species. – Kans. Univ. Sci. Bull., Lawrence, 40(4): 37-
120.
– (1965): (1965b): New Asiatic and African caecilians with
redescriptions of certain other species. – Kans. Univ. Sci.
Bull., Lawrence, 46 (5): 253-302.
– (1968): The Caecilians of the world. A taxonomic review. –
Lawrence (University of Kansas Press), 848 pp.
TAYLOR, E.H. & R.E. ELBEL (1958): Contribution to the
herpetology of Thailand. – Kans. Univ. Sci. Bull.,
Lawrence, 38(2): 1033-1189.
*TJIA, H.D. (1980): The Sunda Shelf, South East Asia. 3 –
Geomorphol., 24: 405-427.
TRUTNAU, L (1980): Über die Lebensweise und Pflege des
Netzpythons Python reticulatus (SCHNEIDER). – herpeto-
fauna, Weinstadt, 2 (4): 26-31.
TSCHUDI, J.J. VON (1838): Classification der Batrachier, mit
Berücksichtigung der fossilen Thiere dieser Abtheilung
der Reptilien. – Mem. Soc. Sci. Nat. Neuchâtel, 2: 1-99.
TWEEDIE, M.W.F. (1954): Notes on Malayan reptiles no. 3.
– Bull. Raffles Mus., Singapore, 25: 107-117.
– (1983): The snakes of Malaya. (3rd edition). – Singapore
(Singapore National Printers), 167 pp.
UNDERWOOD, G. & A. F. STIMSON (1990): A classification of
Pythons (Serpentes, Pythoninae). – J. Zool., London, 221:
565-603.
VENCES, M. (1999): Phylogenetic studies on ranoid frogs
415
Bibliography

(Amphibia: Anura) with a discussion of the origin and
evolution of the vertebrate clades of Madagascar. – Un-
published PhD thesis, University of Bonn.
VOGEL, G. (1995): Dendrelaphis striatus (COHN) neu für
die Fauna Borneos. – Mitt. Zool. Mus. Berlin, 70(1):
147-149.
– (2000): Zur Bestimmung der Unterarten von Boiga
dendrophila (BOIE, 1827) nebst einigen Neuigkeiten zu
Boiga dendrophila gemmicincta (DUMÉRIL, BIBRON &
DUMÉRIL, 1854). – Sauria, Berlin, 22(1): 27-43.
VOGEL, G. & P. HOFFMANN (1997): Über die Lebendfär-
bung von Bungarus flaviceps baluensis LOVERIDGE, 1938
(Serpentes, Elapinae). – Sauria, Berlin, 19(1): 13-16.
VORIS, H.K. & R.F. INGER (1995): Frog abundance along
streams in Bornean forests. – Conserv. Biol., 9(3): 679-683.
WAGLER, J. (1830): Natürliches System der Amphibien, mit
vorangehender Classification der Säugethiere und Vögel.
– (J. G. Gotta’sche Buchhandlung), vi + 354 pp.
WALL, F. (1923): A hand-list of the snakes of the Indian
Empire. Part 2. – J. Bombay nat. Hist. Soc., 24: 598-
632.
*WALLACE, A.R. (1869): The Malay Archipelago – Lon-
don (MacMillan & Co.).
WALLACH, V. & A.M. BAUER (1996): On the identity and
status of Simotes semicinctus PETERS, 1862 (Serpentes:
Colubridae). – Hamadryad, Madras, 21: 13-18.
WALLACH, V & R. GÜNTHER (1998): Visceral anatomy of
the Malaysian snake genus Xenophidion, including a cla-
distic analysis and allocation to a new family (Serpentes:
Xenophidiidae). – Amphibia-Reptilia, Leiden 19: 385-
404.
WANDOLLEK, B. (1900): Zur Kenntnis der Gattung Draco L.
– Abh. Ber. Mus.Tierk. Völkerk., Dresden, 9(3): 1-16.
WELCH, K.R.G. (1988): Snakes of the Orient: a checklist. –
Malabar, Florida (Robert F. Krieger Publ. Co): i-vii +
183 pp.
WERMUTH, H. (1967): Liste der rezenten Amphibien und
Reptilien: Agamidae. – Das Tierreich, Berlin, 86: xiv +
127 pp.
WERNER, F. (1892): Über eine kleine Collection von
Reptilien und Batrachiern von Nias. – Jahresber. Abh.
naturw. Ver. Magdegurg,: 248-254.
– (1896): Zweiter Beitrag zur Herpetologie der indo-
australischen Region. – Verh. Zool. Bot. Ges. Wien, 46:
6-24.
– (1910): Über neue oder seltene Reptilien des naturhis-
torischen Museums in Hamburg. II. Eidechsen. – Jb.
Hamb. wiss. Anst., 27, 2. Beiheft: 1-46.
*WHITEHEAD, J. (1893): The exploration of Kinabalu, North
Borneo. – London (Gurney & Jackson), 137 pp.
*WHITMORE, T.C. (1984): Tropical rain forest of the Far
East – London (Oxford Univ. Press).
– *(Ed.) (1987): Biogeographical evolution of the Malay
Archipelago – Oxford (Clarendon Press)
WIEGMANN , A.F.A. (1835): Beiträge zur Zoologie,
gesammelt auf einer Reise um die Erde, von Dr. F.J.F.
Meyen, siebente Abhandlung. - Amphibien. – Verh. Kais.
Leop. carol. Akad. Naturf., Halle 17(1): 185-268.
WILLIAMS, K.L. & V. WALLACH (1989). - Snakes of the world.
Vol. I. Synopsis of snake generic names. – Malabar,
Florida (Robert E. Krieger), viii + 234 pp.
WOLF, S. (1936): Revision der Untergattung Rhacophorus.
– Bull. Raffles Mus., Singapore, 12: 137-217.
WONG, A. (1994): Population ecology of amphibians in
different altitudes of Kinabalu Park. – Sabah Mus. J. 1(2):
29-38.
*WONG, K.M. & A. PHILLIPPS (Eds.) (1996): Kinabalu - Sum-
mit of Borneo. A revised and expanded Edition – Kota
Kinabalu (Sabah Soc. & Sabah Parks).
*WOOD, J.J., R.S. BEAMAN & J.H. BEAMAN (1993): The
plants of Mount Kinabalu. 2. Orchids – Kew (Royal
Botanic Gardens).
WÜSTER, W. & R.S. THORPE (1989): Population affinities of
the asiatic cobra (Naja naja) species complex in south-
east Asia: reliability and random resampling. – Biol. J.
Linn. Soc., London, 36: 391-409.
YANG, D.-T. (1991): Phylogenetic systematics of the
Amolops group of ranid frogs of Southeastern Asia and
the Greater Sunda Islands. – Fieldiana Zool., Chicago,
new Ser. 63: iii + 42 pp.
YONG, H. S. & I. RAMLY (1987): Conquest of the land: Di-
rect development in a Malaysian Philautus Tree Frog. –
Nat. Malay., Kuala Lumpur, 13(4): 4-7.
YORKE, C.D. (1983): Survival of embryos and larvae of the
frog Polypedates leucomystax in Malaysia. – J. Herpe-
tology, 17(3): 235-241.
ZIEGLER, T. & W. BÖHME (1996): Über das Beutespektrum
von Varanus dumerilii (SCHLEGEL. 1839). – Salamandra,
Rheinbach, 32(3): 203-210.

416
 Index

Index

A B
abbotti, Leptobrachium 91 baleata, Kaloula 125
Ablabes baliodirus. ® Gongylosoma baliodeirum baliodeirum baliodeirum, Gongylosoma 336
baliodeirum baluensis, Bungarus flaviceps 384
affinis, Draco 232 baluensis, Cyrtodactylus 253
Ahaetulla 303 baluensis, Ingerana 138
prasina prasina 303 baluensis, Kalophrynus 120
Ahaetulla caudolineata. ® Dendrelaphis caudolineatus baluensis, Leptobrachella 89
caudolineatus baluensis, Megophrys 106
albofuscus, Lepturophis 344 baluensis, Occidozyga 155
alfredi, Sphenomorphus 283 baluensis, Rhacophorus 206
Amblycephalus laevis. ® Internatus laevis beccarii, Tropidophorus 291
Amblycephalus nuchalis. ® Pareas nuchalis bicolor, Calamaria 317
Amblycephalus vertebralis. ® also Internatus laevis bivirgata tetrataenia, Calliophis 386
amoenus, Philautus 179 Boiga 308
Amolops amoropalamus. ® Meristogenys cynodon 309
amoropalamus dendrophila annectens 310
Amolops cavitympanum. ® Huia cavitympanum jaspidea 312
Amolops jerboa. ® also Meristogenys nigriceps 313
orphnocnemis; Meristogenys whiteheadi Boiga dendrophila. ® Boiga dendrophila annectens
Amolops kinabaluensis. ® Meristogenys kinabaluensis borbonica, Leptophryne 80
Amolops orphnocnemis. ® Meristogenys orphnocnemis borneensis, Microhyla 131
amoropalamus, Meristogenys 149 borneensis, Phoxophrys 247
Amphiesma 305 borneensis, Stegonotus 366
flavifrons 306 borneensis, Stoliczkia 367
sarawacense 307 borneensis, Trimeresurus 375
Amphiesma sarawakensis. ® Amphiesma sarawacense bornensis, Gonocephalus 241
Amphixestus Beccarii. ® Tropidophorus beccarii Bothrops gramineus. ® Trimeresurus popeiorum sabahi
angulirostris, Rhacophorus 201 bowringii, Riopa 281
annectens, Boiga dendrophila 310 Brachymeles 271
Ansonia 63 apus 272
altitudinis. ® Ansonia fuliginea breitensteini, Python 389
fuliginea 64 Bronchocela 227
guibei 65 cristatella 228
hanitschi 67 Bronchocela kinabaluensis. ® Hypsicalotes
longidigita 70 kinabaluensis
platysoma 72 Bronchocela nigrigularis. ® Complicitus nigrigularis
®
spinulifer 74 Bufo 75
Ansonia leptopus. ® also Ansonia spinulifer divergens 76
Ansonia minuta. ® also Ansonia guibei juxtasper 77
Aphaniotis 225 Bufo asper. ® also Bufo juxtasper
ornata 226 Bufo biporcatus. ® Bufo divergens
Aphaniotis nasuta. ® Aphaniotis ornata Bufo biporcatus biporcatus. ® Bufo divergens
appendiculatus, Rhacophorus 204 Bufo biporcatus divergens. ® Bufo divergens
Apterygodon 270 Bufo fuligineus. ® Ansonia fuliginea
vittatum 270 Bufo leptopus. ® also Ansonia guibei; Ansonia
apus, Brachymeles 272 longidigita
arayai, Leptolalax 98 Bufo penangensis. ® also Ansonia guibei
Atropophis borneensis. ® Trimeresurus borneensis Bufo spinulifer. ® Ansonia spinulifer
aurantium, Philautus 181 Bungarus 382

417
Index

fasciatus 383 nigrigularis 230

flaviceps baluensis 384 consobrinus, Cyrtodactylus 255
Bungarus flaviceps. ® Bungarus flaviceps baluensis conspicillatus, Rhabdophis 363
bunitus, Philautus 184 Cornufer baluensis. ® Ingerana baluensis
buttikoferi, Ophisaurus 250 cornutus, Draco 233
Coronella baliodeira. ® Gongylosoma baliodeirum
C baliodeirum
Cacophryne borbonica. ® Leptophryne borbonica Coryphodon fuscus. ® Ptyas fuscus
Calamaria 315 cristatella, Bronchocela 228
bicolor 317 cyanolaemus, Sphenomorphus 284
grabowskyi 318 cyanopunctatus, Rhacophorus 207
griswoldi 319 Cylindrophis 380
lateralis 320 ruffus ruffus 380
leucogaster 321 cynodon, Boiga 309
lumbricoidea 321 Cyrtodactylus 252
modesta 322 baluensis 253
schlegeli schlegeli 323 consobrinus 255
schmidti 323 ingeri 256
suluensis 324 matsuii 257
virgulata 325 Cyrtodactylus consobrinus kinabaluensis. ®
Calamaria baluensis. ® Calamaria grabowskyi Cyrtodactylus consobrinus
Calamaria bicolor. ® also Calamaria schlegeli Cyrtodactylus marmoratus. ® also Cyrtodactylus
schlegeli baluensis
Calamaria brachyura. ® Calamaria virgulata Cyrtodactylus pubisulcus. ® also Cyrtodactylus ingeri
Calamaria lautensis. ® also Calamaria modesta
Calamaria lumbricoidea griswoldi. ® Calamaria D
griswoldi Dasia 272
Calamaria pendlebury. ® Calamaria suluensis olivacea 273
Calamaria temminckii. ® Calamaria lumbricoidea Dendrelaphis 329
Calamaria vermiformis. ® also Calamaria caudolineatus caudolineatus 329
lumbricoidea pictus pictus 330
Calliophis 385 dendrophila annectens, Boiga 310
bivirgata tetrataenia 386 Dicroglossus kuhli. ® Limnonectes kuhlii
intestinalis thepassi 386 Dilophyrus grandis. ® Gonocephalus grandis
Calohyla sundana. ® Metaphrynella sundana Dipsas. ® Boiga
Calotes kinabaluensis. ® Hypsicalotes kinabaluensis divergens, Bufo 76
Calotes nigrigularis. ® Complicitus nigrigularis Dogania 399
caudolineatus caudolineatus, Dendrelaphis 329 subplana 399
cavitympanum, Huia 137 Doliophis intestinalis. ® Maticora intestinalis thepassi
cephalum, Phoxophrys 248 Doliophis intestinalis var. everetti. ® Maticora
Ceratophryne nasuta. ® Megophrys nasuta intestinalis thepassi
chalconota, Rana 159 Draco 231
Chaperina 116 affinis 232
fusca 116 cornutus 233
chaseni, Ovophis 373 fimbriatus fimbriatus 235
chrysargos, Rhabdophis 362 maximus 236
Chrysopelea 325 melanopogon nigriappendiculatus 237
paradisi paradisi 326 obscurus obscurus 238
pelias 327 quinquefasciatus longibarba 239
Chrysopelea chrysochlora. ® Chrysopelea pelias Draco cornutus. ® also Draco affinis
collaris, Pseudorabdion 358 Draco fimbriatus. ® Draco fimbriatus fimbriatus
Coluber flavolineatus. ® Elaphe flavolineata Draco formosus. ® Draco obscurus obscurus
Coluber oxycephalus. ® Gonyosoma oxycephalum Draco formosus obscurus. ® Draco obscurus obscurus
Coluber pictus. ® Dendrelaphis pictus pictus Draco maximus maximus. ® Draco maximus
Complicitus 230 Draco obscurus. ® Draco obscurus obscurus

418
 Index

Draco quinquefasciatus. ® Draco quinquefasciatus Gongylosoma 335

longibarba baliodeirum baliodeirum 336
Draco spilopterus cornutus. ® Draco cornutus Gonocephalus 240
Draco volans. ® also Draco obscurus obscurus bornensis 241
dringi, Leptolalax 100 grandis 244
Dryophis prasinus. ® Ahaetulla prasina prasina Gonocephalus bellii. ® also Gonocephalus bornensis
dulitensis, Rhacophorus 209 Gonocephalus bellii (borneensis). ® Gonocephalus
dumerilii, Varanus 294 bornensis
Gonocephalus borneensis. ® Gonocephalus bornensis
E Gonocephalus liogaster. ® also Gonocephalus
edwardinae, Megophrys 109 bornensis
Elaphe 332 Gonydactylus. ® Cyrtodactylus
flavolineata 333 Gonyophis 337
taeniura grabowskyi 334 margaritatus 337
Elaphe erythrura. ® also Elaphe flavolineata Gonyosoma 338
Elaphe erythrurus. ® also Elaphe flavolineata oxycephalum 339
Elaphe melanura. ® Elaphe flavolineata grabowskyi, Calamaria 318
Elaphe oxycephala. ® Gonyosoma oxycephalum grabowskyi, Elaphe taeniura 334
Elaphe taeniura. ® Elaphe taeniura grabowskyi grandis, Gonocephalus 244
Elaphis grabowskyi. ® Elaphe taeniura grabowskyi griswoldi, Calamaria 319
Elaps octolineatus. ® Oligodon octolineatus guibei, Ansonia 65
Elaps tetrataenia. ® Maticora bivirgata tetrataenia gunungense, Leptobrachium 93
Elaps thepassi. ® Maticora intestinalis thepassi Gymnodactylus. ® Cyrtodactylus
ellipsifer, Xenelaphis 369
Emys spinosa. ® Heosemys spinosa H
erythraea, Rana 160 hanitschi, Ansonia 67
tetrataenia, Maticora bivirgata 386 harrissoni, Rhacophorus 214
Eumeces bowringii. ® Riopa bowringii Hazelia picta. ® Nyctixalus pictus
everetti macroscelis, Rhacophorus 210 Helicopsoides typicus. ® Opisthotropis typicus
everetti, Oligodon 349 Hemidactylus 262
everetti, Pedostibes 82 frenatus 262
F garnotii 264
fasciatus, Bungarus 383 Hemidactylus mutilatus. ® Gehyra mutilata
Fejervarya 134 Hemiphyllodactylus 264
limnocharis 135 typus typus 265
fimbriatus fimbriatus, Draco 235 Hemiphyllodactylus typus. ® Hemiphyllodactylus typus
finchi, Limnonectes 141 typus
flaviceps baluensis, Bungarus 384 Heosemys 397
flavifrons, Amphiesma 306 spinosa 397
flavolineata, Elaphe 333 hosii, Rana 161
Forest Hill Tortoise. ® Heosemys spinosa Huia 136
frenatus, Hemidactylus 262 cavitympanum 137
fuliginea, Ansonia 64 Hydrablabes 340
fusca, Chaperina 116 praefrontalis 340
fuscus, Ptyas 360 Hyla chalconota. ® Rana chalconota
Hyla erythraea. ® Rana erythraea
G Hyla leucomystax. ® Polypedates leucomystax
garnotii, Hemidactylus 264 Hylaplesia borbonica. ® Leptophryne borbonica
gauni, Rhacophorus 212 Hylarana (Rana) chalconota. ® Rana chalconota
Gehyra 258 Hylarana chalconota. ® Rana chalconota
mutilata 259 Hylarana hosei. ® Rana hosii
Gekko 260 Hylarana luctuosa. ® Rana luctuosa
monarchus 260 Hylarana microdisca palavanensis. ® Limnonectes
Geoemyda spengleri spengleri. ® also Heosemys palavanensis
spinosa Hylarana nicobariensis. ® Rana nicobariensis

419
Index

Hylarana signata. ® Rana signata nieuwenhuisi 275

Hylorana nicobariensis. ® Rana nicobariensis lateralis, Calamaria 320
Hypsicalotes 245 latopalmatus, Staurois 170
kinabaluensis 245 Lepidodactylus 266
ranauensis 266
I leporinus, Limnonectes 144
Ichthyophis 219 Leptobrachella 88
monochrous 219 baluensis 89
Idiopholis. ® Pseudorabdion parva 90
indeprensa, Mabuya 276 Leptobrachella baluensis. ® also Leptolalax arayai
Ingerana 138 Leptobrachella mjobergi. ® also Leptobrachella
baluensis 138 baluensis
ingeri, Cyrtodactylus 256 Leptobrachium 91
ingeri, Philautus 186 abbotti 91
Internatus 341 gunungense 93
laevis 342 montanum 95
malaccanus 343 Leptobrachium baluense. ® Megophrys baluensis
intestinalis thepassi, Calliophis 386 Leptobrachium gracile. ® also Leptolalax arayai
Ixalus guttatus. ® Staurois natator Leptobrachium gracilis. ® also Leptolalax arayai
Ixalus latopalmatus. ® Staurois latopalmatus Leptobrachium hasselti. ® also Leptobrachium abbotti;
Ixalus longicrus. ® Philautus longicrus Leptobrachium montanum
Ixalus natator. ® Staurois natator Leptobrachium montanum. ® also Leptobrachium
Ixalus petersi. ® Philautus petersi abbotti
Ixalus pictus. ® Nyctixalus pictus Leptolalax 97
J arayai 98
dringi 100
Japalura nasuta. ® Aphaniotis ornata
maurus 102
Japalura nigrilabris. ® also Phoxophrys cephalum
pictus 103
Japalura ornata. ® Aphaniotis ornata
Leptolalax gracilis. ® also Leptolalax
jaspidea, Boiga 312
arayai; Leptolalax pictus
juxtasper, Bufo 77
Leptophryne 80
K borbonica 80
Kalophrynus 119 Lepturophis 344
baluensis 120 albofuscus 344
pleurostigma 123 Lepturophis borneensis. ® Lepturophis albofuscus
Kalophrynus pleurostigma. ® also Kalophrynus leucogaster, Calamaria 321
baluensis leucomystax, Polypedates 194
Kalophrynus pleurostigma pleurostigma. ® also limnocharis, Fejervarya 135
Kalophrynus baluensis; Kalophrynus Limnodytes luctuosus. ® Rana luctuosa
pleurostigma Limnonectes 139
Kaloula 124 finchi 141
baleata 125 kuhlii 142
pulchra 126 leporinus 144
kinabaluensis, Hypsicalotes 245 palavanensis 145
kinabaluensis, Meristogenys 150 Limnonectes leporina. ® Limnonectes leporinus
kinabaluensis, Sphenomorphus 286 Liopeltis baliodeirus. ® Gongylosoma baliodeirum
kobayashii, Megophrys 110 baliodeirum
kuhlii, Limnonectes 142 longicrus, Philautus 186
longidigita, Ansonia 70
L Lophyrus bornensis. ® Gonocephalus bornensis
Lachesis gramineus. ® also Trimeresurus popeiorum luctuosa, Rana 165
sabahi lumbricoidea, Calamaria 321
laevis, Internatus 342 Lycodon 345
laevis, Occidozyga 157 subcinctus sealei 346
Lamprolepis 274 Lycodon albofuscus. ® Lepturophis albofuscus

420
 Index

Lycodon melanocephalus. ® Sibynophis cf. kinabaluensis 148

melanocephalus melanocephalus cf. orphnocnemis 148
Lygosoma alfredi. ® Sphenomorphus alfredi kinabaluensis 150
Lygosoma bowringi. ® Riopa bowringii orphnocnemis 152
Lygosoma kinabaluensis. ® Sphenomorphus whiteheadi 153
kinabaluensis Meristogenys jerboa. ® also Meristogenys
Lygosoma nieuwenhuisi. ® Lamprolepis nieuwenhuisi orphnocnemis
Lygosoma olivaceum. ® Dasia olivacea Metaphrynella 127
Lygosoma teniuculum. ® Sphenomorphus tenuiculus sundana 127
Lygosoma variegatum. ® also Sphenomorphus Micrixalus baluensis. ® Ingerana baluensis
cyanolaemus; Sphenomorphus multisquamatus; Microhyla 130
Sphenomorphus kinabaluensis borneensis 131
Lygosoma whiteheadi. ® Riopa bowringii petrigena 132
Lymnodactylus sp.. ® Cyrtodactylus baluensis misera, Pelophryne 85
mjobergi, Philautus 187
M Mochlus bowringi. ® Riopa bowringii
Mabuia. ® Mabuya modesta, Calamaria 322
Mabuya 275 monarchus, Gekko 260
indeprensa 276 Monitor Dumerilii. ® Varanus dumerilii
multifasciata 276 monochrous, Ichthyophis 219
rudis 278 montanum, Leptobrachium 95
rugifera 279 multifasciata, Mabuya 276
Mabuya multicarinata. ® Mabuya indeprensa multisquamatus, Sphenomorphus 287
Macropisthodon 347 murudensis, Rhabdophis 364
rhodomelas 347 mutilata, Gehyra 259
macroscelis, Rhacophorus everetti 210
macrotis, Polypedates 196 N
maculatus, Pedostibes 83 Naja 387
maculatus, Xenochrophis 370 sumatrana 388
malaccanus, Internatus 343 Naja naja. ® also Naja sumatrana
malcolmi, Trimeresurus 376 Naja tripudians. ® also Naja sumatrana
margaritatus, Gonyophis 337 Naja tripudians var. miolepis. ® Naja sumatrana
Maticora intestinalis. ® Calliophis intestinalis thepassi nasuta, Megophrys 112
Maticora intestinalis nigrotaeniatus. ® Calliophis natator, Staurois 171
intestinalis thepassi Natrix chrysarga. ® Rhabdophis chrysargos
matsuii, Cyrtodactylus 257 Natrix flavifrons. ® Amphiesma flavifrons
maurus, Leptolalax 102 Natrix maculatus. ® Xenochrophis maculatus
maximus, Draco 236 Natrix murudensis. ® Rhabdophis murudensis
Megalophrys. ® Megophrys Natrix saravacensis. ® Amphiesma sarawacense
Megophrys 105 Natrix trianguligera. ® Xenochrophis trianguligerus
baluensis 106 Nectophryne altitudinis. ® Ansonia fuliginea
edwardinae 109 Nectophryne everetti. ® Pedostibes everetti
kobayashii 110 Nectophryne guentheri. ® also Pelophryne misera
nasuta 112 Nectophryne maculata. ® Pedostibes maculatus
Megophrys abbotti. ® Leptobrachium abbotti Nectophryne misera. ® Pelophryne misera
Megophrys gracilis. ® also Leptolalax arayai Nesobia mjobergi. ® also Leptobrachella baluensis
Megophrys hasselti. ® also Leptobrachium montanum nicobariensis, Rana 165
Megophrys monticola. ® also Megophrys kobayashii nieuwenhuisi, Lamprolepis 275
Megophrys monticola nasuta. ® Megophrys nasuta. ® nigriappendiculatus, Draco melanopogon 237
also Megophrys kobayashii nigriceps, Boiga 313
Megophrys nasuta. ® also Megophrys kobayashii nigrigularis, Complicitus 230
melanocephalus melanocephalus, Sibynophis 365 nigropalmatus, Rhacophorus 215
melanopogon nigriappendiculatus, Draco 237 Norbea beccarii. ® Tropidophorus beccarii
Meristogenys 147 Norbea mocquardi. ® Tropidophorus beccarii
amoropalamus 149 nuchalis, Pareas 355

421
Index

Nyctixalus 176 bunitus 184

pictus 176 ingeri 186
longicrus 186
O
mjobergi 187
obscurus obscurus, Draco 238
petersi 189
Occidozyga 154
saueri 191
baluensis 155
Philautus aurantium montanum. ® Philautus
laevis 157
aurantium
octolineatus, Oligodon 350
Philautus aurifasciatus. ® also Philautus
Oligodon 348
aurantium; Philautus longicrus; Philautus
everetti 349
mjobergi; Philautus petersi
octolineatus 350
Philautus bimaculatus. ® also Rhacophorus
purpurascens 351
angulirostris; Rhacophorus cyanopunctatus
vertebralis vertebralis 352
Philautus petersi. ® also Philautus mjobergi
olivacea, Dasia 273
Philautus pictus. ® Nyctixalus pictus
longibarba, Draco quinquefasciatus 239
Philautus spiculatus. ® Rhacophorus everetti
Ooeidozyga baluensis. ® Occidozyga baluensis
macroscelis
Ophisaurus 250
Phoxophrys 247
buttikoferi 250
borneensis 247
Ophites albofuscus. ® Lepturophis albofuscus
cephalum 248
Opisthotropis 352
Phoxophrys nigrilabris. ® also Phoxophrys cephalum
typicus 353
Phrynoglossus baluensis. ® Occidozyga baluensis
Oreobatrachus baluensis. ® Occidozyga baluensis
picturata, Rana 166
ornata, Aphaniotis 226
pictus, Leptolalax 103
orphnocnemis, Meristogenys 152
pictus, Nyctixalus 176
otilophus, Polypedates 197
pictus pictus, Dendrelaphis 330
Ovophis 373
Platydactylus monarchus. ® Gekko monarchus
chaseni 373
platysoma, Ansonia 72
oxycephalum, Gonyosoma 339
pleurostigma, Kalophrynus 123
Oxyglossus laevis. ® Occidozyga laevis
Polypedates 193
P leucomystax 194
palavanensis, Limnonectes 145 macrotis 196
paradisi paradisi, Chrysopelea 326 otilophus 197
pardalis, Rhacophorus 216 Polypedates appendiculatus. ® Rhacophorus
Pareas 355 appendiculatus
nuchalis 355 Polypedates signatus. ® Rana signata
Pareas laevis. ® Internatus laevis popeiorum sabahi, Trimeresurus 377
parva, Leptobrachella 90 praefrontalis, Hydrablabes 340
Passerita prasina. ® Ahaetulla prasina prasina prasina prasina, Ahaetulla 303
Pedostibes 81 Psammodynastes 356
everetti 82 pulverulentus 357
maculatus 83 Psammophis pulverulenta. ® Psammodynastes
Pedostibes altitudinis. ® Ansonia fuliginea pulverulentus
pelias, Chrysopelea 327 Pseudoboa fasciata. ® Bungarus fasciatus
Pelochelys bibroni. ® also Dogania subplana Pseudorabdion 358
Pelophryne 84 collaris 358
misera 85 Ptyas 359
Pelophryne brevipes. ® also Pelophryne misera fuscus 360
Pelturagonia cephalum. ® Phoxophrys cephalum Ptychozoon 267
Peropus mutilatus. ® Gehyra mutilata rhacophorus 268
petersi, Philautus 189 pulchra, Kaloula 126
petrigena, Microhyla 132 pulverulentus, Psammodynastes 357
Philautus 178 purpurascens, Oligodon 351
amoenus 179 Python 389
aurantium 181 breitensteini 389

422
 Index

reticulatus 390 everetti macroscelis 210

Python curtus breitensteini. ® Python breitensteini
Q nigropalmatus 215
quinquefasciatus longibarba, Draco 239
R angulirostris
Rana 158
chalconota 159
erythraea 160
hosii 161
luctuosa 163
nicobariensis 165
picturata 166
signata 166
Rana (Hylarana). ® Rana
Rana (Limnonectes) kuhlii. ® Limnonectes kuhlii
Rana (Limnonectes) limnocharis. ® Fejervarya
limnocharis
Rana blythi. ® also Limnonectes leporinus
Rana cavitympanum. ® Huia cavitympanum
Rana chalconota raniceps. ® Rana chalconota
Rana decorata. ® Rana luctuosa
Rana finchi. ® Limnonectes finchi
Rana hosei. ® Rana hosii
Rana kuhlii. ® Limnonectes kuhlii
Rana leporina. ® Limnonectes leporinus
Rana limnocharis. ® Fejervarya limnocharis
Rana macrodon. ® also Limnonectes leporinus
Rana macrodon var leporina. ® Limnonectes leporinus
Rana microdisca finchi. ® Limnonectes finchi
Rana microdisca palavanensis. ® Limnonectes
palavanensis
Rana obsoleta. ® Rana signata
Rana palavanensis. ® Limnonectes palavanensis
Rana paradoxa. ® Limnonectes kuhlii
Rana signata signata. ® also Rana picturata
Rana tuberilinguis. ® Staurois tuberilinguis
Rana whiteheadi. ® Meristogenys whiteheadi. ® also
Meristogenys kinabaluensis
ranauensis, Lepidodactylus 266
reticulatus, Python 390
Rhabdophis 361
chrysargos 362
conspicillatus 363
murudensis 364
Rhabdophis chrysarga. ® Rhabdophis chrysargos. ®
also Rhabdophis murudensis
Rhabdophis chrysargus. ® Rhabdophis chrysargos
Rhacophorus 198
angulirostris 201
appendiculatus 204
baluensis 206
cyanopunctatus 207
dulitensis 209
gauni 212
harrissoni 214
pardalis 216
Rhacophorus acutirostris. ® Rhacophorus
Rhacophorus bimaculatus. ® also Rhacophorus
cyanopunctatus
Rhacophorus colletti. ® also Philautus ingeri
Rhacophorus everetti. ® also Rhacophorus everetti
macroscelis
Rhacophorus hosei. ® also Philautus ingeri
Rhacophorus leucomystax. ® Polypedates leucomystax
Rhacophorus leucomystax linki. ® Polypedates
macrotis
Rhacophorus leucomystax sexvirgata. ® Polypedates
leucomystax
Rhacophorus leucomystax var. quadrilineatus. ®
Polypedates leucomystax
Rhacophorus macroscelis. ® Rhacophorus everetti
macroscelis
Rhacophorus macrotis. ® Polypedates macrotis
Rhacophorus otilophus. ® Polypedates otilophus
rhacophorus, Ptychozoon 268
Rhacophorus spiculatus. ® Rhacophorus everetti
macroscelis
rhodomelas, Macropisthodon 347
Riopa 281
bowringii 281
rudis, Mabuya 278
ruffus ruffus, Cylindrophis 380
rugifera, Mabuya 279
S
sabahi, Trimeresurus popeiorum 377
sabanus, Sphenomorphus 289
salvator salvator, Varanus 295
sarawacense, Amphiesma 307
saueri, Philautus 191
schlegeli schlegeli, Calamaria 323
schmidti, Calamaria 323
Scincus multifasciatus. ® Mabuya multifasciata
sealei, Lycodon subcinctus 346
Sibynophis 365
melanocephalus melanocephalus 365
Sibynophis geminatus. ® also Sibynophis
melanocephalus melanocephalus
signata, Rana 166
Simotes. ® Oligodon
Sinomantis latopalmata. ® Staurois latopalmatus
Sinonatrix trianguligera. ® Xenochrophis
trianguligerus
Sphecodes albo-fuscus. ® Lepturophis albofuscus
Sphenomorphus 282

423
Index

alfredi 283 Tropidonotus flavifrons. ® Amphiesma flavifrons

cyanolaemus 284 Tropidonotus maculatus. ® Xenochrophis
kinabaluensis 286 maculatus. ® also Amphiesma sarawacense
multisquamatus 287 Tropidonotus saravacensis. ® Amphiesma sarawacense
sabanus 289 Tropidonotus sarawacensis. ® Amphiesma
tenuiculus 290 sarawacense
Sphenomorphus aquaticus. ® Tropidophorus beccarii Tropidonotus trianguligerus. ® Xenochrophis
Sphenomorphus cyanolaemus. ® also Sphenomorphus trianguligerus
kinabaluensis Tropidophorus 291
Sphenomorphus sabanus. ® also Sphenomorphus beccarii 291
multisquamatus Tropidophorus mocquardi. ® Tropidophorus beccarii
spinosa, Heosemys 397 tuberilinguis, Staurois 172
spinulifer, Ansonia 74 Typhlops sp. 392
Staurois 169 typicus, Opisthotropis 353
latopalmatus 170 typus typus, Hemiphyllodactylus 265
natator 171
tuberilinguis 172 U
Stegonotus 366 unicolor, Xenopeltis 393, 394
borneensis 366 V
Stellio salvator. ® Varanus salvator salvator
Varanus 293
Stoliczkaia ® Stoliczkia
dumerilii 294
Stoliczkia 367
salvator salvator 295
borneensis 367
vertebralis vertebralis, Oligodon 352
subcinctus sealei, Lycodon 346
virgulata, Calamaria 325
subplana, Dogania 399
vittatum, Apterygodon 270
suluensis, Calamaria 324
sumatrana, Naja 388 W
sundana, Metaphrynella 127 wagleri, Tropidolaemus 379
whiteheadi, Meristogenys 153
T
taeniura grabowskyi, Elaphe 334 X
tenuiculus, Sphenomorphus 290 Xenelaphis 369
tetrataenia, Calliophis bivirgata 386 ellipsifer 369
thepassi, Calliophis intestinalis 386 Xenochrophis 370
Tiaris miotympanum. ® Gonocephalus bornensis maculatus 370
trianguligerus, Xenochrophis 371 trianguligerus 371
Triglyphodon jaspideum. ® Boiga jaspidea Xenodon purpurascens. ® Oligodon purpurascens
Trimeresurus 374 Xenopeltis 393
borneensis 375 unicolor 393, 394
malcolmi 376
popeiorum sabahi 377 Z
Trimeresurus chaseni. ® Ovophis chaseni Zaocys fuscus. ® Ptyas fuscus
Trimeresurus popeorum. ® Trimeresurus popeiorum
sabahi
Trimeresurus puniceus. ® Trimeresurus borneensis
Trimeresurus sumatranus. ® also Trimeresurus
malcolmi
Trimeresurus sumatranus malcolmi. ® Trimeresurus
malcolmi. ® also Trimeresurus popeiorum sabahi
Trionyx subplanus. ® Dogania subplana
Tropidolaemus 378
wagleri 379
Tropidonotus chrysargos. ® Rhabdophis chrysargos
Tropidonotus conspicillatus. ® Rhabdophis
conspicillatus

424