A Field Guide To The FROGS OF BORNEO PDF

,f1bout th e 13001\
A ,"' jcld G uide 10 the f rogs or Borneo

introduces the non-spt:·Ciali:.1 reader to the
natural history of this diver.;e, colourful, and
sometimes bizarre- looking section or the fauna
of Borneo. AboUi 150 spc<:ics of frogs are now
known from Borneo and the number continues
to grow. Many of these species are unique to
Bom(.'O. and a number of them arc ~tiJJ known
only from one or two sighting... A.. these
animal .. Ih"e in alJ sorts of environments from
stream bank!; 10 forest canopy. from lowland
rice field:. to monlane forests, they are likely to
be encountered by hiker!> with an imcrc~l in
natural hi~tory. This book covers the specie..
most likely to be seen and identifiahle without
the use of a microloCopc. For each i>pccic:.. a
description and summary of natural hi~lory are
presented. 81 !>pct"ics arc illustrated by II colour
photo. In addi tion. introductory chaptch
summarize the general biology of frog~ arid
their ecology. enabling the reader to undclY
land how these animals iii inlO Ihe complex
!CO!)yslcnL\, of Borneo.
Front cover: RhacQpllOrtls reiml'Urrltl. one of

Ihe gliding frogs of Borneo.
Above: Rmw Iws;;. (Photos: R.B. Sluebingl
'1: O
~ _ _ _ _ _.2.00",====4(~I() kr n ~o
below 200 ill
200 - 500m
500 - 1000 m
1000 - 2{){)O In
2000 - 3000 m Balldar

;lhm" .~()O() In
-Ii
BELITUNG
'"
M<lp of B om~o.
A FIELD G U IDE TO THE
FROGS
OF BORNEO
Second Edition
Robert F. Inger and Robert B. Stuebing
Photographs hy
R.B. Stuebing, R.F. Inger. c.L. Chan. Bjorn Lardner.
J.c. Murphy. W.M. Poon. H.P. Hazebroek. Tee Kim Ling.
M. KOlleial, A.ye. Chung. Kelvin Lim. S. Reimer and I. Das
JIIustrations h,'
Tan Fui Lian
Natural History Publications (Borneo)

Kola Kinahalu
2005
I'ubli:./red b.1
Naturalllistory Pllblicatiol/s (RomfO) Sd". Rhd. 1 ~1t>1;()7· X )

A913. 9th FlOOr. Phase I. Wisma Merdeb.
P.O. Box 15566.
88864 Kota Kinabalu. Sabah. h-lalaysia.
Td : (608!!) 233098 Fax: (6088) 240768
E-mail: info@nhphomco.com
Website: www.nhpborneo.colll
Tcxt copyright (Q 2005 Natural HislOry Publications (Borneo) Sdll. Bhd.

Photogmphs and illustrations copyright <D 2005 as credited.
AI! rigins rescrvcd. No part of this publication lIlay be reproduced. ,tored in

a retrieval system. or tmllsmillcd in any (omlor b) any mean,. cledmnic
mcchanical. photocopying:, recording. or othcrwisc. withoU1thc prior
pennission of the copyright owners.
First publishcd 1997.

Scnmd re,i,cd edition pnblished March 2005.
A Fi{'"ld Guide to the Fro!!;s of Romeo (S{'"cond Edition)

h) Robert F. Inger and Ro~rt B. Sluehing:
[SB~ 983·812·085-5
Fmnti,picce: Map of Borneo.

Contents
Preface ............ . . ... vi

Preface to the Second Edition ............................................ . . ... vii
General Biology of Frogs ....... "'" .•.•.......................•....... 1
Frogs in Human Tradition and Culture .......................... ..... ........... 7
Di stribution and Classification of Frogs of Borneo .............. II

Key to the Frogs of Borneo. .. ............ . .15
A Checklist of Anuran Amphibians ........ ..... ..... 2...
Ecology of Bamean Frogs .................................................................... 29
Conservation ...... -+ I
family Bombinatoridae ............ . . ...................... ........................... 43
Family Megophryidae .. . ............ .... ............... ........ ................. 45
Famil y Bufonidae (Tme Toad:\) ........................... 67
Fam ily Microhylidac (Narrow-mouthed Frogs) .......................... . .... 93
Family Ranidae ........ ...... . ................. . . ... 111
Family Rhacophoridae ...... 159
Further Reading .............. . ..19-4

I\cknowlcdgcmcnb ............. 195
Index ....... ............ . 196
cf>rejace 10 th e § irst edition
I has been j ust under ten years since a discussion on the veranda of (he
I Danum Valley Rest House resulted in the publication of Frogs of Sabal!.

our first attempt to introduce Bornean frogs to people ol\!side a small
circle of amphibian biologists. At the lime, we were hopeful that such a hook
would provide a convenient window for Malaysian students, the public. and
overseas visitors to have a closer look at Sabah's rich frog fauna. Happily for
us, the little book seems to have succeeded at its task. But Sabah is just one
fraction of BurneD. What about the frog fauna of the rest of that large island?
For quite some lime we did not feci up \0 Ihe challenge of taking on the fauna
of the whole of Borneo. Fortune has again been kind. however. and we have
been able to continue working in Sabah and Sarawak. as well as receiving
assistance from generous colleagues who have done field research in
Kalimantan. At last we feel confident that there is sufficient infonnation for
a book with a much wider scope.
Much of the basic text of this new book is, of course, based on the
original Frogs of 5aball. Howeve.r, all of the chapters have been revised and
expaoded. and the coverage of species accounts increased from 56 to 97.
Most Bornean frog species are covered by the key, except those for which
identification is too difficult for alJ blll a frog taxonomist. A lso. since most of
these species are so difficult {O find that only a handful of people would ever
have the opportunity to see them alive, we have restricted the colour plates
in the book to cover 67% of the known frog species of Borneo. We have tried
(0 counteract some of the diflicullies of identilication by providing general
descriptions for most of the genera.

Complicating the task of identification for the general reader is the fact
Ihm new species cominue to be discovered. In fact, 13 new species have been
found since publication of Frogs (~r Saba".
We have participated in the excitemelll of these discoveries and in the
unravelling of the secrets in the lives of Bornean frogs. Above all. however,
we have enjoyed living and working in Borneo. with people who have shown
us great kindness and generosity.
RObel1 Inger
Robert Stuebing
v.
C{JreJace to the Second eclition
11 the Preface to the lirM edidon. we ,said that new spec ie:, of rrog'
I cOlllinue to be discovered in Borneo. Study of this part of the rich fauna

of Borneo has not dim inished and three new species have been
discovered within the Jast live years . But research on this fauna has /lot been
limited 10 discovery of new species. It has also led to increased
understanding of the relationships among the species. In a lew cases we have
come 10 understand that what herpetologists originally thought were two
distinct species afC actually a single spec ie:. and. conversely. that what wa~
thought \0 be a single spec ies appears to be two d istinct Lhou~h simi lar
species. Two developments are especially encouraging: First. the scope of
research has expa nded to include rnOVC IllCIl U: o f frogs ilnd tadpole,. the
impact of differing types of logging on the frog assemblages. gcnetk
variation within ~pecie~. and the evolution:!/)' relations among specie':> and
genera. Secondly. and even more imponam, an increasing number of local
biologists. some from local universi ties and some from National Parks and
other government agencies. are engaged in these ~\udie~. Both of these trends
will lead to greater knowledge and understanding of this fauna. As that
understanding i') fundamental \0 ultimate conservation of this rich and
beautiful reSource. we look to the future with confidence.
Roben F. Inger
Roben B. SlUebing
2005
\ ' 11
Grl1 rral Cf3iolog y of /lrog s
"eryone can recogni,l;I.' a frog. which is a good indication of ho\\
E distinct and relatively unilom1 these animals arc. Their distinctive

features include: (I) no tail: (2) a short. onen ~Iocky body: (3) long
hind legs and short front ones; (4) large bulging eyes. and (5) a \'cry wide
mouth. Not obvious from the outside. bm equally characteristic. are an
extraordinarily short backbone (no frog has more than nine \\!rtebrae ).
virtually no ribs. and a very short digestive lrat'!.
The long hind leg. the propulsive organ in a frog's leap. has a ~pecial.
extra joint. Hesides the J..ncc and ankle connections common to all terrestrial
vertehrates. frogs have another movable one across what corresponds to the
middle of" our foot. This extra hinge provides additional \evcmgt' fur
launching the body rapidly duritlg a leap. A ~hortcncd backbone that i~
hinged to a long pclvi~ helps the frog to keep its e)e~ fat'ing directly ahead
during take-off and landing. A lew frogs ~lfe not so athletic. but in~tead take
~hon hops. or just wltlk. These unusual ~pecies. which include the Bomeall
Horned Frog (MlJRophrys nasllfa). ha\c rather short hind legs. but slill
po~~c~~ the extra joilll (Fig. 2).
A frog' s large tleshy tongue is another organ with unuwal features. It is
generally very broad and soft and. unlike the tongues of birds amI mammals.
is attached <II the front of the lower jaw. When feeding. a frog f1ip~ it!> tongue
Oll! at its prey. then f1ip~ it back in with the prey unhappily stuck on thl: end.
That brings us to the die!. which consists exclusively of animals. Inset·1s and
other invertebrates are the u~\Ial prey. though large species of frogs may
dl'YOUr other frogs. or even small snakes. small birds or mammals. Only one
species of frog is known to feed on plants. though ~omelime~ rragmellls of
planl~ may be ~wallowed accidentally. 1 This exclusively animal diet has
resulted in a short digcstiye tract. In fact. generally speaking. vertebrate"
from fishes 10 1TI1lmlllab that feed on vegetation have long il1!estine~. while
those that feed exclusively on animals have short ones.
1 There j, one "pedes of frog on the Indian ~\llx:ontin~nt that i, reponed to con~ume
aquatic \cgetatioll a~ part of ib diel.
Fig. I (opposite). A ~mall :.Irc<lnl in hill) 10\\ land rain forest in Sarawak. a \ari~ty of
Boml;"an ~Jk6~", of frogs hreed at thi~ kind of cn\ironmcnt. (Photo: H.P. Hazebrock )
GENERAL BIOLOGY OF FROGS
parotoid
heel
extra
joint
Fig. 2. Outline ()ffrog showing anatomical feature,~ referred to in text. (Drawn by Tan
Fui Lian)
Another distinctive characteristic of frogs is their complex life cycle.

With few exceptions. frogs begin life as aquatic larvae. or tadpoles (Fig. 3).
and undergo a complicated series of changes called metamorphosis. as they
assume the form of a frog. In general shape and way of life. hardly anything
could be as difTerent from a frog as a tadpole. Tadpolc~ havc an egg-shaped
body and a fumed laii. They are truly aquatic. taking oxygen from the water
by means of gills. have an extremely long . coiled intestine and feed mainly
on microscopic algae and fungi. Teeth are replaced by a pair of horny beaks.
while outside the mouth wide lips support rows of tiny tooth-like structures .
or dcnticles. Most tadpoles feed by using the lips. denticJes and beaks to
scrape or stir up material on the bottom of a pool or stream. forming a small
cloud of line debri s which they swallow. Elaborate structures inside the
mouth are used 10 liltcr out the organic matter in the sU~jX'n~ion that they
have created.
A~ Ihey grow. their hind legs begin to develop at the end of the body. The
forelimbs actually develop a bit earlier than the hind ones, but are hidden b)
a fold of !)J...in. and thus 1I0t visible. Some tadpoles. particularly tho~e Iha! li\'e
2
Fig. J. Slagc~ of dc\ .:Iopmcnl of lad poles. (Drawn by Phillip Yong after Tan i-'ui Li:tn )
3
in swift flo\\ing "aler. ha\c mor~' ... lender or nallcned bodie~ ami more
hea,"i l) muscled mib than those Ih ing in pooh of ~tanding \'O\ler. Some
,pecic . . have enlarged ~uction "cup~" on the belly or fumlcd by enlarged lip~.
:1110\\ ing them to cling to rock;; JIl !,trong currents. r.."O~1 tadpoles have no
a<.:tiw dere[1(.:e~ again'>! predator. . '>uch a~ li~h and aquatic jn~e(.'l~. but a few.
including the Janl.\c of ~e\'eral 8 0rnean frogs. ha\c poison glands thaI arc
known 10 make them toxic. al le:!'.\ to n.. hes.
!\lctamorphosis. or the change to adult body form. tal..c . . place after the
limn'> h:l\e fully de\elopcd. wh ich in a fe\\ Bornean ... pec.:h!s lTIa) require
onl~ a fe\\ \\ceks. B) th:lt lime. lungs haw a]<,o developed :lIld the tadpole is
read) [0 breathe air. The forelimbs pop through the ~k in em elope, the
internal gill~ degenerate anti the tail is slowl) ab~o rhcd, h i~ a strange
proces~, invoh ing many intern;lll'h::mgc~ and it is hazartlou~ for the tadpole,
froglet ~ince thi~ odd ereature b tt'mporarily not adept ;11 life l'ither ill the
water or on land, f Or!unate ly, llletamorpho<,i~ i~ a rapid prol'e~~, and i<;
u<,ually completed in u r.IU) or two.
Frog" depend o n \\:I1er. not JUS! for breeding . but al<;o for ba~ic ~uf\i\al.
The ..,"in of a frog is thin, not ~cal~. <'0 that \\:lter tend~ to ra<;, through quite
fred) in either direction. Bcc:lu"c of the ease \\ith \\ hich \\aler is lost
through their pernlcable skin. mo')! frog<; acti\'ciy a\'Oid dr) nc~ ... \\ hene\'er
po..sibk, FUTlhemlOre, when re~ting in \\ann, dry places buch a:. the forest
canopy). they adopt a tight crouching posture to reduce the amount of ~"in
exposed to the ai r. Interestingly. \\ hen frog~ \\ i~h to drink. they rarel) do "0
hy mouth. but ill'-ttcad absorb water directly through Ihe belly ~kin from a
pool or other ~ouree.
Frogs OCCUP) a wide \aricty of habitats. No :'JX·~·ie .. actually lives in the
M!i.I. although one frog (Fl'jl' n·(j/!£llwuTjl'Ora) found in Borneo can tolerate
~alinl' \\:lIer<" and one of the:.e \\a') actually found :.\\imming in the <;url" at
the \\'Itcr·" edge off Pulau Tiga. S:llJah, In reality. frog~ Ol"l"Ur in almo"-t all
terre:.trial and many aqualk en\ ironml'n l<' from the cold edg~' of the
pernwfro:.t in the far Ilonh to tips of the ~oulhcm cominems. A few :.pecie<,
li\e III de~cn~. more live in grasslnnd~, but the majority live in warm. humid
forc<,t" _SOn1l" .. peeies live on the ground (or under dead kaves). a fe". burro\~
in the "oil. ~o me p..:rch on shrolls or trees at varying height.; above the ground
whilr: other" live in o r b) bodie, of fresh water.
Thl? feet and hand~ uf mo~t '>pecic, provide cJue~ 10 their habit" and
habitah. Frog, that live in Iree~ anu ... hruhs im'ariabl) ha\c th~' tip" of Ihe
digit" enlargeu into distinct p.. d~ or di:'l" Ihal ;;ene as clinging dc\ ire", Man~
arboreal frog ... ha\c <,ome webbing bct\~Ct:n the fingers as ~\ell a~ bclwet:n the
toes. Aquat.ic.: or ground dwelling species rarely have di!:o,cs at the tips of the
fingers. Aquatic ami semi-aquatic frogs have extensive \l,Iebbing on the feet.
whereas those that burrow or live on the ground away from water usually
have little wehbing.
Regardless of the hahitat. all hlll a few types of frogs lay their egg, in
water. and the kind of si te used for Ihis purpose is peculiar to each specie!lo.
Some frogs Jay eggs in small rain pooh.. others in large ponds and !lotill others
in large streams or rivers. (In a later chapter we will describe the many
differenL Lypes of egg-laying siLes used by Bornean frogs.) Dependence on
water for reproduction rorcc~ man y species to move relati ve ly long distances
from their usual feeding and resting sites. Although in Borneo's \vell \\alered
forests the distance to n body of water is usuall y not very far, iJ1 drier
cllvironmcnls such as certain areas in the temperate zones. frog:, have been
known to migratc a kilometre or morc (0 tind thcir hrceding :,i te~. In rhe case
of Bornean s pecie~. reprod uction (:an in vo lve ve rtical migration as a Frog
moves from high in the canopy 10 lay eggs at a pool on the forest tloor.
For most species. breeding activity begins with males at breeding ",ites
calling. either singly or in groups. Each species has its own distinctive call.
Males usuall y have vocal sacs in the throat which they innate with air 10 form
cfTective resonating chamber". which greatly incrcase the vol ume of the
so und produced . An egg-bearing female move~ towards a calling male.
occasionally even bumping into him. The male climbs onto the female's back
and clasps her body with his forelimbs in a behaviour called ··amplexus··.
common to all frogs. If the pair is not in precisely the right spot. the female
will move the short distance necessary while carryin g the male. which in
most species is the smaller of the two. As the female expels the lipe egg:; into
the water, the male releases hi s spem1 and the eggs arc rertiU zed. After a day
or two. the tadpolcf!. hatch anulhcir aquatic life begins.
One other general feature of frug~ deserves to be mentioned: the facl that
1110st have only tin y teeth . and no claws. The Taiwanese frog.
H op/obmfaclllls rugll/oslIs. has sharp. hard toes which can inllicL minor
scratches. but would be ineffective against mOSt predators. Thus. they h(j\'c
rather limited defence against most predators such as "nakes. birds and small
mammals. Their besL defence LacLic is a suddcn. powcrfulleap . For Lhose thaL
cannot leap. camoullage b an allcm utive. The Bomean Homed Frog
(Megoplll)·s nnsl/ta). for example. closely resembles the dead leaves on
which it ~it~.
A number of the frog; of Borneo have skin secretions Lhat probably do
offer so me protection from predators. For example. the rufous-sided stick.-y
5
frog (Kalol,hrynlls pfellrtNiXlIla) produces grcal quantities of annoyingly

sticky mucus. which ~mall predalOr~ undoubtedly find ex(rcmcl~ unpleasant.
;\ "mall .. nake might di.-.co\cr thai it:. mouth has become glued .,hut after an
cncoumcr v.ith this specie.'>. The poi.'>OIlUU.'> rock frog (Raila hosii) produces
a ~lJn toxin that quickly kilb other frogs and can also cause serious eye
irriwtion in humans. We SU~pe(;1 thm a bird e'lting one of lhc<;c frogs might
never live to eat another olle. When disturbed. the giant river load (Bufo
jl/\"I(lIper) secretes large amounh of a foul-smelling. toxic white lalex-like
<;uh<;lance. A dog biting one of these !OOId... would be in serious !rouble. So. as
a .'>implc precaution. it is always wise to wa:.h hands thoroughly aft~r
handling ::In) frog.
6
/Frog s inJfuman 'Tradition
and Culture
e~pilc a divcr~ilY of about 150 specie,s in Bomeo. frogs do not play
D a partit'u!arly prominent role in the folklore and traditions of the

many human cultures of the island. The evidence for Ihi~ lies mainly
in the number of names given to a type of animal. Birds. for example, feature
prominently in Ihe culture of the Iban people of western Borneo. so Ihm a
wide variety of species have specitit'. regional names. Among the ('oasta!
peoples such as the Malays, B,~au and others, literally hundreds of different
names distinguish particular members of the large diversity of marine and
estuarine life. The purpose of sllch a system probably originated primarily 10
distinguish food from non-food. and dangerous from h'.llmless species. Since
frogs are neither con~picuous (like birds) nor a staple food item (such a~ li~h
or prawns). specific common Illlmes for frogs are few. Cultural information
about frogs tends to be limited 10 minor folk tales and to uses of rrog~ in a
few horne remedies (or magic spells) or as occasional additiom to the diet.
Thus. though the average person in the interior of Borneo would be fairly
knowledgeable about the natural hi~lOry of frogs. he or she will often pay
them linie attention. Thcre is an old belief from inland Sabah that a frog that
bitcs will not release its grip until a rumble of thunder. Despite the faci Ihat
the giant river frog. LimJlOlleCles leporillils. and some related species pos<;ess
fang-like projections on the lower jaw, few people in Borneo believe a frog
would ever bite in the first place. The "fangs". incidentally. are uscd in
counship battles, e.g .. only between members or the same species. As far a~
snapping at objecb. only prey-~i/ed ones (mmtly insects) are targets. and no
frog has yet bt'cn found 4ui1e large enough to snap up a man. There is. in fact.
a rather pointed. philosophical saying in Malay. "10 die from the nip of a
frog". which in context means thar a proud or important character can be
brought low by a small. insignificant person or event. The most common use
of the word "frog" in everyday life in Sahah often is in reference \0 any
unprincipled local politician who change<; hislher political affiliation for
persollal gain .
In Borneo. the projected "character" of a frog can. amongst other thjng~.
pep,onify anything from timidity to small-mindedness. Lack of resolve i~
7
FROGS IN HUMAN TRADITION AND CULTURE
Llemon~trated by (he wa) frogs nee. panic-stricken. from a ~n akc . A

pro\ incial pcr,on is de~lTibcd, appropriately. as "a frog under a coconut
~hcll", \\ hile someone who is out of his depth i!'. !'.aid to be lik" "a frog caught
by a droughf·. Those who arc overly oplimiqic are likened 10 frog~ who
"celebrate at the me,re hint of rain", and if they aspire (0 things Ihal far exceed
their ahililie~, they arc "!ike a frog who walUs to be (a~ big as a) bull". The
Engli.~h ~aying that a leopard cannOI (" hange his spots is in~lead rendered. a
toad will ne\er have smooth ~kin no mailer how much it rains". Finally. to
suffer in a manner that le:l\'cs no hope of suni\al is "to die like a frog".
Mall~ people- in Malaysi:m Borneo will remark that frogs can I.~a ll the
rain. and this notion i<; not without foundation. Though the frogs obviousi)
do not bring rain. many spedes. including the Bomean Ilorned Frog
(Meg()l'hry,~ na,wra) and bullfrogs (Ka/uulll spp.) certain I) sense the
decrca~e in barometric prc~~ure thtl1 pn.. l.·edcs a heavy rain stunn. In Danum
Valle). Sahah. a chorus of horned frogs frequently bur~t~ forth:t few mjnutc~
in advance of a thunderstorm. A B:lIlded Bullfrog (Kaloll"! plllchm) was
heard immediatcly prior to a stonn on Gaya Island. near KOla Kinabalu. The
hypnOlic influence of rain all frog:, in g~neral is used (0 describe pcrson~ who
are daydreaming or dazed. who an: said to ha\t~ the same blank ;"tare as "a
frog sining under a spray of water" .
Though frog/) arc rarely regarded as Ill} stcrious or hazardous, ,>ornc
ta boo~ (iJe rkellall) are as,>ociated with them. Pregnant women are not 10 look
upon certain crea ture~. for fear the lalter will influencc the traits of an unborn
<:hilt! . Similarly. a man who~c wife is pregnant should nOl kill certain
animak for the same reason. Thc~c animals. including frog;;, are as~umed to
have power to influence an unborn chi ld, ~o that the offspring of a man who.
during his Idfc's pregnancy kills a frog, may find that his newbom haby
looks like a frog.
In the folklore of [he Dusun people of the \'!ounl Kinabalu area, a star)
i~ told of 11 hunt!.':r and a giant frog. (The /)loT) wa:'. rel:ued by Mr Anso\\
GUIl~alam of Kinabalu Park.)
"MallY years agu. a man lil'illg ill a I'Wage IU'{I/' MOl/III

Ki/wb(l/II lI"en/ 0/1/ hl/ming. bur it won mined so hem'ily tllm he II 'a.1
fora(1 10 rakl'l·hefter bellealh (/11 o\"a/wngin!; mck, He SOOIl felt a
ehiff.lo h/' fit himself (I jire (I/ld began 10 (ook 1I III/'al. Suddenly
that' \\"tIS a rumblillg 100ll/d, followed by a lant!llide, (lml 11'11(,11 II('
lookelJ up he lI'as terrified 10 liml that the OI'erlrlllzl{inft boulder
III/tin \I'hid, he had rakel/ \he/u'I" lI'as aClUally (I llll,l{e liu,C., COI"('reti
8
FROGS IN HUMAN TRADITION AND CULTURE
Irilll /1/0.1.1'. Tile /lwn fled j(JI" hi.~ lifi'. huf 0/1 hi.\· Imy hOllle fwd Ihe
good fortlllle /0 capilire a .WJ/lllg monkey, and rhlls did 1101 rerum
empty-hal/ded.
Whell he refilmed. he found a //Iarriage ceiebralion \\"(1.\ l!"ell
IIllder\l·a.\'. \\"illl mallv people alreadl' ~!l"IIlIk all rice wille (tapai)
while rhey beat rhe gongs and danced rhe sWlla;::au. Allla;::ingly. to
the delight of (III, the monkey begall to dance arolllld as \\"ell. {II fUll.
the dl"l/Ilkell lIIell dressed him lip ill children '.~ clothes. pairillg him
Irith (I similarly dressed ('(11. aw! urging him to dance sOllie lIIore.
The noise of the gong.l· gre\\" louder (lnd {olldel; (llId el"erwme
lauglln! hilariously. The bride. hOll"evcl; was /JO! amused. alld well!
for lI"afer (/{ fhe Irel! nearbv. Wilen she looked ill 10 {(Ike I\ 'atel; she
lI'lH shocked to .filld a !?ial!! fm!? sitlillg ill.lide. and el'ell lIIore
stllnlled I\"hen he wid her {(} llIke her hllshand and )'eek to esc(lpe Ihe
place, for disaster ll'(H illllllil/nll.
Gilly momems after the bride alld groolll lOok 10 Iheir heels in
terror. a huge .flood appeared from /lOII'here lim! engulfed the
rel'ellers. a/JrllJ)lly silencing Ihe peals of Illughier lind the boomillg
of Ihe gong.l. Aftel1mrt!.l. whell .1'urI'im/:\" dug for tlJe bodies of the
rietillls. lheY/Olllld ollly deadfmg,·. TIllis. 10 rhis day rhe local people
I·temly 11'(11"11 their children //{'I'('/" to laugh at anilllals. for fear (~r
great misforTlllll'."
Knowledge that the skin of toads is highly {Oxic seems almost universal.
In fact. some old Malay charms for revenge do make use of toad skin in the
prescribed mixture. Nevertheless. although Muslim Malay~ and other~ never
eat frogs. toads and their larvae arc eaten by a number of the other traditional
peoples of Borneo. Kada/.an-Du~un in Sabah fry tadpoles (of any species) as
a goumlC( delight. while in Sarawak. the Bidayuh people cook them with
femlented durian pa~(e Ul'lIIpoyak) l:lS a delicacy. For adult toad~. however.
the Kadazan-Dusun. Iban. Bidayuh and others are alway.~ careful !o rcmove
the skin before dining on the meat. Some people also avoid eating the bones
of a toad. which are regarded as poisonous.
The species of frogs most sought-aftcr for the !able include the
Lilllmmecte.1 leporilllls group ("S·ai·· in Dusun. ··pamak" in Iban ). which
ineludes L Ingeri. L lIIales/anlls and L iballol'lI/lI. Mos! arc fore<a frogs
which grow to a length ofabolll 10- 15 em and have heavily mu~ded legs. In
coa"'al area~. the nab-eating ~'l angro\'e Frog. Fejer\'llrya ('(II/crimm. i.. the
frog of chuice. ami !ocrveu up in Chinese restaurants as "thl'cn kai·· or "padd)
9
FROGS IN HUMAN TRAD ITION AND CULTURE
chicke n". In SlIbah, an exot ic species culled ··bongkok Taiwan·· or Taiwanese

Frog (Hoplobmraclw .\· I"IIRlIloslIs ) was introduced. probably in the 19fiOs. It
ha.:; since become v. itlc-spread throughout disturbed habitats all over the State.
,lOti hll:. become:l popular source of frog'.:; legs for Ihe table. Since il grow>,
to nearl) three rimes the :.i.l.e of the mangrove frog. il!:> popularity has gro\~ n
to the extent that in the mid-19XOs in order to purchase the meat in some local
fill/Ill (market), a person would have to be on the .:;pot before u,e in the
moming. since the supply could easi ly be sold out by six. More recently in
large towns such ,\S Kuching and Kota Kinabalu. fanning of the American
Bulll'rog has been underway for a few ycar~ and has begun \0 ~upply most of
the local demand for frog.:;. Since the abmc o!.pccies is larger and meatier than
any Bumcan ~pecieo!. (cH'n L leporil1l1s), exploi tation of the latter. at least in
the major tOIl ns, may decline.
Frog':. leg:. llre now routinel) a\ aibble in coffee :.hops and restaurant~ in
Malaysian Borneo. The most common way frogs are prepared locally is to fry
them either in b:tllcr (Fig. -1-) or with ginger and tall gey (bean sprouts),
though people of the interior swear th:"11 boiling the frogs wi thout removing
the :.k in provides a meal fit for a king. Indeed, chicken comes OUI second be~ t
to the fine and delicate flu\our of frog meal for tho:.c of us willing (0 forget
the latter'~ humble origins.
Fig. ~. Plate of trog" ~ Icg ~ fried in batter.
10
c{)istribulion and Classification
of fProgs of C/3orneo
bout 150 species of frogs occur in Borneo. The exact number is
A uncertain because new species are discovered every year. Thiny-

seven years ago when the firs! comprehensive reviewal' the frogs of
Borneo was published there were only 92 species on the list. Thirteen
completely new species have been discovered since 1990. We ('<Ill Oll!y guc~s
how many more remain 10 be found. but the total number will undouhtedly
top 160.
TIlese frogs belong to six families: Bombinatoridae. Mcgophryidae.
Bufonidae (Ihe so-called "true toads"), Microhylidac, Ranidac (the so-called
"true frogs"). and Rhacophoridac. Except for (he Bombinatoridae. these
famjlie~ are widely distributed in Southeast Asi<l. There are no good common
names for these families: "(me frogs" and "true toads" only mean that the
few species found in England belong to these families. which is surely an odd
definition of "true".
The amphihian order to which the frog~ belong (the Anura) has exi!>ted
for at 1ca~t two hundred million year!>, a~ the oldest identifiable frog fossil
wa~ found in rock:. of that age. The mo!>t widely held opinion on the
evolution of frogs pl(Jces the families Ran idae and Rhacophoridae
(sometimes called the Orienlal tree frogs) close together. although they
appare ntl y became separate groups at least 75 million year~ ago. The
Microhylidae (often called Narrow-mouthed frogs) arc considered to be an
earlier offshoot of the ranid-rhacophorid line , The other three families
represent quite di!>linct and separate lines of evolution.
Each of the Bomean families is divided into groups of related species or
genera (plural for '·genus··). There are 32 genera. but the species are not
evenly divided among them. Four of the genera contain almoq ~O '(( of the
specie:. and ten of them ha\'e on ly a single Bomcan ~pecies, (See the
checklist at the end of this chapter.) Some of the gt'ncra containing a :.ingle
species probably represent peculiar evolutionary pathways- species that
have developed peculiar anatomical feature:. adapted for an unusual way uf
life, On the other hand. the genera with many species seem to he example,
of body plans tha! have been brollClly ~uccessful in a variety of habitats.
11
DISTR IB UTION Al'lD CLASS IPICATlON OF FROGS OF BORNEO
If we examine the anatomy and behaviour of adults and tadpoles in !>omc

of the larger genera. certain <;pecie!> !>ccm 10 form related group~. Within the
genu, Ummmec/l's. the 1""'0 guardian rrog<;. L poiomnensis and L finelli.
(.:on~tilUtc a di!l>tinctive species group characterized by reduced webbing and.
most remarkably. the habit of a male 10 guard thc eggs and then to tran!l>port
the tadpoles on his back to waleI'. In fact. these two frogs are so similar that
they were once thought to he race~ of a <;ingle species.
Tree frogs of the genus Rll(lcop'lOru~' fonn several group!> of relmed
!l>pcdell. The medium-sized to large species hav ing fully webbed finge r!.. ror
example. Wallace's Flying Frog (Rlwcophorus nigropuimuIIH). Reinwardt"!.
Flying Frog (R. reiml'llrdli). the Harlequin Tree Frog (R. [}art/alil). and the
Jade Tree Frog (R. dulilellsis). all have .. moOlh-edged flap:. of !.kin along the
outer edge of the forearms, all have the ability to parachute Of glide. and all
have large tadpoles that live in small pools of standing water. Three !l>mallcr
tree frogs. Rhaco{Jhorus (/Ilgufim.llris. H.. cyollopwU·/(l/U.I, and R. gallfli.
con~litutc another group. The!l>c small species have only partirtlly webbed
finger!> and have a pointed projection of .. kin near the hed. Their tadpoles
live in !>mall . dear. rock-y-bonomed !l>t.rcams.
Generall}.taxonomists arrange !.pcdc!l> into groups based on th ..~i r !lhared
chrtrrtcteristlcs. Once [hat is done. pattern!. of evolution and relat iomhips
become apparent. Those pauems. in tu m. may enahle us to predict the
behaviour and life-cycle~ or frogs we may have identi licd hut whose habit:.
wc know little about.
The geographic di!l>tribution of this large Bornean fauna occurs in clear
pattern ... TIle mos\ striking thi ng anout it is that it is different from any other
fauna in the general region. Almost two-th irds of the spede~ (60% or IJO
specie:.) have never been found outside of Borneo. Twenty-one of thc!l>c have
been seen only once. so it is no surpri:.c that they are nOI known from
anywhere ebe. BUI most of the Bomeo "endemic:," (as ~pecies confined to
one area are called) have been found many times- for e,(ample. the Dwarf
Litter Frog (Leprabrachella mjober!(i) and the Marbled Tree Toad
(Pedosri"t'.~ I"IIROSllS). Perhap) more collecting in adjacent land,-Peninsular
Malay~ia. SUmatr.l. or the Philippine ..-may rum up som..' of the Bomean
endemic~. but it is nOI likely 10 affect lhe general picture: that the Bomean
frog fauna i~ mostly unique to the hland. Thi!l> uniqueness implies that !l>Omc
groups have been isolaled from thei r related species in other pan!l> of
Southea ..t Asia for many milliom of years. despite the fact that geologists
hah! demonstrated that Borneo and the Malay Penin!.ula have probably been
connected for much of the pa:-'I ~5-50 million year:..
12
D I STR IBUTION AN D CL ASS IF ICAT ION O F FROGS Or- BORNEO
Ncverthek~~. Borneo's fauna does show links !O adjacent areas. Fony-

five Bornean spe(.:ies have been recorded in Penin~ular Maluysia. -1-2 'pecie~
also OI.:cur in Slllllatra. and 1-1- species are aL~o iound in the Philippine IsI:md:-.
Twent} -Il\'e specie~ are known from Thailand. and a few Homcan ~pecic,
o(.:cur throughout much of Southeast Asia. This situation is not surprising
... ince these connections are consequences of past geological histor).
geographic:11 proximity. ami similar vegelation and climate. 8 0mean snake~.
bird .... mammal.... and fre ... hwater lishes show the same kinds of relationship<;
to the faunas of adjacent lands.
Earlier in this chapter we noted that one family. the Bombinatoridae. i...
not wide ... pread in Southca'-l A... ia. In facL the genus to \\'hi(.:h the Hornean
<;pccies belongs . Barbollrufa. i... known from jll ... t two species, the Bornean
species (known from a single small area in Kalimantan) and nne from
Pala\\an and two small islands in the Philippines just nonh of Sabah. The
neare ... t family relative occurs in northern Vietnam and <>outhem China.
The . . t:llelllent that a parti(.:ular ~IXTil·.'> o(.:eurs in Borneo should nO! be
underslOod to imply that the species has been found throughom the island. In
fact. our knowledge about the diSlri bm ion of frogs in Kalimantan. the large~t
portion of Horneo. is very incomplete because so little collecting ha~ hcen
done there. :"olonetheless. enough work has been done to cnablL' u.'> tu see
~omc pattern". What appears to he true is that there arc no major differences
among larg(~ par\~ of Bomeo. :"'1 o~{ of {he ~pc(.:ie~ living in hilly tcrrain of the
10\\ land rain fore~t . . eem to UCl'ur throughout Borneo. For example. the Giant
River Frog (LiIlIllOllectes feporillllS) ha~ been found whercver the frog fauna
ha~ been sampled below 600 metres above sea level. except in peat swamp
forc.'> ts. The Rock Skipper (Swamis !mo/){/!lI1l11US) is known from c\'cry pan
of Borneo that has rocky streams with swift ClirrelllS. 1-I00\'e\"er. there arc a
few lowland species that seem to he restricted to (.:ertain regions. For
cXlllnple. the Rough Guardian Frog (/.iml!()lI('ctes ]iI/chi) has been found ~o
far on ly in Sabah. ue . . pite inlen.'>ivc ~alllpling throughout Sarawak. We
\uspect that it also o<:curs in the ea.stern pans of Kalimantan. The Rough-
backed River Frog (LiIllIlOllectes iballorlllll) has been found in many pl:lce~
in Kalimantan . Sarawak. and Hnmei. bm has yet to be discmereu in Sabah.
The frog species that live only in montane fore~t (higher than I(JOO
metre.s above sea len'l) haye limited regional di<;lrihution' in Borneo. "imply
because montane ell\ironments do not exi,t throughout the island. At one
tillll·. it \\ a~ thought that man~ of tht· . . pecie~ found originally on Mount
Kinabalu \\ere found nowhere cbe. but thm has been shown to be untrue a~
a result of rCe(~nl l'xpioraliull of high ele\utiolls on Mount Trus Madi amI
!3
DISTRIBUTION AN D CLASSI FICATION OF FROG S OF BORl"lEO
Mount Lumaku in Sahah, and 011 Mount Mulu in northern Sarawak.

Unfortunately lillie i!) yct known of the frogs living above 1000 metres in
montane parts of Kalimantan. TIlere is certainly potential for many new
discoveries in this vast southern and eastern part of Bomeo.
FiA. 5. Recently emerged frog let of Reinwardt"s Flying Frog (RhacophofllJ

rcillwardTi ).
14
KEY TO THE FROGS OF BO RNEO
Key to the Frogs of Borneo

I. Conspicuous webbing on hand. at least between two outer lingers ........ 2
No webbing nc.twcen outer lingers .. ............. .. 17
2. Webbing bem een linger;, thid.. lleshy ............. . ................ Pefvphr\"l1e l

Webbing between lingers thin. membranous ............ 3
3. Colour black with irregular. small light spotS or lines. ............. .

.............. ................. .. Staurois Imopa/lllallI S
80dy not black ........ ................. ................................ -J
-1-. Top of head and ~ides extremely rough and wany .. .... Thdol/erllla horridum
Not a~ above ................ .... 5
5. Body green .................................................. . ...... 6

Body not green ......... .... . ................ . ......................... ...... . S
6. Adults less than 25 mm. snout pointed and webbing limited to base of
fingers ...................................................... . ....... Rhacophorus /...ajau
Adults more than 25 mm .................... . ......................... 7
7. Sides black. with blue and golden yellow spots. . RIIlIt'ophoruJ H';lIIl't/rdli
Sides yellowish or greenish ..................... Rhacophorus lIiglVpalmoIus
8. Webbing red ... .. .. ... ...... 9

Wehbing not red 10
9. Forearm with flap of skin along ouler edge . . RhacoplwTUS partillliJ

Forearm with no skin nap ... .. ....... . ...... Rhacophom.\· ruftpes
10. Heel with conspicuous pointed projeclion of skin .... Rhacnphorus ha/uellJiJ
No such projection ...... . ................................ 11
11. Outer edge of lower leg wilh row of poimed projections or frill of skin .... 12
Outcr edge of lower leg olherwi~e .. .. ... . ... . 13
I) Pl'lophnnt' arc 'Illa!llo~d~ (u>ually k s~ Ihan 25 nun). E,,:n ~o·(;alled expcns flJld il dilTiculi
10 <;cparate Ihe sl~ci<,s.
15
KEY TO T HE FROGS OF BORNEO
12. Top of head and hack \\ ith spiky proje(·tion~ ................ Rh{/copiIOUIJ ('\'erelli
Top of head without spiky projection\ ............. Rlwcophorw appt'lu/iclI/mus
13. Pak blue spot.~ on \ide~.

groin and inner thigh .
...... Rhacop/lO/"lIS C,wlOpwlCwms
Sides and thigh without blue spots ..... ................................ 14
14. Small (less th.m 40 tmn) frog with ~horl. blunt snout. and a white spot under
the eye ....... ...... ............................. ............. ..... Rlwcophorus gmllli
Notasabove. ................. ....... ........... .................................... 15
15. Sidc-~ yellowish with black spots. . ................ N/wcophol"ll.\' allgu/ilV.lfriJ

Not a~ abow ............................. 16
16. Snout \harply pointed. with conspiellous sharp ridge between eye and nmlril
.. Rhacophorus /laniHDlli
Snout blunt. v, ithout sharp ridge. .......... RhacopllOrusjasciatl/s
17. A large con\picuou~ "v, art" or gland (parolOid gland) behind eye in shoulder
area. as large a~ or larger than the eye IR
No large gland behind eye. ................... ...................................... 2-1-
18. Top o f head between e)e~ with a pair of narrow. raised ridges. 19
Without ~uch ridges ................ . ....... 20
19. Parotoid gland n:lrro\\ and elongated: colour often orange.

... .. ... .......... . ......................................... ................ 8/10 quadriporcaTlis
Parotoid gland oval or triangular ...... " .......... BI!fo dil'el:r:ells
20. Tips of fingcr\ natlencd and squari;.h in outline ....................................... 21

Tips or fingers rounded or tapered ....... .................................. .. .. 22
21. Parotoid gland rounded: hody green \\ ith brov. n or reddish spots.
.............................. .......................................... Peilo)/ibes rugOH15
Parotoid gland elonga\l:d. narro\\; bod~ brown. or black with irregular
ycllow line, ................................................................. Pedmlilics hnsii
22. A bollY ridge curving around eye to parotoid gland: \\arts usuall~ with 51T1all
hlack dOIS ..... .. 811(i) lI1e/anosliNIII
WithoUl >.uch a ridge. ...................... ..... 23
16
KEY TO TilE FROGS OF BORI\EO
2_' . P~ru(0iJ gland rount! or slightly (wal. ahout the size of tbe "'ye ... 81tfil (lSI'('/"
ParolOid gland ",Iongal('d. mllch Ion ger than the eye .............. BII'!) jllrfl/lPCI"
2-1.. Conspicllous projection of s kin from 'iurfan; of upper eyelid .... ...... ... _.... 25
~o ~uch proJ(,L"tion .
25. Skin of back wi th one Of more nal"l·ow. elongme ridgcs . .___ 26

SI-...in without long ridg('s. but with many "piky prujection~ .
................. . Hego/J/II"y5 nh'-({I"dillaf
26. Projection on C) did an m 'el'hanging triangular nap of ~kin . ... 21

Project ion on eydid a short. narrow cone ........ .... ..... Mcgol'luT.I holuoD;s
..... ........ .... .... i\-Iegol'hrn drill:':;"'-
27 . Conical proJ('ctioll of , kin from tipof snout .. M(,~i!l'hIT.\· 1l1lSIIIil

No projection of \kin frolll tip of ~no Lit . MegopirlTs kobdl"llSirii
28. Eardrum \-i,ihil: (rig. 21. .... 39

L:ardrum not \'j"ibk . 29
2Y. Lnder~ide oj foot with ('nlarged. ra ised spade- li ke proj('ction .

Lndersid(, of foot not as aboye . ._.i 2
.1U. Tip~ offing(,I'~ ('xp~mdcd into hlunt pm_h. ...... . ............ . ~I

Tips of tillf'(,fS not expanded . CuI/wi/a'
J I . To(', wchhed . _ ..... .... ..... Koloula hrile(l(o

To('s \\ithout wehhing .. ..... .. ....... Kalollia 1lllIch/"o
32. Snout long and pointed. twice a, long a~ Ihe di,ullL'ler of the eye. pllinted .
... ... .. .. .. Gu.111"O/,l!rYlloides /;O/"ll{'CIII;1
Snout nOI ~o long. ......... 33
~ 1,1I<!~(!JI"n' b,li1<dl \j'" '" fJ.,- l;:no" n ,)lJI ~ from ~1('ull1 KinJhalu and ha< <.t1<' <1 k~' <lIllIlar
«I 11. " Il."'/II . '\I<'~"I, I,,-\, dri"gi 11 " , " , far 1>"::"::11 fnunJ '-"l t~ on \l nunt ~I u lu "'lei "3 t"II~ -I~g~~d
'I'.:"~'.
3 1 Sp(~":JC< Ll J Cdlll,'111' .Ir~ hurn'''~'' .n,d '''rd _
' t:II~(lLJnl~r~d Di'"m:liull> .lIllLln~ I h~"l arc'
Ji<<."ll\ .,~d III Ihl' [nt.
17
KE Y T O THE FROG S OF BORN EO
33. First finger either not visible or much shoner than second finger .............. ~4
Firs! finger about as long as second ................................................ 36
3-l. Fir~! finger missing .............. ...................................... Micruhy/a petrigt'llu

............. .............. . ........................... MicrohYia pefpan-a~
First linger distinctly sticking out from palm ............................ 35
35. Third and fifth toes webbed to Ihc lips .................. . ... MiclVhyla bertimon'i
Third and linh toc.~ not webbed to the tips ................... MiclVhyla borm't'llsis
36. Front tip of lower jaw (inside) with a ~ingle point Of cusp ..... 37 5
Frolll tip of lower jaw with several CliSPS ....... .................................•.... . ..•. 3l)
37. Chc~t and abdomen whitish, without ~pots ....................... Occido::yga luel'is
Chest and abdomen with lllany dark SP{)I~ ....... .......... OccitiO::YSll bull/ellSis
3R. All toes wehbed to tips ...................... ...... ... ..... ... .. ... ... Lilllll()/leetes kuhlii
Toes not fully webbed .................................... .. Lill1ll0/leCres latieeps
39. Fourth or outer tinger less than half (he length of (he first finger .............. 40
Foufth finger almost as long or longer than the lirst finger ........................ -l2
40 Spots present on side of body in from of hind leg ... 4.1

No spots ill fronl of the hind leg ...................................... Ka/ophrYllli s Spp.6
4 1. Spots blui~h or whitish ............................ ........... KlI/ophrYllus lieterocliifll.\·

Spot(S) black. ... .................................... ....... Ka/oplirYlllo p/ellfusriKlI/li
-l2. Eardrum sunken, within a cavity......................... ..... Huia C{lrirYlllpUllIIlII

Eardrum not as alxl\"e ..... ............................. .. .. .. ... .. ........ ........ 43
-l3. Back cinnamon-coloured or red. speckled II ith II hite; a white spot on the
upper c)did ................ ................................................... Nycrix(l/uJ pictus
Colollration not <IS alxl\'C' .................. . ............................... .. ............ ........... 4-l
4) These tun ~P<'L'i~, an: ran:l~ ~een and can be dlslingui"hd from ~ach mher on l> Wilh Ihe aid
of" micru,c0P<'.
5) To see Ihis feawre It j, necessary 10 gently pry Ihe mouth of the rrog op.:n .
6) S~\eTal ,mall 'P<','ies of this genus are eXTITmety difficult to ,~parale. S",e d~'L'nplion, in
le..\1.
18
KEY TO T HE FROGS OF BOR.!"lEO
-+..j.. Tpp of lirst two t\JL:~ while or hlu.:." . -+.i

Toe" nOl as ahove . ..... -+6
-+5 . Top or head and body w;111 dark to black ~pot~ on a greenish hackground .
... ... . ... .. . . .. S/I/llra is IUI/OIUI"
Tpp of head and body dark green to hlackish I\ith a few ~maillight SpOl\ .
.. .. ... ....... .... .... . ... ...... ........ ... Swumis /II/wrilingl/i.\
-+6. l Tndersidc of fingers with enlarged pads. broader than the fmge!" tips: tn:e
ho le rrog . ........... .................. ... ...... Mnaphrl"uclfn ~1If1dllllU
Fingers without such pad~ . ... -17
Hcll~ golden yellow wilh hlack nct\\ork. ..... . C/wpcrinu .li/I /YI
Bell) not as abo,'e ... ... ... .. .. ....... .. .. . ...... . .. .. .. .. .. .. -+8
-+8 . Back dvx:olate bro wn ollnkred by ;1 thin pall' blue or II hili~h linc. sides
grey 10 black . ....... . ...... .. ............... . Rlllw /1I("tllo.\(/
Coloura!ion otherwi ."c .... .. .. . -+9
-+9 . Bod} gr..Tn Ilith a light Yi:'llowi sh stripe do\\"n each ~ Ide of the back .
. ... .. Nann t'I"Ytlirac {I
Co]oumtion otherwise .. ....... .. ........ .. .......... .... ..... ... 50
50. Body dark bro\\'n or black. will1 ycllowi~h (lr n."ddish spots, al\\"ay~ II ith
lighl edge to upper eyelid . ... .. 5 1

Cnlouratioll (ltherwi~e .. 52
51. A continuous yelll)\>, ish Ilr orange stripe on each side of back from eye 10
end or body .. . ... .... ... . .. . ....... Rww !J ig!1at{l
YdIO\\ or orange SpOb on back newr forming continuous stripe .
... ... . .... ROll a pic I/Imtu
5 2. Bod) green , usually \\'ithout "POl',: upp<."1" lip wl1il~·.

Body \l~1l3 1 1y Ilot green: if gr<."c11, Ii p 1101 \\ hile .
53 . F0unh foe (the Inn)!<.",!) with 'It ka,t tllO joinb J"rl' l' o f \\ch: unJers ide of
thigh u"iu:.lIl y reddi,h .. ... .... ... .. . ... ROl1u ("huklll/o ta
f ' llIrth to.: \\ ilh \Icb rl';!Ching base of expamkd lIP: lJnder ~ ;d<.': of thigh lK\l'r
redebh . .... Rww I' llsii
19
KEY TO T HE r-:ROGS Or-: BORt'IEO
54, Fourth toe (the longe;.t) fllll) \\ebbed 10 base of tip ....................... 55
I-ourth tnc \\ ith at least t\\O joints free of full web ..................................... 51)
55. A ~cric, of ~hort ridges down the back: throat always with dark pattern
.......................... .......................................................................... ~
Back \\ ithollt many ~hort ridges: tllront may haYe dark pattern ................ 57
56. Throat \Iith dark pattern or streak down centre: back always with black Spob
..... .... ,. . .. Hoplobmmclw s ruguloslIs
Throat without dark central ~treak: back uwally without visible spotting .
... UlllllOll('("/n ihallOrJIm
57. Entire upper ~urface covered wi th 10\\. rounded wart;. .

............ ................................. .......................... PJl!lId()hl~f() .HI/JaSPI!r
Uppcr ;.urface with ,ome tuh.::rei.::s and ridge~. hut not compictely rough .
......................... . ............................................................ ~
58. A dark. ~trip.:: oct\\ecn eye and nostril on ~idc of ;.nollt: tOp and sid..: of~nout
form a sharp angle ........................................ Lilllllollecre.\· lepo/"illll.\·
No dnrk stripe on side of snout: a rounded surface formed by lOp nnd side of
snout, ..................... Lill/nunn'I('s inJ;t'ri
59. Toes not wehhed ................ . 60

To.::., at least partially wcbbnJ .................. . ........................................ fi5
60. Tip~ of lingers and toes distinctl~ pointcd .. .. 61

Tips of fingers and toes rounded or blunt .. 62
61. I lead plu~ body less thim 2.5 em . ....................... Leprahmchclla !>pp J
Head plus bod) more than 2.5 CIll .............................LeprohmchillllllliXmps
62. Ikad not \Iider than body ......................................... Lepra/a{a.l spp.s

I-kad wider than body .. .................................................... 63
63. Belly with dnrk network .......................................... Ltp/ubrachillm abbolti

Belly not so marked ................................................................................... 64
7) In.: ~mall 'P<'ci.:~ in thi, g.:nu, art' uiJli,'uh to '.:parat~ [rum on.: JnOlh~[ "ithout J
mlcro,copt". 5.:.: ut',cnptlons 1Il1<:.~t .
S) Ta.~{)nol11ish :If(· ~tilltr)ing to ,on out the ,pt"(ic~ of thi, fcnu., anu III the procc" IlC\\
~pccic, an.· t-cing ui'ico'cr~d 't("auily. Sec (k~cripllon' IIltcxt.
20
KI:.Y TU THE FROGS or RORNI:.O
6~ . Bell~ light ill colour. 'pco.:I-.kd Ilith tine bbd, doh .

. -...... . Lt'{lfOhmrl/lwlI Ilo1</rid. I, 'III
R(' II ~ gr~':: found abolc 1000 m :litltLld.;'_ .... Lt'pro/1nlc/rilllll 11/"11/(11/11111
6:' . .--\n II1\lT[ed \·-,I].lp~·d ridgc on hud, b.;'ll\.;'enllw ~ll\Iuldcl" . 6(,

~(l ,ud] ridg~' .. __ ...... _ A7
.... LI!Ilnfll/(Y/t" rlilm 'UIlt'Il.,is

i\o contlilUOll' I·old. bad. rough with 'icaHered round or (),a l l'ldgc~"
. 1.11II1I01I<'Un lililili
67. Tips oJ'toe, II idellc'd in[u di'lilk'l, nlllnded p:tds ordi'l', 77

Tip~ oj' toe ~ not II ilk-ncd inti) di,tinct pa(\, . . (,~
68. ,--\ .,mall. bla,'I-. X in o.:l'ntrc of b'ld. ...... Lt'/'loplr'TJlf /J(!/'h"I/1((/

1\0 'Ilch Illarklll!,' 6 t)
(,l). Head and hod~ dark broIl n or hJ.td.,: a dio;rllKI. pale or:mgc or ydl()\\I,h
marking hdll.;'.;'n ~houhkr' . .. .. • I!/Ionia '{'/llit/i/(T
ColOllratioll ()tht'!\1 i~e . 71)
70. Entire lIppcr ,urfac'c ofhnd) mu!"h and 11.ln\ .. 7 1

L pper ~urral·o.: \\ith ridgc, Ilr:l {.;'II !'(lund pmicl·ti(l)1,. (11 \mooth ............. 7-1
71 Tip, l>f finger' I\ider thall i;1,tioint .... , III ~Ol lill /tllllil'l /tl
..... " IllI'ilillll {1//l(1 IOI/!,I
llll (!l1/ll /luidiKd'

Tip, (>ffinger, 1101 II ider th:m ]a,[ joint _ 7"Y.
,\ II hltc Ill' lT~all1·cololired 'lrcah. rrul1l c~c to armpit .

. .--1l1villia al/JOlllti, Itlliftl
73 lhll'd ~,nJ fifth t\IL'~ \Ichhcd to till> .... ..11/10111<11:111"1'1

Third und lii'th lO~" Itot 11,'bbed [,> liP' I/lII)llill 'pp. l
,/, PI'UIl~UI,hil1~ ~TIl"n); lh~,,' thr~,' " .( I", k fM l.""llvllll,t, Th~ r,r,t 11<,) .I" IH" l!u:Uf b<-h)\\
')I~IIll"lr~' "Ix"',' ' L'" k,tl
[°1 hlUr 'I'<" I~' ,>1 -\IIWn", th,,] .,n, .l,lli'lIll tv '~p,Ir.(r ,' ,h ,(,.~ II", ,'h"r,"'ltn'l1, <;(~
.t(·'~npll "Jh In t~11
~ I
KEY TO THE FROGS OF BORNEO
74. A series of short ridges on the bad. ... 75

A few rounded rough area" or back smooth .............................. 76
75. Three joinb oj" the fourth toe (the longest) with no \\ ebbing. frequently with
a light stripe down the back ..... FtJcll'illya limllochnris
Only one or two joints of fourth tne \\-ithout webbing, no stripe
.. .. ........... FtJell'arya ("(//I{'ril'ora
76. Usually a dark ~treak from eye (0 nostri l on ~ide of snout; u~ually entire
eardrum t:u\en::d by large dark marking: head narrow ..... .. ... .... .. ....... .. .. .
.......... .. .. .. . .. ... ..... ......... . .................. UlIlIlOlleCres (Jaramacrodoll
1\'0 dark ~tn::ak from eye to nostril: usually ahout half of eardrum t:ovcred by
dark pigment: head wide .... ................ LilllllOllecres malcsimllB'
77. Head narrow. ~n()ut long, distance between eye and no~tril longer than eye
diamc1cr ........ ................... ... Raila lIirohariellsis
Head wider. .<.nout ~horter. diameter of eye longer than distance octwcen eye
and nostril , .. .. .................... ..... ..................................................... 78
78, A conspieu{)u~ ~a\\ -edged bony ridge above the enrdrtllll .

................ .. PO(lpedares mi/op/lIIs
No ;.uch ridge .. ......... .............. .............. ....... .. 79
79. Eardrum covered b) dark bnnd extending along shoulder to armpit ,

....... ...... ......................................... .......................... Pot.\pcdare~ lI1acmris
Pallcm olben. ise ... ... ..... ... ......... ....... .......... . ........... .... .. 80
80. Outer t(k: \\ ebbed to ba;.c of pnd 81

Outer toe not II ebbed to pad .. ... R3
81. Third toe webbed to base or pad on bOlh ~ides ............... . ............... 82
Third toe not 'Webbed tu pad on both sides . .................. . , Phi/millis hosii
... .. ..... Phihmllo il/gerill
82.. Baek and sides bright green with dark ~p()h ...... JH£risrngelly.l· killahallll'llsi.1
Hack and sides brO\~ n or blaek , ....... ".... , Meri~logel1y.~ Spp.12
II ) Phi!twl", lum' i, confinl."d 10 ek\atiu", beluVo 750 mt'lf':~ amI P. ingai to ",!e\ation< aho\"~
lUOO meiTt"
12) Th.: f.:rnrllnda of the <p~cie, in thi~ !!cnu, af~ dlffinilt 10 'eparal .... ':'t'n fur l:nunumi<t,
Cun,u!t dnniplinn<; in t"x!-
22
KEY TO THE FROGS OF BORNEO
lB. Tips of finger~ not much widcr than next joint ... .... ........... 84
Tips of lingers abouttwiec width (lfncxtjoim . ........ .... . ····· · ........... ... . 86
8 ~ . Sidc~ with enlarged ~kin gland~. producing: a rough or pebbled ~urt'acc

............... ........ . .. ........... . Raila g/andll/oul
Skin of ,ide not as above .................... ... ... ........... 85
85. Upper lip with a whitish line cO!Jlinu()us front below eye 10 below rear of
eardrum. ........................ . . Rmw /merima CI//ata
Upper lip barred with black and white .... Rww haramica
'6. Pad of longest linger aboUl same width as eardrum ...... .. .... Philalllll S spp. 1.\
Pad of longest tinger much nnrrower thnll eardrum ............... .. .. ......... 87
87. Legs rather short: skin rough ................. . ....... . lngl'l"(//w haflll'luiJ
Legs long: skin smooth ......... ....... ......... ... . .......... 88
88. Snout sharply pointed .............. Pulypedmes cof/c Hi

Snout nm poimed . ... Pofypn/all'J 1l'lIco/lI.nHn
1J) The~" ~malJ 'ipecic< lusua lj~ ks< Ih:m 25 !Om) po,.: ,.:riOllS pfOblcm~ c:'\en lor
h"rp<'lo!ogi'I~. s,," d('<cripti,)n~ in lnl.
A CHECKLIST OF ANU RAN AMP HIBI ANS FROM BORNEO
A Checkli st of Anuran Amphibians

cUlTc ntl y known from Borneo
Family Rombinatoridae
Harbollrula kalim(/I/[(InelHis I~kandar *
Family Megophr:\:idllc
Lepwhmchella hall/ellsis Smith'"
L brel·i(TII.I Dring '"
L. I/Ijobogi Smith '"
L. pallllata Inger & Stuebing*
L. pan'a Dring'"
L. .l emMmae Dring
Leproh/'{/cliilllll ab/mlli Cochran '"
L gummgensis MalkTllllS*
L hendrickson; Taylor
L. II/OI/WI/IIIII Fischer*'
L niRrops Berry & Hendrickson
LeplOlalax araWli Matsui *-
L tlringi Dubois'"
L gracilis (Glinther)
L hamidi Matsui '"
r.. mallr£l Inger, MaJ..larin, Alirn & YamDun *
L. piau,l' Malkmus'"
Megopflrys baluensis (Bou!engcr) *
M. dringi Inger. Stuebing & Tan '"
M. i'dllw'dinae Inger*
M. kobamshii Malkmus & l'<latsui *
M. IW.I'II/() (Schlegel )
Family Hufonidae
AI/sollia (I/hOl/well/ata Inger'"
A. ll/lotis Inger, Tan & Yambun '"
A. jilligillea (~·ltX'quard ) *
A. fJ llihei Inger*
A. IWllit.l'chi Inger'"
A. /witlil'ca Inger'"
14
A CHECKLIST OF ANURAN AMPH IBIANS FRO M BOR.'lEO
A. leproplIs (Gunther)
A. IOllgidi:;:ira Inger*
A. mimI/a Inger"
A plCltYWJIIIlI Inger*
A. spilllll//er (Motquard)*
A. lorrell/is Dring '"
Rllfn {I.I'per Gravenhorst
B. din'rgell s Peter"*
H. juxrasper Inger
B. melt/no.niclUs S('hneider
R. qll(ldripo/"callls Boulenger
LeptophrYI/(' /mrhollicli (Tschudi)
Pedosribes ('I'erelf; (Bou1cngerl *
f'. !/Osii (8 oulenger)
p 1I/(I("/dmll.1 (Mocquardl*
P mgo.H/I· Inger'"
Pefophrylle api Dring*
P. gllcm"eri (Buulenger)*
f'. lIIisera (Mocquard)*
f'. rlw{Jophilills Inger & Stuebing*
P sigl!ma (Boulenger)
Ps('udo/Jufo .I"/Ihwper T~chlldi
Famil:r Mkmh:\' lidae

Callilella brooks; (Bouknger)*
C. pam Kiew *
C. .I/Ililhi (Barbour & Noble) '"
C/Ulperill{l jil.l{"{/ Mocqllard
CasrlVphrYl!oides homel'II.lis ( Boulenger)*
KalophrYlIlI s bO!l/ellsil' Kie\v *
K. ('o~ Da" & Haa~ *
K. hetemchirus Boulengcr*
K. intermedius Inger'"
K. IIl1hico!{/ Dring *
K. p/elllrJl/iglllll (Tschudi )
K . PIII/Cf(/WS Pl;ters
K. sllbr('rresrri ~ Inger*
Kalol/fa balf'{lf(/ (l'.1Ulkr)
25
A CHECK LIST OF ANURAN AM PHIBIANS FROM BORNEO
K. pll/chra Gray
Metu{JIIIYllellu slim/ana (Peters)
Microhy/a hl!!dlllorei (Blyth)
M. bomeemis Parker
M. lIIacl/lifera lnger*
M. pelf){l/"l'a Inger & Frogner*
M. petrigl!llll Inger & Frogner*
Family Ranidae
Fejerl'arya cal/crimra (Gnlvenhorst)
F Iill1l1ocharis (Boie)
Hop/oba/raclllls mgulosus (\Viegmann)
Huill (·m·irYIl1f1l11IUr/l (Boulenger)*
Ingerana baillensis (Bou1cngcr)*
Liflll/ol/ccres asperata (Inger. Boemli & Taufik)*
L. ./iI/chi (lnger)*
L. i/)al1omm (lnger)*
L. ingeri ( Kiew)*
L. keneplIien.\·i.\· (lnger)*
L. kuhlii (Tschudi)
L. laticcps (Bou1cngcr)
L. leporillils (Ander~son)*
L. maiesialllls (Kie w)
L pa/al'allensis (Boulenger)
L. pa/"olI1acrodol1 (Inger)
L. rhllcodus (Inger. Boeadi & Taufik)*
MerisfogellY.I· mnomp{I/amlls (Matsui)*
M. jerbo(l (Gunther)*
M. killabaluclIsis (lnger)*
M. maclVphthalmlis (Malsui)*
M . orphnocnemis (Matsui)*
M. pliacomerus (lnger & Grilis) *
M. poeciitu (Inger & Gritis)*
M. Irhitehelldi (Bou1enger)*
0("ci(/0:Y8{1 ha/llellsis (Bou1cngcr)
O. laeris (Gunther)
Raila baramica Boettger
R. cllOlconota (Schlegel)
21,
A CHEC KLI ST OF ANU RAN AM PHIBIANS FROM BORNEO
N. erythmea (S<"hlegel)
H.. g/ulldlliom Boulenger
H.. hosii Bou!enger
H.. larerinll/culata Barbour & Noble
R. luell/om (Peters)
R. lIicobariellsis (Sto!iczka)
R. pictllmla Boulenger
R. sigllala (GUnther)
Staurois laropalll1aTus (Boulenger)*
S. lIataror (Giinther)
S. fIIherilillglli.~ Boulengcr*
Fami ly Rhacophoridac
Nycfixalus pictus (Peters)
Phifallflls aCullI.l· Dring*
p. all1oenu.v Smith *
I'. all/wltilll" Inger*
P l)lInifllS Inger. Stuebing & Tan*
P disgreglls Inger*
P. el}'fhrophfhalmlls Stuehing & Wung*
P hosii (Bou!enger)*
P. infieri Dring*
I~ kerangae Dring*
P IOllgicrus (Boulenger)
P mjobergi Smith'"
P peters; (Boulenger)
P refugii Inger & Stgebing*
P satleri Malkmus"&' Ricdc*
P. lee/II.1" Dring*
p. IIl1lhm Dring *
Polypedates col/eui (Boulenger)
P lellcomysrax (Gravenhorst)
P macrO/is (Boulenger)
P O/iloplllls (Bou!enger)
RhacopJlOl"IIs ollglllim.l'tri.I' Ah!*
R. appendiculutll.l· (GUnther)
R. balllell.l'i.l· Inger*
R. cymwplln('fOl/lS Manthey & Steiof
27
A CHECKLIST OF ANURAN AMPHIBIANS FROM BORNEO
R. dlllilell.li.\· Houlengcr
R. ('l'ercfli Boulenger
R. jasci(ltlls Boulenger*
R. gmilli (lnger)*
R. /wrrissolli Inger & Haile*
R. k{ljwl Dring*
R. lIigmpalllltlllls Boulenger
R. pard(lli.l' (iUnther
R. rei/llf(/rtiri (Schlegel)
R. ru(ipes lnger ~
Thel(Jderlllo hOITh/1I11l (Bou1enger)
Spe('ie~ endemic to the island of Borneo.

ecolog y oj C
{Yorn ean /}I"og 8
here an.' two major groups of frogs in Borneo based on their general
T habits and habitab. The ~mal1er of the two groups consists of species
closely associated with human economic activity. frogs that live in
padi fields or other types of cultivation. kampungs. towns or CilY habitats
sllch as drainage ditches. or vegetation around buildings. Nine ~pccics fall
inlo this category: the Taiwanese Frog (Hoplobatrachlls rugu!oslIs). Green
Paddy Frog (R(lI1a eryrhraen). Mangrove Frog (Fejerl'llrya callcrimm).
Grass Frog (Fejen'{//ya fill1llocharis). Cricket Frog (Rww lIic()hariellsi.~).
Black-spotted Toad (Rufn lIIefwlOSliclt/s), Four-Lined Tree Frog (Polypedares
leucomys/ax). Banded Bullfrog (Kli/ou/a pulchra) and Brown Bullfrog
(Ka/oll/a ha/caw). Not only do the~('" species live close to humans: they even
depend on us to create favourable environmental conditions such as puddles.
pombo \f"ig. 6) and ditches within large open. disturbed areas. Only Ka/ou/a
ha/eata. Halla lIicobaricnsi.l· and Polypedares lelicolllysfW: can be found in
~econdary growth or ncar primary forest. All nine spe(:ies breed in standing
water: drains. ornamental pools. temporary rain-filled depressions and
flooded rice fields. Only one species. P(J/ypecialcs fCIIC(JfII.l"S/{IX. is a real
climber which perches on tall grasses ami scrubby vegetation. though Rww
eryrhmea males are known to perch in 10\\ bu~he~ in some instances. Two
species are burrowers-the sticky bullfrogs. Kalol/fa Jlllfchra and Kafou/(I
ba/eaTa: the fonm:r i~ frequemly found among the dewJ leavc~ dogging a
drain which is being cleaned. RlIIW erylliraea and Fejerrarya callcril·ora
rarely 1110\·e more than a i"ew metres from water. whereas Fejen·mya
Iimnoc/wri.1 and Halla lIicobariellsis arc great travellers. moving through
damp grass and shrubbery often at some distance from water.
Many. or perhaps most. of these ··town·· or kampung' · species arc quite
resilielll and tough. They can withstand fairly high temperatures and some
dryness. They probably arrived in Borneo a~ stowllwllyS. inadvel1ently
brought in by humuntravcllcrs in cargo such as lumber. agricultural products .
crates. etc. Both Ka/ollfa pu/chm and HOjJfohall"tlc hlls /"IIRII/a.llIs appeared
relati\'ely recently in Sabah. and pos~ib!y was heard for the first time in
Sarawak ( Kuching ) in early 2CHJ..t. Bllfo IIIC/(lII()S{ict/I.l· i;; common in Sara\\ ak.
bUi so far has been found only in one kampung (\ illage) in Sabah (near
Kampung Kibunu!. on {he west coast). indicating that it i~ ~Ii!! ~lowly
29
ECOLOGY OF BORNEAN FROGS
~pn:auing. Hoploblllmchll.1 1"II!':lIloslIs (""Bongkok Taiwan'") wa~ introoun:u

in Sabah a~ a "ouree of food . perhaps as early as the 1960s (See p. 7).
The great majorit) of BOOlean frogs are confined either to forests or to
forest edge~, One or two ~pccie~. ~llch 3S the Saffron-bellied Frog
(ClII/perilla Iasea), occasionally make a go of it in ~hady garrJl'n~. Generally.
however. Illost frogs in Borneo are forest dwellers, and take up one of the
following life styles:
Fig. 6, Pond with frop.
1) Lifetime stream bank sitter: Some frogs never leave the banks of
streams. Their tadpoles always develop in those streams (Fig. 7) or rivers.
and when the newly transformed frog lets emerge from the water. they stay in
the vicinity where they feed and grow-if they can avoid predators such :I.:'.
other frogs. snakes or liwrds. Frogs of this type include the GianI River Frog
(Liml/ol/cai'S lcporilllll"). Kuhrs Cr~·cJ.. Frog (l imllol/cC1es kllhlii). and the
Black -~potled Rock Frog (Sraurois IImawr). Frog~ of wetland area~ ~hould
also be included. one~ ~uch as the Greater Swamp Frog (LiIllIlOIU'Clt'S ingeri)
and the Yellow-bellied Puddle Frog (Occido:.yga lacl·is).
30
ECO LOGY OF BORNEAN FROGS
2) Adulto; only stream bank sitter: Certain ~pccies of frog~ commonly

found along the banks of stream~ live away from the water during their
juvenile or pre-reproducti\'e stage. The Spotted Stream Frog. Rmlll pierllmlll
and its close relative R(IIUI .~igl!a/{/ . as well as the poisonous Rock Frog. RwUl
1/O.lii. are good examples. Tadpoles develop in quiet areas of small ~treams.
often in hrown masses or accumulations of rolling vegetation called ··leaf
drifts··. When the larvae metamorphose into tiny froglets about a centimetre
in length. the~e juvcniks hop away into the forest. sometimes as far as
~eve ral hundred metres from the waler. There. again if Ihey r.:an avoid
numerous predators. they will feed lind grow to about 2.5 r.:m long. After
reaching this <;tage, they will return to their stream . where they will breed and
remain for lhe duration of their lives.
Fig. 7. Stream in hilly lowland forc~t.
31
ECOLOGY OF BOR...'oIEAN FROGS
3) S trram hreeder/rorest wander e r : A thi rd group of c;pecie" u,es ~ln:am"

only for hn.'cding. T~ picall). onl' of the.')\! frog:> (for e'l;ampll". the BrOIl n Tree
Toad. Pn/(J\fiiJel hmii) l'merge.') u~ a tiny lOadlet from the "ide pool of a
.,tr~·am ;lnd hop.') ;muy into the fore~ t. 10 lile thcr~' for the re~t of it~ life.
r\!tllrning to the ~Iream only at intcnal,. w breed. :-.iumewus frog~ in Borneo
di~pJay thi~ life style. including the famOlls horned frog~ (i'Vfegopllrys ~pp.).
the liner frog" (Leproh/"{/clliwlI ~pp.). the White-lipped rrog (Raila
c/w/tOl1l1wj and th(' Harlequin Tree hog (RlwcopllOl"/fs I'ardalis).
~) Forest Ilandcrcr: ~ l a n} of the frog ... of Borneo "pend their lile.')

\\;mdcring \\idely through the fore . . t. ;md :Ire rarel~ encountered along the
banJ..~ of .')treams. The majorit) 01 the.')e .')pecies forage Ilithin thl' for\!.')t floor
liller tFig. R). J\lOSI of Lhl'Ill la~ thdr eggs in ~mall pooh on the foreq floor
IFig. 9). I\hik "ollle u. . " \Iater·filled tree-h(lk~ and other~ depo~it their eggs
within the leaf iiller where there may he n~) . . wllding II ater. hut II here lhe
hurniJily i, high and ~t:Jble. Forc'-\ poo1" Gm be formed in cal'iti..:, t:r..:aled by
uprooted tree~. or by large animah ')U(.:h a~ wild pigs (or 111 c;1riier time~. the
rhinOl.:ero~1 that exealatc 1\:11101\.') in areas of ~oft mud. Clll1op)-dweJling
Fig. K. Searching through leaf hUer lor Iro£" in lOll land fnfC'i
.l!
ECOLOG Y or B ORNEAN FROGS
froc'>. such 3S \Vallace's rJying Frog (Rlw("opho/"ll.1 lligropalmwlI.I).

ReinlI3rdt'~ Flying Frog (HllOcophorus n:illll'ardri) and others. dL'~cend to
hang their foam ne~IS 3t the edge of these pools. usually attaching the frothy
egg rna~~ to ovcfhanging vcgctatio n (I-'ig. 10): Terrestrial forest <,pecies "uch
a~ the Saffron-bellied Frog (CilOperill(ljill"("{l) Of thc Mahogany Frog (Raila
IIICII/O.la) also breed ill such pools.
Arboreal. or tree-living. frogs thai breed in Ivater-containing tfl'C hole~
se<:m La recognize lhe diJkrenec: between large and small trcc,. The tiny.
piping Tree-hole Frog (Jl>'lernphrYllc//a sI/lu/{//w) scck<; out holes in trees or
Lia!l>ls that havc tmnks or stems usually no larger than about 12 em. In
contrast. the Cinnamon Tree Frog (Nyc/i.wllls pic/lis) alw>l)'s deposit, its
eggs in holes located in trec ... having a trunk diameter ah"'ays larger than
aboul 30 CIll. The Brown Tree f rog (Rh{/('ophol"l/~ IUlrri.l"wmi) u~e" nOl only
holes in thl' trunks of large trees for breeding. but deposit~ ib egg~ elen in
the ··(rough~ ·· lomlcd by fused buttress roots of giant menggaris C·wpang·· )
trees (KoolII/Jassio "p.).
Several of the spel"ie~ of bu~h rrog<, (genus Phi/awl/.I) lay only it small
number of eggs (6- 12) under dead leaves. or bcne,!lh dead log~ or layers of
damp moss. The lar\'a develops in an unusual \\ ay. undergoing the pnX"ess of
Fig. 9. Pool in lowland fore.;!. breeding ,ile 01 many trog~.
33
bCOI.OG ) or BOR;\bA:" FROGS
de\ o:1npmelll \\ Ithin Ihl' C'gg. not 3!- 3

t:!dpllk. blll <I" il lllt~ fmgkt \\ jlh .! tad.
The I r(,& let r~'Tllal!h \\ ilhl11 the .:gg
until all Hllerna! lIrgan~ t~ P!c:l! of an
auult frog hale fl1rnll.'u. /-rog, 01 thi ...
t ~p~' ckarl~ halC' 'lcquir.:J <! lite c~r.:k
that ! ... l'omph:I(" I ~ imkpC'ndenl 01
bodi.: ... 01 \Ialer. They ~ll':l·.:("d e~pt'~
('iall~ l\l'lI in high-altitude Il$)nl:llle
t"oreq . . II hne a bJanl.. ..·t of II el. "'rl)llg~
!lIO,,, {:ole.>r'" many e\pu ... cd "urfar.:.: ....
T~pe~ of hl(" c~de . . haw JJllk 10
do I$(h a frog' ... lel1t.:al lIr loraging
zone.> 1tl Ihe fme ... , There ar~' .trlmreal
and tt'rr.: ... trial "pe.:io..:'" 1Il each (It the"t'
t~pe ... 01 Ii 10..: cycle ... frog ... III Lllg un tht'
ground U"'': d lari ... l) or n11crohabililt ...
or ... uh ... lrale.>"': Ihe Grealer S\I"mp Fmg
lLiml1l11/t'lNS Illyen) Ihually "ih on
muJ. "here:1!> "'" larger r.:latllC'. the- FiA. 111. ro.UTl ne'" (It tn:.: frog
G iant Ril er Frog (LlllllllllltTle.1 er-hanging rhlllo II aUOII
III
/t'I'om/llI) 1110'-1 frc4u('ntl~ i, found on

gran'lor ... ~\1ld. TIll! Rock S kipper (SulI/mi, faropafll!{/fl/f) .:m.:rge . . at night
10 peI\'h on large rod, . . In miu·"lredll1. ,\rl-'oreal frog~ \<try in thei r ~eJectiun
of pen.:hl'''': the Cinnamon Tree !-'rog (,\'l"(/i\(/IIIS 1,ItTl/I) and Ihe Fnlled Trel'
Frog (H.hacophol"l/\ app('/u/inillllIlS) ~ it on the leall" of ~hrub~ and young
lr.:(' ... 1\ her",:!.., other.... p.U1i~·ularl) larger ..,pc~·I~' ... <;ueh a, th~' File-eared Tn.'e Frog
(Pohl'n/tlles ori/oflfll/I) and the Bro\ln Trl'c Toad (Pi'dlmihel hosii) perch
on \1\ Ig'. braIll"hc ... or elell trunk, of 'Illall tree~. One of Ihe per... i,Il·111
mY~l.:ril'''' of Ir",c i'rof!" concern' the hdghh ilt II hleh Ihe~ ked rind re~1. A
human ()h~ener (ltl the ground r.:an "POI fn.!g' only to a n:'rlain height.
perh:lp... up to eight llletre .... S...eing a frog al a Ui,t.:Hlce further than thaI.
(''''fX'eiaJJ~ \\1111 ont'· ... hnt' of 'ighl hmt.lered b~ hranche~ and k'3W'. i~
':\lr(,1!lel~ dirlicul1. :-;e\erlhck",. \Ie kl1(\\1 (rog~ dolil(" higher up hccame
\I\.' ha\(" heard Illl'lll calling Immthe C:lIKlp~.
The "pecie ... til IrO!!' li\ ing at ground 1..·IL'l :11"'0 dl'rLl~ a \arid~ of
beh;l\ iour.... though II llh I e~ 1"1.'\1 eH'eplion ... all ar... Illk.:tuma1. A fell 0)
them. for l''\ampk'. Ih ..· ~padd~'ot frog, I g<;'l1U'" rllffllf'/fo I. hUrT(m, Olher,.
ECOLOGY OF BORNEAN J--ROGS
"llCh a~ the ~tiek~ frog~ tKalophr\'lIl1sL ~pI."nd mo~t of the time under the
dead lean~ .. of the fore~t floor. r-.lale~ of the guardian frogs (LilllllOllccles
Fllchi and L. pU{U\"{IIIClLli.l) call from under dead !em·es and then hop acro~s
the de:.ld k:.l\e~ :.I" the~ carr) their tadpole" ~ll \~ater. The horned frogs
( Mcgophrl"S) forage for pre~ alTO~~ the leaf litter.
The microhahitat~ of tadpoles. though not as \"aried a~ tho .. 1.' of adult
frog ... -.till em I.'r a hroad rangl.' of condilion". DI.'... pill.' the n<..'ce~~ity for living
in \\ater. l<ldpole~ pm~e~~ :.I tremendou~ number of specializations within
panicular type .. of aquatic habitats. Ewn among ~pecies that Jive in a single
~trl.'all1 therl' can he major diffl'rences. ror example. there are tadpole .. round
onl~ in the ,\\ift or lurhlLlent portion .. of rod.) ,tream .. (Fig. 7). ~uch a~
tadpole~ of the gem'" JlcrisrogclIYs. \\ hich c1 ing to rO(:b in extrt"mely strong
current~ by ll1ean~ of an abdominal sucker. Tht"se same .. tretchl's of water arc
al .. o used h~ tadpoles of the Spin~ Slender Toad (AI1\olliu 1/7illll/(tf.-r). \\hi<:h
dill\:r h) u .. ing their C\panueu lip .. a .. a c1inglllg uel icc. an adaptation seen in
a Ilumbl'r ur BOnlC.lll rrc,>Il\\~lcr li~hc:. . Other k..ind~ uf tadpole~ may be
pre~ent in the rapid~. but the~e 1\ riggle duwn intu the cre\·ice ... between nx::ks
and gravel at lhe bo!!om . induding larvae of the slender litter frog ..
(Ll'{Jlo{a{lIx ~pp.) and the Short-Ilo<;ed Tree Frog (Rlw("opl!o/"ll.1 gWlIlil.
Aecul1\lIl:ltion .. of dead k:l\c'\ ma~ form in ... tream') \\here the
tOJl()gmph) lTc~tc~ eddie'. po<:J....~·t~ ()\" liltle or n() ClllTCnt at the ~ide of a rapid.
Thc~\;' leaf drilb form the prin<:iplli minohabitat uf uther kinds of tndpoles.
lIlduding those of the Gi:1I11 Rj\·er Frog { Lllili/o/I('crc.~ {cporillu.O. the Spotted
Stream hog (Rulla "icrum/tI). the Brown Slender Toad (AlI.lfmia Icpropu\")
and the Brown Trel' Toad (!'cdo.\ Jihes 1I000ii). The<;e four .. pecie~ are
sometimes found in quiet. <;heltered side pnols CUI olf from the main l"lment
f10\\ h~ :.l gran:l har. or rod ... Thc"l' placid pooh 1\ ilh thei.r si lt-laden
bnll(llll~ :Ire al .. o the main habitat of a.nuther ~et ofwdpole species. including
thu",c of the White-lipped Frog (Rmw ellaleolloral. the Crested Toad (RI~f()
dil·t'IRt'1I51 and the Harlequin Tree Frog (Rfwcopho/"ll\ pcmlali.lj.
Some qream tadpoles. such a~ those of the large-c)ed lilln I"rogs
(l,t'fJ/o/Jrucliilllll <;pp.) <;el'ln to thri\"e In a variel) of microhahital~ no llWlter
\\hat the currcnt .. treng:th or hottom t)pc. Soml'hu\\ Ihe:.c large . fat lad poles
l"all mainl,lin Ihc111 .. elle .. Ilithuul "qrcamlining ·· or ~pecial clinging
adaplatiom c\en in '>t rek"he ... of moderalel~ ~trong current.
Si1t~ poob on the fore~t floor ~lre the main type of el1\ ironment for
tadp(}le~ of ~e\eral tree frogs and Ih o~e of se\"eral incoll'picu(lu", dllarf
.. peLil'~ ~uch a~ lhe Bornean .'\aITOII·mouthed r rog (Minvfn·/a hOrllC'clIsis).
Usual1~ a rhino or pig II aIJO\1 I Fig. 10 1. or isolated fore~1 rain pool II ill hold
35
ECOLOGY OF BOR.i\'EA~ fROGS
\CI('ral kind~ of ladpoles: a tree frog wdpok and one or two terre~trial dl\arf
"p('cies. In the~e ~mall comn1llnitie~. the 11\0 or three types of tadpole~ differ
greatl) in size and mode l1f f('eding. The tree frog tadpoles (R/wcophorus
lIi~rop(//I11{1/lI\ or R. reilll\"(/I"dri. for ('x3mple.) are at lea~ltl\i..:e a" long a~ the
others. and r('('d hy "rooting"' or "grubbing" 1!1 the decaying organi ...· mailer at
Ih(' bottom. The ~mall('r tadpoles UHicrohy/a bOI?l('(,lI.lil") hover in mid-waler
and filter out food par1kk~ as they pUIllP water through till' mouth ea\'it)' and
ul'er tIlt' gilb.
Tiny p001s of water. ~umcliml'~ Ill) more than I cm deep and 10..:111
acro~s. whidl collect on the surfa\.·e of rolll.:11 logs or el'en all large. dead
leaves can prm ide suitahle habitab for tadpole~ or thl' Rufous-sided Sticky
Frog IKa/opflrYIII/\ p/(,lIm,lli~IIW) or of one of the ~m:jll fon:~t lOad~ of the
gcnus f'elo{!!irync. Tadpok~ uj the, .... species do not feed. but in~k'ad
complete their del'e!opment b~ ab~orhing the large alllount of yolk within
relalild} large egg~. lJelelopmelll to metamorphosi~ take~ place in 7- 10
day~ . and 1I11Cll the ;.oung froglet~ hop :j\\a~ from their hirth pool. they are
hardl) as large a ... a I11cdiulll-,ized ant.
Pockeb of 1\ ala ahow the ground in tree ho!e~ or butt!"l."~s trough~ arc
the mierohabi!:lb of anolhn group of tree frog~. The~e eleqllcd
microhabitats illu~tra t e 011(' of the major differences between tadpole:. and
frog ... Ohliously. tadpoles canllot "lind" a plal'c tn live in the usual sen~e nor
c:m thl·) migrate from one habitat to anolhcr. Thl'~ dc\elop wherever the
egg~ ar~· laid. Thu<;. if eggs are placed in tree hole~. lhat i" where tlw tadpo1e~
mu~tlilc or dic. NClertheless. a situation could ari~e I\ithin a ~trea111that i~
prone to tlooding. in which theTe could be some ill\'olullt:Jl") 1110lC111ent
between habitah_ Although huk l~ known for certain. gil en lhe oneil
powerful bUI ~horl-liled nature or tlomh in Borneo·s hill stream~. tadpole~
\\,1<;hed out of a ~heltered pool probabl~ ar..- not likely to sllfyi\"e their
lurhulelll journey to a new one.
On..- of the strange~t ~1I1d 1110"t intriguing of 11K frog Ii!"c cydc~ til Borneo
i~ that o!" the guardian frog.s ILi,lllw/I{'crt's(inchi :lIld L. pa/(ll"tllwl/li,). Both
of the~..: \pccic~ la) eggs 011 the fore~t tloor. \Ihi<"h me inilial1~ guardcd h~
the male (Fig. 11 J. A" thc tadpolc~ hatch. they somehow all:Kh to hi ... had.
IFig. 12 ) lIml ar..: carried around for.an llnknO\\n period of time. perhap~
"L·\cral da;.~. until the parenl frog L·n":()Ul1tcr~ a ~llitah1e pool. The male
mOl c~ inlO the \Iater :lIld larlae SI\ im ,1\1 a~ ami complet(' their de\elopment
in th..: pool. The purpo~e of thi .. odd beha\iour l11lhl .....:rtainl) he. protection
ror the lanae. but the conditions that led 10 il~ L·\olutioll an~ not let
ulldcr~t()od.
36
ECOLOGY OF BORNEAN FROGS
Like other frogs of Southca .. t A .. ia, the frog.'> of Homeo are entirely
cami\'orou", taking: onl) animals a::. fooJ. Overal!. they prey primarily on
insects and related iIl\·eI1ebrates such as spiders. Mosl frogs Cal a wide
\arid) of pre) wi thin certain size limits, hUI, ~uml' h'lve rather specialized
diets. The Rufou s-~id cd Stick) Frog (Knlophrnllls plelll"Osligll/{lj feed ..
mostly 011 anb and termite~. so of course most of the prey are small. The .. e
rnoJest-sized frogs can eat from 50-150 ants or lennite.'> at a single s itting .
Til.: frog will often park: itself al the entTane.: 10 an ant or tennite ne~1. or b)
a ··paradc·· of thc .. c food item~, and pack ih tummy at a lei .. urdy pacc. The
Cre~ted To<ld (Buto (/i\"t'/"gells) while feeding on a wide as~onmelll of in~ects.
<llso leam to\\ard a diet ri ch in ants and tennite ... Stomachs of thi~ .'>peeies
\\ 11en examined contained on a\erage about 6-1-lJ. allls and 3 1(,c h:nnites. and
an u\erage of -1-."1 pre) itcm~ per ::.tomaeh. The Giant Rin'r Toad (Bufo
juxlwper) rullow~ a ~imilar pattern despi te its large si7c. Thc main differerll·c
belwe\!ll the [\\0 lOad~ lies in the prey size. with the river toad eating Jarg\!f
pre) (:t\erage 7.-1- mm) and the crc~ted tnat.!, 'Ill<lller (a\·erage -1-.0 mm). It i~
110 Illystery wh) ant" and ternlile~ 1"01111 <l major diet componcn t for ~ome of
these forc~l I"rog~, .. im;e the.'>e il1\'eI1ebrates form the !argl"~l elemelUs of
animal hi()llla,, ~ in SOU1hea~t A~i:Hl tropical forests.
Tho.: majoril) of Bornean frog~ are. hO\\('\er. gl'neT<lli~b where their Jich
arc cOfll·erned . evcnthough ants still rcmain an impOf1anl food type for man)
of them. The Smooth Guardian Frog. a small (about 20--30 mm ) ~p~'l·le;,.
Fi!-:. 11. l .il/lllont'cff'5 jinch' male gu;,.rdillg egg.;
37
ECOLOGY OF (30RNEAN FROGS
Fig, 12, Lilllllolll'l"l<'Jjillcili male carr;'lIlg ladr()k~ ,
('Oll~lnne~ ~pidef~, bt't'tlt's, I-..m;.did\ and other onhoplefan~ , but aJ1t~ arc
ne\enhele'i~ mort' than ."We ofih pn:~ . The generali~b tend to eat larger prl'~
:md to inge~t fewer pr('~ p("r meal. For "",\<Imple. fi\t~ stomach, of thl? Ahho\\'\
Littt'f FfClg (LeIJfo/JmciIiulil (/bbofli) eontaint'd crickt't~. 'pider) and <I
cockfCIach. all \\ ith an al eruge length of 17.S n1l11: the a\ l'rage number \\'U~
onl;. I.S pre~ itl'm" The Bornean Horned Frog ( Meg(lphry~ I/(/SI/((/) with its
imprc\,i\cl) II ide mouth eats e\'en largt'r prc: (,!wrage 32 mlllL and is
\\illing to tucl-..le and swallow ..j.~cm snaib and 10-cm ~eUl"piom. The largest
pre;. of all are eaten b~ the Ciant Ri ",.:r Frog (Lilllllollcclcs /c[!orillllY). th("
Peal Swamp Frog (LiIIIl/(!IleCTI;' I' lIlo/csimIllS) :md the Greater SI\amp Frog
ILiIllI/O/ICC/C.1 ingail. \Oml' or nomeo' s largest species. These cal rrog~,
snake\. crabs. fort'st millipedes. :lIld scorpions: almost an;. \mall animal
passing too close to one uf lhe~e adult frogs is lil-..("I;. tn di,uppl'ar in a single
gulp.
Th(" indi\idual breeding and ft't'ding hahil\ oj" i'rog~ in Borneo have led
to illtere~ling pal1crn~ of dj~trihutlOn among them. fool" example. as in SOI11("
other lenebrate gruup~. dj~tjnel c(lmmunilje~ of frug~ exiq at \ariou\
altitudes_ undollht("dl~ cau\l'd h~ radicall~ diffr em hahitats in dilrcr~'T1t
tupograph;. and at different allilude~. Recognizable frog cOmmullilie~ OC('ur
in mangrO\(' \~hen:. hec,!u\e rnml frog" ("an not tolerate ("\en milt.! ~:dinil~.
tlnl~ tht' \ langHlH' nr Crab-eating r rog t Ft'ie/"mrya ("(II/criw!/"(/) and
ECOLOGY OF BORNEA~ FROGS
(Kcasionully the Tai\lunese Frog (Hoplobarraclil/5 mgll/oSIIIJ l'un be found.

Other di~tillet communitic<; GIn be seen in peat Q\lllllp. Ilith numerou"
"'lump frogs If.i1ll/1()I/('C!eI· lIlah'.lilllll/l and L. (Wrlllllacrodoll). SI\ amp TO:l.d~
( BII/i) (ju(ldripoI"CllfII.l"i. and Rough -~i lh.'d Frog~ .(RwlU glundl//o.lu); a much
richercotllillunit~ can. ofcour~e. be found in the hill) 10011and~ (Fig . 7). \\lith
nllrneroll~ Jllember~ oj" the genera Mcrislof{t'nr5. Rona. Kafophrynll5.
/\nwnia. Hl/(/I"opflOl"IIs and oth~'r<,. Intere<,tingly. this cOlllnllll1ity i~ again
diiTerent abow ahout 1200 metre~ in elo:\ at ion. Perhap~ hecau<,e of the steep
IOpograph~ the meandering qreams with their nunKrou~ microhabitah urc
rncreasing.ly rare. Stronger currents caused by more extreme gradient~ le:td
abo to fewer oppoJ1unitie~ for org:anic material to collect. and 10 more
··C;.lt;]slfophic·· qre;]111 flows. \Iashing out (ktrilUs (as well as an) resident
tadpoles!). Thus. at higher altitude~ there i~ :1 ~igllilic;]nt ahsence of frogs
fro111 certain families or genera. Abo\e 3000 rnetre~ there arc no
tllt'g()phr~ id,>. fe\1 bufonid, and fell microhylids. The mo~t abunu:lIlt specie~
are Iree or bush frog, of the genera RII(I("opho/"ll\" and Plii/al/ilil. The rea<;ons
for \l1i\ appear to be direct!;. re!;lled to brccding:: tht' trog<; that remain are
thu~t' thut do not depend on ~tream~ to bre\:'d. Sum..:. ~uch :I~ thc mo~t
conmlon g:t'nu~. Phi/al/tw. ~hO\I tendencie~ tOIl ard~ direct develuprment of
the taupole~ \Iilhin the egg. eliminating the free-sv.unming tadpole ~tage
entirely. There are aho int\.Te~ting oppormnist<;. ~uch as the tiny (a\erage
length about 15 mm) Blad. Toad Pelophrynl! mill!m. which deposits its eg.gs
in the water-filled resenoirs of pilchcr p1allh. Tim'>. there i~ here a hint that
frog communities may be crafted not ~o much b) \Ihatthc adults can do. htll
under II hat conditions their tadpoles can ~un i\e .
All ur thi~ ecological information can actual!) be ~unmlarizt'd ruther
~illlply: frog~ hal~' wcc("eded in exploiting almo~t every imaginable nook
ami ,:ranny of Bomeo'\ rmcq<,. and to u<;e in one way or another essentially
all the in~ecb or other il1\ert..:bral("~ round there. Tadpoles expand further the
range of elll"ironmenls and fond re\ource~ lIsed h) the adult frog<;.
Furthermore. the species making up thi, rich fauna dil ide und ,han:
resource,> of '>pace and food. forming a critical pun of thc c(lmplex st ructure
of the tropical min forest.
39
Co n S {J r l5' Cl' i017
wing the early 1990 .... the attention 'of biologi ... ts \\as suddenly
D dra\\tl to\\ards the decline in numbers or e\'en the disappearance.

either gradual or abrupt. of \ arious ~pecies of frogs in locations in
North and Central America. Europe and Au.'>lraliu. No one was surprised that
one or ~l'\cral "peeies disappeared when the environment \\a.'> drastically
disturbed. !>ay. by Ihc clearing: of forest or the conversion of nmi\"e grassland
10 wheat fields. But \\hal concclTled hiologist~ wa<; the declines thai took
place in protected. and apparently nol ~eriously di~lurbcd. em ironlllcnl~.
Frog populations in a variety of habitat type .... from mountain ponds (0
coastal Slr<.:ams. dl'cJincd or disappeared. A number of causes have been
suggested. including pollution huch a~ peqicides. herhicide~. fertilizers. or
acid rain ). geneti(' damage cau~ed by incn;a~ed cxpo~urc to ultraviolet light
(implicming the loss of the earth's ozone l<l)erl or even pathogem (such as
lethal \ iru~6 and fungi) . So far. however. no single factor linking all these
apparent dccline~ ha~ heen identified.
In Borneo. ba~ ed on our \\ork and Ihm of our colleagues over the lasI40
years. there is no e\·idence of a dccline in Bomcan frog~ within lmdisturhcd
forests. Fluctumion~ in populations. both up and do\\n. ~omelimes dram<.lIic.
ha\ e heen obsen"ed. bUl no long-term trends are apparent. Though the
hunting of frogs for food has occasionally been blamed for a loss of these
amphibian~ from their hahitats (as in the ricefields of Bangladesh in the late
1980s ). human c()n~umplion of frogs is rarely a problem in most cases.
compared at lell~t to human l110diiiCluion of natural hahitat~. The only nati\c
species Hhat is. those living in undisturbed forest habit(lh ) that aT<.: probabl~
affected by hunting are large. long-liYCd. slow-gTo\\ ing species. such as the
Giant River Frog (Lill1llol1eCle.1 leporillll s ) and its relati\es.
Fc\\cr than ten per cent of Hornean amphibians. and not a single
uniqucl:- Bomcan <;pecies. can thri\·e in habitats greatly modified for human
u~c. Though ~Ollll' qudic<; <;hO\\ that well-managed seJecti\'e logging is one
of the human ac!i\ itic~ ka~l d<.lmaging to anuran communitie<;. mo~l oj" the
forest exploitation (·urrentl~ practised doe~ l·ause noticcable ··colllmunit~
shifts'·. in which certain specie~ increase and others become much more
difficult to find. For example. lhe silted ;,treams of logged area~ in SaraRa"
shO\\ an lIlcrease ill the Giant Ri\·er Frog (Lilllllollcc res /cp orillll s) . \\hi1c
some species of Torrent Frog s OlerislOgc lIYs spp.\ become scarce. Tho.:
CO:-':SER\ATIO~
e.'l;plaliation lic~ III Ihl: Tt"prwucti\"e habit ... of the,c 'I'leCle:.. The Gi,ml Ril er
Frug I' nOl 'l'n,l!l\e \() turbjdil~ and ...ek'ch ~h:Jllo\\ ,and bar, 10 I'l~ ib i,:g~"',
while Ih tadpule, feed Oil organ1\..' debrh ,uth a ... dt:ad leale~. Ton·em frog'.
on Ih~' other ham.l. need clear Ilall.'"r for ,ucce"ful hrt"eJing. and their
tadpole, li.:cd on tht: thin film of micru,I,.'opic organi~nh g:rO\\ lIIg on thc clean
:.url'IC"t" 01 roch and houlder,. In 10gg:l'd nrea~, n)('k~ in :.t rcanh become
coal"u \Iilh a thill lu)l'r of ~ilL and the: 100d ~uppl) of the,>c tadpole~ i,
de~tru)cd. ;-.re\cnhele~~. re,ean,:h ha" al~o rCI"aleJ Ihal 1llan~ !>pccit"<;. e\t'tl
<;en~ili\l' one~_ loan relUm ,f Ihe :,uTT"ollnding fore;,t i:. allO\\ed 10 remain
undi:.turlxd for Iill' (0 ten ~car,.
:\grkulturt" \"T~'ate ... spedal problem'> InT the nalil\,.' frog fauna. :'>lot unl)
dll...', agrkulturt", lil-.e ,:omplete tllllbt"f e\traclioll. remuI\.' the natural habitat
and IIlcrea~e Ihe ,ilt luad III 'trt:am~: il ahu IIltrodul,.'C", man~ !,:hl'm icah.
pe~ticidl''>. herbil'ide,_ fel1l1l/cl"'_ etc .. that ,eriou,l) h.um thc faurl:l. With the
increa,>\,.' III ,ize 1,.)1 tht" hUl1lan populilti~lIl. il i~ incI Itable that 11lnre l;md III
Borneo \Iill be ~'umerted to 3.gricullure. which 11111 lead 10 increa'>cd
,iitallon, l,.-hL~IIlIc;11 pollution. and mereawd runotf and Ilooding: .Ilter hea\~
raill~, :'0 that lrog I,.-UIl,cn atlOn IllU,t he p.m nl an~ iand'>capt"' ,'on,cflmioll
pmgr3.lllme.
If we I\anl \0 pre~en ...' Borneo\ irog fauna_ I~hich i~ among lhe rich!;" ... !
all~\lhl're in Ihc \Iorld_ il \\ ill be necc"3.r~ 10 prc,crve:l \anct~ of n:.ttur.l1.
original habilah al 3. \ariet~ 01 rlace~ 111 Borneo. Th.1I is. of COlll"C. the ~ame
recipe for pre'>er\ in,\! almo~t all of i:3ornI'O',> plant and ;'lI1i mal ~pccies.
Jalllily /3ol1llJillalorida e
hi;; fami!} m:curs in ea ... tern A~ia and in Central Europe. The genus to
T I\llich the Bornean ~pccic ... belongs is found olll~ in Dorneo and Ihe
Pa!:l\\an group (Phi1ippin~ b.land~). This j, one o f the aIde .... groups
of frog~ and Blirhollmia is unu<,ual beC"au~c i1 seems to lead an entireh'
aquatic ('w,l!.:ncc. (See .. ]",0 p. 19.)
Barhollmlll ka/imll lllallellSis Iskandar

Fbt-hc,tded Frog (Fig. 1.1)
Description : A medium·"j7cd frog \qlh a llanened head. rdalively ~rnall

(')c .... and broad. rounded snOLl\. Thi~ frog has a numher of distincti\'!.'
,""atllres. The fingers and tOC, arc full) \\ebbed to tlw [ip ... and tile tip ... of buth
lingen and we, arc .. lightl) ,>wollen. A di~{inct !lap of ... kin run ... along the
outer finger. 111(' third anu fOlll1h toes aft.. ahoul the same length, a~ are the
~econd and third lingl'r~: the effect 1\ 10 mal..L' both hand and 1'001 look like
rounded paddh:~. There i!. a <;,mall cOnl.:al projecllon 01 <').,111 Ju~t below the
from thml of Ihe eye. The eardrum I' not \isible. The hat.:1.. b rough. \\ i[h
flWn) ,mall. !>piny or rounded ,).,in projectioll<;'. and tho:n: are fold, of ,).jn
along Ihe ~jde~ of tile hod) and along Ihe thigh :lI1d l(mcr leg. The enlire frog
i, dar\... brown or black. \\i[h lighter mottling on lho: under~jde.
;\ lales: 68 mill.
FemaJe<,: 77 mm.
Tadpole: LTnkno\\n.
Ha hits and habitat : Onl~ 1\\0 ~JXcimens of thi ... odd ~pecie<;, ha\c C\o:r
been found. both cauglu \\ho:n boulden. \\ere O\crtumed in ~hall(}\\. ~\\ifL
rod;.- 'Ireams. Both <;,~eimcn~ \\ ere caught in the ~ame 'Irea. The ')pecie!> is
1.: le:lrl~ full) aquatic. hut nothing el<;,e i~ kllO\\ n of it~ habit<;,.
nis triblilion : KnO\\ll onl) froLlltlte middle oflhe Kapua, Ri\er b.. ,in in
we 'tern Kallln:mlan.
-B
BOYlHINATORIDAE : BAKBOURULA
Fig. IJ. Hurh(lurula ~alilllalll(lm'II'i.\ 1~";lIld;lr.

.J'alllily M egophryiclae
he Bornean specie~ u~ed to bl! c\a~~itil'd a~ mcmbcr~ of the Family
T Pl'1ohaLidae and weft': so listed in Frogs of Saba". M ost live on the

forest floor. :lmllenO to hid\? during thL" day under leaf litter or. in the
case of Horned Frop . ~malJ burru\\ ....
Genus Leptobrachella
D\\ arf Liller Frog"
he genu .. LeproIJul( Iwl/a Include;; the .. rnallest adult frogs in the
T fatluh r-Iegophr)ld.lt:. ,mo 1.., \...no\\11 onlv from Borneo and the
BllnQ:ar.1ll l ..omellmes c.!Iled Natutl<l:.l b],md, There ar~' ~IX
in Borneo and three. Ll'{JlObrachl'lJ{/ II/jobergi (Sanl\\ak). L. bafllellsis
spcl'les
(Sahah), :lnd L (/{lITO (Sahah). are relatively cOlllmon. ~one have been
reported ~cl lrum Brunei and onl: two from Kalimantan. but this weakness
in the rel'ord mere!) n:lll:cb the wcakne., ... of field exploration in those areas.
Adult~ and .iU\'enile~ \\<lnller o\('r Ihe l"ore~1 noor kat' lith,'r. wh..:r..: Ihey
forage for \'ery small inse('ts and other i11\ ertebrate~. Adult~ are most
cOllllllonly seen along the banb of clear. rocky ~treams aboul 1-2 metres
\\ ide. \\ here they congregate to breed . '\1a[es ca[[ \\hile siuillg on dead leaves
on the hun\.. or p..:n::hed on leave~ of herbs J 0- 20 cm abO\"e the ground. For
their ~iLe, Leptobrac!u'l1(1 .. pecics are .. ome of the loudest frog~ in Borneo.
They ha\'e an extremel~ inten .. ~'. high-pitched. metallic Irill. 1.arge number..
of breeding male~ along ~treams ('an (\ro\\ 11 out e\'en SOTlle of the night
insects of the fore<;\. Tadpoles of D\\arf Liller frogs li\e in crevices between
gr~l\el and ~rnall roch on the bed of riffles. ~haring Ihi<; en\"ironmem with
tadpok~ of tlw Sknder Liner I-'rog~. These frog~ have been found frotn 150
to 2200 rneln,::, ahm..: ,ea k\cl. with 1\\"0 species. L. hU{Ui'II.lil and f ..
bn' l'icms. restricted to i:!C\ali(l\l\ aho\..: 900 l11etre\.
Descriptiun- adults : All "p~Tk." ar~· tin~ (Ie ...... than 25 mm). \\jlh the
head ~lightly narrower than the bod~. The Tllo."t unu~ua[ kature of these liu[e
frog" is the polmed. a[mo ... t aITO\\ head-~haped. tip~ of the fingers. The
eardrum i, \ isiblc. 3nd the "kin is smooth. AII"'pel·ies excepl L. palmUla lad..
:" 1 ~(jOI'H/{YI D AE L~PTOBRA(,IIELLt\
\\ !;hhing on the fel.'l_ Tho..' gcneral colour,lllon I' dar~ brown or black \\ Ilh
dark marking ... ('n thl.' b;ld. U,uall) there are 'lllull hlne\... 'p(\(' on Ih ... ,ide .. .
Colouralioll of the b... ll~ \arie, among the "pecie~.
:" laJes: l5-191ll1ll
FL'lllale,>: 18-~~ 111111
Tadpole: Lar\a...' ar.: !!rc~l,h and e\tr... mel~ .:lol1g,l1(,u like the t,ldptlk"
of Skmll'r Liller 1-rog'>. :\lthou~h tadpole,> of bolh group, ha\\.' lu."a\ ~ blat'k
bea~~, lho,!; of the O\\arf Litt ... r rn..l!!" bd the rO\n offil\e.lill~ tceth prc,ent
on th ... lip" oj [aupole, of th ... SIeIH.1er Litter }-rag" \\'h ... 11 r... ad~ lU
11leta11lorpho~e. Ll'llTII/nm-llI:ffa tadpole, Ilh:a,ur.: . .W--I.? 11IIll.
Leplobrac/H'lIa Im/llefl ,'ijis Smith

Kinabalu [h\ ari" LIltt'r Frog
Description : A ,mall. ~kllu.:r "pecic'> lacking \\ebbing un lhl' fooL The

,I..in I' ,mooth on Ihe bad.: the "jUL'" h;J\L' a fell light-coloured gland ... Thl'
~'oluur i" broIl nor hlack, \\ irh seIL'ral black 'JlO'" 011 the hack :md a 'h;lrpl~
defined. blad ,trl';lk m-er the ",antrum. Tho..' ~iUl" hall' l---llarge black ~poh.
rile beJl~ i~ dJr~ gr('~ \\ ith small light ~pol~,
,\l;lles: 15~19 mm.
females: 18 ~~ I mm.
Tadpolt': See d.:~niplion under gemh.
Hahits lind habitat: Thi~ "pccic .. ha .. been found onl~ ill ~ubmonlanc
ami montane fore~'" from 900 to nol.) Inl'tn.-', above ~ea le\'el. See abo under
g~'nll'.
Call : Scc- under genl1~,
Distribution : Tll1~ ~p",cie~ has been found in th~' 1l10U11taith of \'.e~tern
Sabah, northc11l S,lra\\ ak. and nonhea<;tern Kalimantan.
Leptobrach('/Ill mjobergi Smi lh

t-. ljobeq{ ~ D\\arf I.ittcr Frog I Fig:. l~)
Description : :\ "mall. ~Iendt'r "p~'L'iL" lacki n£ \\ehhlllg on th", foot. Th~'

.. ~in i, 'lllOOlh on thl.' back. \\hil", the ,id('~ ha\ l' L'long;Hc gtand ... forming ,I
10\\, llarrO\\. hrokcn ridge. The colour b brO\\n or hl;ld. \\lIh ,e\eral black
'1"It.!'" on thc hack TI1L're is no black. "trL'Jk O\<.:T IhL' .:ardrum T he che,t I"
~ I EGOI'H}{YIl>Al::.: LEPTOBRAC II ELLA
Fig. 14. I.('pwbmc/rel/ollljolil'n:i Smith.
tkd~J \\ ilh bro\\ n or ha~ a brol\ n nt."l\\ork: Iht." re~1 of Ih~ h~ll) i ~ u~u:llly
unm.lr"~d.
:'-1:.le,: 17- .20 mm.
Felllale.~: 18- 21 nUll.
Tadpole: See dc,cription und~r gellu~.
Hahits a nd habitat: Th., 'l)Cdt'~ IiI es in hilly lowland fore~h rrom 150
In 500 lll~tr~'~ abo\(: ~ea Ieyel and breeds along smalL clear. nx:"y ~Iream~.
S~~·al,o Ulllkt gt'nus.
D b tribution : Known from Sar3\\ak. and reported from northeastern
Kalimant:\Il.
Leptobrachella palmaIa Inger & Stuebing

Weo-fooled O\\arf Litter frog
Description : TIl!, I ' (Ill" "malkq ~pecies inlhe genu,. and thl.' only one
\Iith \lcll-tI.... lek1ped \\ebbing on Iht." fool. The back and bl'lI) art' ~moolh.
\Ihile the "ide:. haw ~<:alleri;'d round glands. The upp.:r "llrfa(.·t'~ are medium
onlll n. II ithout marking~ except for dar" crm~har .. on thl.' limb,. Th(" .. ide:.
h'lIl.' a dark. net\\ork encio,ing the \\ hit(" gland,. The middle of lh(" bl'll~ I~
light. II ithout marking:..
\ IEGOPHRY II)AE : LEPTOBRAC HELLA
~Ltle': 1~-17 mill.

I-"em.t),.. ,: L'n"n()\\Il.
T;tdpo lt,: l'n"nO\~n.
Hahil !l and habitat. The only "nown '~nrncn' I~ere found along a
~lr~'at11 h:lIlk in hilly forl!~l:1I J50 metres abm·e __ ea Jel\!!.
Oh,tribution : Kn 01~ 11 onl) fWIll;1 'ingk ~i!e il11he interior of Sab;lh.
Leprohrachell(1 parm Dring

ROllgh·,ided DI\ aTf Liner Frog
O(',cripl ion : ,\ ,mall 'pel..'it', lad,.!ng \Iehhing. The ,ide .. tUl\e . . malL
roumlt'd. 'l·all ...'rcd gland, \fl:lI1ened blimp. . ). I\hik th ...' hac" j, ~moOlh. The
general co[ourmion j, ,imilar In L. hll'/I('II~i-, . but the ~i(k~ 1;ll·" large hlack
,po". II10ugh there ma:;. be .. mall hlad, dOls. Some indil idu;d~ ha\e a dar"
ar\',1 O\w the eardrum. but il i, not ,harpl:;. defineu [Ike the \[l"eaJ... in L.
l'III'/('l1lil The hclly I~ dar"
~bk .. · 15- 11,l1ll1ll.
l-em:lle': 19 mill .
Tad,)Olc: $ ...'...' under g ...'nu,.
I.... bib and habitat: I.I"e othe,", 111 Iht' genu~. Ihi, ,pecie' lilt', in hill~
fore\[ ... where dear. gril\..:! or weJ...·oonomcd <,tre:lIlI, (,\j,L It, kno\\n
cll'"l1Inna[ range h 15010 750 meln:, "hml' ,ea kId.
Di<;lrihlltion : A[! of $abah (lnd nurth ...']"n rarl III Sar;l\la".
Leptnhracliella ,~eras allae Dring

Slrir~'d D\I arf Lil1cr hog
Dl'!>cription : To.: .. \\ ilh lillie tlr no \1 ('ht>lng. The ,ide, hale il ft'\\ 'Illall,
rounded gland ..... Lq/lobrfllhd/(/ \(·/"(/\{IIIl1I' 1\ IhL' nn l:;. ,pt'cie\ III the geml"
h;1\ lug il cOlllinuou3 bl:lCk ~Irt'ilk fWIll Ihe t:~ \..' 10 the gmin.
:\I..tlt: ... 17· Ilj mill.
rL'malt:,: 20 mill.
T.ulpo le l nkm1\\ n.
Habih find hahitat · AUld" halt' been founu on the b'IIl'" 0/ .. mall
'l[('all)" :lIld :!mid~l Ihl.' k.tllIl\\'r III 1111l~. !O\~ !an t f(ll"e~1.
Di <;trihlltiun ·ln BOn1l!o. ~l1U\\n " I tar{)nl~ 1r01ll ,outhea,lern SaLl\\a".
Thh 'pcne, a!,o \..',.' ...·ur~ lill Pul,1 S~'Ta,an. 11l the Bungaran h[and,. ont:;. SO
"til ott the CO:l'l of BUIl1t'o.
MEGOPHRYIDAE : LEPTOBRACI-IIUM
Genus Leptobrachiul1I
Large-eyed Liner Frogs
re\"iou~l~. the 1l10~t \~ idely di~tribll1ed ~pecie~ of Lcprob/"(/chiu/II 111
P BornC'o \\"as referred 10 a~ i.eprohr(l(:hiwlI I/IOI/{(IIIIIIII Fischer. Field

\\orJ... m("r IhC' Ia<;l ten ) ears. ho\\("\\."r. has rC'waled consistent
differC'nces bet\\een lo\dund and highland (above appro:..:. 1000 metreS)
fOlm .. of .. L. lIIon/mllllll ·· " femaiL'" of thC' high ele\·atiom, are ~maller. and all
adult~ imariahly show a chalk) white eye-ring and haye plain. gre)ish
hellie~: 10\\ land fonm han" larger female~. and all adult~ have a bold pattern
of hlack and \\hite on the underside and no di~cernibk q .... -ring . A small
perccntagC' of Sarawak rrog~ from thC' lowlands have no bell) pattern. Thc .. c
and other diffC'rences bet\\een Ihe kl\\ land and highland forms h:l\'e required
re\'ision of thl' .. p~'cies names. Based 011 t~ Pl' ~pcciml'ns collected more than
half a century ago. lh~· names as~igned to Ihe~e ~pe-.:ie~ are l .epTOiJr{U"hiulII
(lhholli for the 10\\ land fonn and Lep/ obrachilllll 1I/OI!fo/IIIIII for the high
e1e\ation species. Two other sp\.'-.:ie ... L. hnulriL"l. . .\(!ni and L. lIigmps. are
kllO\\1l on l) from coastal areas of Sanl\\a"-
Adult~ andJlllcniks live in the fore~tlloor karlittcr. \\hcre the) feed on
rather large in~ech. The frog'i sit peculiarJ) erel·l. but bO\\ 10\\ into it crouch
when disturbed b) a heu\ ~ foolstC'p or a tlashligh!. With their ~knder. .. hurt
hind limbs. the~e frog~ arc t'apahlc only of shan hop~. The c<Jlb of mo~t
"pecil's arC' unmusical and loud (an:-ording: 10 '\If R. Malkmus ). Adult~
migmk to ..,mall or medium-sized str. . . alll .. to breed and lay their eggs in quiet
pools. The ladpoles remain in side pooh or opcn pooh of ~trC'alT1'. hiding
under web during the day. Tadpole ~ feed on dead IcgTtation. \\hich Ihe~
snip into bib 1\ ith their large. hC'a\"y beaks.
Description- adult : All 'pecies are of moderate ~ize . with female.~
larger than male~. All are ~tOt·J...) rrng .. with slender. short hind limbs and a
head that i~ slightly wider than the body. The eardnllll i~ visible. The tips of
th~' tinger.., and toe~ are rounded. ex.cept ror , .. lIigmps. \\ hich has di<;[inclly
l·uned and pointl'd fingers. The slin i~ ~mooth. \\ ith i.I CUT\l'Ll ridgl' o\'cr Ihl'
eardrum. All ~plTie.., ha\'e dark brown 10 black -.:u\ouration. uwally I\ith a
fain!. comple'( darker paltcrn on the top of the head and bad,.
"Iales: 35 75 mm.
Female .. : 37- 95 mm.
'I~'ldpole : The tadpole~ of all .. pl·t·i ...... arc brge. hea\'~ -bodied and oval in
bod~ .. hare The tail i~ long. ~hapl."'d li1...;.: a narrO\I leaf. and taper~ graduall)
10 a hlunt tip. The tadpole~ bet·ome heal il) "polteLl II llh hlad as thC'y grnl\ .
49
.V!EGO PHRYI DAE: I.E PTOA RAC HI U:\!
The ;t('1Uai patl ... rn of ~pottil1g \arie~ \\'ilh the spt>eies. Ju'-.\ hefnrc
m ... t:lmorpho"i~ Iht' tadpoles reach 70 90 mm in total kngth.
Leptobracltill111 abhotti Ccxhran

1.0\\ land Liller frog (Fig. 15)
Description : A ~toek~. broad-headed frog with hLllging C~C~. The legs

afC ~hor\ and ~Iender and ~eem almo<;{ too ~Illall for a lrog of this size. The
lot's ha\e almo~t no \\·ebhing. The ..kin i .... moolh. The h...ad. back. ~Hld side ..
are dark bro\\'n 10 hlaek. \\ ilh a f:.linl dark pattern u"u ally e\'ident onl~ on lhe
lOp of the head. The hcll~ i~ hea\ il) marked \\ ilh a hold hlad and \\ hite
mOll il ng. SOTlle indi\idlJaI frog" from S~lf<l\\ak lTla~ ha\e ,I \\hite or gre\
unmarked bell)
t-.lak .. : -U-75 mm.
Ft'male~: 60 95 mm.
Tadpole : The t3dpok .. ar~' Jargt'. re:lching 75-l}() llllll h~ lhc linw they
are read~ to 1l1clamnrpho .. e. At hatching the) 3r... pak brO\\l1 to stra\\-
Fig. 15. Lepfobmcliillll! Ilhh01l1 Cochr;1n .
50
MEGO PH RY IOAE : LEPTOR RAC H lU~1
(:olnurcd. hut gradually darken into medium brO\~n and develop lurge black
~pots 011 the body and rail. The fir~t ~pOb to appear arc tho~e at the end of the
body or base of the tail.
Habits and habitat : Ju\enik~ and adults li\·e \\·ithin the dead leaves and
liner on the tloor of primury and old sCl·ondary· growth fore~t~ bclo\~ 1000
metre~ ,lbove s("a k\(~1. The species has no! been found in s\\ ampy fore~t.
Their food consi~ts of large insect, and other invertebrates. Adul ts mm·e to
small or medium-sized ~treams to brecd und la~ their eggs in quiet pools. T he
male~ do not form calling aggregations. and it is unusualtu sce more than one
or t\\"o maks on u given night.
The tadpoles hidlO' ll11der roc\...<' at the hotlom of pools in the centre or at
the ~ides of stream~. The) feed on dead and decaying \'egetation falhng into
the ~tn.'am. and can ~nip off large bib \\ ilh IhlO'ir heal, ~ bcab.
Call :.1\ .. ingk. re<,onnnt. squawking note.
Distribution : Known from all parts of Borneo below 1000 melres abO\e
~(;a le\"el.
Leplohracliiflln hendricksolli Taylor

Spoiled Litler Frog (Fig. 16)
Description : Like the Lowland LitleI' Frog. \\ ilh \\ide head, large IO'}CS.
short leg ... and almo .. t no wehhing. The species is more bro\I, nish or reddish
than the LO\\ lund Litler Frog. bUl also lacks a compicllollS dark belly paltern.
Tht' llnder~ide i ... \\·hilt' or creurn -colollTC"d \\ith many small black dots.
,\-1ale s: -J.5- 50 mm.
Ferna1e~: 0:\ 1l11l1.
Tadpole : Large and hea\")' a~ in lhe LO\\ lund Lilll'r frog . Body and tail
puk hro\\ 11. \\ ith many small black dot~.
Hahits and hahitat : So far this "pecie..; has only been found in flat.
"lo\\amp~ fort'~ts . Tadpok .. 11\C In quiet pools of slow flowing streams.
Call : UnknO\\I1.
Distribution : Kno\\11 onl~ from area~ ncar thc coast of Sara\la\....
,'Ithough not ~et reponed from Kalimantan. It pmhahl~ occur, in the
,\\;llllP~ 10\\ lands that cross the \\·e~lern and southem parl~ of th:ll sUlIe. [t
mu~ abo occur in the peaty areas near the ~outh\\'e~lern ,:oas! of Sa bah and
in Brunei . Thl' .. pccie~ al~fl occur~ in Penimular :-' Ialaysia.
51
\ IECOPHRY Jl),..\f: LEPTO BRAC HI C \I
L eplobrachillll1 1110 llfalllll11 ( F i"'l'h~rl

\ l l1nt.ul<..' Lil1t'r hog ( .... ig. 17)
Ut'o;cril)liun : Lil-..: thl' 1.111\ t.l1ld L iner hog in h..:.ld 1\ il.hh ~nLl hoLl)
... halle, \\1th Ihe ...ame und..:r-'l/l'd hlllLlllrnh.... The ... kin i~ ,nKIOth. ColllmatlOn
j, III-l' th..: LO\\ bnd Liner Fwg l'\t.:l'plll,uall~ hghler and more bWI\ n lh.m
blad Th..: dark pattern on Ihe bad, and head I~ IllOTe (,llIhpiCllOth than mlh..:
11.)\\ land Illnll. t\ naITO\\. bluj,h \\hit~ ring l'un~', around lhe eyeball and i,
normall~ li'ibk at the edge of the e) did. T h..: b..:l1~ i .. aillay<; a dirty gre)
"ithout 11l~lrling"
~\a\e,: ~6 ·63 mm.
Femille,; 50 65 mm.
Tadpole: Like Iho,e of thl' Ill\\ IJ.nll lorm.
Hahit " and habitat : Thi, '>Pl'l'il:' 11\ l" ill ,ubmollwne :llIll montane
for<;"h highl'r than 900 melre, a~lll' '-l'a k\l~1. Thl' highe .. 1 re..:ord lIe I-no\\
of i.., 1680 ll1..:trl" O lher h;lbil' a, In 11.)\\ land r,)m1.
Coill : A ,,-,ri..:, of louJ. ,qU.111 king nule ....
Uistributiun : R":P(ll"I..:J i"rolll 1l0nhern. nonh\\e,t":ll1. and ,()uthlll'"tern
I3mneo. Pre .. urnabl) lll"l"Ur.. tht"ilughnul Borneo al'on' 900 llll'lrl·'.
'lOl l': \ 1"I:'('t'ntl~ Je)lTibl'd 'pl'..·Il". Lt'{J[(Ihl"({('hilll ll !.!lInllll!.!<·II,·/\. UJ-
O(."l·ur.. ~~ilh L. lI/ol1lalllflll on \ \nun! "IIl..lhalu and h ...liLl to dd"fl'r from ,_
II/OI/f(1II1I1II (lnll b\ it .. ('alL
MEGOPHRYlDAE : LEPTOHRI\CIIIU~\l1
Fig. 17. rl'f)I(J"radlillllllllolllmlllllllri~..:hcfJ,
Leplobrachiul11uigrO{JS BelT) & Hendrickson

BlaL"k-c)cd Liller frog (rig . 18)
Description : Slllali~r and II ith narrower head thnn other "pecic". bUI ~lili
a Slock~ frog wilh ~hort hind leg~. As in the other ~pccie" 01 [hi" genu~. the
tOe<; arc almo~l without II·ebbing. The finger~ halc a tii'lincl. ~harp ClllTe
llL"ar lhe lip~. whieh nre pointed. The ... harp, hOll) linger tip~ can a..:runlly hurt
a p~r~on gra~pjng one of thL"~c frog", S"in ~Illooth or II ith low ridges. TIll'
l'olour i... gre~ to hroll n. II ith a compicuous pattern of black . OIal SpOh on
upper ~urL1cc~. The ~'~e i~ d~cidedl} b1:J~·k.
~ l ale~: :\:'---1.0 111m,
Fe1l1alc~ : 37-50 mm.
Tadpole : Shaped like those of other Lt'flto/Jr(l('liillll/. D i~ti nguished b~ a
golden "tripe dOlln the centre of the back. i\l:l\imUlll ~ize 75 mIll.
Hahit.!. and hahitat : Thi~ "fX,Tic\ ha~ bcen found in both flat and hill)
fore ... t... and in ~\la111r) terrain . TJdpok .. li\~ in leI";' slO\dy mO\ing: \\all;r.
Ca li : LTnkno\1 n.
Uistrihution : In Borneo . it i~ uncommon and ha<; hL'~'n r~pol1ed on l;.
from C(la"lal area~ of Sar:mak. The specie" abo on:ur . . in I'enin~ular
:'I131;\\ .. ia.
I\lE(lOPII RV IDAE : LEPTOLA LAX
Fig. lit L('/'/ohm, liimll 111,,1"01/1 Herr: k l-kTHJrick'on,

- -- - - -- - - - - - - -- -
Genus l.eptolalax
Sklldcl" LitleI' Frog..,
1 lea~t tJ\'e ,pecies 01 thl~ gL'nu, occur In Romeo. All ha\'e the same
A general appear;:lllce and thL' ,amc gcncrul habib. T he,':- ~lre frogs of
the forest noor. tho ugh ()cca,ionall~ an indiyidual \\jJI perch on a
herh or a 10\\ hranch oj a \hruh. The~ feed on "mall il1sect~ and oth~T
lll\cT!ehrates: thL' pn:~ j, ran:l~ more Ih~lI1 10 mlll 10ng.l\dulh n\O$' 10 !>11la1J
$r m('Jiulll -\lIcd . ..:lear ~Iream~ to breed. There Illa~ hL' man) adult.., al a
",n:am on a particular night. hut they do not group together. Tht' calb ('omi,t
llf a , ..Til', of pul,ating . high-pitched notes. \;U~ lllg ,lightl~ in the number of
nOle:'> per calJ and in pitch laccordmg til Prot'. ~1. t>. labui). Although egg~
ha\e not been found . \\e pre,ullle thalth('~ ,H(' bid under rocks. The tadpole,
\\!"iggle into the C'fL'\ lCL', h('t\\":('l1 nl<.:b on the bottom of riffle" al1ll are
reltlarJ...ahl~ c!\)ngate and ,knder.
Description- adult..,: Slender frog~ \\ ilh the 'wad not lll lll'h \\ i(kr than
tIll: bod~ . ThL' t'~e i~ \er~ large. ib dWJIleter u\uall~ longer than the ~nOlll.
Th~' eardmm i, \ i,ible. The limh' <He slender .md the hllld limb i, \er) Jong.
Finger, and too.:, h'l\e rounded tip.'- 3nd thl" t()L'~ an: all11l),1 \\ itholll \\ebbing.
~lEGOPHRYIDAE: I.EPTOLALAX
The ~\"in ()f lhl: had, and .. ilk .. i.. u.. uall) ft1ugh \\-ith ~mall rotllH.led or
elongate lump~ and ridg ... ~. Th ... bell~ i~ ~mlloth. The colour i~ gre~ 10 dar"
hrown. usually with a pa!lern of black ~pot" on the back and sides.
\1;\lc .. · ~6- _1LJ fllm.
Fermllcs: ~ 1-----l8 111111.
Tadpule : T:ldpole~ are long and "lender and h,l\e a hea\'ily Jl1Ll~c1cd tail
\1 ith \'ery 10\1 fins. The eyes are small. ThL'" entire tadpole i.. Jar\.. gre~
\\ithoul mar"ing ... Although lhc~ hah' hca\). blac" bca"~ like tadpole~ of
LCplObr(/chilllll. their rol\-~ of teeth ar ... I1lw.:h reduced. Jmt before
rnel:!morphmis the t:ldpoJes are 50- 55 mm long.
Leptolalax drinK; Duboi~

Dring', Slender Litter rrog (Fig. 19)
Dt'scription : ,\ slender. large-e) ed frog. II ith moderatel~ long ... lender

hind limhs. Toe .. lac" \-\-ehhing. Tlw .. c frog" ha\~' ~mal1. rollnd lumps Oil the
bac" and .. itk ... lind c..,pcciall~ on the lOp or the ~noUl. The back and lOp of
head are grc~ ish brol\ ll. often II ith tinges of reddish brown. There are usuall)
blad, markings Oil the back and head. but often these are obscured h~ the
Fig. IIJ. Lt'j!IOfllfll_1 dnngi Duhoi'".
55
/l.1EGOPHRY IDAE : LEP'rOLALAX
dark backgrOllJ1d. pani.:ularl~ on the head. The .:he~t and bdI) ha\~' ~lllall
olack ~pO!s. The elbO\\ ma~ be light ... r. but i~ not !>harpl~ !>et off from the
colour of the reS! of the forelimb.
l\ l ak~: .'(>-3) 11\11\.
rcmale~ : :;7--1.8 111m.
Tadpok: See under genu ....
Habits and habitat : This species Ji\es in primary and old "ewndary
feoreQ, in hill~ to 11\oulllainom ("oullIry from 200 to 1KO() Illetre" aoO\.: <,ca
k\ cl.
Dis tribution · Leprole/fa.1 drillgi ha~ been reponed from the northern pan
of S:lr,mak. from the \\hole of Sa bah. and from nonhea~tern KalimalHan .
LeplOla/ax g racilis (GUnther)

Sarallah Slelld~-r I.iller Prog tf ig . 201
Desc ription : Shape similar to the preceding ~pecie<;. Wehbing olll~ at

ba~e of toe .... Sk.in rough abOl·e and all ~idcs. 1\ ith ~l1lall. round lump, and
.. hon. 101\ ridges. The frogs nre dark Oroll n aoon":: and (1Il "ide,_ II ilh oO".:un.-
dark spot;; on back and he-ad . The- "idc~ hal L' light and dark .. pob II ith light
Fig_ 211. L"I110/<I/<I.1 l.mICili, IGlintheri
56
MEGOPHRYIDAE: LEPTOLAI.AX
cenlre~. The l'hesl and belly are hea,"ily marked v.. ith dar\.. brmvn or hlad
.. pot~. The forelimb has a light cream area at the dbo\\. ~harply contraMing
\\ ith the dark upper and 10\\ L'"r am1.
J\ lak .. : J )-.19 mm.
remales: ";0--..+8 mm.
Tadpole: See l!nder genu~.
Hahits and habitat. This spe(: ies l.ives in primal)' and old <'enmdar}
growth rore~b in hill} (·ountry. Breeding takes place at small to large .'>!reams
with clear water and sandy to rocky hottom~.
Distrihution : Thi~ ~pecic .. ha~ b<.."cn found throughout Sarawak hc1o\\
500 metre ... and in north celllr:ll Kalimantan.
Lepto/a/ax hamidi Mats ui

White-bellied S1end('r Lilt('r Frog (fig. 21)
Description : O\eral1thi~ ~pecies i.'> quile similar to L. pierI/I (sec p. 58).

The lo~'~ art' \\ebbed at the base. The skin is <;Jllooth. The eolour:ltion i ... abo
.'>i..milar 10 that of L. ph·III.I. with light-bordered hlack markings on the
FiA . .!1. 1.. 'pIU/II/1I.1 hUlllidi 1'I]al'oui.

:-' lEGOPHRYIl)Al:::: LI:.PTOLALAX
flll'UiUIll I1rOI\ n had.. and h<.:aJ. Th~" bo.:ll~ i~ Ililite or cream-o.:oloured I\'ithout
~p\lh . Thl: I()fl:limh U(ll:~ flot hal~" a ~harpl~ defineu light area at the dbO\\.
\1ak ... : 2Y-31 111m.
Female,: J7--J.l mm .
Tadpoll': L"nknrl\\ 11 .
Hahil ~ and hahitat: Thi) ~ pecie~ li\ 'e~ in h111~ prilllar) and o ld
~L"nl11dan lore,t fr~)m 100 to 250 Illetre, abOl'(' .. ea Ino.:1.
[)i <;t~ibulioll : Thi~ ~pecie' ha~ been eolkttl:d 'l.n the ~outhern half of
Saral\al.. and in een!f:ll Kalimantan.
Leptolalax maura Inger. rVl al,darin. Alim & Yambun

l\lolltanl'" Sknuo.:r Li\\l;'f Frog
Desc ription : Thi, l~ a rdatild) ~tock). ~Illall frog. The toe, aro.: Ilebbed
"nl) at the ha'e . The bad; ha!> many 10\\-. rounded lump~ and ~lightly larger
\Ian ... on tho.: ,ide ... . The ~·01our i~ \-ery dad.. hrm\11 \Jr black: there i... no \'i.,lhk
ratlo.:lll on the ba..:k. This i... the on l) ~rL"eie" in Ihe genu~ that is ulllfnnnl)
darh brov.n 011 the underside of the head and hind limb~. The foreilmh d() . . ~
not hale ('ontraqing light and dar!.. l'olouralion.
'\-1ale: 26 mm.
Female 32 mill.
Tadpole : l-n!..Jl(l\\ n.
Hahit~ and habitat Thi~ specie., has heen ('olkcled onl:- once . making
it uirti("uit 10 drall condu~ions abollt hahit, . Th.: on 1) ('ollecting site \Va" in
monlan.: rore~t at 1850 III on l\ lnunt Kinabalu.
Distribution : Kno\\n onl) from :-'I ourll Kinabalu in Sahah.
J,eptolalax pictlls rvl alkmu ...

Painted Slender Littt'r Frog (Fig. 22)
Dcsni ption : Similar iII ;;hare 1(\ I .. g/"ociIi5. -"he toe~ are \\l'hhL"d al lhe
ha:.e . The ~kin is ~l1ln01IL \\llh al mo~t a fel\ lump~ on the ~idL"~ . Tho.: L'lllllUf
I~ light to medium hrO\ln. \Iilh ("on~picuou~. light-edged. dar!.. marl-..ing~ on
58
r-.IEGOPII RYIDAE : LEPTOLALAX
had, and head. T he .. ide 01 the h~ad ha~ .'oharpl) defined hlackbh bra\\ n
'pOb. The heJJ~ j, \\ hill" \\ ithuul "'pOb. The forelimh lack .. cUllIra ... ting light
and d.:lrl.. coi()uration.
:o. 1;lIc, : 31 - 36 mm.
Fema!e~: .~9~ 7 Illlll.
Tadpole : L'nJ...no\\ n.
lIabit.., and habitat : Thi, 'llCCic<,. 11k thc other .... lhe~ in prirnar) and
old ,econdar~ grO\\Ih fnre,1. [t dOl'" mil I.X:CUl" at 10\\ eie\-mion ... 11<1\ ing been
found frorn S(X) to 15..10 ml·ln:~ .
Dis tribution : \\'e.'otern pan .. of Sabah.
59
I\IEGOPHRY IDA I:: : I\ l EGQPI1 RYS
Genus J\1egopltrys
Horneu t·rog:-
Il 0 ka(ure~ ot the D(lmean "pecie, of HOnl .... u Frog" q:lI1d out: large.
T II ide hl·au .. Jr~~ pOinted pr:lljl'I'Unn" o~ "kin. from tllL'!f eyelid ... Ihe '0·
called "hum" T he ~pecJe, alt..' mIl Idenllcal III Iht' lurm w!..en h~
(hew ··horn .. ··. but :111 'Ik(ie<i hale (hem. Th..:ir large he:ld .. and \er~ IllUl'
mouth .. \lIgge<;[ :In ilbili(~ 10 W\...e In rel:llil"el) largl' pn':~ - llhl{'h i~ exactl~
II hat (he~ do. Snail, \Iith ~helh 50 111111 I\ide are no prohkm for their
('"nOf1110U, mouth .. L:lrge pre~ item~. mo .. \I~ in~ect;;; but aho (:entipcuc, and
,,\.:prpioll~. are the: nrk lor Ihe~e ;;;pede".
The <ilender It'g'' an..' incapahle of lilUnchrng the"e heal ~ ·bOtIit'u
lerre"trial frog .. all il~ from a pOh:ntral enemy a~ the Huml'u Frog<i call onl~
l10p .. hol1 <li"l;ln(e ... Since a largt' kilp '" OUI of the que.,tiol1. lhe~ rel~ Oil
call1Oll fbge. All hale lilt' genera l brol~tl ur n:uui"h hrown colour of thl' dcau
leale<; amid"t Ilhkh llw~ "pend their lile ... SCI..:ral .. pccie~ hal'e long. kl\\
ridgl'" on Ihe bac!... II htr:h Tl'''eTllh]e the leill!» on de:rd 1..:;\11.'''. \Ihile the .. harp
"hom,," and "nout t,r .\lI'~uJ'hr:'o \ I/I/II/Ill re~t"mble tht' ~·ur1cd. pointed leaf
Ilp~. ,\In;oplrry\ !"I/H"{//~/illac hOl" ~lI.ld proJcction" from 11ll' "I.,ln of the head
and bm:k. bn.. akrng up the outline ot Ih..: hud~.
ju\enile, and adult .. Ih'e Oil the fon.: .. ! nuur in primary and old ~e,:ondar~
gn1\\th fore~h from (10 10 ll)OO metre .. aboll' "~':J le\el. During the da).
jUlt'nilt"~ retreat benealh dead ka\e". while adu1t~ hide under dead log~ or
ttrge rocb-. Adult ~ mUI": 10 Ihe hank .. of "m:111 to ml'diu1l1 -"i/l'd ;;;tream .. (O
hr..:..:I..I. The<ie migralloll" 1l1u'l tx, rather <;10\1. cOIl~jdt..'ring hOIl far from
",tfl·alll .. lhe adult ... ro;rm ;md Ihe ... hmllh"t3nce Ihe) cOler in a "Ingle hop. The
loud honk ot ,lh',!:o/Ihn<. /wsl/Ta in aUlance of hea\~ min l~ Olll' \)]" the
char:h,:teri , tic "{)lInd~ of tht" 10\\ land r;lill lore~1 III 131(' ;)f(ernoon or el"ening.
J::£g~ are laid in poo).. a\ tilt" edgt" of "tn~arm and the tadpolt'~ li\'e in riflle,
o r quit"1 !>ide pOI..II". Thl·"e t3dprok" h:l\e a remar\...abh... II;I~ of feeding: Ihe~
hang from the ~ul"fa~'l' \\llh Iheir fUllnel·ll1011lh~ bee bl'l(lII) "pread and filler
~'ul ,mall blh of organic maHn cauglll III Ihe "'llrface IiIm.
Descriplitlll ;ulults: .lfcqol'hnl arc medilllll- .. izt'd 10 large trng ... \llIh
\er~ largl·. Ilid(' head .. :ltld ... lend..:r. relatf\l'l~ .. hon hind Jimb ... T h..: tUl·" arc
not \,ebbed In :.tIl hut (lne Hornean "pl!L"1e .. Clf. i1ril1~i). the he.rd \1 idlh l'qu:rI ..
all110 . . t half Ihl' eomhrncd ht'"ad·plu,-btl{l~ lenglh. TIll' diameter of the e~ e j..
J.bnut equal 10 Ihe knglh 01 Ihe 'nou!. l he upper l'~l'1id h:h a projeclion o f
. . \...in 111 ,ome 1\lfIll. l.rr~lI1g amon~ the ~Jk'eie,. Thll!..('l1cd ~kin pal1iall~
60
MEGOPI IRYIDAE. MEGOPHRYS
obscures the eardrum in sc\eral "plTie", Mo~t species han' thin. narrO\\
ridges of skin at leaS! on lhe i"ronl parI of the back. All specie .. ha\e a strong.
curved ridge or fold of ... lin from the eye, cun'ing mcr the eardrum and
ending at the armpit.
Males: 39- 1O..f mm.
Females: 55-125 mlTI,
Tadpole : The tadpoles are slender and dark \\ith blackish JI1arking~ on
the tail. Their most remarkable feature i'> the mouth. In all Borneun wdpoles
but those of Megophrys. the mouth points downward and i.. al the bottom of
the <;nout. In tadpole ... of MCNOP/1Jy.l. the lips are \\ idelh:d lmd turned up\\ard,
so thaI the mouth forms a funnel at the ~ame Ie\ <.'1 as the lOp of the head. Total
length at metamorphosis i~ ..fO---+:1 llllll.
Call: The \nice of all th .. ~pecies recorded IS Imld, har~h. and usually
cnn .. lsl, or a ~erie~ of notes emitted at intcnab (according to Mr R.
r-.lal"-.lllu~).
Megophrys baluellsi.'i (Boulcnger)

Kinabalu Horned Frog 1Fig. 23)
Desc ription : A ~tocky ll1edium-~iled to large frog. with short, slender

hind limb ... The head \\'idth i<; almoq e(jualto half the head-plus-bod~ length.
Thc eardrum i~ \·isihle. i\ short. narro\\. triangular "horn" project.. from the
upper eyelid, which aho ha .... e\Tral ~horter. \'enical projeclion~. The back
has one long., interrupted. thin ridge of ~kin on each side and a fe\\ scatlered
short ridges and round Illmp~. The upper ~urfacl'\ are brown or rcddi"h
hro\\n. uwall) \\ith a dark triangular mark hl't\\een the eyes. The side of the
head i.. dark brown. sometimes \\ith 3. llaITO\\ ~ ello\\" stripe at the edge of the
~nou1.
-+1---+5 mm.
r-.lale~:
remale~: 55-70 111111.
Tadpole; See under genu ...
Hahits and habitat: This species 1iw~ 111 montane forest~ fwm 1200 to
1900 mctr<.'~. See :llso under gerHl".
I1btribution : KnO\\ n from lll()untaill~ of \\e~tern Sa bah and northern
Sara\\ ak.
01
~IH;OP HR YlDAE. i\ IEGOPII RYS
Fig. 23. Ihrwphnl ha/ul'ltIis (Huukngcrl
,\/egophrys drillgi Illgl:f. Smcbing & Tan

Dring'~ Hornl!u Frog
D('!>criptinn : A ",l(ld,~ frog. with . . k'n<kr !lInd limb .... Thl! \\ iulh of the
head i... ,Ibout one-third th.: h:nglh of heall~plu .. - IxxI~. The eardrum i ... panl~
hIdden b~ ... "'in. The upper ~~l'!id h~h a nal'J'(m ... hun. projectmg "horn" , A
... hoft. \\e~ll-. fold oj" skin run~ ba.:1-. from lhe eye p:lrl \\a~ down lhe bod~. and
~omcLil11e ... a pair oJ \\cnl-. l'iJge~ form ... a V on the front or th ... IOr"o. Coluur
gn:~ i"h or redlli"h bnm n. sometime ... \\ ilh ;\ dark triangular mark betwe~n the
e~e".
:'>.1,lle~: ~3---4 7 mm.
F... mal ... ~ : 55 !l11Il.
Tadl)ok Cnl-.llo\lll .
Hahib and hahit:tl : Julian Dnng. the di.,cO\.:r.:r 01 thi, ~pet· k ... , found
auu!h at the ...dg.: ... of a ... 1...ep. "'11 if! ,m:am 111 mont;ml' 1",lrc ... ( at 1800 mc1n.~'"
Distributio n ' Krl{\\IIl"'O faronl~ Ir01l1 i\ lollnt :'>.tulu. Sara\lak.
MEGOPHRY[I)AE: MEGOPHRY S
,\1l!golJhrys edK'ardillae Inger

Rough flomed Frog (Fig. 24)
Description: A <;toc\..").mediunHill.'d frog . \\ith -;Iendcr ll'g'. The width

of thl' head equal, hall" the he:lu-plu~-body length. The l':.IrdrUlll i<; mo<;tl~
hidden by "\"' in. The ··horn·' h a drawn-out. ~kndcr. triangular prnjct·tion
from the l') elid. The hl"ad :lnd bod) ,Hl" em crcd \\ith many Irregular. round
or u bit e1ongatl"d. ridg~" or projet·tiom of ~kj ll. including olle pointed
projet'tion at thl" comcl":> of lhl' mouth :lIld sewral \ ertKal ones on thl" l") clid.
The general colour i~ clay bro\\ n. wi th blad hal"', on the ~j de of tlte head and
J.ero" the [imb~. A large dar\.. mar\..lIlg u<"uall~ cmer.. much of the back.
:\ I.t!e): 39-0: mill .
Female ... : 7()...H :! mill.
Tadpole : See under genu,.
Hahits and hahitat : Thi~ ~pecic, li\ l" in hill) or ~tccp l()wlllnd rain
fore,!. Ju\eniJes and adult, !cell nn Ih" fores t tloor ami hilk under dead
1c;'1\~" and log:<. during the d,IY. Auult!o mo\e to small. clear. nx.')"y streanh to
breed.
Fig, 2.'- \Il'~(lI'''ns edl!<lrdmae Inger.
63
;...IEGOPHKYID,\E : MHiOPHRYS
I)j ..tribution : Sarawak ;md \\~:'-tern S;lhah. ben\l:~11 .:!()() and 600 metn:~
abO\..: . . ea kId.
Jlegophrys kobayashii ~ I alkm u <;, & r- hll\ui

\ Innw.ne Homed hog
I)csnilltion : A ,tock~. 1.lfg.: <;,ped~~ of frog. \\lth .. hon, rc1ati\'el~

~kl1fJer leg~. The \\ ilhh of lhe head I ' half thl' Ilt:ad-pluybody length. The
"hom" i~ a I\id..:. pOltlted. triangular proJ~c t ion I'romthe edge oflhe e)did.
The tip of the ~nollt is poilllL'd. but there i.. no projection oj ~kin a~ in
JIe-.:ophrn 1ll/.III/fI, Th~ eardrum I ' paniall~ co\'t"red by ~kil1. Th.: back ha,
\III) Jong. narrO\\ fold, of ,kin, Th .... colour i, medium to dark bro\\11 on the
b:h.:L lhuall~ Illth a fell 'l'al1ered bl:ld ~pot:-.. The under . . ide fades from
broIl n II lth darh'l' 'P0b on th.: throat a nd eh..:,!. 10 light..:r gre~ i . . h \)11 the rear
of the alxlomen. U,ually then.' i, a dark ,m:a around the l'~ e on the: . . idC' of the
head.
t<.lale,: 93-101 nUll.
rl'l11a1c,: 99- I09mlll.
Hahilat : Thi~ large memher of the genu~ ha~ been found onl) in
mont an..: fore~1. In)1I1 IDO tll 1675 melre, ahO\e ~ea leH'l. lntljliduals ha\!:
been ~L''':11 1\ ilhin Ill.: leaf lilll..'r of the 110m of tho~1..' l\Jrl..'<;{~ and along the
hal1k~ of rocky 'tr~'am, flO\\ ing through them.
Db trihulillll ' Kml\\n ('Inl~ from the mountaHlOU~ \\e..,tcm pan of Sahah
from :-' Iounl KIIl;lbalu ,outh\\arrJ through thl' Crocker Range. Probahly
occur, in norlhea,tern KaliTllant;m .
.lIegophrys IIf1 Slf{(l (Schkgd)

Bom~,111 HOl1ll'd Frog (Fig. 2:'i)
J)('<;c ription ' \ \er~ ~ tv!.'k~. medlUllI-;;lzed 10 large frog, \llIh "hon,
~Iendl..'r leg.,. The \\idth of IhL' hl'ad i, h;1Ir the head-phl'-hod~ length. T he
"horn" h a ler~ llide. pointed. triangular proJl'l'tion from Ihe edg..: 01 the
..:~('hd. Thl' .. peeie., al,o ha,;1 'lmilar. pointed rrojection from the tip of the
"nou!. rhe h;.ld 11;1' 1\10 pair" of long, narro\\ fold, of 'kin_ The colour h
fight (:kl~ to r..:ddi,h broIl n, "omellllll', \\ IIh one or 1\\0 dark ~POIS on Ihl'
ba.:k. U<;uall) Ihere l ' a II idl'. darl-. bar on Ihe ... idl..' (It the head helo\\ Ihe L'~,,:,
6-1
i\'1 EGOP I-IRY IDAE : \ IEGOPHKYS
i\1.lk,: 70- 105 Illlll.

h:"nlaie .. : 90 125 mm.
Tadpole : See under gl'nu,.
Habits and habitat : Thi, ,pecie~ appear, to be the mo .. t \~idl'1~
di~tribllted und common one III (he genu,. It li\t~~ in flm (0 ,(eep rain rore"I~.
from near ,(,3. Ie\cl \() abuut 1600 metre:.. The call is a loud honl... given by
male .. a( the approa(:h of a \lOrm. Adult<; br(,ed :1I 'tn.·.lIlh ha\ ing \\ eak to
mOlkra(e CUlTelll~. Tadpole~ h\e III riffle ... and are frequently found hiding
where root m:lIs of hanl.. \ egctalion trail in the wmer. See .11<;0 under genll~.
Oistrihution : Found throughout Borneo. It al<;o OCl:ur, in SUlllmra and
Pcnin,ular \ f ala~ ~i:l.
65
!Palllily ifJuJonidae
True toads
Thi" family ha~ a world-wide distribution. hut 8 0111(0 and Soulhca~l A~i<l
have u dislim:liH' ~cl oi" genera amI species. All h'l\c rough. \\aJ1y ~k.in.
Genus Allsollia
Slender Toads
hese relatively small. slender toads have the dry. rough skin of typical
T toads. hut lack the large parotoid gland behind Ihc eye and above the
cunJrum characlcri~lit uf most loatb. Allhaugh the genm occurs in
the Philippine Islands and Peninsular Malaysia. Borneo is where most
species (12) occur.
The species of Al1.\OlIia are remicled 10 primary and old secondary
forests. including selectively logged forests. in hilly to mountainous country
from Ihl' lowlands to 3000 metres above sea level. Adults live on the foresl
llooT bUI are occasionally seen on le<lves of herbaceous plants. Ants and OIher
small insects make up the diet~ of the four ~pecies that ha\'e been studied and
probably all the species haye similar diets. Adults mow to dear. rocky
streams to breed. where they perch on rocks. small shruhs. or herbs ncar
rapid~. Tadpoles. with the exception of tho<;{' of A. /epropus. live clinging 10
rock!. in ~\\in t·urn·nt~. ~itu<lliom requiring streamlined ~hape.
Description-adults : All ~pecies are rehlli\'ely small . with a maximum
size of 6.+ mm. Males are smaller than females. 20-50 mm \'ersus 25-04 mm.
Most species ha\'e rather slender. elongate bodies and relatively long legs.
The snout usually projects heyond the mouth. TIle eyes arc large. their
diameter equal to or longl'r than the snout. The small eardrum is \isible. The
tips of finger ... and toes are u .. ually rounded. bUI the oUler fingers ha\'e
~lightly \\idt'ned lips in ~e\'eral ~pet"ies. Webbing of the toes varies greatly
among ~pecie~. All species have rough skin. most species being co\'ered \\ jlh
small. rounded warts. There are no enlarged parotoid gland~ hehind the eye
and above the eardrum. Colouration is usually brown or black. \\ ith indistinct
pall ern.
67
BL"r:O~1DAE : ,\:\SO:,\I,\
Tadpole: Tadpole .. \)f all ~pt'cic!'. art" .. mall ImO~lj~ je~~ than .:!O nUll) and
haw tlattt"ned bl)Jil.:~ and clIp-,hapl'd lip~ forming a ~uckcr. All bu t one IA.
Iq![(JJlIII) ha\t" distiI1L"II~ ~lr~'al1l-1in~'d hodil'\. Till' u<.ual cniouratioll i~ bbck.
Olll~ a ft'\\ ~pC'cit'~ llfe Ji"el~ to be I.:ncuuntcrcd. \Y~' prc"cnl no!!:, on
Ji~lincli\c jl'J.1l1rl'~ nl tho~t' "pC'ciC'",
AI/sollia albomacfI/afa Inger

\\"hik'-1 ipped Sklllkr Tuad (Fig . .:!f,)
Desl'ription : ,\ \tock~ . ~rn3.ll "pecit'~ . \\lth snout projectlIlg beyond

mouth. Tho.: tip" 01" all linger .. arc roundl'd. ;llld lhl' fir~l fingl'r i\ diqillCl1~
Fi g. 26. Alllf!llI<I uIIJlmli/, 'u/ui<1 Ing<'"r.

nur-ONIDA!::: ANSONIA
~hOller Ihan Ih ... <,ccond. ThL' IO~~ an~ almo~t full~ \\ ... hh~d and Ihe tip~ are
rttunded. Thl~ toad i\ r~ddi~h bro\\ n abo\e. u~uall~ \\ Ilh a thin dar!.. X~,haped
1ll,Irking O\('f lhe ~houlder\. A ~lalHi ng lIghl hand rum from the comer of the
e)e to the ,U"tnpit. TIlt're i~ oft~n ;l Itghl ~plll belu\\ the eye.
i\ iaie,: 10-2S 111 Ill.
rl'l11ah:,: :;0-.15 mm.
Tadpole : The tadpole, are ,~arJrop~.,haped . ~mall (maxImum knglh 15
mm~. and black \\jlh aliglll LTO"~ on Ihe head and bod~.
Hahit .. a nd ha bitat : Thi., h a primary rain forl''''' '>pe":ie,,. occurring
tN.'lm\ 400 melre:. :Ibmi' "en le\el. Adult ... mO\~ 10 the banJ..., of rllcdium-
.,izt'd 10 large. clear. ~\\·ift ... In:''1I1'' 10 brl'~d. Calling male" are !"t)und ..eallered
along ... tre"lll bank~. hut l'alling group~ h:l\ ... not heen "... en So Jar tadpole<,
haw hC'C'll f(lunu on!~ in large ... tl"e:llllS.
Ca ll : A high· pitched metallic Irill IhJll'an be heard Je'pite the nol ..e of
rhL'J' rapid,.
Dbtrihlltion : K ll()\\l1 lrum Brunei. Kalimaman. ami Sar:I\\;l k. but not
Sahah.
A lI,Wllill j uligillt:CI (.\'Iocqunrd )

\ Iont:me Slender Toad
Dl'scription . A 'Illall. ,kmkr 10 ..tocky toad. \\ Ilh long kg .... The toe~ arc
incornplet ... !) \$'hhed: lip~ of the fiN two to(" {thl' ,hort!;"t olleq hard)
project h':) lmd the \\ebbing, the third a nd fifth toe~ art;' abou t ha!f-\\ebbed
nnll the lourth (the longe~tl toe le"~ \\chhed. Thl' li[]ger~ hu\e rou nded tips.
Ilke tho\L' of the toes. The cnlu"C' upp;.:r surfa('e i, di'n~el) c$\ ('red \\ith
rounded oj' oval tub ... rc1t: ... or \\;Irh. Adult male., have a clu .. lcr ollurge black
~rint'., on Ihe fir'l I ulIwTl ling!;'r The upper ~u rfacc, :Jrl' medium 10 dark
hro\\ n. The ... ide, ;.mu 10\\ t;'r ,urfaee" are dark hro\\ n or b l 'I~·!.. .
\ l ale~ : 32-36 mm.
remak, : J9-t--t mm.
Hahits a nd habita t : Thl' "PC(:IC'. \\bleh i.. hllO\\n onl~ frnm de\:Jtion ..
bet\\L'en 1600 and 3-tSO 1lletrL", li\e, in montane IllO,,,) 10re'l and in the
,tull1L'd for ...... t ahmc that tOlll' rrog, ha\e be ... n 'CC'1l \)11 the ground and on
\el') 10\\ " ...getation ..... llhlJugh taupok" are unknO\\ n. judging b~ tadpok, or
other 'pecic, l!l thl' genu ... the~ probahl~ !J\L' on ,tnd among rod, In 'mall
IllrlUll1ain ... trL'alll'.
Di<;trihutio n Knm\ n '0 I:lr onl~ Imm .\ llmn! Kjnab:llu. Sanah.
69
BtJFO\,IDAE: ANSON IA
A nsonia gllibei Inge r

Guibe'~ Slender Tuud
Descr iptio n: A <;mall. slender toad. II ith n;!ulilcl) long leg~. The toes in
males Ikmaks arc unknown) are fully webbed (ex<:ept for thl' fourth. Ihe
longesll 10 the base~ of thl' ti r~. I~ hich are slightly swollen and round. a~ are
the tips of the fingers. The l'ntirl' upper ~urf::tce i~ <;ct with many round or o\'al
\1 arts or tuhercle ... There are no ridge~ on the lOp or the head. \ 'tales have a
densl' dll'.tL'r of \'er)' 'lmall black spines forming a pml on the lir~t lingl'r.
The) abo h<lll' a Ibnened. halloon-like swelling on the rem of the thigh on
each side of the anus . .\'lediu1l1 lO dark hrown on the upper surfaces. with
irregular lighter bro\1 n or yelloll i~h ~pot~ on the top of head and hack. TIle
lmdL'T<;ide is light grey with indistinct dark SpOlS .
.\1aks: J l-J~ 01111.
Females: 3-+ 111m.
Ta dpole: Tadpoll's url' black and h:ardn)p-,hapcd. as in 1110<;\ "pecies of
the genus. Maximum length is 30 mm.
Habits and habitat : This species ha~ been found on the forest floor and
on hanb of clear. swifL rocky streams in montane fore sts from 1560 [0 1800
Tm-'tre~ ahow ~ea level. Sometime'l it appears in large numbers along the
banb or ~tn.·aT11\. prc<;wllahly in hur'lts of hreeding activit),. Tadpoles at times
are con;,picuousl~ abundant dinging to rocks in the c1car water.
Distributio n: Known only from Mount Kinabulu and \-fount Tru~ Madi
in Sabah.
Ansonia J/{lIIit~jc hi Inger

Kinabulu Slemkr Toad (f ig. 27)
Desc ription : A ~mall ~1cnder. 'lome\>. hat nat-hodied toad with :1

projecting snout. The tOl'~ an.: about thn:c-rour\h~ wcbhcd and halC rounded
tips. The tips of the outer fll1gers are ~lighlly widened. lmd the liN linger I'
definitel\" shoneI' than the second. The back is c(l\'ered with ~lllaJJ rounded
and;] fel\ o\'al I\art~: usually a few wans form short. slender ridges o\'er the
~houldcr~. These toad .~ are dark greelli~h grey [0 reddi sh browll. often with
blw:k diagonal han or spots on the back. The limbs have black crossbars.
7()
BU FON IDAE: ANSON I A
l\1<lles: 20-28 mm.

Females: 28- 35 mm.
Tadpole : Tadpoles arc black and teardrop shaped. ~1,L'(imum length is 20
mm.
Habits and habitats: This species is common in rore!>t~ of mountainous
coumry. m ele\'mions higher than 950 metre~ above sea level. They arc mo!>t
oftcll scen on the banks of sm:lll. rocky mountain creeks. Occasionally. large
numbers of males are heard calling.
Call : A high·pitched chirp or ,hon trill.
Distrihution ; 50 far this specie~ has been found in the ll1oun!ain~ of
wc~tcrn 5abah. northern Sarawak. and the northca"lcrn corner of
Kalimantan.
FiJ!. 27. , lnsollia /wllitschi Inger.
71
BL'FONIDAE : ANSON IA
AIIHmil1 lepropm (G limhc:r l

Bnm 11 S1enu~r Toad ~ Fig. ~8 1
Dl'!>('ription: A .,Iemkr-hnulcu ~pe\.·ie.'> \\ilh long. thin hind kg.'>. nl~

toc., ,Ire abuut lhrt'e-follJ1h~ \\l'bhl'U 111 males . .'>Iightl) k.,~ than that in
fl'111i1lc~ . The linger'> ha\(' rounded IIP~ li ke tho<;(' oftht' toe.'>. Th~' l'l11in:! back .
... ith:~. and upper .'>urran: ... of the limbs art' rough. \\ith lllan~ .,mall. rounded
\\arb. TIl\.' top of the head u.,u<ll1y ha . . 10\\ ridge.'> . t-. I<lle ... have a ... ingle ru\\
o f ... mallorange or bra\\ 11 .'> pUll' . . under Ihe chin. T hi.'> load h ullIformly brO\\n
on the b,K)" and top of the head. and ~lIghtly dar).,er on the .,ide". The
unJef<..ide of the hl'ad i, da rk grey \\jlh a I;I1'O\\nl,h tinge. and Iht' I:hl,.,l and
1)I.:11~ are gre~ \\ ilh darkn 1ll(lt11ing.
l\!ail'.,: 3()- 40 mill.
J-ell1:dt'~ :45- 65 111111.
T.ulllUIl': Till' tadpole, are '>mall Inol l1l0rc th an 10 mill lung) and light-
hndied. with a bl<lc\.. ( w ...... or X on the upp..:r . . u1'facC' .
l'lallits and habitat: Till ... I" a to,ld of hill~ prilllar~ fore . . t o r old
,>enlt1tbr~ gro\\ lh belO\\ 600 1llC'lre" abO\e .;.ea le\el. Adult ... 1lI0\l! 10 clear.
med iulll-,j/cu '-Iream.'> for bre~'ding . t-. I:lle<, u<,ua ll y call in group . . lrom "Iream
Fi~. 28. -\ 1/\1111111 "'JI/('/1I15 l (junlh ~ r ~,

Bt;FO:\'[DAE : A:'\SO:.' IA
banks or from rocky bars in mid-"tn:a11l. T<ldpoles li\'e mainl) in

accuillulations of dead k;l\c~ in medium-~ized clear stream<, olkn ill
a~~ociati()l1 \, ith tadpole~ of Raila sigll(lfa, LimlllJlll:("/I:.\' feporil1l1s and
PCdOSlibcs hosii.
CaH: A high-pitched trilL
Distribution : Wide!;. lbtributed in Borneo in hill;.. but nOi mountainous
countl"). Abo known from Peninsular Mala)sia.
Amwl/ia /ollgidigitll Inger

Long-lingered Slender Toad (Fig. 29)
Description: i\ toad \1 ith long hind lilllb~. The ~noUl projects beyond thc
l11outh. The toC\ an: about half II ebbed. The finger" haw fOunded tip~ like
thc tOI''', The libt linger i~ at least as. long as the second. Soml' of the warts
Oil the llpper ~lIrface of the body. espenaIl) thosc at thl' ~houlder. are capped
\Iith small spines. Adult mak .. hUll' thnT or four rows of black. conical
~pines under thc chin. Thi .. toad i~ cia) brollll 10 black. The limbs have dark
Fig. 19 . . Ills,mia irJIIl!idigi/ti Inger.
7J
BUFONIDAE: ANSON IA
cros~bars on a cia) brO\I n had, ground. TIle belly is cream-coloured . heavily

~pl1!led llith brown or black.
~ I ales: 35-50 mm.
rcmak..: -l-.'i-70 mm.
Tadpol e: T;.uJpoll'\ are black. teardrop shaped . and about 20-25 rnm
long.
Habits and habitat: Thi~ toad li"e~ in primar} amI old ~econdary or
~clecti\'t'I) logged foreq~ in hill y or mountainou~ L'uuntr} from 150 to 2200
lllclrc~ ahO\C" ~C"a ll'lel. Adults mO\'e to clear. swift. rock~ ~tream~ to breed.
r.. l ale~ fOnll loo~c ealling: aggrC"gations. Tadpnle~ are most often seen in riwrs
on large ~moolh roeb hal ing a ~lron g laminar Ihl\\. The) arC" frC"qucntly
found with tadpoles of Torrell! Frog~ tgenus J '1crisro~cIIYs).
Call: A long . l11gh-pitched trill.
Distrihution : This ~pecie~ ha, been found in the we ,tern and central
pmt~ oj Sabah. in Rrunci. and mall) pl3.ces in Sara\\aL It ha~ heen found in
one IDeality in upper 1l0nhL' a~lcm Kalimantan. It ha~ IlC\L'r bt:cn ~CCll (lul\idc
of Borneo.
Am'ol/ia milll{ta Inge r

DII arf Slender Toad
Description: A ~mall ~lender speeie~ \Iitb long hind limb~ and snout
projecting II ell beyond mouth. Tip~ of outer fingers are II·idened. The first
finger i~ distinctly ,honer th:lIl the second. The toes are fully webbed in
males. ahout th ree-fourth., wehhed in females. The upper ~urface~ of hody
and l iJ11h~ arC" ("mered \~ith ~mall. round wart~. Adult malL'"~ hale (II!) or thrl',-'
ro~\~ of ydl(J\\i,h ~pin.:~ under th.: (·hin. Thi~ toad i~ bro~ln Ilith in-cgular
~l1lall orangt" or ydlO\\ ~pob or ,Ircak'- on Iht" back. The bt"lI) is pale with
black ~poh and yello\\'i~h dOiS. The ~i de of lhe head has dark and light bars.
with a 1\ hiti~h area belO\I lhe eye .
.\-Iales: 20- 23 mm.
Females: 23- 28 mm.
Tadpole: L'nknown.
Habits and hahitat : Lillk i~ knm\'ll (}rthi~ ~r'~ei,,'~ l"\e"'plth;jl itlilc,> in
hill~.lo\ll:lt1d rain forc'>t bL'IO\I 700 mctr.:,> .!bule ~e;j le\'e!.
Call : Unknolln.
Distribution : rlHl11d ~o far onl~ in Surallak. and Kalimantan.
7.
Bt;FO;-\IDAE : ANSO:-.JIA
A " son;a sJlilwlifer ( M o(.:quard)

SPIll~ Slender Toad (Fig. 30)
Description: A comparativel) ,1l'nder ~pecie~ with (he ~Iloul projecting

... light)) beyond the mouth. Tip,> of all linger~ and toe~ arc rounded. and the
to\-" are almmt free of \\cbbing. The first finger j" a~ long a~ the second. The
entin.' hack and hind leg~ are cO\'ered \\ ilh clln~pil·uOU~. large. "piny \\an~.
whkh I,:an ea~il) be felL The lOp or Ihe head and bod~ i" bl:ld, with a
~ellO\\bh or whi tish mal or di:lmond-~h:lped patch Ix'tv,,-'en the s houlder~
and a light "trcak along the ~ide of the bod). The bell) I!> black \\ tlh ,>mati
I.:n:;nll-('nloUTl'd :.put:. or cream-coloured \\ ilh heavy black marhling.
~ l ak~: 30-W mill.
t-enwle,>; ~O---lS mm.
T:ldpole: Taupuk:. are teardrop·,haped. hIJ.:k. and do not ell:cecd 20 mm
in length.
H'lbit ~ .md habitat: Thi~ toad ha:. been seen perched un ~ hrllb~ and
h"'rb~ otll~ along ,\\ift Ilc)\\ ing. de;!\'. rocky stream, in primar~ forest helO\\
700 metre,> abmc ~ca Ind. Male,> do nOl fonn calling group:., Tadpole . . cling
10 large rock . . in 'trong l·urrenh and feed 011 the film of ;11,gae grm\ ing on the
rock~.
Ca ll : CnhllO\\ tl.
Dis tribution: KnO\\n from all hill) parts of 10\\13I1u Romeo \\herc-\cr
\Uil;lble en\'ironmelll C\i~h. NOI knowll outside of Bom~'o.
Fig..'". AIII/I/Wl ~/'illtlliJt'r (~Iocquan.ll.
75
BU I-ON]J)AE.' BLTO
Genus Hllfa
Toad~
he,,, :Ire the ,o-calk'd "tru!! lOnd~". frog~ \\jth \\~ITl> '~II1. a large
T p'.\fotoid gland behind tho.: o.:~~' and ahmt' tht' eurJruJl1. ntlll<:r ,hoT!
hind limh,. amI mllall~ nano\\ bOll) cr~',h on the head. Al though the
par0toid gland and \\;!Tl, ~ecrel(' a !ll!lk~. late.\-lik.e to\in. ~'\lIdcd \\hen the
t~I;ld I, di<;lllrbed. fonlln'lId) it 1~ not dangerom to hami!e Iho.:'~' animal" a"
long ;I, on(' \\a"he'> one'" h;lIld, atll'T\\ard". The po j"oll. ho\\Io'\<:1". I'>
dangt'rtlll" ttl "mall mammak ,uch a~ l'at~, thai mi,ght t~ 10 pre~ on fo.IlI ....
T he Bumcan "pl'cic" of 81t1il. ll J.. e tho~t' ebe\\hl'rc In the \\orld. are
ielTe,tnal. ,I !though Ol'l':.t,illll;lll) Ihe GIant Riwf Toad (lillNI )l1.Hlllp('/') rna)
l'h1l1b up a <;Iopmg dead ~I\lmp, Diel, or all "peeie" are ~imllar. Although Ihe~
l';lt a grl';)t \:lriety of in"e,·t~ und OIIH.:r l1l\l·rl~·hrale". Ihe princIpal food i~
:1l11'. \\'ilh (lTle l'\Ceptl oll (B. IIIclilllOl'ficflts!. the R (l!'l1Can 'l)Cell!" liye in
primal') or '~· .... ()mbr) tnrc"t~. 8ulil I//c{wul\lic/IIS i~ a camp tol101\ cr of
mankind. li\ing in emimn111Cllt" l'n.'ated h~ Inunan, \iJlage~. 1(J\~Jh. and
ronJ'Jde~ and appear" to tlL' ,I r~'~'l'l1l i1l\adcr of Borneo.
BlIfo (lspa Gra\enhllr.,\

Rl\er 'loaJ (Fig. 31)
Description : Thi~ i~ a larg~' toad, \\ 1111 t~ pieal load-like rough. \1 ar{~

,,~in. Then! are llO ball) ae~b on Ihc ho.::'1(J. The ('ardrUIll j, \I~ib le . The
IX!HllllHl gland hehind the eye i~ ll\UaJl) round or O\aL hut Ihe length i"
al\\a~, Ic\~ than [llie(' the \\idth. All the 10t'~ except the kll1rth (Ihe longc\11
are \\ebbed 10 the tip'" The general colourntioll i~ dark bro\\tl or bla.... k
~Inle': 70-100 mm.
Felll:tle ~: 95 I-lO mm.
Tadpolt' : The tadpole~ are "mall ( I ~- 1 5 mm hclorc Illetamorpho,i,» :ll1d
hbd. The hod~ IS 0\':11 ,llld ~ligIHI~ l1allo.:m·u. ,md Ihe tail 1\ ~haped like a
tlarro\\ Il:ai \\1111 a rounded lip, The lo\\el' lip i~ \er~ \\id~·.
lJabits and hahital : TllI~ ;;;pel'je~ liles in primar~ and o ld ~I.'l'nnuar)
fore~1 H1 the k1\\ lamk It '" l1nll~e mn"t lOad" in th at jtl\enile, a11l1 adulh do
!lot \I:tnder through Iho.: rorc~1 hlll remain along: "tream ban~~. Adlllt~ du nol
mOIL' mudl. ~o th;1I one ma~ \el' thl.' ~a11lC lI1dll idual toad "itting Hllhe 'ame
'pIli on :t ,Iream bank hlr m~lIl1h, . The lad pole, al~o differ from the u"llill
'"
BUFON IDA E : BUFO
toad pattern. living ill ~trong currents and apparently clinging to the bottom
with the enlarged lips.
Call : A raspy. chirp. sometimes repeated.
Distribution : Widely distributed in Samwak. Brunei. and Kalimantan.
In Sabah it appears to have a patchy distribution. ha"ing been found at only
three or four scattered localitie~. Outside of Borneo. this species occurs in
Java. Sumatra. and Ihc Malay Peninsui:l.
Fi/!. JI. RlIj(lll.I{JI!f Gra\'enhor~t.
Bujo j uxfasper Inger

Giant Ri"er Toad (Fig. 32)
Description : This is a "ery large lOad. with a broad. blum head :.InO
t)picallOad-li~e wart) ~kin. Like B. wpcr. it ha~ no bony crests on the head.
The eardrum i~ visible. The parntoid gland behind the eye is long. usually
l---l. times as long as wide. and olkn i~ tapered towards the rear. All the toes
e.\t'ept the fOllrth (the longest) are \\ ebbed 10 the tip~. The general colouration
i~ d:Hk gn.'). brm\ n. or black . sometime~ \\ ilh ub~l'un: darker "puh on the
back. In addition to ils large size. this ~pe('ie~ i~ remarkable fur ih bad.
almlht di"gu"ting. smell.
77
BUFONIDAE: HUrO
~b.k,: YO 120 mm.

Fl'lllall:~: 125 215 mm .
Ta dpole: In general fOITH :1111.1 IIp,. thi~ tadpole i!. li~o.: that of B. mpa.
The main differences are in ~iLl' and colouration. Ju~t before mo.:tamorpho~i<;
the t:ldpolc~ or R. jux/mper rnea~ur!;' 15-20 mm. The under·,jde of the heud
In B. jl/lw.lper tadpoll" i, dark: in tadpole~ of B. asp('1' the 11l1der~ide of the
ho.:au j, \\hite or pink.
Ha bits and habitltt : HI/fa jll.I/l/I'jU' 1' can be found wandering through
primar) and old <;econdar) fore::.t from ncar 't'a level to 1600 metre!>. Thi~
<;~cie::. i!> not a'> tiell to ,Iream b:Hlk~ a!> i::. B. asper. 110\\ l:\er. Ihe'>e two
'f!Ccie<; are alike in brco.:ding "tl: and habitat of tadpole~ . BIIJ(} jlf.\/mper
,o.:elll' to breed in s\\ ifter l'um:nt, than B. lilper. and we ha\e found tadpole,>
of B. jll.\/(fI/Jf.T in strong rapids .
Call : A hoaN~. squeaky croak or chirp. In Sabah. tho.: Du~un n:Hne for
this specie!> i~ "bu·allg ~ ut". \\hich irnit:nes the ,>ounl! of nOI Olll'. hut 1\\0
toad'> calling al night. rrl'4uo.:nll~ wads of two different size!> (and.lhu,. IOlle<;
of voice) will sit within se\eral melre:-- of Ollt' another. When the first calls.
Fig..U . Blllo jll.ltmpl' T Inger.
7R
BurONIOAE, BUFO
the other rapidly :lns\\er<;. rc'tuhing ill a two-syllable ~o und Iha1 appears to
originme from one calk-f.
Distrihution: KnU\\ n from all pan~ of Borneo. The di,lribulioll in
Saruv.a" i~ ~potty. while ill Sahah lhi~ toad is extreme!) abundant in lhe
[owl:lOds.
811/0 dil'ergells Peler:.

Cre,,[cd Toat! (Fig. 33)
Description : A ,mall stock) load ..... ilh man) scallered. SPill) "anS on
the had, and ... idt's. The snout is l1arro\\ and project:. beyond the mouth. There
is a pair of narrow, rai<;ed crew. bcl\H!'en the eyes. 111(' eardrum is yi<;ihk.
The paroloid gland behind the eye b longer than wide. tapered lov.ard Ihe
['e:lf. and is u~t!ally followed by an oblique rot", of war1~. The length of the
gla nd j<; Ie,,~ than three limes its width. i\' une uflhe toe" j<; fully webbed. The
general colour is monied clay \0 rcddbh brown. often \\ ith ;1 ~crie s of hlad..
c hl'vron ~ on the back .
~Iale~: 18-13 mm.
Fl!tllaJes: 36..-55 mm .
Fig•.33. Hurn tlin'rgt'lls Peter<.
79
I:HTO;\'IDAE : BUfO
Tadpol e: The tarJpok~ are (l\;tl. b!ad, ami <;mall. J ust before
meta illurph osi~. the) arC' 15-17 mill long. The~e afl~ t) pkal toad tadpoles.
The) do not h:1\ e expanded lip~ li!"l' tadpoles of H. (/~P('/' and R . j!lXf{IIPl!I'.
H :t bit ~ and h a hita t : This i, a tuad t'! ror~st floor le:lf liller. It O<.Tur, in
primal) and old "'C'l'ondar) forest belo\\ 700 metro.: . . ahO\e sea lewl. Adult~
brl;'\.·d at \~l) ' hallo\\. ~mal1. in tCTllllllem ~tream~ o r III rain pool<; 011 the
fore~t !loor. ~Iale ... usuall~ fonn !loi,) calling: groups. Tadpok~ fro m egg:"
laid in ~mall "Ir~'atlh usually are found \\ h~'rl' the eurrent is \\eakest.
C all : A queruloll ..... raspy. rising trill.
Uistribuli on : round throughout Homeo. Ebe\\hcro.: it i~ known from
Sumatra.
BII10 l1I e lall o~·tiC: lm· Schneider

Common Sunda Toad (Fi g . J-' I
Descri p tion ' " ,tock). medium-siled 10 larg~ toad \\ Llh a fe lam el)
"'Ill,tll head and ,hor! hind limb:.. LOlli,! dark LTC'~h border the e)e1id:. ;lIld
extend dO\\ll\\anl . . on either "ide of the e)e, Another. thil· !"er (:[('<;\ runs from
the eye to the pawwirJ gland. The gland i<; la rge. l1\·al. and about a, lon g as
C)C and snolli combined. Tho.: bad i . . con'red \\'ith round \\art, of \llrying
"iA'''''. o ften ~mrounded b) darh-r piglllo.:nt and c;)pped \\ jtll till} d~lfk ~p in l..·"" ,
Th~' general colour i .. £ reyi~h or rl'dJhh hro\\ n. u~ually w ithout m arlin~ )
e\("I;'pt ror the \\ art) a rea~.
~1;I[e~: 57-83 mm .
I- emale~: 65-85 1111l1 .
Tadpole: Shapl;'d li !..c tadpok" 01 B. dh·crgl'lIS. Bb(.'k anrJ ~1ll:l11. nOI mer
15 l11m long.
H a hits a nd h;lbil al: A~ ;llre:ld ~ Illentioned. thi<; IS the o nl ~ Bornean toad
that doc, not Ii\e 1Il fore>,ts. It hob ;Idal)tcd to the ~JI\iron l\lenta l di~lUrb;lIlt'l'~
l..'re<lted b~ man~lIld and occurs throughout trnpical \~ia III \·illage ... and
to\\ ns. Adult.... arl' tNI,l[[~ found either at temporal'} T;.tlll pool" or drain,. or
under ~treet bllllh \\!1l..'rl' the~ feed on the in~eet~ dr0pping from th~ hghh.
Tadpoles de\'e!op in ,tanding \\ aler.
C all : ,\ 10\\ r:ntling Irill.
Dis trihuti on : Known fmm to\\ Ih III Sar:m al.. and KalllHalltan. In Sab.lh.
it ha . . heen found "'0 f;tr onl~ in onl..· '111;111 \ ll1ag:(' near Kota Ki nabalu.
80
BUFONlDA[ : BUFO
Fig . .It Rufl! mi'lwlII.I/ic/ltI Schneider
81110 quadriporcatlls Boulenger

Swamp Toad (rig. 35)
Descriptiun : This i~ <l ~lllall tu lll,,:UiUlll-:-.il.e-d. stod'J ' toad \\ ith a pair of
short . raised ridges between the eyes. a!> in n. di\"(-"/gell.l". The snout is llalTo\\
and project". beyond the mouth. The eardrum i!> \'i~ible. Thl' parutuid gland i~
long. compressed from side to side. and distinctly raised. It is more than three
tim..:~ a" long <\<, \\ ide- and ends in a row of large . spiny wans. The baek has
low. smooth \\Iart~, whereas the ~ides have rn~lIly small. spiny warts. None- of
the toes are full} webbed. The colour ranges from orangeish brO\\n to almost
black. usually without marking".
Males: -1.8- 50 mm.
females: 49---6:2 mm.
Tadpole: UnknO\\n. but probably very ~imilar 10 that of the related H.
di''CIgells.
Habits and habitat: Thi~ ~peeic~ ~eem~ to be confined to peat fore~l~
ncar the t·oa~t. It is \t'r~ abundant in ~ueh e11\ irorHlle-n\<; bllt has not been ~ccn
81
BUFONIDAE: LEPTOPHRY NE
in hilly counuy. It almost surely breeds in the pools of standing \\ater that arc
so COlllillon 1Il swamps.
Call : Unknown.
Distrihution: Thi~ ~pccic<, ha~ been n:ported from Kalimantan. Sabah.
and Sara\\ ak. It probably abo occurs in Brunei. which has large areas of
appropriate environment. This species al~o occurs in SumatT:) and Penim,ular
Malaysia.
Fig. 35. 814ft} lIIWt/l"i{lOJ"("(I/11I Boulcngcr.
Leptopliryue borbonica (Tschudi)

Cro~s Toad (Fig. ~6)
Description : A ~mal1 .. lender-bodied load with long leg .. and a shon .

tapt'"rt'"d ~Ilout \\hidl projects "Iightl~ o\'er tht'" mouth in profile. The e:u'dmm
i~ 0\31. anu about one-third the ~ize of tht'" C) e. Th('re is no parotoid gland. 0f
an~ diqinct enlarged glands OWf tht'" upper side~ of the body. except for :l
~1ightl~ rai"ed fold of skin that rurh d(m n the side of the back. The tip~ of the
linger~ and toe~ <Ire rounded and hlunl. Webbing is restricted to the ha,c 01"
BU ....ON IDAE: LEPTOPIIRYNE
the tOC~. The colour is greyish with brown markings. u,ually fomling an X
in the middle of the hack. The throat and chest are dark brown. while the
belly is grey to yclkm· i~h. The undersides of the legs are abo greyi~h or
yellowish with brown spots.
J\ lales: 27- 31 mm.
Females: 40---47 mill .
Tadpole : Unknown.
Habil" and hahitat : These ~mall toads arc mO~1 often found in the leaf
litter of ~eepag(' areas in forests below 400 melres abme sea level.
Call : Not recorded.
lJistrihution: Known from all parl~ of Borneo. The species also occurs
in soulh('a~lcm Asia and Sumauil.
Fig. 3ti. l..i' pU/{Jhnlll' /JI!rhollico ( T ~rhudil,

BUFONIDAE: P[DOSTlBES
Genus Pedostibes
hi~ long-legg('Ll group of rnedium- to Iarge-~iJed fore!>l toads are
T probably the onl~ trul~ arboreal 1ll0;:1llbers of the Bufonidae. They

occur lllll<,tly in llJldi<;turbed forests. though they rna~ hI' encounlcn:u
in old ~econdar~ habitab. Their iingl'r~ are long and di~tinctl) shaped.
probabl) an adaptation 10 l'limhing. Some indi\'idullb ()f the Br(mn Tree
Toad. PcdOl"tihes hosii. lla\'e been found a~ high as six metre<' up in tree~. The
onl~ tadpok~ known an: of this ~pccic<;. occurring in accumulations of dead
lea\ ('~ in the quid <;ide pooh of fore~l !>lrearns. Adult frog~ of \hi~ gl'l1l.1S haye
a dio;:\ that is mo~\l~ ant~.
Pedostibes I/Usii (Bouknger)

Bn)\\ n Tree Toad (Fig. 37)
Descriptiun : A large. ~tllut toad with a thick bony ridge behind thl' eye
conn"'<"led to a small parotoid gland. The eardrum i~ \isibk . The toe~. e:,,(.'(:pt
for th ... founh (iongntl. are completely \\"ebb~d. and the tips are slightl)
widened . The finger tip ... are expanded into stubby pads . :md ther ... is a slight
webbing at \h ... hase~ of the finger~. Thl' top of th~' head is ~mooth. and the
back has a k\\ ~eatlered \\arts. Allmaks and aboul half of the kmales are
uniform light 10 chocolate brown. The n:mainder of the females are black or
dark purple \\ ilh a pattern of ~el1o\\" spot:. limning a den~ ... network.
j\[ak~: ."i3- 78 llllll.
Female~ : X9- J05 !l1111.
Tadpole : The bod~ is oval and the tail is ~haped like a naJTO\\ leaf wilh
a rounded tip. The colour is dark. bro\\ n above and on the sidc~. without
markings, Total length rea~'hes 19-20 111m.
Habits and habitat : Thi <; arboreal toad li\'e~ in primary forl'~t in
10\\ lands belO\\ 600 metres abm'e sea level. Adults come to rnedium-~iJ.ed.
clear ~tream~ 10 bre ... d. Breeding i~ not sea~llnal. bUi occur:. in spurts or pulses
with many indi\'idual" coming to brc~d for one or two nighl~. then retiring ttl
the fore~l for se\'eral \\ceks. The male" do not fomi calling aggr~·gations. but
~ome nighl~ there are so Tllany pre~el11 that they ~ .:cm to be e\'el") \\ here along
the bank. Adult~ feed on a \ ariel:- of in~ ...ets . \\jlh ants being the dominant
l·ompnnent. The tadpole~ li\l.~ in quiet ~idc pools and in masses of dead lea\"t·~
on the botlul1i~ of stre:Ul1~.
BUFONlDAE ; PEDOSTlBES
Fig. 37. Pl'do.\/ibes !to,\ii (Ruulcnger).
Call : A grating. slurred ~4t1aw\.... ri~ing in pitch slightly from beginning

to end.
Distrihution : All pans of Borneo. The species is aho known frolll
Sumatra. Peninsular Malaysia. and Southem Thailand.
Pedostibes rugosus Inger

Marbled Tree Toad (Fig . .18)
Dl'SCl'iplion : A medium-~ized to large. ~(Oul toad. \\ ilh extremely rough.

warty ~k.in and a large egg-shaped parotoid gland behind the e~c. The
eardrum i~ \·isihk. The tip .. of the lingers arl' .. light I) \\ idened. but blunt. All
the lOe~. except the fourth IIhe longest). arc fully \\ebbed. The colour i ..
85
BC I-O:'\IDAI::: . PEDOSTlRES
bright green II iell br{)\\Il or n:Jdi,h broIl II ~po!\. From and hind Icg~ IJme
bro;uJ broIl n cro~" bar,.
I\!<lk,,: 74-77ll1m.
Fern<lle~: SO 95 Illtll.
Tadpole: L·n~J10\\n .
Habib and h"bilal : Thi~ 'pecie" Ila, been fuund onl~ in hilly primal!
fort"1 from 15010 1050 nk.. tre~ ahme ~l"a kleJ. \\'~ ,u~peCt that adult::. arc
arboreal and di'rer.;e Ihrough the Inrest. All the adult, lIe h<lw "cen ha\~
heen taught at brcl'ding ~ilc' along ,:\car. rock.~ ' tream~.
Call : Unkno\\ n.
Dis tribution : KnOll n from Saball. Sara\\·ak. and \Ieqem Kalimant;m .
Pre!>Lllllabl~. Ihl' "peCI':" OCCUP., throughout Borneo. II i" not kno\\n from
el'e\\ here.
rig . .l8. Pl'dosrii>n mC:t>S/lf Illg.... r.

BUFON IDAE: P ELOPHHYNE
Genus Pe{ophry ue
D w~rf Toil d s
hc~c are small mao" \\ ith smaH, ~('ane rc(\ w:ln~ on the hack and head ,
T blll wjrhOLu a large parotoid glund behi nd the eye, All have the nnge r~
p:lrtl) \'ebbed, Bel'allse they ha\'e not been observed \ery often, very
little is kIlO\\ n of Iheir nalUral hi~lory. Female" of !Ill s pede~ ha,e very ~mall
d utehes (fewer than 20, of large. whiti~h egg". Tadpolc~ of one specie" were
found in a shallo\\ pool: they c!l'a rl ~ wer,,: no t feeding. a~ they lacked the
lIsu:11 la(\'al mouth pnrl~. :lJ1d \\ere completi ng dl'\cloplllcm ba"ed on Ihe
large amollnt of yoH.. l'olllained in the egg.
S~cie~ of Pdoplrryj]{:' h:\\'e ne\'er been ~cel\ oUbide of primal) \)f old
~econtlar:- forc,!. and an: I...nown tIl occur from near ~ea Je\cl to 3100 metre'.
1\10\( indiyiduab han' been found o n the forest floor. bUllht') also dimb into
10\\ \,egewtiotl.
Pelvphrple misera (Mocqwml)

Kinabalu Dwarf Toad (Fig. 39)
Descriptio n: A \W~ ~mall toad. \\ ilh the hand wehbed. incl uding the
em ire ti r~t linger and half Cit' the o\!tcr ringer", The tip" of thl! nng.er~ .w~
n;lrrow. Webbing on the fCiot reaches the lips of the lin.t lhrt'(' foeS. The
('antrum i~ ,bibl('. The back and ~idc~ h:I\'e "mall. ~callered, round or OV:II
\\<II1S, Th~' l'olour b black or drlrk brown, with \\hile hlotl'he" on lhe
under .. idc,
Male .. : 16-~1 J1l1ll.
Fema1e~: I S~13 111 1ll ,
Tadpol e: Lnkno\\ 11. but probably a .. descrihed und('f genus.
Il a hiis and ha bitat : All krlO\\n record" of tllh ~pecic~ arc rrO\11
ei('\:!!ion<; between 1600 and 3100 rnetre~ abmc ~ea 1c\cJ in 1l10~sy montane
rore~l. Thb ~pecic,> has h('cn :.een rllu~ t l~ at ground Incl, bllll\\O aduh~ \\erc
found ill the ClIp of a pitcher plant on \ \0\11"\\ Killabalu. Femalc~ ba\\,.'
dutchc,> of 6-16 egg~.
Call : a ,cries of high-pitched note ...
Oistrihutio n: :-'1\1unt Kimball!. Sahah. Therl' is one repon from \ Iuunt
1 1urud. Sara\\:tk. Prnhabl~ the ~pecie .. O('nlr~ on other muuntai n,> in Bomeo.
87
BCFO.\'IDA!:: : PELO PHRYNE
F ig. 39. i'!-/Ol'/"-Hlc' miS(,nI It1. locquanJ).
Pe/opllrYl1e rllOpopJtififl .~ Inger & Slllcbing

Climbing D\\:lrf Toad
Uescrilltion: A \-ery small toad. \\ nh thid, \\ehhing on th ..' hand reaching

Ih", cnd~oflhe 1\\0 inner finger~ but only Ihe ba~..:~ of the t\\O oul..:rones. The
lip~ uf the finger~ an,' \\ ldened into hlun! pad~. Thid-. \\ ebbin~ on the fool
lea\C~ ju~t the lip~ of till.' three inn..:r llh,':' free. The cHrdt1.11ll is \'i~iblc. The
hnck ha~ man) ~lllall. round wart~. e;'II,:h of which ha~ a Ull) whilish con..: in
the cemre. The hack i~ brlJ\\n or black. otkn \\lth dark mar!..ings outlining a
broad X. Th.: under,ide of Ihe hod~ b \\hile with slllall bl::tck ~pob Ihal
OI.·l·Upy Je~~ th:m hnlf the lolal area.
~Iak,,: 21 - 24 mm.
Femak~: 23 mm.
Tadpole : Un!..no\\n.
Habits and habitat : Thi~ ~pede~ h;J' hl'(,ll C ,lIe':ll'd JU'I Iwice. On one
occa,ion. ~i\ male, were ..:alllTlg. from ka\I." of bll~h..:~ in a <;ubmolll:lI1l.'
forl'" at S 15 I1l1.'lr('<; abo\e .....:J IeH'1.
Call: NOI d"'~l.'nbed.
BUFONIDAE : PEL OPI-I RY NE
Distribution : Known only from southeastern Sarawak and neighbouring

western Kalimantan .
Pelophrylle signata ( Bou!enger)

Lowland D\~arf Toad {Fig. -1-0)
Description : j\ remarkably tiny toad . with thick webbing on the hand

reaching Ihe tip of \he first fingl:f but leaving half of lhe outer fingers free .
The tips of the lingers are widened into blunt discs. Thick webbing on the
foot reac hc\ the tips of the first twO toes only. The eardmrn is visible. The
back ha~ ~mall. round warts and is black or darl-.. brown with small black
spot ~. Some indi\iduals have a \\ide. dark X-shaped mark on the back. There
i~ a yellow or cre:ltll-colollrcd band froll) belo\\" thc eye passing above the
armpit to the ~ide.
~1ale: 1-1--17 mm.
Female: 16- 18 I1IIll.
Tadpol e: Linknown. but presumably as described under the genus.
Habits and habitat : This spec ies lives in nat or hilly primary forcst
from near 'lcu le\'el to about 1000 ITlt'trcs. We han' found it on the ground and
Fig. -w. Pe/ol'flr;l1t' ,1<';1/(1/(1 (Houlc>nger).
89
Bt;FO;\ II)AE : PSEt..:DOBUFO
in 10\\ vegetation. Female:. h:.I\e ... mall clutch\.':. o( large egg:. thai arc aoout
one-Ienth the length of Ihe boJ~ .
Ca ll : UnknO\\ n.
Distrihutioll : All rarl~ of Borneo. Thi, <;{X'cie~ ha~ aho hecn reponed
from the Bung-aranlsland~ ami Pellin~ular Mala)sia.
Plieudohufo sllbm,.per Tschudi

Aquatic S\\amp Toad (fig . ~I)
Desc.-ipliuu : A large. <;Iock) lO<ld \\ Ilh a "mall head. and nO!>lril~ all the
lOp or Ihe ,nOlll. The flll,!:!er ... arc long. :.lender and hlunt. v. Ith slight \\cbhing
at the ba:.c .... The toes are full) \\ chbed 10 Ihe fjp~. Thl' hack j<; covered \\ ilh
large. round \1 art~. and there are dbtinct parotoid gland~. The hack and side ...
arc hro\\ n. with a y(' llowish to ) ello\\ -orange stripe along \\ h~'rl' hack and
~i dc, meet. The belly i~ ~dlowi~h . In mature femak~ the throat b yellowish
and in males. black.
Fig. 41 . "\{'Udllhll/o mixuprrhchU(.h
90
BUFONIDAE : PSEUDOBUFO
:'vlllles: 77- 94 mm.

Females: 92-155 mm.
Tadpole: Unknown.
Habi ts and habitut: Adults are u~ually found in pools among nOaling
vegetation mainly ill coastal and peat swamps associated wilh large rivers
(such as the Kapua~ in Kalimantan ). The cryptic colouralion and lethargic
habib of lhi~ load make it difficult to find. Males have been reported 10 call
from mats of vegetation overhanging water. into which they promptly dive if
disturbed.
Call : Unk.nown.
Dis tribution : The only Boml"an specimens have heen found ill
Kalimantan. The ~pecic~ i~ aho known from Sumalra and Peninsular
Ma!ay~ia.
91
JClI17ily Microhylidae
Narrow-mouthed Frogs
Thi~ f:unily is ('~<,cnlially tropical in its di~lriblltion with only a few ~pecies
Jh IIlg in temperate climates. i\'l.:1l1) ~pc(:ie ... .'oecrete a thick. <,tid.y mucus from
gland ... in the ~kill .
Genus Calluella
Spadefo/J\ Frogs
11 the frogs o f thi" group (Ife burrower~ and. bt-(::wse of that. the
A Bornean ~JX'cic, ;m: rarely ~een. Onl~ one -"pede ... ha~ been collet· ted
more than thn:e limes. and onc '>pcl'ie ... i ... known from a ... ingle
.'>pecimen. Nothing i ... ".l1own ahoul the breeding habil~ of the Bornean
-"pecie", hut tho ...e from Thailand :mcJ o lher pans of the continent breed in
~mal1 tCllllXlrary ponds.
Dcscription- ndulls: All of tllese frog.. h;1\(' rather flat bodie .... \\ide
healh. hidden cardmm ... and 'mall eyes. The tip'" uf the fingcr~ nnd loe . . are
not \ddened and the lUt'~ h<lve \ery little \\ebbing. All of them htl\e a large
compressed !lange under the foot \\ hieh. be"ides being the Tea"on for the
~'omJllon name. i~ the "Iruclure tl"cd in digging.
Tadpole: The tadpoles of the Bomean specie~ arc unknown.
Call1l ella brook.fii ( Buulenger)

Brooks' Spadefool Frog
Description : A mediuITHi/cd. "tot'ky frog wi[h an extreme]) hroad.

:.hort head lind "hort, thick leg', The M10U[ is stuhhy. wi[h Ihe nostril, al ils
\er) end. The toes arc narrow I) webbed at the ha~l' , AI! of the UPP'!T ,urfaees
:Ire medium brow n. \\ ilh many scattered, ,mall bhl(:k ~pOI", The throat of
male~ i" black, thaI or]'enl:lte~ \\hiti~h , Tht, l'he~t and :Ibdornen are \\ hitish
or ~ dlo\\ bh.
93
MICROIl YULJAE : CALLUELLA
Mak~ : 51-55 nUll,

Femak: 60 mm.
Tadpole: Unknown.
Habits and hahitat : Then: i~ little known about this speeie~ other than
ib hurrowing habit\ . All record:. are from fore~t~ al <;,k\atiom close 10 ~<;,a
le\<;, I.
Call : Unknown.
Distribution : Recorded from Kalimanwn and Sarawak.
Cafluella sm;thi (Barbour & Nohle )

BlotclHided Spade foot Frog (Fig . .:+21
Desuiplion : A <,mall stocky frog \\ ilh ;1 wide head and ~hort. thick legs.
TIl<;' "nOll! is nalTo\\ ~·d ahm'e the ja\\. with the noqrils Ilear the lip . The toes
are narrO\\ Iy \\ebbed at the ha~e. The "kin of the back and top of the head i~
smooth v.ith a few ~callered. round humps. The throat of the male i~ tinel)
granuhu and the belly of both sexe~ IS ~mooth or finely pebhled. The back
and top of the head are wn or medium hroW11. The side~ an.' a lighter 1311 or
yt'llowi~h. \\ itll a linger of the lighter coluur projecting up onto the back. The
lower sides and hidden ~urf;1ces of the Jeg~ h,.\\e large black blo t che~. each of
Fi g. 42. Cn//iI('I111 \/lII lh i ( Barbu uf & Nuht"l.
94
M IC RO HYLillAE: C HAPE RI NA
which has u narrow red hord ...'r. The throat of Ihe male i~ black, while thai of
the f.:mule i~ brown \\ith ~e\eral yellowish 'pOis.11l(' abdomen i~ yello\\i~h
1\ ilh brown spot-. or molt ling around Ihe margin~.
,\-Iale: ~3 mm.
Femak~: 38--W mm.
Tadpole: Unknown.
Hahits and habitat: Thi s specie~ i~ known from primal)' foresb ut low
ele\'ation~. The) haw somelime~ been found under dead lea\'es or logs on the
for("~t 110or.
Call : UnknO\\I1.
Distribution ' Recorded only from Sahah and Sarawak.
Chaperilla jilSca Mocquard

Saffron-belli ed Frog (Fig. -U )
Descriptiun : A vcr) "mall frog with a budy the ~iLe and shap(" of a large
almond, The tips of the lingers and toe ... are slightly widened. and the to-.:~
hale no webbing:. There is a tiny tkxible ~pine al the elhO\\ and heel. The
buck is greeni~h blm:k. while the underside of the body and legs has a black
Fig. ~ J. Chaperillaji/I{"l/ .\hx:quan.l.
1)5
r-.IICROHYLIDAE: GASTROPHRYN()IDES
Ol·twor\'; t:nclo~ing: hright )ellm\ ... pots. Oudly.the }cllC:1\\ l"olollJ" \\ill rub ofT
onlO human finger ... qUite o.:asily.
r-.I:tk . . : 18-21 1lI1ll.
Fclllak,,: 10--14 mm.
T:I(II)OI~ : TIK· ...e are ... mall. o\al. .and . . tightl) Ilal\cncd above. The tail
tapo.:r . . abrupll~ near the end In a rounued tip. The wi) is hlac\.; :Ibove. lighter
helm\. The tail fins an: colourk~<;. Total length reache~ 20 mm.
Habits and habitat: This ~pecies l.in'" in primary and <;ccondar) fore~1
and will l'\en live in gankm. It i... c<;pel'iall) abumjallt in [l::il cOlllHr). bUI
al""(l (\Ccur" in hill~. Allhough Ihis ~pccies i:. c . . <;cntiall) an inh:!bilalll of Ihe
fO~"'1 110or. il <;01llt'"li1l1c<; dimb . . inlu Im\ q~gL·l3lion. t-,Ia!e:- f01111 calling
g.roup .. around . . mall. \cnlporar~ poob of \\ ::lIer. Egg: . . ilIa} bc laid in dear or
turbid \\al('1". ~oJl1clillle;, in \':lIer thm i!o r:Jnk \\ilh decaying mailer. The
tadpoles ho\\.'r in miu,wmer. lillering microorga1l1~rns.
C all : A . . hort. iIN':l'I-ltke bulZ.
Distribution ' Ol\'ur" in all pans of Borneo. II abo i . . found in the
...o lltht'11l Philippine bbnd, and Pl'mn:-ular ~1 3Ia~,ia.
Gastruphrynuit!es bUrl/eel/sis I Boulengerl

L ong-~I\outed Frog
n ~s(" ription : 1\ ~Olall to n1t'"dillrn-~ill·d. ,toc\.;) frog. \\ ith ... hon. Ihiek
legs. Tho.: he:J.u i~ . . haped Ii\,;(' a dr;mll-olll cone CUt oil at the lip. with the
,nom mali..' lh:m /}Iiet'lhe knph of In!.' I:'"ye dj;JPwlt>J .,nd rr~i [,c ljllg bt>yond
Iho.: gape of Ihe mouth. The eardrum i:. exposed. but po~iliuned lo\\cr than in
mo~t fro£~. being almost at thc !C"\·et of the upper lip. The lingers arc ~hon
and ralher :.t ubh) . The toe ... are almo,,! w ithout webbing. The lip" of tht' toes
are ~lighl l ~ swollen. lht' upper ~urfacc" are \cl)' :-moolh. Thb frog i..; grl")"
brown \\ ilh small \\ hilt'". ~e;I!tl."red :.pol<". The underside: i ... \\ hite.
r- Iales: t.:n\.;nO\\Il.
Female~: 3().....41 mill.
Habib and habitat : Vel) lillic is kllO\\!l aboul the habit . . ofthi!o frog:. It
has olll~ hun ~tcn in lu\\ land for.: . . \'o. les:. than 500 n·II.~!re5 ;tbo\c ~e" ].:\e1.
Individual, ha\e bee I, found under dead lea\...:" on Ihe fore~1 noor (llltl in
\\:llcf-tilkd Iree hok ..... Oot' \\;\<., takell ;1\\3.\ fw .n:l GianI River Frog. which
was ... \\ allo\\ in§! it.
[)istribution : Thi~ frog h'b lx-en ~een on l) in Sara\\a\... hUI is lih'l) to
be morl' \\ide . . pread.
.\ II CROI-IYLlDAE: KALOPHRYNUS
Genus KllloplirYlllls
Sticky Frogs
KafopJ!lylllls are small to tin) ~tieky frogs of the leaf litter with pointed nosc~
and ~hon. ~Iender hind 1cg~. Three h3\'e spob in the groin (in~ide the thighl.
Their habits are quill: ~imilar. AJlli\(' in the forest floor liller where they feed
mainly on all\!. and termites.
Ka/ophrYliliS baluellsis Kiew

Kinahah.l Sticky Frog (Fig. -l--t )
Description : A small. ~tock) frog. with 3. rclati\'ely llarrO\\ head. pointed

!'.noul. and shan . slender hind legs . ringers and toes are Ilot webbed. and the
linger~ are long. \~ ith ~lightly flattened. blunt tips. The eardJum is not visible.
The skin i, finely granular. with a raised fold acro~s the head helween the
eyes. The head i~ brown. and the back j, bruwnish with irregular [ighl
patches. Th ... sides of the h~'ad are yellowish with dark spot~, while the ~idl"S
of the body are light hrown. There is a dark brown ,treak running from the
Fig. ~~ . K(I/opl1l1"IIJ1.' /lldul'I1sis KICw.
97
r-. IICRO HYLI DAE: K.....\LOPIlRYNUS
snoul 10 tbe groin on ~'ach ~ide of the ba<:~. The bell) i~ g:reyi~h \\-hile \\ ilh
seatk'fed bro\\ n ~PO[~ and enlarged \\ hili~h bumps.
:-'lale~: 3f! mm .
Female' : 3(') mm .
Tadpol e: L'nknU\\ 11.
Habits and habitat: Thi~ i~ a frog of Ih<.' kaf liner ~nown onl~ lWlll
prilllar~ mOntallL' f0fe~1 on I\ \ounl Kinabalu at '-Ihout 1.500 1lletre~. When
handkd. It exude~ 3 sticky mucu~. I t ~ habit~ are probably SOIl1\:\\ hal ~irni lar
to Iho~.: ufthe beller kl1o\\11 K. plcllr()sri~lntl (see pagl' 100). r-. 1a Ie~ call at te]
he3YY rain" and breed III temporar~ pools.
C all : Cn~nO\\ 11.
Distribution : KnO\\l1 on ly from :V1ount Kinabalu in Sabah . It proha bl~
abo occurs on thc higher mountain,> in Sarawak and Kahmantan .
KafopilrYlllIS heterochirus Bnulcnger

Variabk Slicky Frog: (Fig. --1.5)
DescrilJtion : A ;;malL ~tocky frog. \\ith 3 narrO\\ head , pointed ~n(l lit.
and ~ lcnder hind kgs. The ou1enno<;[ toe and finger arc lery short. and tbe
!Oes have lillie \\ebbing. The ,kin is fllle l ~ granular. and glistem "ligh tl~
v,hen tbe IfOg is handled. Adul1s arc msty orange In dark brown abO\e . with
~irnilarly coloured !>idc~ ~e1 off from the ba<:k b) 3. fine. dark-bordered li ghl
line running diagonalJ) from the ~nout to the groin. From one 10 ,e\'eral dark-
bordered cream-coloured ~pots mil) uccur both in the groin area and on \hL'
inner Ihigh of each hind kg. becoming conspicuou~ \\ hen the i"rng jLllllr~·
The throat and l'hcq are cream) white \~ ilh dark mark.ings.
" l ales: 2-l.-27 mm ,
Females : 30-33 mill.
Tadpole : Lnkno\\n.
Habits and habitat : This i~ a frog of the leaf litter knO\\"11 from pr iJll ;1f~
and secondar~ fore~\ from near ,ca le\'el to 1200 metrc~. Like (1tilL'1
Ka/ophryI1115. il c"ude~ a ~tick~ . glue-lih' !Tlucm from gland~ in Ihe skin o!
the back. The <:alling site~ ul" males are probahly smalL temporar~ pool~ OIl
the fureq fl oor. ur in rolling logs. Ju\cn 'cs and adult, feed on t1n~
l1wenebratc~. such as ~mall ~pecic s of ant~ and tcrmile~ .
Call : l 'nkno\\n .
Distrilmtion: Sahah. Sara\\ak and Kalimantan.
9S
MJCROHY LI OAE: KA LOPHRYNUS
Fi~. J5. };alopltrnw_\ heleITK{,in" Ruulcngcr.
KalolJhrYl1l1s intermedius Inger

Intcrmediate Sticky Frog (Fig . .J6)
lJe'\cription: A small frog. with 3 narru\\ head. rounded "noU!. and ~hon
Jimh~, The outermost tot'" and linger ,m.' \cry shon. and the tOC!l have liule
\\-chbing. The skin i~ coar<id~ gr>lnular. "lth a di"IIOCI gland behind the ear.
Adul", are brown anO\c. '(Jnll'lime!l \\ Jlh ob"cure dar].. nmrk..ing' on lhe had ,
The .. ide, are ~ellO\\i'h IU ue.lnl-coloured. meeting the l.:olour3110n of the
h'll·k at a ~harp bounda!). The Ihr<>"11 ha, \"0 irregular, dark longitudinal
hand". The lhrO:ll b pjnki~h, \\ hile the Chl',", and bell) are cfcam-(.'ulouro::'d.
There arl' a I"e\\ dark ~pot" 011 the 3hdnml.'n near lhe chest.
\ Ialc,; 27 mill.
Female!>: 38---l1 mill.
Tadp(Jle: Unkl1{l\\ n.
Hahit.. and habitat: Thb i~ a frog of the Ic;).1 lilll'r knO\\ n from 10" land
primal) lorl"t helU\\ 300 melre" ele\ation,
e :ll!. L'nknO\\n.
99
MICRO II YLlDAE ; KALOPHRYNUS
Fi ~ . .l6. Kalophnlllf\ 111/('11111'111115 Ing.:r
Distributiun : Safa\\aJ... L i}..~· other .'>mall. sec r("ti\~' ,pt'{:ie, in thi" genu".
thi, 'pecie" probabl} ()('cur, in other parts of Bornco.
Ka/oplirYIlIIS p/eurostigma T"l'huui

Rufou,-.'>itletl Slick) Frog (Fig. 47)
Dl'sniption : A ... mall. .'> Iock} frog. \\ ilh a re latlvel) narnl\\ h.:mJ and
pointed 'nOUI. re.'>ulting in a "Oll1e\\ hat triangular shape. Th\..' outcnnO.'>1 IDe
anti finger arc \er: .'>hort. and the IflC' ha\e litlle webbing. r.~·male:. have a
find) pehhlcd upper ,urface. \\ hile male.'> ha\'e man) lin) 'pin.:.'>. Frog" are
light 10 durl.. brown on the hael.. and rU~ly-orangc nn th~' ~ide.'>. with a fine.
\\ hitbh lin.: funning diagoll<.!lI) frOlll Ihe ~nnul 10 the groin. A hlacl.. 'pol j,
con'picuQu<" ju<;! in i"ront of ea,:h hind leg:. The '\-troal ,mel che't afC rU~I) to
bro\\ni,h ,:ullithe abdomen lighl gfe) to gre).
t\ \.lle,: 17-50 mill.
h:~Jllale~: J5-57 mill.
100
1\HCRO HYLlDAE : KALOPHRYNUS
Tadpule: Gre) to olive tadpoles with an egg-sh<lped hod) and hlunt tip
to the wil. nlCW :..mall (maximum total length <lhnut 9 mm) tadpoles
apparently do nOI feed. bm compktc their development using the yolk
present in the egg.
Habits and habitat : This is <I frog or the le<lf lilla known only from
primary foreq ut low ele\,<ltion~. When handled. its ~kin exudes a sticky.
glue-like mut·u~ from gbnd~ in tht' ~k.in of the had The calling sites of males
<Ire nCar ~mall. shallow temporary pools on thl' forest tloor. or in rotting log~.
T<ldpoles h<lve heen round in very small. ~hallow pooh of water. such as
crevices of logs. Juveniles and adu1t~ feed on anb and termites. The frog
moves until il finds an ant or temute trail or nest. then laps up Ihe prey until
sarisiied. We have found owr one hundred am" in a single siomach.
Call : Probabl) a sharp. repetitive chirp.
Distrihution : Found in all parts of Borneo. It also occurs in Pt'ninsular
Malaysia. Sumatra and the ~nLlthl'm Philippine Islands.
Fig. ..J7. KU/llphnnllS J!/fIllO\li~lI/(/ '1\l:hudi.
101
MICROH 'l'LlDAE : KA.LOPHRYNL'S
h:a/ophrYIlfH s ubterreMri.~ Inger

Le"ser Sticky Frog tFig. -I-H)
Desc ription : A .. mall to lln~ frog:. II nh u naff(}\\ hcaJ. rounded ')noul.

and ~hort limb~. The oulermo~1 10": and lingo;']" are l"er~ ~hon . and the toe~
hal'e little webbing. The ~k.in i.. coar;,.e l~ granular. and gliqens slightly when
the frog is handled. Adults are hrown anOl·('. \\ IIh slightl~ darlo..er coloured
~idc .. ~cparaled il'om th(' haclo.. h~ a line. light line running diugonall~ from
the ~11()u1 10 Ih~' groin. The thl"oul and <:he::.1 urI! du~k~ II ilh light ~pOb or pale
yellolli.'lh II ilh :t dark band under the j:l\\.
Male..;: 21 ~2J mm.
Female ,: 26~27 nUll.
Tadpole : linkIlO\\n.
Habits and habitat: Thi .. is a frog of the leaf litter knO\\ n from primary
and old ~econdm} rore~t of the 10\1 Ian&.,. I\laJe~ haw been collected calhng
at the entrance to burro\\'s. hll"eniles and adult.. fe('d on tim im ~'rt~'hnlle ...
~ueh as .. mall species of ants and term ite ...
Call : UnknOl\"[l.
IJistrihution : Kno\l n from S:'lbah. Santllak and north central
Kallillanian .
Fig. 48. f{%plinlllt5 slIbrt'l"rC,l/ri,1 ing('r.
102
MlC ROHYLlD AE : KALOU LA
Klliouill blllelltll ( M Uller)

Rrown Bull frog (Fig. -1-1})
Dc ~criplion : A ~qual. round frog with ~hon .. thick hind limbs and a short
~nout. The eardmm is not \'i~ible. The tips of the fingers are widened .md
biLml. Thl' tol'~ h'He (lnl~ ~light wehbing and arc blunt at the tip~. The ~ole
of the foot ha s II pair of prominent ridge~ or nange~ m the ba~e of the toes.
The "kin is finely granular. often with ~cattered lumps. Colour of the hack
\aric~ from chocolate to dark brown. often with a lightcr band from behind
th..: e~t: 0$"]" \hl.: ~houldcr to the e1bO\~ . The rump i .. darker than the fore pan
of the b:lek.
l\hles: 50- 61 mm.
Fernak~ : 55--06 nUll .
Tadpole : Egg .. an: laid in the rain pooh l"om1cd in the dl'pre~~iom of the
di~turbed habitab where the!>'" frogs lj\e. Tadpole~ are greyish brown with an
unpigmented tail. with their eyes on the side .. of the head . Maximum length
is ahout 15 fl11l1.
FiA. -19. Ka/o ll /a ba lema l .\lLi.lkr ).
103
.\ lI CROHYLllJAE: KALOULA
Hahit s lind habitat: Thi ~ frog prl'fl?r~ di~lurh~'d or secondary habitats

inland. ~uch as the tlood plain:. of ri\.:r~. ur pond, and ditches produced b)
logging or agricullllral acti\ · itie~. It apparel1ll~ ~ecre":~ it~c1f in plant debri,
such a~ dr~ leaH" during dry periods. emerging rapidly and in largc nUl1lber~
to breed alter extemkd hca$ rains. [I fonns large noi~y breeding gl'Oup~.
calling loudly from hidden ~ite~ ill \cgclation. During: a call. males intlau"
them~ch cs grotesquely. The diet is mostly ants. though uther ~mall cra\\ ling
in':o",d~ arc abo caten.
Call: A luud ~harp honk.
Di stribut iun : Reumkd from Sahah. SaTawak and KalimaIHan. Also
known from Ja\'a. Peninsular \1ala) sill. and Pal,m<.lll in the Philippines.
Kaloliia plIlcltra (Gray)

Randed Rullrrog (Fig. 501
Description : A squaL round frog with ~hort. thick hind leg~ and a ~hol1
rounded :'110Ul. TIll' toc~ h;1\c onl~ a small amOllIll of webbing at the base.
and are blunt at thc lip ... ThL'TC i .. a promincnt "spadc" or thnge on the sole
of each foot be!\\een the hed and the oil,e of thl' lOl'~. The jingcr~ are long.
with no \\ebbing. but ilre ~lightl) \\iden",d and blunt at the tip~. Tht· .. )..in i~
finely pebbled. and there are \\ idely scattered light coloured bump~ Ovel' lhe
lower hack. The frogs are chocolate to dark brown with scattered dark ~po\s.
T\H) \\ ide, irregular. tan·coloured bands run backwards from the eye 10 the
groin. In most individuals a row of dark spots forms a line down the centre
of [hl' bac\.. i"rom th.: top of th.: head In the anus. The snout in front of the eyes
is <llso lUn in colour. \\hi1..: the under~id.: .. of the hcad arc hT0\\,11. The che~L
belly and under~ides of thc leg~ ha\ e ~caller~'d brO\\ n .. pot~, ~omctimes
connected to form a network .
.\-tales: 5~-67 mm.
Females: 55- 75 mm.
Tadpole: Eggs are laid in the temporary pools where the frogs call. The
t;jJpuk ~ are hlac)..ish \\ilh transparent fins. and their eyes are set on the sides
of the hcad. \!aximull1length i~ ahout ~5 mm.
Habits a nd habitat: Thi~ i, ;j frog of human .. ~'!t1cmCllh. pril11aril~ of
to\\ n" and ei l ie~. It E\e.'> under rubbi~h heap~ .lIlt! olher accU!llulatl'd (khri~
hllch as in clogged drains) during dr~ period~. emerging in \;jsl numhch
ahef dO\\ llpour~ of sen"ral hO\lf~ or more. It fo rm s large noi~~ brl'.:ding
grollp~ In tlooded area~ such a~ flooded drainage s~stem .. or la\\I1:.. \ Iall''o
\'lI CRO HYLIDAE: META PHRYNELLA
intlate themselves in a grotesque manner a~ the) call while 1l0011ing III the
~ urfaee of thc-sc- temporary ponds. Kalol/la pl/lchm feed~ <llmo ... t exdusively
on ants . though il will occasionally lake small beetles or other crawling
insects.
Call : A loud groaning honk.
Distribution : Reported from Sarawak and weslem Sabah. T his i... a
relatively recent introduction into Borneo. probably by accident rmher than
deliberately. It is nO! known hO\\ far it ha~ "prcad in Bomeo. The species is
widely di~trihutcd in tropical Asia.
,,
Fig. 50. Kahil/fa III//chm (Gray).
MetapllrYllellll SUlldalla (Pcters)

Tree Hole Frog: (Fig. 51)
Description : A tin) . .\o tol·k) frog: \\ ilh a poinll'd ~nout. The lip~ of the
fingc-rs are widened and the underside of ea<:h tinger ha ... a thickened. Ilcsh)
pad:1t til(" hase. T hc- feet are rather fle..,hy and the toe~ are about half \\ebbeu.
The hack and tops of the legs are covered \\ ilh minute wart .... T he frog~ are
105
~ U C RO H Y Ll D"E : ~ I ETAPHRY :\ E LLA
Fig. 51 . \ / ('1111'/11"111"'/(1 111/1(/1/1/(/ (Pct('r~l.
grey or light brO\\ll. llsllall~ I,\.ith;/ l;/rge d;ul.. bro\\11 mark lhNlllhe ('entre
of the h.Il·I.... The top~ of Ihe tinger" are )dhm.
l\ t llc . . : 19-26 mm.
Fell1..1k,,: 23-29 1111ll.
Tad,}Olc: L/nknO\\ n.
Habits and habit .. t : T hlc, I~ an abundant ~pecic~ of l1a1 and hilly prima~
!ore~1 of the 10\\ hm(b. :'> Ia[e<, call trom \\ :Iter-containing hole .. III ~rn"lItree~ .
lhuall~ 11(,1\\("CI1 one and thrcl' me:lre~ abO\(" ground. Female . . 1110\C abou t.
npp;/rl'1111~ going to n holl' lrom \\hich a male call" \\hen . . hc i~ read~ to
hrL'cd, The piping call oj thi ........ pecie~ I~ 0!1l' of the charactL'ri"lk night <'Oll1l(.b
oj fore,t . . belo\\ 700 Il1l:t re",
Call ::\ <,ingk plpl!lg note at rcglilar intc"al~. The lone: \;\rle,>_ mUlcating
Ih'll lhmen'lon . . or Ihe hole alleclthc pilCh,
Dbtrihution All pan" of Bomen. It b not knO\\11 I.:be\\here.
106
tvll CROI IYLlD/\E : .\·IICROHYLA
Genus J1icrohyla
Narrow-mouthed Frog')
iH: ~rx'cie~
of Microln1(1 an..' nO\\ known from BOnl1!o. Only one
F ~pecie~. M . benimol'e/, occur::. oubiuo: ur Bornco. /\Illi\'e are small to

tiny and ha\!;' lung hl11d 11mb". The) all li\~' 1!1 primar) and old
"ccondar~ forests be!O\~ 700 mctn.:, ahoye sea level. Their gcncra1J~ brown
colounLtion \\ IIh a hroken. dar~ pallcm make them difficult to :.ee on the leaf
lillcr of the forc1>1 noor. and their capacil~ for long: leaps make .. them \'elJ
dillieuh 10 catch. All ~po.:cic, la) eggs III \el} 1>mall pooh of \\:lIer. Most
spede .. ha\c a dark marling Irom hcl\\een the eyes do\\n the ba~'k: the dark
area narru\\., and \\iden~. tbllall~ ~~'ntling pairs of dark arm~ bac kward and
to the ~ide.
Microlly/a berdlllorei ( Bl yth)

Benlmorc', :\':IITo\\ -lIlomhed Frog (f ig. 52)
I)('scription: A small frog. \~ ilh long hllld limb~. The t:'Olnirum \::. nol
\ hiblt' The ~noul is rather blunt. Four linger .. are e\ idelll . The tip1> of aliloe:.
Fi~. 51, \lum/alll herdlllord [Ul~ Ih).
107
j\!1ICROHYLIDAE: :\1ICROHY LA
are widened and all except the fourth (longest) are full) wehhed to the hase
of pads. The skin i~ smooth. The ('olom is light hrov, 11 or g:r~') hro\\ n. \\ ith a
dar\.. hro\\ 11 mark 111 the centre of the had.. and bel\\Ci.'ll the e) e~ and u~uall)'
v,ith a ~lilming dmk bru\\ll band km un the ~ide. The throat is brown and the
belly white.
I'vlales: 24-28 mill.
Females: 17-32 Tlllll.
Tadpole : Cnkno\\ 11.
Habits and habitat : Thi~ ~peeie~ occurs in primary and ~econdar)'
fore~b at 10\\ elevations. Like all members of this genus. adu1t\ li\i.' on the
ground in leaf litter.
Call : Unkno\\I1.
Distrihution : K1l(l\\ n from a fe\\ localities in Sabah. Sara\\-a\.. and
Kalimaman. II probably ha~:1 wide distribution in Borneo along major river
\·alley~. The ~peeie~ i, also known from Thailand. Penin,ular M alay~ia and
Sumatra.
Micro/iy/a bomeell.'I i.'I Parker

Hornean -'arrow-mouthed Frog (Fig. 53)
Description : A ~mall frog \\ith long hind limh~. The eardrum i~ not
\l,ihlc. The 'I1OUt is ~lightly poimed and projecting hC)lllld tht, mouth. Four
linger' arc 1.:\ ideIH. Tips of all toes are slightly \\ ilkned. The tllree outer toes
arc ,tbout half \\ ebbed. The skin is smooth. Thi" 'pecie~ i~ grey to brown Oil
the back.. \\ith a wide . dark hrown or purple, light-edged mark down the
centre of the back. This dark mark ha\ pairs of arms projecting hackward emu
to the side. The chc"t and hell) arc monied brown and white.
i\lale,,: 17-1 H mm.
Females: 19-23 mm.
Tadpole : The head and body appear triangular. Ilat above and rounded
bclo\\ . The tail taper~ gr:lduall) from its hase inlo <J lung narrow filament.
The emire tadpole is grey or hbck. (',\cept under tbe body. Total length
reaches 22 mm.
Hahits and hahitat : Thi .. "pc("i~" i ~ known only from primar~ fnre<"h in
10\\ land .. _ Adult, !i\(' on thl' rore~t nom and breed at small rain puols. We
fnund <,e\er:JI in \\ild pig \\aIlO\\~.
Call : Un\..1l{l\\ n.
lOX
MJCROHYUDA E : MTCROH YLA
Fig. 53. .lIicro/iyla homeellIiJ Parker.
Di stribution : Kn own from Sara\\ ak . Sabah and nort heastern

Kalimantan .
Micro/iyJa petrigella Inger & Frogner

Pothok Narrow-mouthed Frog (h g. 5.t)
Description : A \·el") small specie!> . with long hind legs. This and M.
plTfl(//")"lI are the only Bornean ~pecies of Micmhrla that appear to ha\·ejusl
threc fingcr~. Thc cardrum i" not \ i~ihlc. The t ip~ of the toe, arc \.\ iUl'ncd and
ull lue" cxecpl Iht: founh (I~l1lgc,t) are rull~ \\l'bbcd to the p:Jd,. Th.: "kin i~
"111oolh. The general wluur i~ light gre~ - bro\\n. \\ith Ihe top of Ihe he:ld
slighll~ darker. A dark mark o.:.:upies the celllre of the back. with three
constriction~ and expansions. The throat and belly are dark brown with an
irregular. large. \\ hite blotch in the centre.
109
t-. IIC' ROHYLI DAE: MI CROHYLA
Fig. 54. \!u-rohyla {Jelrl~(,11(1 Inger &: Frogm:r.
Male,: 1,",-16 llllll.

Female ... : 15-18 mm.
Tadpole : The hcad and bod~ are oval. !l allcned :lbme and rounded
hclil\\. The tail I' deep and tapercu abnJptly ncar the end \0 a ~hon. narrm\
lip. The head ami body art' hl'II,:1.. above and on the side .... The !ail fin~ are
,olourles<, (''',cpt fo r a \\ ide. \enical iliad. band near thl.' r:nd. Total knglh
reaches 15-17 mill.
I-Iahits and hahitat: Thi~ i ~ a specic!> of the tloor liller in prilllar~ fo rem
belm' 700 metres allO\e ~ea Ic\l,:\' Adults feed on tiny anl~ and other
JJl\enCOrale!>. Breeding take~ place :II small pOlhole~ on rocJ..~ banks of
,trl.·'IIll"'. The hole, \\here t:'ldpole,> have becn found \\ere Ie" lhan 30 ern in
di,lllleter and dcplh. The tadpole., hover at mid-depth in Ihe.,c lill) pooh and
tilter tin) mgani~m$ fromlhe \\,lIer,
Call : UnJ....tlowll.
Distributiun : Sahah and S.lr~mak There is no n.:a'on 10 behc\c thallhis
"pecic" dOt"~ not occur in Kalimantan ;jnd Bmnei.
11 0
:\11CROH YLlDAE: MICROHYLA
Microhy/a perpan-a Inger & Frogner

Lca~t Namm-moulhcd r-rog (Fig. 55)
Desc riptiun : Very similar to M. f'e//"igt'II(1. Onl) three finger, an:

<:vident. The principal diffcrl'ncc" from pefrigellG are in size and colouration.
MicmhY/(I p('/pan·a ha~ a du~"y throat. but the belly is entire!) \\-hite.
Maks: 10-12 mm.
Females: 13- 15 mm.
Tadpole : Head. hod~, and tail are ~hap\;'d as in tadpole of M. perrigclla.
Colouration aho .. imilur to lhm of M. pelrigellrl. Total length reaches 10-12
mm.
Habits and habitat : This "pecic:'; li\..;s in Ilat or hilly primary forest at
low elevations. Adult" hrccd at rain poob in the forest.
Call : Unl...no\\ n.
Distribution : Thi~ ~pecie~ has been found in Sar;mak at ~cveral areas. bUl at
only one site in Sabah and nonh central Kalimantan .
Fig. 55. Microhrltl/!Nparl"ll Inger & Frog-ner.
111
f((J n ily 1(ClIl icl Ci t'
Thi~ i:. probabl~ the mO~1 familiar group 01' rrog~. It on:urs almost
everywhere in the \\orld and is the largest family in Borneo. where its
members include ri\'er. "Iream. and forest floor dwellers.
FejerJ'orya COflcril'ora (Gra\ enhorsl)

i'.lungroH' Frug (Fig. 56)
Description : A medium-sized ii'og with a long snout and well-muscled

hind limbs. The toes are more than Iwlf-\\-ebbed \\"ith the ends of the toes free
of\\chhing and not ~\\ol1en. Tht' eardnlTll is cOI1<;picuous. The hack and __ ides
hu\c ~horl riJg<:~ and round bllmp~. The rrog~ arc brO\\ 11 to gn.") on the back
and legs . \\jlh dar" markings. Then~ is a dark bar acros~ the top of the head
between the eyes. and fOllf to fi\'e dark bars across the lOps of the legs. The
underside of the head varies from white to white with dark mottling. Ma]('"
J13
RA:"IDAE : FEJERVAR 'l A
h;:i\t 3 d3rk area umkr the cunK'r of the j3\\ in the ,kin o \erl)ing the \ocll
"a("~. Some indi\iJll;ll~ hu\~' uurJ... ~pOh on the chtst and bell), \Iherc oth{'r~
an: cr(,311l~ \\ hite .
i\lak~: 51-7() 111m.
Female,: )3-8~ 1l1111 .
Tadpole: The tadpole ha~ Ull 0\31 hod~ and is dark grey \\ith darker
"pOls. The t3il i.. lt~ .. Ihanlll il'e till' bou~ length.
Hahits and hahitat: Thi~ i~ a frog of di"turb~' d habitat-. Ilear the coast
and in mangrmc. It i.. the on ly \1 alay:-.ian frog Ihat lokrat ... ~ ~ulinc habital~.
Although in Penin .. uiar J\laLl~ .. ia it occ urs in coastal rice field~. it i~ not
knO\I'n from ~ueh pla~'e~ in Romeo. Male, do nO! form c3l1in.? groups. blll
call singly from riler banh.~. ditch ..... ,Illd other l'Oa'-lal IlalC'T\\a~". The diel
nm .. i .. t.. of a I'ariet~ of small iJl\er!ebrate~. induding critb).
Distrihution : Kalimantan. Sabah and Sara\\"ak. Al~ll knO\ln from
,outhern Thaiblllito Jal:\ ulld thc Philtppine Islands.
Fej en'OI)'a Iimlloc/wris ( B oiel

Gr~h' Frog Ii·"ig. 57)
DCSlT iption : A ~mall frog II ilh a long narTfm head and a 'knder. 0\'31
body. The toe, are pointed. le~~ than half I\ebb~d alld lh~' linger.. url' al~()
pointed. The em'drum i~ I·i~ible. The :..k.in i~ fine!) pebbled. \1 ilh a ~ni\.'" uf
1111\ . lIlternlpted ridge~ running dO\ln the back. becoming a line of bumps
both on the rump 3nd the ~ide~. There i~ a fold of skin from the rear of the
c);,: mn th~' L'artlmm. The frog, are niSI: hro\\11 10 brownish gre~' abme.
\\itb darJ...a blotchL'~ Oil the hacJ.... Th~Tt.' i~ usual1~ a LJ - or \\ '-shaped marking
across the ~houlder". i\ lmt indi\ iduah uho hu\\.' u light '-lrcak dowl1 the
middle of the back li'om the nose 10 the :lnu~. The lip~ arc e()mpicuou,l~
haITed brol\n 3nd \Ihite. The 1Illder~ides of the females me complek'l: \\ hite.
J\ l a1c-~ hal'e a black. M-shaped b3nd aeras, the throat.
J\LLk,,: .1:!-50 mill.
fenwle\: -I-9-5~ mTJl.
Tadpole : The t'-1tlpok hu\ an mal hod~. which l~ 3bout half the length of
the tail. The bod: j .. olin;, gr\.'Cll abol e and Ull the \ ide<." ~ped..1ed \1 ith black.
The under,ide is \1 hite . The end of the tail i~ barred Ilith black ur i .. I.'lltirC'l:
hlueJ.... Total length re3ches -1- ) mm.
Hahit s and habitat : The frog i~ confined 10 di~turbL,tl hubilah
~L\"()l"laled \Iilh the> <lctil'ities of manh.ind. such a .. agril'ultural lil."ld,.
RANTDAE: 1I0PLOBJ\TRJ\C H L:S
Fig. 57. fi'jc/lWTII/illll1lw/wri.1 IBoie).
roadsides. lawns and football pi t ehe~. Thu~ it i~ (1]ll' 01" lhe rnml common
rrog~ in \ ill ages and !Owns. It gather;, in large group~ around ~tanding wuler
and l·honl~e' are often heard well into the morning after a night of hea\} rain.
rt eah a \ariel) of in~ect<;. millipedes and. occasionall). snails.
Call : A rasp~ ehirp. rapiJly repeated.
Distribution: Widel) diwibull'd in all parts of Borneo. \\herc\cr the
fore<;ts ha\·e been remO\ed. It m:curs IhruughoUltropical A~ia.
Hoplobatra chw; rugllloSliS (Wiegmann)

Tai\\anese Frog (Fig. 58)
Description : ,\ large. powerfully built frog \\ith a broad triangular he:ld.

The llle~ an: lull) \\ehhed and rounded at the tip<;. The middle two flllger<;
ha\e a fringe of ,kin along the inner side. while the other finger, han' no
fringes. The tip~ of the jjnger~ arc pointed. TilL'" eardrum i~ larg.e. ThL' ~]"in i~
extremel) granular and rough. \\ illl man) ~callcn:J ... mall to large bump'>
along the sides of the back and rump. There arc al~o man) inlenupled.
115
RAN IOAE: HOPLO BAT RACH L'S
r ig. SX. H"I,/ob(lIrodlll\ m~,,'osll\ I \_\~"::.·~~"~'"~"~"~I_._ __ _ _ __ _ _ __ __
pr()]l1llk'llt JlOm) riJge~ of ~ k.in running JO\\ i1 (he had , anti upper ~ lI rt'al'e~ of
(h~ Gjh .. ~. T h ... ~e frog~ are bn.mll (0 gr""~lli"h g re~ 'lhO\\? \\jlh ~l'a!l e red darJ..
"1"\(1" un Ihl' had and leg~. r he ~idc,> afe ~poucd or moult-d. fOfl11lng a
p.lUl'm or IK·(\UlrJ... Inlnrge IIldi"iduab The ullder~idl' (1ll he head I" ,\ h1l1,h
\\ Ilh ;1 di~linl' l dark '-IrcaJ... i..IO\\ n the midi..lle of Ihe throat In '(line lIli..li\ idll.l"
Ihe throat i~ he.l\ il~ ll1(lllkd II ith d.1I"k ,POh forming a nell\OrJ.... The chc~1 i~
hgbtl~ ~p(}!led. and lht' b.:lI) and ulllkr'lIrface<; of the l eg~ are lI'llal1~ \\hil~' .
i'd a l \?~' 7()~ J OO 111111.
PCI11;t!c": 1-:5-1:25 nUll.
Tadpole: nIl' t.ldpole, ofthi, "pe.:ie, are 1,Irgc (up til XO 111m). and hale
he;l\~. , harp. hla("k hen" ,. T he body I' \e r~ darJ... ;lbO\~. ,mi..l ,0IlK'\\hill
lighter on th ...· -.I(k, \\llh Irregular hl;ll'I.. "pOb.
HlIbit ~ a nd ha bitat. Thl" frog h nO! nati\"t~ to l3ofl1e ...). II \\a~ app'.IT("J1II~
t!llported intu Sabah from E a~l <\,ia (ptH, ih le Tai \\anl I!I lhe 1960, U~ a
'{lu rl"~' (If meat. II h:I' nU\\ 'prt!:ld Ihwughuul "ahah trom Kota Bell.Id to
S;mtiaJ...an .:.tnd flI.)"lhl~ 10 \\e,tefl1 cO;l ... l<Il Sar;t\\aJ.... It '1..''':111, to he limited 10
di'lU rbo:d .:.tr.:a~. tmlll rice fielJ, to ("on'iI"lKlion ~itc, The frog' gal her III
breeding gfllUp, ilfI)unJ temporar~ pom\... :1Ilt! Illal.:, l'all Imm Ihe,,~ area, of
110
RANIDAI:.: IIUIA
~t:lnding \\ all'r :.lner heal ~ ,toml~. Thl'" 'pecie" C:.l\!) a II ide \"arict~ of
imenehralc' :lnd probahl~ ahu prey" on ~m:lller frog). In Southea.;.t A..,ia. the
ladpole' of Ihi~ ~peCH:' af<: ~nown 10 !o.:~· d on ,mall cadpolt!':-.. The) :.llmo'>t
ccn'linl) han~ Iho.: ~ame hahit'" in Borneo. Sin~e mv~t t;ldpole~ of Dlher
~pecie, that liH" in Ji~ t urhcd o.:mironmenh abo de\\']op in temporary pond~.
the pro.:li:llOI) behrn lour of n(~1I10.\11\ tadpole,> ma) account for the ~uccc~:-. of
Ihl\ 'pedes as all i11\ adt!'!".
Call : J\ 10\\ ra,py lIole. ro.:pL'aled mpidly 10-20 lime~.
Ui<;ll'ibution : In Boml'O I..no\\"11 onl~ from Sall:.lll :lIld \\('q SarawaL The
~~cio.:, lICl·UfS III opcn. di:-.turbed el1\ ironmeJll~ throughout Southea"t A:.ia.
H"ia cUI'ily mpml/llll ( BoulcngerJ

Holc-in-Iht!'-head Fwg (Fig. 59)
Uc<;c ription : r\ rneJiu11l-,ized 10 large frog. II jth a roulll1t:d ,nout and

long h11ld limb,>. Thl.' lip'> of the fingl.."f' and IOC', are e\.panded into .. lightly
poimcd pads. All 10C.., :lre full~ \\l'bbed \0 the bil:-.e of the pad". Thi:-. i~ Ihe
onl) ..,peeies ot frog in Borneo ha\ing the I.'ardrum wn"- into a depression.
Fig_ 59. IIl1illl"llll/llIIjl(lIlWII (Boukn~<.:rl.
117
RA'\!IDAf::: : L'·G[RA1'iA
T he ... I..in I" ,month c\cept tor ... mall. round hump' on lhe ... idl', and
rougllt'nL'd an.:;}.; on the "IIJt'" of Ihe ill'ad. TIll'" top of th..: head ;lnd thl' had,
ar..: chocolate bro\ln. The "de, of the I1c:ld and boU~ :lnd the upper "urfaces
ofth~' limb, are tan \Iilh dar\... hl"Ol\tl "pol". Throat. (:hl',t and hell) are \\hite
or yt'llowbh mangl'. Th..: upp..:r hall (If the iris i~ light gold and the 10\H~r half
Clllllamon bw\\ 11.
r-.lak~ : ·-1:::-52 mm.
I-"elllale~ : 75-80 mm .
T:ldpol e: The tadpoles (II" Ihl'> "pc-cie' are amollg thL' large"t III Borneo.
re;ldling ,I kngth (If 70 mm . They h'l\ (' an :I!xIominal ~llc\...er ell,lhltng them
10 ding 10 rtle].;~ in ,wili ClIm'm" Th\..' Hl{/~dc of the l;li! i .. d;Irk \\ IIIl JIg-Ill
'pOb. and the lin .. arl' aho d:lrl.. but ha\~· a lighl hand along Ihe ba~(' :Illd light
~p~lt' :llong: Iht' margin,.
Habit s and Imhit .. t: Thi" ~pc.:k, Jive' In primM) 1'01"("( in htll) cuulI(r~
from J50 I\-I 750 metre" aho\(' ,e,\ 1,:\(:1. Auul t ~ \\ander \\Idel) through the
fore"t. hUI il i, nlll \...nQ\lll \\hether the) feed on the ground or ('limh up inlO
\o:'gct;llion. ThL'y breed onl~ along ~m~tche~ of ~\I'ift \Iater in rnedium-siLl'd
Hllargt: 'Iream'. The tadpok' IiI C In Iht' ,lrongt:,1 current, on Iho~c 'Iream~.
dlllglllg to beJmck or hould..:r~ \1 llh Illeir :lhdolllinal ~lK \...t'r.
Ca ll : \ \lcaL high-puched dlJrp
Oi!oll·ihutiu n : Ka!imant;lll. S;lbah and S3r:l\\aJ.... Thi' ~pe\':k' ha~ ne\er
bet'n found ollt!tidl' of Bo01eo.
Iflgerau{I ha/{wl(.f,iJ (Boulengcrl

lngt'r""" DW,lrf Frog (Fig. 60) •
De<;criptioll : A \t'1) )llIall. "lflCl\y trog \1 ith a \\ ide. ~Ollll'"\\ hal pointed
head and ,hurl hind limb~. It ha!t ~hol"l. ,tubb~ lingen with broadelll'd tip~.
The ttl!.:" haH' widened lip~ and ,Ire ahoul half \\ehhed. The ,I..in of the head
and back i" pehhled and Ofll.'l1 sel \\ ilh !thor!. narTO\I ridges. Tho:' bel1~ and
Ihm.:!! aho haH: a pebbkd 'lIrface. Colollralion i~ \;Iriabk. Som~' indi\lullab
are medium bl"O\\ 11 I\ith ,mall. ,..:arten.:J di(rJ.. "POI~ on Ihe b.1ck ;mJ d:!r). har'i
hel\\ Cl'n Ihl' nostrih and het\1 el'l1 the ('~l'" Olhc-r indi\ Idl.lal" are gn!) -bro\\ 11
lin top of tIll' ,nout and nn the ~idt:". hut arc dar\... bfll\\ 11 (In Ihe bal:\...
ix'gilllling bt'1I\eell Ihe e.It.'\. II) ,Ii)) mht'f\. Ihe back lw, a dill"!.. longiludmaJ
!land \\ ilh deepl~ indented margilh. ,\ fe\\ are dari;. br(l\\ n e\~·l.'pt for a lighl
~ Ir..:ak llll e,lch ,ide 01 Ihe hJel... The lilt',t ~nd bdl~ ar .... pall;' gre~ llr \I hite
The thn.at I ~ u,uall~ ht'":l\ il~ JOl1l'lI or marhled \\ 1111 brO\~ 11 .
118
RANlDAE : LIMNONECTES
Fig. 60. 11!!!l:'l"ill/a halllemil (Boulenger).
~ I ales: 20-~5 mill.

Females: 24-30 mm.
Tadpole : LnknO\ln.
Hahits and habitat : Vcr) liuk i~ I-..nO\\n about thi~ fore<;t frog. 1\11
ob"ern:d ~o far were ~een 011 the fore~t floor. in ~Iopillg seepage areas. or at
(he edge~ of :.tremm. The expanded tips of fingers and toes suggest that the
specie~ climbs.
Call : Unknown.
Distrihution : Known ol1h from Sahah :mJ SUT:l\\ak.
Genus LinUlOllectes
his group of species has recentl) been ~p1it off from the genu~ Raila
T (set' p. J-I-J). Although the) ~hO\\ much \ariution in .'>ize. ~kin te\.ture.
\\LOhhing. lind brceding habih. they .'>hure man) features of adult and
tadpok anatom~. The tip~ of the toe.'> are slightl~. but distinctly. S\\ollen and
the toc, arc half to full~ \\ebbed. In colouratiollthey are all shade;; ofhro\\ n
or hlad. Thl' tadpole:. h,l\e elongme m'al bodies. rather naITo\\ taih. and l!
I J l)
RAi\"IDAE : LLMNONECTES
(,,:on .. istenl pattern 10 th~ rows of tin) "leeth" on the lower lip. The~ are all
t,:nT .. trial. ~ome \\anderillg widely through frlfo.'qS and others spending their
enlin: li\(?s on .. tream banb. t\~ a group. they ar~' among the most abund,mt
of the rrug~ in Bornean fon."s\;,. E\ery area of IlWlUre rure')t has three 10 five
of these spe(:ie~.
LilllllOllectes mperata ( Inger. Boeadi & Taullk)

Rough-backed Frog
Description : A ~mall. sto(,,:k) frog. \\ ith 'hon. thick legs. All toes, e\o:pt
the fmmh (the longl'~t). are webbed 10 the ba~".. .. of the slightly expanded tips,
Tht' eardrum is \·i~ibk. but partia!!y covered b) ~kin at it.. uppL'"r rear corner.
The cntire upper ,>utfa..:e i.. wt \\ ith crowded. conical or .. lightl~ o\'al warts.
e\ell on the upper eyelid and ~nout. TIlt? colour in preser\"ati\~' (\Ie ha\c
neler ~e..:n one of these alil'e) i~ sand) bnm 11 1\ ith faint darker m:lrk.ings that
are probably blad-, in life. The underside is I\hitish, \\lIh small brown sputs
on the chin and Ihroat.
\-ta!es: 35 38 mm.
Female (only one j~ known): 3fi mill.
Hahits and habitat : Th".. onl) known represelllati\'es or thi~ .. p ..xie~
Iler~' cullected on the floor uf a klldand forc~t Ihat was being loggeu in
south -(,,:<.:ntral Borneo. though the ~pe~ic~ pnlbahl) has a wider di~lribulion
\\ ithin the i.. land. The only adull female J...n0\\ n had a small number of larg.e
(relatin:' to thc .. ill' ()f Ihe female) eggs. suggc';ling a modified type of
hreeding behaliuur. II may heha\'e like the Smooth Guardian Frog
(Lilllilonecies palmYlllell I·is). II hich guards ib eggs and tran:spurh tadpoles.
Distrihution : Recorded \0 far onl) from sOUlh-central Kalimantan .
Lil1lllollectes IiI/chi ( Inger)

Rough Guardian Frog I Figs. II. 12 & hI)
Description : A small. modnatcl) <;locky frog with reJatilel) lung hind

limbs am! thid.: Ihigh~. The loe:s ar..: unl) partia!]y webbed. and ~lightl)
s\mllen <II the tip~. The eardmm is lisibJe. Thc b,H..'k has a di~tlt1ct illlel1ed
V-"haped ridge in the 11liddk and a serie:s of interrupteu ~horl ridges along the
.. ides. The upper c~"..lid~ have a number of small bump ... Thc frogs are brown
to dark bral\ n. \\ilh dark markings. including a dark har across the top of the
120
RANIDAE: L1MNONECTES
head between the eyes. The chin and throat are darkly mottled. while the
che~t. hell y. and underside of the thigh are either white or ha\e a } \:110\\ ish
tinge.
Males: 32-39 mm .
females: 39---t5 mm.
Tadpole : The tadpole- of this species has a slender. mal bod) and a tail
shaped like a ~lender leaf. The mouth. ju~t bc\o\\ the lip of [he snout. is
narrow. The body has irregular dark spots. and the taillws venical dark bars.
When nearing metamorphosis the tadpole is about 25 IllIll
Habil'i and habitat: This frog live" on the floor of primary and old
secondary forests al low ei.-valiollS. It doe~ not occur in swamp forests.
Males call singly from under dead leaves at night. A fenwle Jays its eggs next
to the male. who remains with the eggs. Usually the eggs are stuck to the
underside of the leaf ~hcltering the male. When the tadpolc~ hat<:h. they
somehO\\ attach Ihemsehe~ to the back of the male \\ho then carries them to
a rain pool or a small intermittent stream where the tadpoles swim off to
complete their development. Adults feed on small insects and spiders.
Call: A short trill.
Distribution : Kn()\\l1 onl) from Sahah. though it rna} occur in eastern
parIS of Kalimantan.
Fig. 61. Limllonecresjincili (ingcr ).
121
RA!\·WAE: LlMNONECTES
Lil1llloll ecle ~· ihall Orlllll (Inger)

Rough-backed Ri\l.~rh og (Fig. 62)
Desc ription: A lurgL'. robust frog \~ith a long. lap.:-rl'd ~nOl.I1. The fingers
;!r':-rounded ::md blullI. \\jth IlO \wbbjllg. allhough Ihert' ar.:- narrO\\ jbp~ of
skin along both edges of the sccond and third lingers. The toc~ are blunt and
full) w.:-hbed. The Cardl1.lIll is \ i~ibk. A serie~ 0[" ~hort. mostl~ parallel ridges
cmer IllOq of the buck and the sJ...in hctween them ha" low radiating lint's .
.'\duhs ur~' u ... ually grc) i~h hrown to bla,:ki~h bro\\n . and th.:- thro:l1 is \\ ruti .. h
\\ ith dad, blotches. The re~1 of the lower surface i" pure \\ hitl'.
\llak'>: 80.-130 \11111.
FeJ11ale~: 80·.... 101 mm.
Tadpol e: The bod~ lS oval and n.:lati\ely slender. The tail is also slender
and aboull\\ice the length of the bod). The overalJ colour i~ greyish brown
with a darkcr mottling thai e\tt'nd,> on 10 the tail. r-. lature tadpok . . are about
J:; mIll long .
Habit.s and hahitat : Thi ~ spccies is common along clear. ro,:k~ streams
10 \0 J() metres \.. ide in primal") and old secondar) gro\\ Ih forest s in hill)
count r) . Adults are !lCVer found more Ihan a few rnetre ~ from stream banb.
Fig. 61. !.i11l/1Otlfern ill(llWfllll1 (Inger1.
I"
RAL'iIOi\E: LrMNO:-.lECTES
They feed on large in~ect~. but also lake crabs. ~mall lizards. and prohabl~
rrags. The tadpoles li\ e in quiet ~idc pools of thc~c rocky strcmm and reed
on organic dcbri~. ~uch as decaying leaves.
Call ; Male~ do not have vocal sac;; amI h<l\'e never bl'cn heard tu call.
Distribution : Brunei. Sarawal... and Kalimanian. The specics does nOI
occur in Sahah ami has neWT been found oUl<;ide of Borneo.
LimllOllectes illgeri (Kicw )

GreateT Swamp hog (Fig. 63)
Description : Thi~ i:- a large hC<l\)-bodied frog. with heavil) muscled

kg~. It is similar to Li,WWIU'cle.\ /epc)/"iIIUL excepllhat the top and "ide of tho;:,
~notll do not rOml a sharp angle as in L. lepnrilllls. but arc instead rounded.
The eardrum i~ very di~tincl. The toes an: fully webhcd and have ~Jightly
swollcn tips. The bad has a fev. ~hurl ridges and the sides ha\e a few
scattl"red bumps. Thl" upper eyelid i~ rough. The upper surfacc~ are reddish
bro\\ n to dark hro\\ n. with indi~lind darker markings. Unlike L. leporil/l/s.
this species ha~ no dark stripe from the eye 10 the nostril. The chin and Lhroat
Fig. 63. LI/IlIIOIIl'Clt'j ingeri ( Kicw).
123
RANIDAE : Lil'vINONECTES
are hca\il~ mottled \\ilh g:re)-hrown. The chest and bell) arc greyish white
\\ ith mJistinct mottling,
:\'laks: 75- 132 mm.
Females: 70--127 111111.
Tadpole: The lUdpole~ of this specie:> look \ t'r) similar to tho<;e of L.
{('poril1l1s and L. Ibo/lOI'lIlIl . II i~ oyal and "lender. \\ith a lail about t\\ice the
length of the bod~. TIll' hody is irregularly spotted and Ihe tail i~ speckled
with dark bro\\ n or hlack.
Habits a nd ha hitat : This species li\'e~ in both primar~ and disturhed
forests. and nw) c\cn he found in rubber plantations. It i~ ofttn encoLintered
in swampy arc ..!'> and along muddy streams. Adult~ of thi~ ~plTic~ arc always
found ~ing ly . Tadpok~ live in quiet side poob of stream~. Adults feed on
large pre). including crahs. frogs and e\'en sm;lil ~nakes.
Call : r-.lak~ han.. no \'ocal sacs ,lIld are not lnO\\ 11 10 call.
Distribution: Bmnei. Sabah and Sara\\ak: nOI I-.nO\\ 11 ~)uhide of Borneo.
LilllllOll ectes kuhlii (T<;chudi)

Kuhfs Cred frog (Fig. 6.+)
Description: A ,hort <;tocky fmg. with heavy legs. Adult mall."'~ h.. t\'C a
grotesque!) broad head. The eardrum is not visible. The toes an.: rull)
webbed to the \Iightl) ~\\ollen tips. [n rnale~. the front of the 100\I,.'f jaw ha~
a p!.lir or large tooth-like projections. which are \"i~ible only \\ hen the mouth
i~ opened. The skin of the back and ~ide~ has man) .. mall. rounded humps.
Frogs from most pans of Borneo ha\e the hind limh .. co\'O;~red \\'ith \'ery
rough warts. each of which has a ~maJl \\'hili~h cone at the centre. In parts of
Sarawak the frogs ha\"e relati\"e!) smooth kg\. The upper surfaces are dark
grey to black with jndi~lincl blal·1-. hlolches. The chin and throat are he,I\'il)
sponed . \\ hile Ihe rest or the undcr\idc i~ usually white. The ilnderside of the
legs may be darkl~ ~polled.
Male~: ++-7'+ mm.
Females: 51--67 111m.
Ta dpo le: (Jenerall) <;irnilar to the tadpole of L. fepor;lIm. \\ ilh an mal
bod) and tail <.Iightl) less than twice the length of Ihl' bod~. The hod) i,
grl') ish bnl\\ n \\ ith a narro\\ dark band aeros~ the bod~ jU\1 hehind the eyes
and a .. ('cond band :1I the end of the bod). The lail ha.. ~trong black \'('nical
hand;;. When full~ gfll\\ n Ihe ladpole\ arc 25-JO mm.
12.-+
RAi':IDAE: LlMNONECTES
Habits and habitat : This frog li\e~ in primary or old secondary fore:.ts
in hilly country from near sea levclto 1600 metres. It is almo~t ne\'er found
more than a fe\\' metres from Ihe banks of small to medium-sized streams
(5-10 metre~ \\idc). [t is common along such stream:.. but usually rare where
the Giani Rjver Frog (L. leporil!LI.lj is abundant. Kuhl's Creek Frog clearly
prefer~ rocky areas with moderate current and may he seen at night sining on
rocks within a fcw centimetres of the water. Adult:. do not form groups.
Tadpoles arc found in very small stream<;. or even in isolated pools of streams
thaI have interrupted flow.
Call : '\1ale<; do not have vocal ~a('s and those Ii\'ing at 10\\ elevations do
not call. Our colleague. Tan Fui Lian. has called our attention 10 the fact that
males Ii\ ing at higher elevations on ivlount Kinabalu emit a soft ··\wee\".
Distributiun: Known from all hill} pans of Horneo. The species also
occur~ through most of Southeast Asia.
Fig. (j-l. Lillmonecres ~lIl11ii ITschudi ).
125
RAN lDAE: L1 MNONECTES
LimllOlleeles la/ieeps (B ouienger)

Rivulet Frog: (Fig. 65)
Description : A 'rmll SlOd} frog with heav~ hind limb~ and:! \\ ide
head. It i" \'er)' ~irnil:!r to Kuhl's Cret'k Frog: in general ~hape and eolour and
in lac1.ing a \ isib1e eardrum. But it is !>malkr. and the toe~ are k~" than fully
webbed. Thl' ,kin of the bilek ilnd "ides is \\ rinkled and has scattered, ~l1la l l
humps. The~e frogs are brown or o1i\O.,:-brown \\ilh irn:gular black mar k.i ll g~
on the back. TherT i, Ll~uall) :1 dark ~trcak bet\\een tht' e}cs and red and
bhu.:k har, on the lip~. The chest. bel!:, and under"ide of the leg, arc lemon
yellO\\ .
M<lles: 18- 35 mlll.
Felllale~: 32--+0 1111ll.
Tadpole: Ln]..mW,Tl, but probably \ery much li ke tho~e of the related
specie;; L. ~uhlii.
Habits and habitat: This "peeies ha~ been foum.1 on the banb of ,mall.
intennillenL "wamp)' Stream~ and near stream~ in primar) forest at elevatioll'
below 500 metre". Little is knO\\ n of its habits.
Call : L'nkllO\\ n.
Distribution : In Borneo knQ\'>-!l rrom Sarawak and north celllral
Kalimantan. The ~fX'cie~ also occur~ in Pel1in~lllar f.. lalaysia and Thailand.
Fig, 65, LiIllIl(JIIl'I"/£' Imieeps (Bllukng ... r).
126
RANlDAE : LlMNONECTES
Limllollecles leporillllS (Ander~sol1)

GianI River Frog (Fi~. 66)
Description : A large frog \\ith a rather long tapered ~nout and large.
pO\\-:rful hind limbs. The top and side~ of the ~n6ut meet at a sharp angle.
and the eardrum j" !;"learly visible. The toes arc full) \\ebbed and slightly
~woJJen at the tip~. The hack has ~l'allered ~hort ridges and bUl11p~ that tend
10 disappear a~ the frog gels larger. The Uppl'r eyelid also ha~ ~malJ hump~.
The frogs arl' reddi~h brown to brown. with darl marlings. There i<; always
a dark streah on the side of the snout between the e)c and the nO~lril and
often a d,uk W-sh,lped mark between the ~houldt'rs. p:micubrly in young
frogs. The underside is \\hite with gre) mottling on the chill and throat.
j\ laks: 90-175 mm.
Females: 85-115 rnm.
Tadpole : The tadpok has a .. lender. oval shape and a long tail. The
momh is just hdow the tip of till' ~nout. The general colour is grey-brown,
with durk mar].jngs. \\ hich foml all alnlU~t zig/ag row of dark spots on the
tail. When rl."ady to metamorphose. tadpoles of thi ... speci.:-" are abolll 36 mm
long.
FiA. 66. LimIlOIll'('/cS ft'porillllS IAndcrs~on ).
117
RANJDAE: LL\'l NO:\I::CTES
Hahit s and habitat: Thi~ i~ a frog ofhoth primary and di .. turht"d fore~\ ...
from ncar ~ea klel to about 750 melre~. It Jil"''' along banb of large or
mediulll-~izt"d "(ITaIm. either turhid or dear. \laJe ~ hUll' no local ~acs and
no "adlerli .. e1T1t"nl·· call. They afC u~uaJJy I\idcly spaced. The male ~coops
out a shallO\I dt"pre~~ioll at a ~and or gral'eJ bar wht"re the remale la~ ~ her
egg ... The ladpok~ live in .. hallO\I ~idc pools and in accumulations of dead
leale". II hich arc a common kalUrt" of the ho!!om of B0111ean fore~1 ~lreams.
t\dult~ ked on a larkl;' of large prey. including crabs and e\t"n other frog'>.
Call : CnknO\\n.
Oistribution : round throughout Borneo. when'l cr suit:lbJe ~lrcanh and
forl"\ occur.
:"lote: In the Fro;:'1 (Jf Sab(lh hook. thi~ .. pecie~ lIas ealled Rmli1 blythii
(nolI knov. n a\ UI/II/(JIlt:rles blnhii!. Taxonomic .. tudy in the 1a~t three year ..
has ~hO\\ 11 that the BonlC'o frog i~ a ..cparate ~pecie~ from L blythii proper.
Idllch appear .. to be eon lined to Penin,ular \Iala)~i ;l :.md Thailand.
Uml10lle('(es malesia"m' (Kic\\')

Pem SII'amp Frog (Fig. (7)
Desc ription : A large. ~tout frog: with a broad ht"ad and thick. ~lOck)
limb~. The toc,> are abolilthret"-founh" Ill'hhed and arc ~lightl) ~ Inlllcn ;ltthe
tip~. Tile eardrum i~ li~ibk. l\;lale~ halc long fang-like qructure~ on tht"
lm\('r j:l\\ a .. in Kuhl's Creek Frog. The ~kill i~ generaJJ:; "ntnoth. \Iith "omc
"callereu 10\\ bump" 011 the b;lck and . . ide ... \ lost frogs arc reddish bro\\ n.
\1 itll jewer ~peeil1lelh a cho ...·ulatc hro\\'n. Se\ eral Ilide black hal'S rtlll from
the c~e acro .. ~ the lip". In mo~t indi\"iduals . there is a fine. II hiti~h Jine
running Irom the tip or the snout. bel \\ een the e) e~ and down the middle of
the back. Therc i" often a '>lmilar fine linc on the uppcr ~urface~ or lhc legs.
The throat and chest haw dark rnonJing. and the under-.ide of the hind limb
i.. vcry dmk.
\lak,,: 70-1::;0 mm.
FelllJ.k~: 7::;-95 mm.
Tadpolt> : Tadpok .. are !>Jim. ola1. and hal e .. lender tail~. The colour i"
golden bro\\11 ll1011kd \\ith dark pigment. Tot:lllenglh reaches~::; mm.
Habits and habilal : Thi .. frog i\ t)pical of p'~ at s\\amps and kl\\ alllll iaJ
plain .. up to ahout 150 m<.:tr(·, aho\'t~ sea kl('1. It OC('Uf'> in hoth prim:..n~ and
diqurhed forc\h. ~uch a., Ilkl mbber plantation~. \\'e hale ob~erll'd it
hrel'Jing on om: occasion (north central Kalimant:ll11 in tht" middk oj a
128
RANIDAE : L1M~ONECT E S
fig. 67. Lim/WII{'CleS main/iII/liS (Kicw ).
broad. ~hallow qream \\ ith a slo\\ !:urrent A pair \\ ere seen ~itting in the
middle of a circular depression in lint" gra\'el. \\ here the female had expelled
her egg~. The diet cOllSist~ of;1 \\ ide variety of il1\enebratc~ and small frogs.
C all : L:nknO\\n.
Distrilmtion : Kllo\\ 11 from Kalimantan. Sabah and Saraw;1\.... Almost
t'ertainly it OCl'ur~ in Brunei \\'here much ~ui table habitat exi~ts. The specie,
aho occur~ in Penin,ular .\Ialaysia.
LimllOllectes pa/al'tlllell sis (Boulenger)

Smooth Guardiun Frog (Fig. 68)
Ilcseription : A ~mall frog. \\ith long sknder hind lirnb~. The I()C~ are
olll~ about hall"-\\ebbed and h'l\e ~lightl} ~\\ollel1 tip~. The eardrum is
vj,ible. The "\...in of the hac\... i~ ~mooth ewept for an im ened V-'haped ridge
in the cenlre and a thin. l"ontinuou" ridg..: from the e)l' to the (urnp. The hac\...
is reddish 10 chocolate brO\\Il. \\i1h ;,} dar" bar acro,~ the top of the head
bl.-'tween the eyes. The imel1ed \ '-,hared ridge 1" al"o edged in black. The
129
RANIlJAE: LL\I NONECTES
fif!o 6K LimIllJIIl'c(t's f"tlul"rIlIt'lI,i, ( Buu/cng<"r),
umkr<;ide of Ihe head j" g.reyi~h \~ hi te. while the chest. beLl~, :l1ld undcr<;ide~
of Ihe legs are usually lemon yellow.
l\1ale,,: 25- 30 1111ll.
Femak,,: 35-40 mm.
Tadpole : The ladpole~ are small. with the tail about twice the length of
the hod~ and tapering abruptl} near the end to a nam::l\\ tip. T he body is tan
or dark brown ami monied \\ ilh darker pigment. T he tail is speckled with
bhH':k.. Total length re ....:hes 30 I11Tll.
Habits and habitat: T his frog: li\"es 011 the tloor ()f p rimar) fore~h from
/)c'<l!' sea Ic~\ d (0 I.~OO Ilk'lre,>. III hahir.' me \ cl}· similar to tlzo\(' of L. /vvlil.
M ale~ are solilar~. calling from under dead leaves. Egg" are laid in a Illannl'r
"iTllil:.tr to thaior I.. Jinchi, and the m:.tk carric~ the immature tadpoles on hi"
back to ,mall rain pools or i"obted pools in ~mall ~treatll heds. The tadpole"
complete their de\elopmem in these ~1l1all bodic, of \\:lter. Aduils prey on
alll~. tl'rmit..:" and oth..:r "mall inn:"rlehrall''' of tiH~ fort:st tloor.
Call : A ~hort. high -pllched trill.
Di ~ tributjon : Kalimantan. Sahah amI S::mJ\\aJ.... It :lbo occur' on
PaJ.l\\an Island in the Philippine ....
130
RANIDAE : LlMNONECTES
UI111l0llectes paromocrodoll (lnger)

Lesser S\~arllp f-rog (Fif!. 69)
Description : A medium to stock.) frog. \yith moder:nely long and

muscled hind limb!>. The "TlOllt ends in a rounded point. The eardrum is
di<;tinct. The tips of the toes are di!>tinC"tl). but !>lightly. widened: the tips uf
the Ii.nger~ are not. All of the toes except the fourth (the longest) are webbed
almost to tht'" tip!>. The upper sllrbc,.... are roughl) pebbled and there are a few
low bUf11p~ on the back. Thc helld and body arl' greyish to reddish brown
ahm c :.md on the sidt'"~. \Iost indi"idu als ha\ e a dark strea\.. on the side of the
snout from the t'"~ e 10 the nostril. T Ill' di~tilletive feature of the colouration is
a dark brown. almo~t black. di;11110nd-~IIaped . rna<;\..-like spot that begins at
the rear of the eye and coveYS Ihe entire eardrum. The rear half of Ihe belly
and thc underside of thl' Jeg~ are lIsU311y ydlo\',.
Male ~: 60-75 mill.
r-emales: 55-66 111m.
Tadpole : Unknown.
Habits and hahitat: Thi~ species has bl'cn found along clay and gravel
han\.., of small stream, in primary ,tIld -'l'kl'lively logged rain forests al low
e]e\atiollS. It also occurs in heath fon:-.ts and in peat s\\amps near the coast.
Presumably il breeds in such plac •.:s as some of the females caught were
Fi.l,:. 69. Limnollecfes IWf<lm(l( mdOIl llnt!cr).
131
RANIDAE : MERISTOGENYS
gra\'id . Adults pre) nn a variety of insect" and in\'eI1ebral~s of motkratc to

13rge ~il(:. One ~lOlllHCh conwlm:d a centipede 75 mill long.
e'lll: L"n\"nown. but ,inee male, lack \'oeal ~acs the ~pe\.·ie, may lack an
ad\crti~emenl call.
Distribution: Known from Bl1Inei. Kalinwnwll. Subah and SafUIIHk. 11
ha" al!>o hC~1l found in Penin!>ular i\'1alaysia ami Singapore bland.
LimllOli ectes rhacodu s (Inger. Bocadi & Taufik)

WrinkleJ Frog
Description : A smull. Siock) frog. \\ ith shon kg~. All the IOC~. cxcept
(or rhe fouJ1h (the IOlJg ... ~(1. are fully "cbbed to the bas ...." of the S\\OL!CIl lips.
The CardnJ!11 is partially hidr.h:n hy the skin . Th~ hack ha~ a long. interrupted
fold Ilf skin at each side and a ~crics of wrinkle .. or fold" running acros~ the
back between the 1\\0 long fold~ . The sides h:'1\c short ridges >lnd round
mbereks. The colour in presenatin: (\\c have lle\Cr <,ccn this speeic~ 31i\'e)
is brown. Jar\"er on top oi" tlw snout. betwe~n the shoulder... and onth ... sid..::s.
The chest ami helly are whitl~ or creatlH.:oloured. with dark hrown spOiling
on the chin and lhroat.
Males : Unkno\\ n.
Females: 22- 2-l 111m.
Hahits and habitat : Very lillie i~ known about this specie~. \\ hich has
been ~;.;..::n only two or three time~ on the floor of 10\\ land forest. One female
had a J"c\\ large (reiativc 10 the size or the frog) egg~. indicating brceding
heha\iour different from the usual type in frogs. perhaps ~imilar to that of the
Smooth Guardian hog (p. 130).
C all : Unknown.
Distribution : KnO\vn only J"rom central and suutheastern Kalimantan.
Genus MeristogenJs
Torrent Fmgs
hi~i., a uniquely Bomeall group of ffllg~: no species ~r Ihi~ genu~ h:l~
T heen found any\\ here e1<>...:. As a group. auulh of MCflsrogeIlY.\ are not
conspicuou~ly different from many spe('i~~ or Rmw. In facl. for man)
yea.r~ lhe) were tn:nted as mell1h<-'r~ of that gcnu~. Howe\er. lh~~ ha,'e
remarkahly distincli\c tadpole,>. prm iding the main reason f(lf regarding
132
RANlDAI::: ~\'lER!STOGENYS
them as a ~eparate group. These frogs arc inhabitants of primar} or secondary

roreq~ but only in hilly or mountainou~ country \\ itb clear. rocky streams
\\hl're there arc ~uitable breeding ~ite~ . Juvcniles and aduhs often wander
through the forest far from the streams \\ here. adults breed and tadpoles
develop. Calling male~ perch on rnels or low \egctation on ",ream hanb.
and <llthough man) males may bc l'!.llling on a given night. they do not clump
together. Eggs are laid near rapids and the hatching tadpl1les quid..!y mo\·e
into the win water where they complet!.: their devclopmelll. Aduhs feed on
medium-:-i:Led to hwgc insects and ol.hel· ilwe.rt"bratc'\. indm\ing sUI.:h
noxious anim.:lls as centipedes anu ~corpiolls. Tadpoles feed on a thin film of
algae thm gn1\\s on rocks in ~\\ift water or clear stream,. relying on he:l\'Y
beaks to \crape off the <llgae while clinging to the rm:k.s by il1cam of an
abdominal sucker (~ce below).
Description- adults: Merisrogenys arc ,mall (males of some speck~) 10
large (feJl1ak~ of some sreci~'s ) frogs. with moderately ~lender hodie~ and
very long. ~lelllJer legs. Tile ~nou t in most ~pecies i~ narrow. but rounded. The
eye is large and the eardrum is \ i,ible. The tir~ of finger, and lOes arc
expandeu into slightl) pt1inted pad~ that are onl) abOlIl half a~ \\ide as the
earurum. The to!.:, me fuJly wehbed except for the pads. The skin is u,ually
lind:. pebbled or ~ml)O\h ,,\ ith a \ow. naITO\\ ridge or fold of skin on each :>ide
of the hack. In mo~t ~peci..:s the back and top of head art' medium to dark
brown. the sides are lighter. and tile under~ide j, \\hite. sometime~ with a
yellO\\ or lime-coloured lint. The Cyl' in ~ome species is golden green in the
upper half and golden brl1wn in the lower. or \\ ith a narro\\ strip of dark red
separating the two.
Adult maJc~ have a small. lhin-skinned poueh at each ~ide of the throat
near the angle. of the jaw ~ . When they call. the vocal sal'~ puff out into these
pouches. In most specic~ the eardrum is eonspicuou~ly larger in males than
in fell1ak~.
Male~: .14--68 mm.
Fem;:..les: 57 ~93 lYI.ffi.
The calls that are knOll n are mostl~ high-pitched. ~ingle squeak, or
chirp~.
T:tdpole: The tadpoh> uf this genu, are speciali/ed for a lii'lo in strong
currents, \\ ilh a hea\) body that is hroadl) rounded :1I the \nout and !lat
beIO\\. The large mouth is on the undl'r~ide of the ~nolll and is [o!1O\\ed h} a
large ahdominal "ul"ker that cO\cr, the abdomen . The mouth ha~ heavy black.
ribbed beaks. The upper beak h i'vl-~haped. \\ lIh a narro\\ gap di\·iding it into
two equal part ~ . Tnt.' !O\\ er beaJ.,. i, V- ~haped and mayor may not bc di\ ided
I{Al\!IDAE: MERISTOGENYS
in till' eemre. The tail i~ ahout one and one-half tiInes the length of the bod~.
:lIld ha~ a \'er~ heal~ mu~ck bUl km Ilns. The~c tadpole~ :Il~n hase small
poison gland~ in duqer~ un the boo) ano the rail lin~. hut the anangement
diffef~ among ~peci~'''.
The tmlpole of Iluill clIrirYIII!JWIl/I/J ha~ the same gcneral form a'>
MCl'i.I{(/\;l'IIYs 13dpoles ano a large ~ud.er. I:hll the l110Ulh of lluia tadpole~
differ in the number of fOII~ of fine leeth and in the \Iructure of Ihe heak.
dewib not t':l~ily li .. ihlt' to ally,lIle without a microscope.
In mo .. ! grollP~ uf frogs. the tadpolt's of rdated specie~ an.: more difficult
10 ~eparate than the adult ... BUI in M('/';\Wgl'I1YS the r('\'er~e i~ true. Adu lts of
.-\fcl'i.l/(Jgl'IIYs dn not differ greatly in rUml or colour. For this rea::.on we are
pre::,cll!ing desl'Tiption~ of unly fOllr ~peci~'''. \dllCh reprc\cTlt the major types
of \'ariation among adults in the genus. Our notes on adults of M.
pll(/colII(,/'/I\ Ilould apply with ler) little change to adults of M.
WHomI'll/Oil/US. 1\1. j")'/)()(f. M. /lIatrop/lI/wIIlIllS and /1,1. /lOcciflls . None of
these species i<, disuibul..:d throughout Borneo. but One or ,e\eral .. pecies
occur in all hill~ and mountainous fore~lt'd pans of BOllKo.
Mer;!ilogeIlYs killllhaillemis ( Inger)

Montane Tom:nl Frog (Fig. 70)
Descripti()n : A mcdium-sized 10 large frog \1 ilh long legs. The triangular

head is about as wide as the hod) . All toes are fully \\-ehhed. The ba(:\... and
top of the hcad are reddi~h brown mix..:d with oli\e grl'en. or mo~tly olive
green \\ilh bnm n spQ!s. The ~ides of th..: head and body are bright oll\1C'
gr..:cn. becoming yellow IOward~ the bell). There is a dar\... ~Irea k funning:
bell\CeTl the eye and the nmlri\' The limb~ are brown with dar\... crossbar!>.
The underside of lhe head and hody are sillery or yellol\i<,h white.
sometimes with brol\1) "pots or mar\"ing:~ on the throat.
\'t a!e~: 58-oH mill.
Female~: 75-')3 mm.
Tadpule: This i, a I;' pical MerislOgcl1.n tadpole. \Iilh a large abdomina!
-"ucker and a !1lu~eular tail. The lower beak i~ an uninterrupted V-shaped
stru..:ture. There an: no poison glands in the t:lil !Ins. The buJ~ IS rnostl~
broIl n or hlack on th..: upper ~urfacc~. Total lengtl] r..:ac-hes 55--60 !1l111.
Habits and habilat : The<:e frog .. hIe along roc\...~ hanks of "Ireallb pI'
~ubrnontanc forests from 050 to 2000 llll'tres abme ".:a lewl. The tadpok~
cling to nx:J...s 111 S\I if! 1101\ ing curren1'-.
13..+
RANlDAE: MERlSTOCENYS
Fig. 70. Men.I'{(IxeIlH /':'illlliJ(llllellsis jlngt'r).
Call : UnkllO\vn.
Distribution : Known only from the mOllntains of wc~tern Sabah.
northern Sarav,ak and the nonheastem comer of Kalimantan .
MerislogclIYS orpllllocllcmis (M msui)

Northern Torrell! Frog (Fig. 71)
Description : A llll.'dium-sized frog . with long slender legs. Thl.' eye is

large, ilS diameter being almost as long as the ... noUl. All the toes are futty
webbed to the base of the swollcn tips. The cardmm is \isible and much
lurger in males than in females. The back and sides arc dark brown. u~llally
with faint light and d:lrk spOh. The chin. throat and chest are \\ hitI.'. usually
with dark bHm n mouling. The reS! of the undcr .. ide is \\ hile. yellO\\ or
grL'cnish yeIIO\\. The rear of the thigh i~ brO\\ n. \\ ith small or large light .. pots
the same colour as the belly.
,\ Iules: J~----W mill.
Females: 55-70 !I1lll.
115
Ri\NJl):\[ : \ 1ER1STOGENYS
Ta dpo le : A, in olher "pecic .. oflhe ge11ll'>. Ihis ladpok has a large ~w:\...cr
on 11ll' tlatlened underside. Both j<t\\ ~heath., arc black and di\ ilkd. There an:
<;mal l glands. appearing a~ dar!.. "pl1h. on Ihe 10\\ er l;:Iil fin and :\1 Ihe <;iues of
Ihe end or th ... hod~. Thc hody i.. dark brown or bladi .. h \\jth irregular lighler
arl'a~. 1\!a.\imulll {Ow] length about 35 In11l.
Ha hits and hahitat. Thi., .. pecies li\"., in primar~ and oid secundar)
furl'''!'> ha\' ing dear. rocky stream .. \\ith modera\<.: to strong current. Calling
Illales perch OJ1 Jo\\' ~bruh<; overhanging hanks of ~tre::lln or sit on roc\...~ at the
water'~ edge. Although it breed~ throughoulthe year. ih activit), le\'e1 \arie ...
.. 0 that on .,ume nighh i1 i .. extremely llUnll'rou ... but \ ..:r) uncommon on
others. When nol hreeding. Ihi, species wand..:r.. \\ jdely through the fore~t.
The tadpole~ ding to rocb in the '>I\ifiest curren!. Thi s species occur~ from
10\\ ek\ation~ tea. 150!l1 abOVe ~ea lnd ) to 1300 m.
Dist r ihutio n : Thi" ..;pecies i~ common 111 su itable habitats in most area ..
of Brunei <.lnd Sahah and ha .. h ... en found in northeastern Kalimantan .
Fig. 7 1. \Jeri\I(>~"lIn urf'/1II01'1If'1IIi!s ~ i\L.thUi I.
13n
Rt\NIDAE: ;-..tERISTOGE:\,YS
M erislOgellYs plweomerus (Inger & Gritb)

Bro\\'n Torrent Frog tFig. 71)
Description : A medium-sized frog with long. ,lender legs and a

triangular heaJ. The eye i, large. All the loe . . are full~ v.ebbed. except for Ihe
fourth Ithe lunge,I), II hich usuall) )1<IS ont.! joinl frt'e of 1\ ebbing. The upfX'r
.. urfaees art.! medium 10 dar" bro\ln, often \\ith :omal1 d;lrker ~pots on the
b;lek. Thl;'re is u~u'llI) a naTTo\\ black. border On the side ju~t ol'lO\\ the bat'k.
The upper surl"at'es of the limo . . h'1\'e dark. crosshar~. The undersides arc
II hitI;'. but in ~Dme frog~ the rear of the hl'll~ and thl' under,ide of the legs an:
lime-yel1{)\\ ofleIllUn-)ellml. r-.lale~ halt' dis l im:tl~ larger t'.lrdrum:o than the
femak,.
~1:lk,: 34~~ mill.
Females: 57-71 mill.
Tadpole : A t) pi,:al Mi"l"il·togeIlY,\ tadpole II ilh a large 'lbdorninal sud.er
on the tlalll;'ned undenide. Bulh heal>.,> hal·e a narrol\ gap in the centre. There
llre no poison gl:md, on the tail lin'. The<;c tadpok' are unique \\ithin the
genu, in hal1O),' a )c11O\lish gn.:en bod) marked ,lith hl:lck. ~pots. Total
kngth .. reach 30- 35 m111.
Fig. 72 . .\Ial'/oilrm I phI/roil/aI/I (Inger &. Grili~ ).
137
II"bil, and hallit,n 111" IWj: " ,'olIlJ1l<>n LD 1>111) I"" l~n.J fOR"" (,1I~r
f'Tun... '1 [>f ,.I.J ..:c.\Il.Jal), A.Jult,.tn· "'nK""ll," f ...... oo f.U" In'1I1 _t .... am •. tllll
u,u;lIl) the} 'IJ} d",,- to Itl.: t>~nh tit n.wIU1l1-,,~~d. d~.lr. n", ~} ,t,., ... Ill,
\Inll'" n~", nighl ~ k" c"Uin~ m"k, ... ,{, .....·Ju.·,.,d "I,m~ Ihe ban l . will> ""
0o.'.'J",,~) ... 1 ni!,Ju in "hll'h lilt) ,II~ I'rc,~m III I"r!,~ numbel'> \I no lillie do
"Jlhn!' IIwl," fonn dell .... ~"'ul" TaJ[><'k. dm!,!<> h<lullkl'> .lIId bc:dwo:L III
'''I''.!' Tadro,k- l", ...... ,,,'n ... lI) f01'1n LaJie <:1",11'1'0 "" ,1"''''111 t>c>uldcl'> III
,h. . I1,,,, "a'~'.il ni!'tll
Call \ hll'h-rltdlo,.·d "'''e.....
Ui~Lril"'liun J... ",-",n Ir\lm J... JlimJnl.m JIhl "l\It~m Jnd "~'Iem
SJT"""'I.
.\li'ri~lU/fj"'.\·S ...·IIilt'lli'ml i ( BOlt 1cllJ;~r J

\\ hi,che,J..r, T,IITem F,,-\~ Ih~ 7.'1
l)oo'>(' rilliion ..... ",.:d,um· ,,,,'II to I"'lie 'pt,;I<' "lIh ' .. '" 'tin!' hind hm!>,
\11 hll" ar.. full) " .. I>...,.J Th.: 1>.... 1. .ul<lIOp of I"" Itt ... d ... n' d ... rl l:>ro\\n
Tl'k'n: " U_UJIJ~ 3 duk 'JrC'al. ,'II IJr 'Hk Ilr 11K' Il.·Jd Irl'lIl 11K' r.\~ II> the
n"'t"I, nIt ,ide, are h~hlcr Ih,," Ihe b ..... ~ JIId ",,,.. II} '1"'''''11 "nh darl
hr"run The uPPO:' ,urf,,,-.·, nt lh.· leI!' ar< dJ,L h'''''" "uh n.!IT"" h!,!h,.·,
"'I1",b.lI'> The ,h.'" Jnd bell) ar.. "h, ,e and Ihe Ihru~( "",'III .. d "lIh hn>'o' n
The 1,'1' h,11 01 Ihe •. )" " t>nj:hl ) elk", "h green 1ne Nrdrum of !]1,,1e, " 1>1.'1
"'-""'<.'Jt>I) I~er ,hJn Ihal of kmak'
_\l Jk' .~ mm.
h·m... k_ ~"'_YI mm
T~dl)Olc -\ I~ 4t>Jo~IlIfl~1 , .... let' COH"T'o IItt uml.: .... ,,Jt III' m..- rod>_ ~
I,"' .. r bea~ I' \'-,Mpt',1 JrId Ulld",ded TaJrode, hJ'" '''1.111 P"""'" gland,
.h.l( JjlllCJr J' '111.1.11 hlJ"l 'pi'll' III lhe I"",·, lin "h..·", II n......·I< (Itt IJII
111" .....·1 .. TIII< I' Ih~ Ia'ge'l 1~.Jp"k 1111"" genu,. "nil IUlul I<'nglll' ",:lCh",!,!
(,... _77 111111
II l. bib and hablll.1 fhl' '1""'le~ I,,~, i" primal) ... 00 old ",,,nnJal}
h>re,1 In nl<IIlIl!JIAoU, ~,,,,m,,, Ill'''''' ~~O "'eu\', ,\dult, hJ'" bttn 1,IIlnd
"nil on (ho: t> ... nL- of ,(,.. am. """", ,"") per,h un Idl}!c n ••:h alld l<l"
,,·t:<'IJU"" Toldpl,k, h,~ III ",.,...1\1,
C~II lnkOO\\n ,
Uhtrihutiun " 1IO"n lrom m".mlain~\u, I'm, "I "e"em SaNh,
I!,>tlhcrn S.,rJ" aL ... nd (he n"nhe~'lcm ,(lnJer 1,1 .... Jlmt ... nIJn
I,\lot
RANIOAF.: OCCIOOZYGA
GCIlUS Oaido:yga
Se.:p and Puddk Frog"
hew ,111.111 Img> li,-( on lh,· 1100r of 1o" I~nd tore,\ U~lIan) In ",epage
T urCJ" <m~1l PI>oj<. nr mur'h~ J!ffiund Bmh 'peele, ~n: "dl

,JIHoutlaJ!c·d. O. ImllleIlSIS!O "~I [XIldl", ot "'ll ;md gr~\d and 0
"kl';r 10 mudd} ,,:ner. Ahh(lUgh the) h~, c -alU!. mu,,-,ubr kg-. (hc) u<u:Jll~
InJ~e 001) <h(.rt hul", Thc l"dptJlel arc u!1Ihu.ll h.'lh in form I 'cc belo" lund
hah!!>, IoNe:11l of '«"rJping ,It the 1>0111>111 Imld of ,lreJI11' (If pond, and tecll",~
on de"3} 1IlJ! plam fr.. gment, anll Jig ..". tIll' Iddpuk' of O<'<'id"~yg" "-,,t on
1IIl' bon"," Jnd p.....} un '11).111 m,,'ch or (heir bl'\ae, "hi<'h Ihe} pmhal>l}
cartur,' II) ma).:'"J! 'k'f1 lung~,
RANIDAE : OCC IDOZYGA
Descriptioll- adults: Small. ~Iocky frog~. with ~horl hind limhs. Skin
hide~ the eardrum. The ~nout i~ Ilattened. making the eyes ~eem more
prominent. The lips of the loes are .. lightly expanded into ~mall. rounded
pads.
Tadpole : The bodies and taih are vel") .. lender. \\ ilh ,"ery 10\\ tail fins.
The mouth i~ al the end of the snoul. instead or just uncia the tip as in most
tadpoles. The lip .. lack the rows of fine hlack teeth that characterize the lip~
in other tadpole~ or this famil).
OccidozYKa ha/lIefisis ( Boulenger )

Seep Frog (Fig. 7-n
Dc ~cription : A .. mall. ~1Oc\.,~ frog with :J moderately poi!lted ~nollt and

.. h(lrl. stoc"~ hind limb ... The toe~ arc about thrl'e - quat1er~ \vehhed. The .. "in
or Ihe bac" i .... moOlh or faintly \\Tinkled . llsuall~ \\ ith a short ridge at Ih..:
•
Fig. 7-l. O('('/do::Y.~(1 ha/ut'lIJis I Houlengcr).
RAN IDAE: OCODOZYGA
napc behind the eye. The middle ofth!.' bad.. has a weak U-~hapt'd ridge oJX'tl
(0 lhe rC;Jr. Colouralion i~ variable. rn mo~t indi\'iduals the top of lhe head
and from of lhe hat·k afC tan or clay-coluured and the rear of the back dark
brown or purpli<.h black. The upper surfaces of rhe limbs are tan wilh dark
cro<.~bars. The under ... ide i~ cream. heavily ~pollcd \\ ilh brown.
Males: 15- 25 mrTl.
Femak,,: 25-35 mm.
Tadpole : Tail more than 1\\ ice the body leng(h. Culour pale tan with a
black streak from the lOlAe[ from comer of the eye 10 the snout and a dark
ver(ical bar belO\\ lhe eye. The tail tJ1u"cle and narrow upper tIn h;J\"e small
d,lrk ~pO!s. Total Jength reaches 26 mtn. See abo under genu~ (ab(we).
Habits and hahilat: This smalJ fores t frog is aimo.',t al\l ays found where
water seep ... frOm a .. Iupe into a small nalli~h area. Its blotched colour pallern
makes for remarkable camouflage in t he~e pat(:hes of reddish clay and Um or
hrown pebbles. Unless it hop~. it i~ almo-'>t i!l\isibl,'. The aduh-'> feed on ~lIlall
insects and other invertebratc~. su(:h a~ ... mall ~pidcrs. The !adpolc~ live in the
same seepage ,Heas and are often in only a thin film of water, leaving their
bodies mmtl) out of waler.
Call : a scries of IO\\-pitched. raspy notes.
Distributiun : KnO\\ n fmm Kalimantan. Sabah und Sara\\ ak. II probably
aha otTUP, in HfLlnei.
Occidozyga laevis (Gunther)

Yellow-bellied Puddk Frog (Fig. 75)
Oescripti()n : A s111al1. squat. stod) frog with short. fat hind limb..,;. The
toes are fully \\ebbed and have round tip'>. Thi~ spc(:ies is \·ery similar in
appl:,mmce to young Lilllllolledes kill/iii except that in o. 100'l'is the distance
bel\\een lht' eyes is about the \ume or narrower than lhe width of the eyelid.
Also. tIl{: inside of the tip of the lower jaw is unlike Ihat of L. kuMii in having
olllya ,ingle (ooth -like projet·(ion insH'ad of a pair. The ~kin of thc back and
upper )urfa(:e~ of the Jimb, ha~ a uniformly cmrugalcd appe:.mmcc wilh
oCTasional rounded bumps. The frogs are uniform dark grq-brown. though
,I fe\\ indi\·iduah /ta\c a \\ ide light qripe down the middle of the back. The
tlnder~idc of the IlL'ad i~ often sped,Jed dark gre). and then.' IS uSllall) a
lemon )L'lIo\\ tinge over Ihe belly and underside of the thighs.
i\-lall's: ~1-31 mm.
Fl.'lllaJe~: J5~8 min.
1-11
RA:-JIDAE : OCCIDOZYGA
Fi g. 75. Occido,-ygo flU-d,1 (uiimh<:r).
Tadpole : 1\ small. \.:r) slender tadpok with a laillonger than 1\\ icc the
hody length. The body and the margin~ of the fin~ ha\c dark spob. Total
length reach ... ~ ~5 mm. Se.: aho under gt'llU~ (ahm'e).
Habits and hahitat : Thi~ forest speci~'~ prefers marshy areas . muddy
pudlill's ~lIld \er~ ~l1lall stream~, and apparenll~ docs not liw In group~. It i~
abo round in and around rhino and pig waIIO\\~ and eyen ill \\ater-filled
elephant footprinb. Frcquemly. adult .. "it or float almost submerged in water.
In thi~ po~ition the ,,:olouration of the adult corresponJ~ close!) to the colour
of turbid \\ater. They fc(.'J on insects and will even ~'at small fn:sh\\ater
pra\\ 11<;. f\ lo\[ ladpole~ ha\'C hCl"l1 found in "hallo\\" mar~he<;. This ~pe<:ic, has
becn found only at low eie\'a(inn<;.
Call : Unlno\\Il.
Dis trihution : KI](I\\Il from Kalimantan. Sah1h and Sara\\aL Although
O. {acl'is h,,~ been reportcd from Penil1~ular .\'I :lia)~ia, Sumatra. Ja\;.t and the
Philippin.: hlands. it i, not certain that all of these frog, are membcr~ of the
same ~pe<:i.:~.
1-+~
RAN IDAE : RAKA
Genus Raila
he Homl.!.l11 'pe(:ie~ of this gl.!nu~ ha\e vel) dhl.'r~c habits and form
T TIll'~ range in ~ize from 'mall to large. Some li\e along the banks of
medium·'17ed 10 large ri\er,. ~OlllC in !>\\amp". other!> li\e on fore!>t
Ilour!>. while sllll o thc r~ Ji\'e in rice tidd!>. Some remain at ground lC\d and
other~ c1imh into ~hrub~ and trcc,. The tadpoles of 'ome 'peci('~ de\elop III
"mall to large ~trearns. other, de\elop in j<'olatcd rain pool" on the floor of
forl."b. and ,till ot her .. tlc\eJup in fl00deu rice fields and d i! t"he~. H ecall~c of
thi~ great di\er,il~. \\1.' do not prc<;l.!lII gl'neralized dc<,criptioll', 3" we have for
other genera.
Raila baramica Boettger

Rrown Mar~h Frog (r:ig. 76)
Description : A ~mall t o mediulll·, ilo.:d frog \\jth a rather ,t ide head and
:1Iar2e pronllllCn! ~')e. The eardrum i) \i~jbJe. 11K' toe!> are Ie" than half-
\\ebbed. 11K' lingt'!"!> are long and the tip~ arc .. light I) enlarged. The !>kin 01
Fig. 76. R(IIIIl ban/mini RO<lIgcr.
1-13
RANIDAE: RANA
the back and ~ideS has seatt.:n:d. small. rounded bU!1lp~, This ~pecies i~ dmk
brO\m abO\e \\ith indistint·t naiTOv.-. lighter areas. Thc . . id\?" . . are yello\\'i~h.
marked \\ ith irregular dark brO\\ n ~p()h. Th(" lips are marl-..~·d \\ ith black bar~
"cparat("d by ~mall \\ hite ~p01s. The 100\cr .. urfaces are whili .. h. \\ith irTcgul::n
brO\\ n or hlack rnarh..ing~ .
l\ lales: 38---l-6 !lUll
Females: -.++-6 7 mm.
Raila barmnim i~ \("r) similar 10 R. g!{//ulu!o\"u. but is smaller: the male..,
ur R. g/wulli/osa an: a .. large or larger than fem<lk~ of R. hUl"Umica. Also. the
\\ <lfh or ltlrnp~ of ~h..in on thc ,id("s of R. r:;lalldlllo5(1 an: much larger than
those of R. hummic{/. A third ~pccic':.. R. /alerillltlCldara. ha .. reeentl) been
~ho\\'n 10 h.: di .. tlnet from. though sup~'rticially similar to. H. bommiCli and
R. glulldllfo5a. R(IIw /merinwculllla i~ ~lllalkr (maks to 39 mm. femalc~ to
-.+7 n1lll) than the other 1\\0 and. be<;ide~. ha~ a light line on the upper lip
l'ontinlIOU~ from belo\\ the eye to belo\\" the eardrum. Thc upper lip in R.
barmllim and R. glallt/II/om is di .. tinctly barred.
Tadpole : Tadpoles of thi~ ~peci~'~ h,1\ (' not yet been found in BOlllCO.
Habits and hahitat : Rww bamlllica ha .. hel'n found in Borneo mo~tl)
in ~\\"atllp fore .. t" n~'ar the coa<;t. Adult:;. live 011 the rorc~t !lOOT. but :llso climb
up into 10\\ \·egel<ltion. Like the related ~pecie~. R. gllll/tilllow. male.\. Cill!
~ingl).
Call: A repetitive 10\\ t·hirp.
Distrihution : LO\\ land coa"tal rorcst~ of Bnlllei. Kalimantan. Sabah and
Sanm aL
Rana chalcol/ota (Schlegel)

White-Lipped Frog (Fig. 77)
lJescriptiun : A ~Jllall to Illedilltn-sized frog with a Ji~tinctly pointed

head and clearly \'i~ibk eardrum. The legs are moderate!) long. and the toes
arc rully webbed. except perhap" till' longest (tounlt). The tip .. of all toc~ and
linger .. are distinctly exp'lIlded. running rounded disc~. The .. I-..ill or the hack
is linel) pehhkd. \\ lth an indi~tinct ridge or fold along each ~ide. The hack
is a rich gr..:en by da) and bro\\ n at night. Thl' . . idc" of the head and bod~ arc
LJ<;ually bfll\\l1 or oli\c. The upper lip i~ pearl) writc. at leaq from beJO\\ th..:
C) e to the rear of the mouth. Thl' undef!,ide of the head i .. monlcd hro\\ n. with
morc . . catl<'"red bJ"lm ni .. h "poh dO\\ n a \\'hitish chest and hl'll) The under"ide
01 the kg" i~ reddi~h in ~\lnl\? populatiotl~. but not all.
RANID /\E : R A..'\JA
t\lale ~: .1J--4-l mm.

Fem:l1l' ... : 46-59 mm.
Tadpole: The 1;ldpole of thi ... "pel'ie~ i~ onl.' of the fe\\ con !>p i cuou~ly
marked I:ldp()I~'!> in Borneo. Thl.' body i<; pall.' ~ ellow or <;t r,m -coloured. "Jth
black har~ ur ~pOIS helm\. in front. and behind the C)l'. On the undcr~ide of
the 0\;11. !>lightl) fl:II1~'neu bod}.lhcn: are man) "maU "hi te g.land!> arranged
in 1\\0 long ratl'he~. one on cach !>iue of the hell).
Hahil.s and habilat : Th i...... pecies \\anders freel) through pnmar) and
di<;(lJrhL'd rore"l~ and m:l} e\en occur in \\ouded gankm.. The u <;ual perching
\ill.' I" in !>hrubs and ... mall tree". \ laIc ... form large calli ng group ... al frequent
inlenah al <;i,k pool!> of medium-sized <;Ircam!> and al Ihe edge .. of pond ....
Adlilt'i (ecd on man) kind.., of in ...eCls and "'pider!>.
TU'!JXI1c . . live in quiet "ater. Illoqly in shallO\\ ... ide pool.., of ... tre~l!lh
\\ here Ihe~ gruh 111 the debri ~ of dead Jea\'es that line the hOl1om.
Colli : A <;1:tct':Ito se rie~ of didJng notes.
Distrihution : COll11llon 111 all parts o f 100\land Borneo. Also fount! from
...outhern Th<liJ<lnd 10 J:1\ a.
Fig. 77. Rmltlilltl/n>llo/ll (Schlegt:il.
145
RA:'\LOAE . RASA
RfllIlI ery tll raefl (Schlegel)

Gr~·I.'n PadJ) r-m~ rhg. 78)
Dl'~c ripli"n : A "mall to m..:<.iiullI-"ized frog \\ JIll long. Tllu"cuiar hin<.i

limh;; and a lung taper~'d "nOlll. The IOC<; are abuut half \\ebbed. with the
long:(' .. t toe (th..: fourth) fn:\,.' of webbing n~';lr the tip. The tips of th..: toe .. and
finger .. are expanded <;ligllll) and rounuc.'d. The eardrum 1" con<;pictlou", The
~kill i~ 'llloolh. thuugh there 1:-0 a \0\\. \\ide ridge on e'Kh ~ide of the bad.. ,
This ~pel'il.'~ i<; bright 10 dark green on the top or the head. liO\\ n the back.
and along the "Ides. Thl'rl.' i<; a \\ide )..:l1o\\i<;h to \\hitl<;h ~tripe from hehind
the t.'~c: back tu the nlmp un each ,ide. nIl! upper lip i .. \\hite alld joill~ a
\\ rut..: "!Tipe running mer the: <.Ln1lpit and along the 100\er part of the ~idc. Tht.'
upper :-ourfaces or Ihl.' limbs :In~ hghl bro\\ n \\ ith fine darl- markings. The
under~idl' I" pear!~ \\hlle. occ;l.'iionall~ with "lighl dark monling I(march the
outer eJgt' ...
;" la!es: .l2--15 mill.
female .. : ~8 - 75 111m.
Tadpole: Ih..: bod) i~ 0\ OIl ami the tail rath..:r deep and l<.Lpcring near Ihl.'
end 10 a rlnffO\\ lip. The hod~ and lail are green or brO\\ n with darl- "peekling.
Hahit s Hnd habilnl: Thi~ b a j'rog of dl~1UrbcJ fre~hwater hahna!<; ~lIch
n~ irrigation ditche~ ;lIlli. e~~ciall~. tllloded rice field ... It i~ extremel) \\a~
146
RANIDAE : RANA
and lhf'1icuh to approach e'en aI nighL ;"I:III:'~ do nOl form calling group,. bm
111a~ be localized within a ,mall an.:a. The) usually perch on grass or rl:'ed!..
\~ hill' females are more onen ~ct'n on pond hanJ...;;:. R(f/w l'ITlh/"l/('(/ cab ~mall
terrestrial in\l'rtcbr:lle~. ~uch a~ small milliped.e~. crickets and ants. nle
w.dro!e~ li\ e in ~tanding waler.
Call : A squeaky \~arhlc.
Distrihution : Brunei. Kalimalllan. Sahah and Sarawak. Thi ':> ':>pecies is
widel) JI,lributed in Southeaq A~ia.
Raila g/(tlu/u/oso Bou lenger

ROll gh-"idcd Frog trig. 79)
Description : A medium·,i/ed III large frog \\ ilh a broad head and

prominent eyes. Thc tOl', an: onl) half \\chbed. and IH1\C thid.ened.
tri:mgular pad~ allheir lip~ . The tingers arc long and have pad, atlhe tips like
tho~e of Ihe \oc'>. The eardrum j, large. The ~k.in is covered \\ilh slightly
rni<,ed. round bump~ that are rno'l pwminent along Ihc ,idc~ of the bod) and
thc wp., or the legs. There i, ~I ,hon rid!!e of skin running bad. from thc upper
c~clid and ending ahO\(' the canlnull. The;;:\! frog., :.m: bro\\n (0 dar~ brm\n
\\ilh dark ~pOTS on the hac~ . The !.ide:. are lighter bro\\n and ha\c large dark
Fij:!. 79. Ralll! f!/'IIIJu/o~a Hllukngcr.
147
RANIDAE : RANi\.
bIO!che~. The chin. throat. cheq and belly arc \\ hitish with man} brown
spots. The undl'r~l1rface of tho..' kg, is white tinged \\ ith hrO\\"Il.
i\. lales: 5S- 93 111m.
Femal es: fl5 - S-.f nUll .
Tadpol e: Tadpoles ha\c nol }et been found in Borneo. Those from \\bt
\ lalaysia h'l\c mal hodies and a tail that i, twice the length urlhe hody and
pointed at the tip. The body and wil ar~ reddi,h hrown and spotted \\ith darJ...
bro\\l1 . t-.lature tadpole~ an: ahout 55 mm long .
Habi ts alld habitat : In Romc'o this species lives mainly in swampy
forest and is morl' common near the <:oa<;1. We have found it as high as 700
metres abo\'e ~ea k\cl. Little is known of ib breeding behaviour but males
call sing l~. Adults eat in~ecl\ and other ill\'eI1ebrates.
C all : A loud resonalll sequence of " \VAHK '!
Distribution : Recorded from Brum'i, Sahah and Sanl\\ak. It prohahly
occurs 111 Kalimantan . El sewhere reported from Peninsular Malaysia and
Sumatra.
Raila " osh Boulenger

Poisonous Rock Frog (Fig . SO)
Description : A medium -si7.ed (males) to large (fem:lks) frog. with a

moderately slender body and long. strong hind limbs. The tips of the fingers
and tue~ arc expanded into pad!>. The tncs are fully webbed. The ~"in of the
back is jillel) pehhled . A weak fuld uf ~"in i~ llsllally present Oil e'K'h ~id(": .
The top of the head and the back are a ri('h. ue\.'p green. The sides are uSllall)
hrown. and the upper ~urface of the legs light brown \\ ith dark crossbars. The
belly i~ greyish or sih'er} while. Allhongh not 3.nUlHIWall} patterned species.
it i!> OIH: of the mo~t handsome rrog!> in Borneo.
Males: -.f5--68 mm.
Females: 86-100 mm.
Tadpole: The tadpoles of this species have not becn de<;cribed.
Hahits and ha bitat : Adult RWIlI hosii are never ruund more than 3. fe\\
metre~ from the edge of rock~ creds or riwrs. Although more common in
prill1ar~ fore~\. it also lives in paJ1iall) logged forest pro\'ided a ~\\in. ('lear
stre:lm is availabk. The frogs perch on rocJ...~ "lr ~hruhs and small trec~
o\crhanging stream!>, p:.micularly where the current is stronges\. If disturbed
\\ hi1c .;,itting on a rol'k. one of these frogs can ~k.ip a(TO~" the ~urface of ,,·ater
!U another rock. l\' lale~ form loose calling groups.
RANlDAE : RANA
l'ig. HO. RallG hosii Bouknger.
The skin secn. tion~ of this specie .. can kill sm31l animals . such a~ other
rrog~. It is not dangerous for human~. However. if one absent-mindedly rub~
one's eyes after handling a Rana hasii. the result is quite painful. Always
\\ash the hand~ thoroughly after handling this ~pecie~.
Call : A weak chirp.
Distribution : Found throughout Bomeo in appmprime ell\'ironments
from near sea le\'ello about 750 metres. It also occurs in Peninsular Malaysia
and SUllla!ra.
Raila /uctllosa (Peters)

.\tlahogany Frog (Fig. S 1)
Description : A small to medium-sized frog. neither stOCky nor ~lellde r.

with a triangular head and rather short h:g~. The toes are less than h.1lf
webhnl and have rounded tips. The eardrum is distinct and almoq 41<; l:uge as
the e~e . The skin is entirely smooth. The colotlratinn i~ unlike that of an~
other frog in Borneo. The top of thL" head and bad are rich chot'olate or
RANIDAE . RANt\
red(h,h !;Im\\ n \\ llhoUl mor"ing" of ally "ind. \\hile the ..,kk~ of lhe head and
[rull)., art' hJad. ...... !hill hglJl Jmt'. fmlll lIlt' lip of The q}(lUI to tht:' end of the
hod~. \epar;lIe . . 'bt· hro\\n and hInd...... urfaee ..... The fore and hllld Jimh . . ha\e
Jet blac" bat . . ~epar:Jted b~ pale blue ;!r~'a ..... The undet..,ith.~o;, are dar".
j\ l alc~: 5" 59 111m.
Fe111ale~: 5.~ 60 111111.
cnulpol l': A rohu .. l. 1ll0deratd~ klrgl' !adpolt'. Th\.· emir!.' tadpok 10;, dark
gre~ or bro\\ n \\ illt irregular darker ~pL'dlinl:' UH~r tl,..· hod~ and wil. TOTal
length readl(.· . . 6:; mill.
Huhits lInd habitat : V\.'r~ liltk I" "nu\\ n aboul lhi~ rore~t ~pecie". The
unly adult i'\\:'r .. l"en <may from a brt'L'uing .. ilL' \\"a~ hiding under dL'ud lea\t=~
on the for\.' . . 1 t1oor. The . . pccle!> breed.. al nun poof". \\ here ma/("" fo rm c:lllillg
~rour". Each male calb Irol11 a . . mall huk' it ha:o. e\(";I\;lIed aJpcenl In the
pool. \\ hether mating take . . pla("e in the holl.' ami the pair then 11111\ e .. II) Ihl'
pool I)r the male emcrge . . \\ hen a femall' appnlache~ i . . unJ..J10\\ n. Thl'
tadpole . . de\ell1p in pool, \I~ually Je"'pcr than ~() em. hUI . . On1e tadp(lk, ha\ e
beL'n found III ,baIIO\\ rh1110 \\ aIlO\\~.
C~III ' A ..... ngk mutt'd. ri<..ing l1ote.
Distrihutiun; X/JOIIlI ~o J~r from S.lP;lh ancl S;lf<l\\·:u... Ou",idc of
~O1l1('O illX'CU1"' ill Peni!N!lar t\lal::iy~ia.
150
RA\lIDAE: RA:\iA
Rall(lllicubariensis (StoliuKa)
Cril'ket Frog (Fig. 81)
Description : A small 10 medium-si/!.'d frog \\ith a long. narro\\. pointed

head. The eardrun~ i~ \ i~ible. The leg,>; are slender. and the linger.. and loes
extrernd) long with slightl~ ~wollen tip~ . The toe~ arc only .lboUI half
wehhed. and lhe IOl1ge~tto!.' \!xtend~ far beyund the webbing. The skin ufthe
back is finely pehbled. and there i~ a di<>linct thin ridge or fold of ~kin along
each side. The h:t<.:k is brown. \\ ith dark spm.., . The !.'mire upper lip is ~arly
white. The lHldersidl'" i~ dirt~ \\ hite \\ ilh grl'"y motlling.
Malc~: 37---+7 IIUll .
females: -I-7-5~ rnm .
Tadpole : Tadpule~ ha\'e an O\'al hod~ and :I tail about tv, ice the lenglh of
the body. The lower lip has long.. finger-like projection". The hody i<> light
bro\\11 mottkd with darker bro\\Il. <Lnd the tail has a d:lrk n!.'lwork . Total
knglh reaches -1- 7 nun.
Hahits and habitat : A \\ idely di~tributed frog of di~lurbed habitah.
found hOlh along logging roads and .. treets in town~. The har<>h call of the
lSI
RANlDAE : RANA
male\ i~ one of the characteriqic <;oumb of \'vl."t grassy area~ and roadside
ditc he~, Male s are often \'ery abundant. blll do not appear 10 call in groups.
The s pe(:ie~ seems 10 breed year round. The tadpok ... d~Yelop in the shallo ....
water of dilche~ and temporary pond~. Adults wi!! eat most kinds of insect ...
of moderate ... ilc .
Ca ll : Six to len ~harp "ke"" notes in rapid ~equence .
Di strihution : Reponed from Kalimantan . Sabah and SarawaL It almmt
~ure!y occurs in Brunei. The ~pecie ... is found from Thailand to Hali.
Raila picturlltll Boulenger

Spotted Stream Frog (Fig. 83 )
Raila signata (Gunther)

Striped Stream Frog (Fig. 8-1-)
We group the~.: ~pecies becau..,.: they are ~o similar in habit... and form.
ex<:epl ror details of {:o}ouration (see Fig. XJ and 8-1-) .
Desc ription : Small 10 medium-sized rrogs with a triangular head ubout
same width as body. The cardmm i~ \i~ib1c. Bmh ftnger~ and toes ' havc
"lightly ~\\ullen tips and the toes are slightly more than half wehhed . The
skin of the back is \'ery tlnely pehhled and the upper surfaces of the legs are
almost smooth .
Both specie ... aTe generally dark hrown or black with lighter markings.
The underside is dark grey with smull hghter marking~. Both species ha\c
red or orange markings on the upper surfaces of head and hod}'. A common
feature i.. a light orange ur red <;tripe from the tip of the snout to and along
the edge uf the upper eyelid.
In the Striped Stream Frog ( R. siRIIOW). the ~tripe on the !.."dge of the
eyelid is continued to the end of the hody without interruption where the side
meets the back TIlere may be a kw light spots of the ~ame colour in the
,:entre of the bac".
In the Spotted Stre;)m Frog (R. pict lll"{lf(/ ). the stripe rrolll the eyelid may
not appear on the side of the hack. but iJ it i<; pre~ent it is alv.ays interrupted .
The centre area of the bad, i<; alway,- marked .... ith a numb.:r of irregul:!r
orange or red .. pOlS .
Ma1c~: 33-1-7 mnl.
females: -1-9--68 mill.
152
RAN IDAE : RANA
.·i~. ~J . NlIIW p iCllfraUl Boulenger
Fig. 1(.1. Hm/(/ si~lwla {Gunther).
153
RAN[OAE: STAURO[S
Tadpole : The tadpoles of bOlh ~pecje~ have rdal in.'l) ,lender hodies ,md
tail;, .md are almosl black. Small glands that appear a~ tine light dots are
<.catten.:d m·er Ihl.: tail tim and are grouped in .. mall patclk'" under Ihe bod~.
TOlal length, rea~:h ..+ I mm.
Lanae of the 1\\0 .. pecie~ diller in the arrangement and number of gland~
untkr the body. but the~l.' featllre~ cannot b . . ~een withoul Ihe aid of :1 good
mil.."ro~cope.
Habits and hahitat : The .. e frog~ li\(~ along banks of ~!llall to medium-
.. i7ed strcmm in primary or ~lightly di~turhed foresh at elevation .. from near
~ca leyel to ahout 750 Illelre~. R(///{/\i!:llof{/ .. eem~ more common in heath
fun:~ls and ...·oa<;ta[ peat .. \\arnp~. Males call from irregularilie~ in the hanks
of ~lrL"amS or from "mall rools that project !"rum the banks. i\lmo~1 . .·\ . . ry
night a lew .. oJilar~ male~ tan he heard. \\ hi Ie at odu inten·ab a large calling
aggreg,llioll will ["orrn. Taupo1es li\c in shaJl(l\\ ,ide pools or in
accumlilali()ll~ of dead leaves thaI form in the bottom .. of Ihese .. Irearn'.
When Ihe) transform into frogJeb, Ihe) lea\ e the qream and Ii,·e in Ihe foresl
lear liner. Thl') refilm to the <;[ream a .. Ihey approach adult ~i/e.
Call : The call of R. pictl/raw ton~i~t~ of four 10 six grating note,>. 01"
,harpl) decrea:.ing illtemilY. The. call of R. \·igllow ha.. not been n:cordell.
Distrihution : Rww piC!lIml(1 and R. .l"ignUf{/ are Found throughout
Borneo. Both ~pecic~ apparently also o(.:cur in Sumatra and the ~ I alay
Peninsula. although identification-. of ~pecill1el1s from tho<;e area .. are
ul1L'ertain .
Genus Stallrois
he spe...·ie .. of this genu .. are stream~ide frogs . rar . . ly fOllnd more than
T a fe\\ metrl.."~ from dear waler. The) are common in the sen~e Ihat
they are \\idc .. pre.ad in :.trl';l~ of Ihe appropriate emirollment. two of
them (5. luw[Jlllmllflls and S. l1(1{uwr) belo\\ 1000 mdres and one tS.
rubt'rilil1~uil) from about ..+00 10 1800 melres. Th . . y differ from one another
in the kind~ or perches Ihc) use. Sraumi, low/Ja/llwfII.1 ~it~ on rol'k~ ne:o.! to
the .. !ronge~1 (·UITel1tS in rod~ ri\'ers. S. /lulalOr perches Oll 1\\ ig~ of shrubs
m'erhanglllg ~and) or rock) .. [reams. and S. III /erilillguis i.. u.. ually seen
perched on ~Iem~ anu lea\·e:. of "mall herb .. or ferns gro\\ing on rocks ill
midsrream. De~pite their wide di"lrihmion in Romeo. lillk i~ knO\\n of their
ladpoles. The one 1~ pc of tadpole kno\\ll. Ihal of S. I/{/f(/[O/". has been found
[5-1
RANIDAE. STAUROlS
in masse~ of d<'aJ kaves in pool~ within denr qreams. The a!'."ocimion of

tht'se species \\ ilh dear water is '>0 ~trong that allthc hundred\ of specimcns
recorded ha\c come from hill) country: nOlle ha\e come from flat or flood
plain 10rl.'"ls. which imariabl) ha\"("~ turhid, silted stream ...
Description-adults: Small to lllt'dium-sii<,d frogs with long hind
limbs. The tips of the fillgt'r~ and toes are expanded into wide pads ha\ ing a
blum <'nu . The pad, ar<' 0\ al rather than round. All roes arc rully webbed to
the pads. The eardrum i.. visible. TIl<' ~J...in of the back i) coarsely granular or
.. ct \\ilh round aml o\'al swellings. Two species an; mllTow-bodieJ and long-
nosed. \\'hi1<,' the third is squat and pug-no<;ed.
Stallroi!llalopa/lIlatlls (Bouknger)
Rod Skipper (Fig.S) )
Description : Thi . . i" a medium -si/cd species. with a body that i\ neither
qOl"J...~ nor ~lender. The snout is \<'ry .. 11011. rounded. and slopes !lack from it~
upper smfal'e 10 the mouth . The kg~ .ue long alld \Cry heavily muscled. The
Fig. MS. Srmll vis /lI!Ul'a /maIll J IUolllenger l
155
KANID!\E : S TAL' ROI S
t\\O out~r finger<; are ahout h:llf \\L'hhed. The enti re upper .,urr~ux ha"
l'oar"eJ~ ),!1"lIlular ~kill. "hik lhe bell~ I'> "momh. All UpJ)\:T "urface~ arc: blad.
\\ ith ~hon. narrO\\ yelJo\\ i~h or \\ hiti,h marking". The undcr"ide I~ \\ hill" or
p~lk l"fl'am,
f.. lak,: -IO-SO mm.
Fenwlt:, : 60-70 mm.
T;ldpole: Unkno\\ll.
Hahit ... and hahital : Thi~ 'pecie, i~ found only on (:lear. "\ltl. rock~
~ Irl'~ml' "here it p..:rche<; on rol'k". l',pecially lho,c \\I\h a \enkal 'urfacc.
eith~r (Ill the bank. o r in m id-qream. II I.h<;lurbed. it kap" from rod. to rock.
or llla~ 1>'\\,.'11 leap into ~\\ 1ft \,ater and ,kip aero<;s the ~ur ral.:e tn another rock.
Although 1110,t rrt"quentJ~ wen III primar~ fo rl·'1. il can abo lhL' on '1ream<;
11m, Ing lhrou,;;:h loggt"d area~ il ,ollie tree, remain and if the \\~ltl'r 1<; clear
allt! Ihe hmtotll fro.:c of "ilt.
Ca ll : :l ~hort Irill.
Dblrihutiun : KIl U\\1l IrOIll Kalimantan. Sahah and S:lnll\ak.
Stollrn;s nola(or (G iinther)

Blw.:k-"pOIted R(x:k fro,;;: (Fig. X6)
OC"'lTiplion : A small fwg \\ IIh a pointed ~nout. naITO\\ booy. and Inng.
"lender hind limh". The ~kin o f Ihl' h3ck i~ di~titKII) pt:bbled and ~~t "ilh
,mall round. unii"onnly sized ~\\elling" The bell~ is l'mlr,cl) granular. The
upper ,urface~ are oli\ t: green. \\ lth large black "Pot~. Thl' 11l1der~ide is bright
~dlu\\hh green or lemon )ellllw. Th~ upper surface of thl' tlm~e inner loe~
:md 1111' \\ebbing betwe.... n them I~ turquoi...l· bluC'.
~ I ale~: ::!l)-37 111m.
Fenwk,: 44-)) mill.
T.. clllOle · nlC' bod~ i~ 0\<11. .. hit flanened .•lIltl le~, than half till' lail
length. The no,tril, art" JUS! ;lbo\c the end of the ~nouL The eye' are "mall
and 111 YOUll£ tddpok, arc cmereo \\ ith thick ~kin. The l~tI h;I' \ery 10\\ lin'
and 1..:111.1.. lt1 a broad tip, The tadpolt:, aTC eolour1e~~. but the hi:lIie .. appear
pinU,h h(·ci.u<;e the cirl'ulatlllg: hlood can bl..: "l'l'n Ihrough the thin ,kin .
Habib and hahitat : Thi, i, a frog of prilllar~ foreq in hill~ l'Olllllf).
Ii, 11Ig. along "mall rocky ,Ireallh. TaJpole, ha'~ b.... l·n found in acnl1nulation;;;
(It dcad lea\e, in ,Ircam ...
Call : A ,harp l·hirp.
ISo
RANIDAE : STAU R01 S
Dis trihution : Ka li mantan. Sabah. and Sara\\ak. It b ~o wide<;prcad in

Iho<;c qale, thai il mu!>1 abo occur in Brunei. It ha!> abo been found in the
Philippme h land~.
Fi g. Mh. Slmmlll "m(lwr (Gunther).
Staufois tuherifillglli.\· Buu 1e nger

Grecn-spol1ed Roc ~ Frog (Fig. 87)
Dl'scription: A ... mall .... 1f:nder frog \\ith a poinlC'u "'nuul and long. "lender
hind I!!nlh. The ~kin of the b;lc[... i~ finel~ pehhled \\ ilh many round or mal.
unequal-'l/cd l>\\cUing!>. SOllle of the <;\\elling:. ma~ form an Interrupted
ridge from the eye to the ba<;e of the thIgh . The upper surface~ art' a .. lightly
mctalJil' hue of brown \\ 1111 irregular greeni .. h patche<;. There i ~ a greeni<;h
\\hile patch behind the c~c anti a l>llnilarly coloured mouling ;,long the side.
The underside of the h(,';u.I i.. light blue-green. \\ ilh dark ~pob, The che~t and
belly nre hlu; ... h II hite. The IOp~ of the three inn":f toe:. and the upper \urfaee
of the \\ehhing are chalky bhll~h \\ hill'.
i\ ]ak .. : 2.1-27 oml.
female ... : 29-J6 mm.
157
Ri\:'\[l)AE: STAU K0 1S
Tadpolf' : Linkno\\Il.
Habits and habitat · Thi~ i .. a frog o f primary forl." .. t in h1l1: coun tr:
i"nl1l1 about 200 to 1800 llll'tro;'~ ~lbO\c .. ca 11."\1." 1. It I~ Tl10~t cOlllmon aboYe 500
mctrc~. It i~ Ill:\ Cf found :1\\ <I) from ;,mall rocky ~trearn ... One frequentl) .. ee:--
~ma1l1l1ixed group .~ of jll\enik~ and adu1t\ all clinging to one ,mall ~tC11l or
frond gru\\ ing on rod.. in mid- .. trcam.
Call : A thin mewlii(: "queak.
Oi stdbulion : KnO\ul from S;jhall. Sara\\;j/'" and northca .. tern
Kalinlantan.
Fig. 1'1 7. SWllm/1 IIIherilill>:lIil' Rnulengcr.

J'WII ily i{lwcoplwridu('
hi .. I~ ,limos! an exclusi\c1) A~ian family. \vilh only a fc\\ genera
T ]j\lng III Afnea and Madagascar. Almm\ all the specie .. li\(' ahon;
gJOund 10 hu~hc') Oi trees. '\·10S{ of them breed in sLanding water. A
few Bornean splTie~ do not have rrcc-~\\ Imming tadpo!c".
Nyc/i.mllls picflts (Peters)

Cinnamon Frog (Fig:. 88)
Descriptiun : A small frog with a relali\cly long snolll and long hind
limbs. The eardrum i .. \i~ible :md a hit ~lllalll'r than the diamch:~r of the eye .
The lips of the lil1g~'r~ and toes are e"(panded into round rad~ that are smaller
than lhe cardnull. The toes nrc about half-webbed un" the finger') la(:k
\\·ebbing:. The skin of the had.. h~'ad. ,md the upper ')urfaces of the limb') is
rough. \\ ilh many slllJ.ll ~pin~ bumps. All the upper ~urface s and the sides
tend tu be cinnamon hn)\\ 11. though SOllle indil iduals are red or even orange .
SC<Jllered oVer all arc ~m:t11 glossy II hite ~pot ~ that form a broken line from
the edge of the \110UI. along the edge or the upper eyelid. ,md conti mIL' pan
way dO\\ n the ~ide of the back. 1111: upper half of the iris of the e)',,-: i~ abo
white: the 100Ier half is brown. Thi~ i~ ont' of the mo~t distinct I) coloured of
Bomcan frogs: it cannot he mistaken for an) other ~pecies .
Males: 30-33 mm .
Females: 31-3-1. mm.
Tadpole : The body is oval. almo,t as \~ide as long. and slightly rlatt..-:ned.
The tail i<; lC'~" than twice the length of the body and Lapel'S to a roundt~d tip .
The bod~ and taillllllscle are purpli~h bro\I'n and the tail fins slight!) lighter.
TotallC'ngth~ reach -1.-1.-53 mm.
Habits :lnd hahitat : This ~pecie s 1i\(~~ in primal') and old secondar)
for..:~t'< in flat and hill} Il'rrain. and from ncar ~ ea lelel lu !650 metres. It
"~'em s to be abund:ml and \\ide<;prL"ad in Borneo. a~ \~e hal e seell it alJl1 o ~l
e\ el') place II e h:ll e II orked . j \Jltlh hal e been founLl 0 11 le:l\e~ of "hruh~ and
~ma!l tree~ nne 10 three metrc .. abOle ground. bUI presumabJ) reach higher
klels . T\\ o to rour m:l!e;;; Illa) galh..:r around 1\<11ef ~'ontaining tree holes or
159
RHACOPHORlDAE : ~YCTIXALL'S
Fig. 88. \\aiwlils phil" (Peters).
~"l\ dies in log". T~dpole~ dcvelop in ~uch c<l\iries. and once we found
tadpoles in a rolling hollow fruit lhat contained \'.lIer.
Call : Unkno\\ 11 .
Db lrihulion : R~'ulrded from all parI , of Homeo C\ccpt Hrunei. \\ here it
probabl) occurs. Thi, 'f>Ccies abo Ii\\!, in Sumatra anti Pemmular \ lala}"ja.
loll
RHACOPI[ORIDAE: PIIILAUTUS
Genus PhilaltlUS
Bu!)h Frog!)
he-M: ~rn311 frog .. arc among the rno<;( difticul! for llnyone. (:\en
T e.\pen~. 10 identify to spcc iC' ~. }\ fc\\ of the Borne;1lI "'lX'cies ;lre

relati\cl) easy !O ~eparmc. from the others. but mo~t are tro uble<;ome
even with the use of ;1 micro~wpe. Om' distinctive char;I('teristic of these
frogs i.e; thai they lad. :l frec-~wimrni n g tadpole. Fell1alc~ lay a few (Ie')~ than
20) very large c gg ~ and tlK' embryo ue\'e!op'" within the egg into a lin) frogle\
wit h a tail. J u~t before h;!tching the: tail i<; ab~orhcd,jusl a .. in other tad polc~.
j\·lost <;pI.'(-ies li\"e ~calle rc d through primar) forest"', from ncar !>ea k\el
W abo\'!.' 3000 I1lct re ~ 0 11 ~ lount KinOJ.balu. A I night. male", of most species
calJ fWIll le;-n'e, of shrub:. olle \0 three nk'tres nbo\c the ground. often in
small areas \\ithin a forcM where it is undear \\h) the frogs \\ould have
cho~e n that pal1icu lar ')ite to call. A matcl.i pair dC~('ends to the ground whl're
Ihe large cgg~ arc depo~ited under dead lean~~. Eggs o f Phi/limits havc ;11:'0
been (ound in pitt:her planh. bUi it i:. nOI certain \\h ich ~JX.'c i es \\(1:' the
parent. One ~pc(.: i e~. Phi/I11I1I1.1 /I(Hii. calls from vegetation on :.tream b,m\.....,
and appear~ to :.tick it ... l'ggS OlttO le ilve.~ o\"crll<lllging water.
Becau<,e of the diflkultic<, in idcmif) ing mo..,t of thcw spec i c~. we \\ ill
gi\l' gener;llized dt'~n i ption~ th:ll lit group<' of them. Severa l featu res.
ho\\ever. are common to them all: the tip~ of the lingers and t oc~ are
expanded illlo round pads. and the toe:. arc u~lLally about half-webOCd. As
Philmlf/ls lack.., rree-<'\\ imming tadpoles. Ihal part of the descri pt ion i<,
lacking in the following at'Courl1~.
1~lliialitus {lurflllfium in gl!f

Golden-legged Bu<;h Frog ( Fig. 89)
Dcscriptiun : A \ef) sma ll. ~Iod.y frog with a wide- head and moderate I)
long l eg~ . The eardrum is partl) hidden by ~\"'in. The outer fi nger:. ha\e rather
1,\ ide pad.~ :It their lips and a narro\\ web al their ha,c~. The tir~1 loe ha, ver)
lillie \\cbbing. \\herea, the outer toes arc a bit more than half-\\ehhcd. The
upper !>urface, are <,mooth. c.\eept for a l'ur\"cd fold o f <;kin from the e)c to
the annpil. The undcr ... ide of the bod) !, t'oarsel~ pebbled. T he colour j.., pale
,and~ or pinki'h bro\\ n. The bad, u:.uall) has blac k marking:.tilat t'urw from
I fi I
RIl ACOP HOR1DAE : Pll1LAUTUS
Fi,::. WJ. Pllilmlfl/I iumlll/il/l11 Ing<'r.
Fi ~. 911. Phi/mlllll hllll/fIIS Inger. Smcbing 8: Tan

RHACOPI-IORIDAE : PHILAUTUS
the centre to\~;ml, ~at'h groin. The hidden wrface:. of the Ihigh arc golden
yellO\~ or orang~.
l\ l ak~: 24-28 mill.
remale ~: 26- 27 1I11ll.
Hahi t .. and habil .lt : Thi, ,pecic, lh e~ in primary or ,ek~'lh'e l y logged
forc,\ from 750 \0 10-lO metre, abo\t= ~ea level. l\-tak~ eaU from 1ca~('~ of
... hrub ~ and lrees bel~\een 0.5 ;lIul 3 metre, :lhO\(' ground.
Call : ;\'01 de<:cribcd.
Dislrihution: So far known only from \\e~lern Sahah.
Philatlftls blllliftis fnger. Sluebing & Tan

Green BUi;h Frog (Fig. 90)
Oescripliun : A ... mall. <:I()(' k~ fmg \\ ilh a wide head. u large eye. and ;\
\ i.. ihk eardrum. The pad ....\1 the tip~ of the OUh,'r lillgl;'\,~ are larger th;lIl Ihe
o.:an.lrum. The flllgcr, ha\e a narrow weh at their ba~e and u<;uall) a narrO\\
fringe of "ki n. The \tl~'~ ;Ire aboUl Ihrce-founh:. \\ ebbed. The h(.':td and had.
ha\'e smalL 'cal!cl'I,,'d bump:-. and thc untl,,;r~ide i.. coar... eI~ p;.'bbled, The h(';lu
:lIld hal'k arc lighl green wilh hlack ",pi.'l'kJing thai fonn, an itldj~litl c l pallcm.
The top~ of Ihe inner finger<; anuthe \\ebbing of the inner toe<; are )('110\\ or
orange. The belly and Ihe und('r~ide of Ihe inner lingcr~ und IOC<: :Ir~ or:lI1ge
or pale greenish orange.
l\1ale~: 35-l1 tnm .
r:emales: -l+----46 mm.
Hahits a nd habit:.I : Thi" ~pccic .. li\'e~ in ~llhmnntan~ und mnntant=
fore<;t~ hcl\\cen 1350 and 1600 mClre~ abO\'e ~ca 1c\cl. Both llla1c~ and
femal e ... perch o n l e~l\e<; of .. hrllb~ and ~mailirec ... from 0.5 10 -l mClre~ ;.bove
grounu.
Dhl rilmlioll : KIlt,\\ n only from Ihe mOulll:Jin~ of \,,'c<:tCnl S<lbah.
P"i/(llllll ~' er},llmJplltlfCllmlls Smcning & Wong

Red-e~ ~d l1u~h Frog ( Fig. l) I)
IlCSlT iplion : 1\ ,mall ~I~nder frog \\ilh a ... mall broad head. hlunt 'nou\.
Jnd large reddi,h eye .... Th~ fingers ha\(' .. mall. \\eakl) rounded dbc ... at the
lip ... and nalTo\\ \\ ehhing al the base of thL' Ihird and fourth Ijngcr~. The toe ...
163
RHACO PH ORIDAE: I'HILAUT US
are about one-hHlf 10 two-third ... \\cbbed. and their dist·~ ... Iigluly smaller than
those of Ihl.' fingers. The eyelid sh.in is granular \\ ith no bump" or projec:tion,.
\\ hik the s~in of tlie back i<; ... Ellooth. There an: no fringes or Ilaps of sIJn
along the edg~' ''' of the aml ... or legs. The upper surfacc of the bod~ is light
green. and whitbh on the side<;. with :1 faint dark nel\\orl... The bas\! or the
upper anTI. front and inner pam orthe thigh :lrc golden )10'110\\ . The eyes an..'
~cI1O\\ j,h 10 orange. occupied \\jth.:l dark red nc\\\ork.
Female: 16.1mt1l.
Tadpule : I;nl..nO\\n.
Ha hits and hahitat : Thi., 'pecie:-. is \...no\\11 only from upper montane
forest ;1\ an alti tude of ahout 1550 III abO\'e sea Ic\e l. in 10\\ \ ....-gcl::nion along
a ;;.[re;Ull.
COl li : Unl..no\\Il.
Uis tribution: Th is spl.'t"ie<; ha~ been collected onl) once. 011 Guntlng
Muruk t-.l iau. it rugged area nellr Long l'asia. Sahah.
Fig. 91. Phi/mllll' enrhlllplllhl/lmlH Stucbing & Wong.
16-1
RHACOPHORIDAE, PIIi LAUTUS
PhiJallllls has;; (Bou lcnger)

Hose\', Bush Frog (Fig. 92)
Description : A mediullHized frog ""jib a ~harp-edged ~nuuf. The

eardrum i~ \i,ihk The lip!> of thl' fingcr~ are c\p;mded into wide, round
pad~: the pad of the third finger I!> wider than the eardrum. W~bbing un the
finger~ i~ limited to:1 nalTO\\ fringe at Their b:l:.e. Th\,.' lOe" are aJmo~1 full)
\\ebbed. The skin of the ha~'k is lind)' pebbled. Tlll're:.lre 1\\0 weal.. ro\\" of
.')mall hUlllPs fomling an open V bet\\CCIl the ,Imulder, and ,e\eral .. mull
bump.') un Ihc upper L"}e\id. TIwre are no flesh) ,pines althe hcd. The colour
j, ) diem "h 10 dar)" ~alld) hrown. u ... ually \\ ilh a d;1r!.. triangle between (he
c}e~ and a thill d~lr\... X-shapl:d Illark on Ihe b<ll,.'k. Tlw .')ide<; have faint ~trca".')
of yel1o\\, The upper third of the irh is pale gold :!Od the lower POl1ioll light
gre) l~h bJ'O\\ Il.
Male!>: 4()......49 mill .
Female,; 5 1-63 11111l.
Ha hits and h:tbitat : Thi " 'pecie" live!, ill primaT'!- or old secondary
fore,!". frolll nt.'ar "ea k\elto about fiOO melre~ in bOlh nat and hilly terrain .
Mo<,t individuab hah' been found perched on leaves or twig" of shrub!> and
5m3!! tree~ O.5-l metres .lome the ground. ~ l jles call ::.ingly. We haVe found
Fig. 92. I'll/lUI/IUS 1I0.,i; (Houtengt:r!.
165
RHA('OPHORiDAi:.· Pl-li LAllTUS
~rn,lll ,gr(lup" 01 ('~£. .. aLlhering 10 It';!h'~ n\erh:mglll£. qr":,II1l~ Ih>!1 \\l' bdieh'
,Irt' Iho~e (If till' ... pecit.· ...
C;liI : \ ,tan',no lllu'Il';'I1 Iriil of tho;' llr ,I', nOll'"
lJi!> ldhuliUIl. Kllo\\/l Jrom m:m~ pl;Jl'c" III Sara\\a" ;'Inti Sahilh.
PIIi/allfll S ;ligen' Dring

Sharp-,nollted Bu,h !-rug IFig. 931
Ol'!>c ril}tiUlI' ,\ ,mall tl1lllediwll-'IZc'd lrog, \Ijlh a poim..:d ... harp-edged

.. nOli!. Thl' C'ardrum I' \ J... i!)lC', -[ he tip~ of lilt:' finger,> arlO l'xpandeo I!H('I II ide.
fllll!1d Of ll\al rad ... nlt"re i .. a n31To\\ \1 C'b:11 th..: h;1<;e of lilt' fingo;'r, and a thm
fringe ('If .. kin runnin,g OUi the linger... tV Ihc pad~ . The foul1 h loe tthl' hl11ge'l )
i ... h:lll-\\ebbed, hili th..: Limo and litih IOC" are .. lInll,L fllll~ \\chhed. Thl' L\lp
of thl' hend ;md nnd drl' fine!~ ~hbled, Tilt' uppl'r l'yeJid ha, tllO or Ih,w
.. mall I:>Ull1P~,mtl ,('\eral uLher~ .m: ,cattered 011 the back. fher..: are one or
1\10 .. horl. tle"h) 'pme .. :n till" heel. Thc colour '" pale ,and) brml '!. Jighlt.'r
un the ,itk, :lnd II ilh a ,harp dark . . tn:a" on th..:: .. idc nfth.: ~nout b..:tlll't'!I Ih ...
... ~c :lnu Ih..: nn,!ri!. nlere h u"l.Iall~ a unr" triangle bcll1CCIl the l'~(:' ~lJ1d
ri l;!. 9.\. "111111/1/111 III£!I"I'I Drin,!,!
166
RII ACOPH ORIDAf: : PHILJ\UT US
~\'meliml'~ OJ conc;l\l'-<;idl'd dark marl.. in the cemf': of the n;Id.,. Th~ upper
thml of the ifi, i~ pale gold ,md thl' Im\t:f two-thmh bro\\ n.
~1a1l',: ..,6 mm.
felllale~: 47-50 mill.
Hahits and hahitat : Thi~ ~peci('~ hw., been ~(,Cll onJ~ ill r110lllane rNe~h.
frolll 1300 to ISIIO 111~'lf':'> :IbmI.' '>l'<l level. nl\,' frogs p('rch on o;hrub) I - S
mt:tn..'~ above th(' groum.!.
Call : ,\ of Ii\c ~of1 note~.
"eri~~
Di stributiun : Thl' 'p.:de, i, I-..n\)\\ n from lhe 1ll0l1Jl!ailh of \\ ("tem
Sahah and nOnhea"ll'rn Sa]';lwaJ...
PhilUliftiS peters; (Bouk:nger)

Bro\\11 Blhh Frog !Fig.lJ4)
Ut'scripliulI : A ,mall. ,tout frog \\ilh a \\id~ head. The ~noUI j, rounded
III mak~. but III km,lle' il cnd, in a c.Ji<;\IIll:1. ,ome\\hat h;lrdencd cone. The
cardrum i" \j,ibk. The finger') ha\'e \\ iUt:. rounded pad<; al th~ lips ano.
Fig. 9-1. Plrihlll1ll 1 lld(" ,' (Houkngen.
167
RIIACOPII ORIDAE : PHILAUTUS
narrow II ehhin£ at the ba~c. Pad~ at the (ip~ of the toe ... are a bit ... maller than
lho,,~ of the finger~. The loe<: are h:lif (0 thn:e-fouI1h!> \\ebbed. The ~JJn i~
fine!) pebbled on (he upper wrfacf'~ \1 ith "catl~' rcd bump~_ espedall~ on the
eyelid_ on (he top of till..' ~noul. al Ihe !>idl'~ of the b;lck. and upper ~urface of
the limb~. C<:uall) there i~ a <:ofl. ~pinc-like projection::lt the hcd. The wlour
and patlC:nl ::Ife highly variable. Th~' back and tOP of the head may be !"it'll
dnnamon brOIl n \1 ith a black "tripe l'un ing a long the bad. and sidl..·. or dark
greyi"h brOIl II II illl ~lllall black and yellowish ~pol<:. gre) i~ h hroll n on Ihe
he'ld and fore part of Illl' hack and dark brown or black in thl' rear. TIl<: ~ide
of the head ha~ dark bar~ mdialing from tbe eye. The !imb~ have dark
l"roSsbar~. The front and fear race~ of the thigh arc reddi~h.
~ I ale<:: 21-24 mm.
Female,: 29-J~ mill.
Hnhils and hllbitat : Thi~ ~pi:cie<; live" in 'iubmontane :md montane
fore:.b. from iOOO IO.:!OOO metre!> aho\·e ~ea bel. AI night IIduh ... feed and
call from ~hmb~ bc(\\een 05 and ~ l11etre~ ab{)\e the ground. Thq hide
under dead kan:.'~ duri ng the da).
Ca ll : A ... hort high-pitched trill.
FiA. 95. Phi/lIlt/uI IIrjohN!!1 Smith.
""
RHACOPIIORIDAE : POLYPEDATES
Distribution : Reponed from Sarawak, wc ... tern Sah;,jh and the

mOllntainou:. area .. of llorthea ... tern Kalimantan ,
Noll:: With :.light Jllodiflcat i on~, the de:.niption aho\'e could appl~ 10
Philalffll,~ lIlIIoelllf_~ (Sab:lh), I~ IOllgiCl'lIs (S;.bah and Sar:lwak), p. mjuberrd
(Sabah and Sarawak) (Fig. 95), P. refllgii (Sara\\ ak) :lnd P umbl"{/ (Sar:mak),
The la<,t :.pecies i~ ~ligh tl) Jarger than P /wf'rsi and thl' second one ... Iightly
smaller. The females of mo~t of Ihese spccic~ do nOI ha\'c :l conc on Ihe lip
of the snout.
Genus PolYIJedllleS
hree of Ihe Bornean specil..·~ of thi~ gl!llliS li\c in primary or "'ccond,u)
T fOrl''''!:. in Ihe 10\\land~. T hc exceplional spet· ies. Polypelillres

1t'lIcomYIH/x, li\'es :llmo:.1 C"t'[) when:.. excepl primary foresl~-in
\Own". \ iltages. :lgricullUral field". road ~ide~. and <.:iC:lred forests. Adult"
spend their lime abo\'e ground in ... hrub<; and Irees. though how high they Jive
ill Iree~ i<; 1I0t ~llo\\'n. Again. r. ll'IfCOIIIY\/(/.\' is an cXI.:eplion: il i ~ often found
ill ground Ie\ el. ei lher in gra~~ or under, or e\'C'n in, house!). All have similar
hreeding habits. M:lII!!:. call around bodies of ~h"ltow ~I:md in g water. The
rema le \\ hen cla~pc.·d by ~. male produces a foam ni.·~1 thai i ~ made by
churning mucu:. willl the hind limh... and tl1al i~ attach~'d 10 ka\·es of 1)hrubs
overhanging \\aler. The foam ne"t uf P lel/wIIIY.\Il/X is often placed on the
ground al \\'ati.'r·~ edgc. After hatching. the tadpole .. w riggle free of the ne~1
and. if the fo:ml llla~s ha~ not a lrcady fallCI1 on to the poor" ~u rfacc. drop inlo
the \\ ater,
Description-a dult.~ : The ~e arc ~lllall to large frog~ wilh relati\ely
... lender. l1allened bod i e~ and long leg~. The eardrum i<; visible. All ha\'(' l:lrge
round or oval pa(1) :ttthe l ip~ of the lingers. \\ hkh arc long and lack \\ebbing.
The toc~ also h:l\e enlarged pad" al the tip ... , but thc~e are sm:liter than tho<;e
of the linger". The tOC, are alnl\N fully \\ebbed. The skin i!:. smoOih or finel)
pebbled on the upper ... urfaces. The bell~ b coar...cJ) pebbled. EXi.·cpI fo r one
~pccies (P, COllt' lIi).lhe ~ kin hcl\\een Ihe e)'e~ i, ru:.ed 10 the underlying "kult
111 larger individuab.
Tadpoh.': Thc wdpoli.' ... Iw\'e fal. :l lmo,1 globu lar. bodie~. \\ jlh Ihc eye"
~el at Ihe :.ides of Ihe head. enabling the tadpoles to "ee up o r do\\ II . The tail,
h:t\'e rc1atheJ) high fin, . The moulh is jU'>l belm\ thl.' lip of Ihe snout and :-.(,1
wilh <;Irong. h!ur.:k bea~ ....
169
RII ACOI'HORIDAE: POLY PEDATES
T\\o fea1Un;~ <;eparate the~e ~JX'(:ic" from the next genu~. RlllIcophorus.
The adult~ of Pol.'pedatl's ha\\! no \\ ~'bhing o n (he linger~. lind thl' !:ldpole~
of Po/.'pnltlll's ha\l' the e) e~ .. e! at the ..ide. All :Il.lul!, of R.1I(I{"(lphol"l/S
.. pecic::. h;l\ e nO!ice:lb1c webbing on (he hand and in all wdpole'i Ihe e~ e .. Me
::.cl on lOp of the head. nOt :II (he ~i dl''' .
Poiypetiafes col/elli ( B ou ien!! t!r)

Collen', Tree frog l Fi g. 96)
Ue-scription : ,\ ~mall I lll :lle~1 I() large (fema]..:" ) tree frog \\ith a
tri:mgul.:lr heao and d i<;( incti) pointed :-.110U\' The <;kin b .. mooth except ror a
fold o\l'r the eardrulll. The .:.lJn on tOP of the head is nOI fu)ed to the )!.u ll.
The!oe frog .. are u.suall~ light Ian (I I" plllki:-.h bru\\ n \\ ith a cOIl::.pituou<; d:lrk
mark on th~' hack in thc <;hape of an hour-gla::. .. or an X. though ,Ollle
imli\'iduab :!f!,; unmarked. The ,ide~ hu\e .. c:lttered dar!. .. pots and Ihl' limh"
haw dark cro::.~b ar ...
:\ Iak): -'+-52 mill .
rcmah.; .. : :;9-77 mill.
Fig. 96. I'"hpt'dmes ol/klli (BoukngerJ.
170
RHAC(}PIIORJDAE: POLYPEDATES
Tadpule : Although ladpoh:, a:.:.umed 10 he those of P colletti have bc..:n

wllecled. their identification remains uncertain.
Bahits and h:lhilat : The specie,> ti\e~ in primar) and o ld ~econdal')
Im\ land and peat ~I\ amp forest ... from near "C" h.'\ eJ 10 650 metre". Adllll~
de..,!:cnd from lrcc~ 10 breed around small pool; of standing \\;\t ('r.
C:III : A :,crie!> of har,h l'f<J!:kling 1l01~·!>.
I)i slribulion : Rcwrded from Sarawak. Sabah. and north central
Kalimantan.
PolJ'p edale~' lelt co mJ'slax (Gravenhorsl)

Four-lined Tree f:'rog (~ig. 97)
Description : A :ol11all to medilllll-si/cd frog \\'jlh a :olender bod) and

lung. :olender hind limhs . T he :oide. . of the ::.llom arc ..,harp. bm the lip is not.
The skin j, !>11l00th C'cl'pl I'or a CUf'l'd fold mer the eardnlm. The~c fT0g!>
"ary from light beige 1U dark tan on the he:ld. hack and legs. :-"Imt indiliduab
h':l\e four !l3ITO\\ dark siripe, running dO\\1l the back. I\hile the rc!>! han:-
<,calh:red bm\\ n 'pOb.
Fig. ,}7. l'O/l'{l('lllI/t' l h' IICO/l/II/II.1 (Gr.11 .:nhor.. [).
171
KHACOPHORIDAE: POLY PE LJATES
l\ lale:.: 37-50 mm.

Fcmak-:.: 57-75 mm.
Tadl)o ll': The bod) i<; ~Ii!!h t l~ tlauened ahme; the (.:olour light buff or
bro\\ n \\ it h irregular m;lrking!o.. Tutal length:. n.:ach 50 min. See al<;o und\:f
genu . . (abo\ e).
lI a hi ts ll nd ha hitat : Thi<; is a frog of di:.turbcd habit:lh. found
c\"ery\\hcrc. including the in:.id(.: of hou:.e~. hut rarely l'ntcring primary
fore . . !. i'- l a1c~ form (.:alli ng group:. around ;111) :.tand ing water. We have e\en
:.ecn pair:. mating <II the edge of :.mall. water-filled road nJl:.. Adults eat a
\'aricty of im.cch and other imt'rtebr;Ut':'.
Ca ll : A 10\\ -pitehed na:.;d l[uack.
Dis tri h utiun: Occur,,- throughout Borneo in dislUrbed ell\ironmenh a:.
high a:. 750 met re:. "bon! . . ea [e\d . This specie,> i, widel) di . . trihuted in
. . outhern A-:.ia.
~o t t' : Difference,,- in -:.i/e and pallem indicate that"P' /i'llcomysln.\·· may
IIlcludc more than \)f1C "-pecic:..
Polypedares macrot;s (Boull:ngc r)

Dark-eared Tree !-rog (Fig. t}~)
Descri ption : A medium-sized to large frog wilh a triangular head and

large e) e.':>. The skin b :.moolh. though ,Ollle frogs han.' \cry small bump' on
the b;J('k. Orten there is:l narrow. \\hiti,h ridge of .':>kin ,Iiong the outer edge
of the fore:I011. This spedl..'') i~ tan to bro\\ 000 the head. back and legs. Some
indi\idu:lis ha\C" a pai r of \\ide black sl ripc" down tht' back. The re j .... a
di'linct dark bro\\n band from the eye. co\ering the eardrum and hecoming
narru" CT as it continue" along till' .... idc. The lJndeT,ide of Ihe head IS heavily
moltlt'd \\ith brO\\n.
1'. la le:.: -i5-57 mm.
Female:.: 66-1-:5 mm.
Tad pole; Tadpole:. h;.]\1.' larger. dee-per bodies ;lIld tkcpcr lail fin, than
ladpole~ of P. lewwIIYs/o.\. The hod~ i:. dark green abo\1.' "peckled \\ ilh
lighter necks. and the uJ1der~idc i<; ,ih'er) gre}. Total length, reach 55-60
mlll. SCI.' abo under gellus (ab{)\e\.
H abits ~lIId ha bita t: This b a frog ot" primar} and disturbed f(Jrc<;!S of
the 10\\ land . . . It gathers in large group . . in \egetat ioo o\cr "-tanding \\mer at
fore<;( edge ..... [ t ll1a~ e\I.'O hrced at dilt·hL·' ,urrounded b~ ,hmbs. Adult, Cat n
\\ide \arit'l) of "mall iml.'t"ls and otha imcrtebrate:..
172.
RII ACOPHORIDAE : PO LYPEDATES
Call : A long dr~m n out r.:hurkle. v. ith the Iir~1 nOle louder than Ihe re!>l.
Ois lribulinn : K~1imalllan. Sabah and Saraw.:lk. It :drnoq n:rtainly
OtTUrs in Hrunci . The !>pecie ... ha~ abo been reported from SUlllatr:t and Ihl.'
...outhem Philippine I.. land<;.
"i~. 98. Pollp<,<ill/('I IIWtlV/i.1 1Bou!engt'r).
17J
KHAlOPHORIDAE: POLYI'EDATES
Polypedate!, oli(ophlls (Boulenger)

File-eared Tn:~ Frog (Fig. 99)
Description : A large tree frog. with a ~aw-edged bony ridge mer the
eardrum. The head i~ triangular with a protruding. ~harp point at the angle of
the jll\\. The toes are onl) half-webbed. The skin is smooth. except for a
pointed projection at the heel and a ~lllal1er one at the elbo\',. The rrog~ arc
tan or yel1o\\i~h \\ ith thin dark. line~ running dm\1l the back and head. The
rear of the thigh i~ marked with eight to t\\eh-e hold hlack bars 011 a \\hite
hackground. The ullder,ide i~ a dirty white. This ~peeics ha..;, a di<;tinctive .
mu~t) ~tench. which mml people find unpleasant.
Mak~: h+-XO mm.
Femaks: 82- 97 mm.
Tadpole : The tadpoles are \W) large (up to 60 nHn long) and are
ye!!O\\ish green. The Lail fins are quite deep. See abo under gCllm (ahove).
Habit.s lind habitat : nli~ tree frog JiH~s in both primal") and di~lurh....ct
ror("~t~. as well as in tree pl:mtation~. It has heen found from near ~ea k\eltu
1(XXJ mclrc~. \bJes call from \egetation around pool<; of standing \\ater.
Adult ~ feed 011 a variet) of insech. but e~peeiaJI) on large tree crickets.
Fig. 99. Pn/\l'edarf S OTllof/hl/I' (Rnuleng~r ).

RHACOPH ORlDAE : RII ACOPH ORUS
Call : A ...eries of slurred. ra~ping nO\I'~ foJ1o\\cd b~ s(':\'cr;11 loud

·\:-hud... ··.
Distribution : Kno\\1l from Kalinlilnwn. Sah;lh and Sara\\:lJ.... It aho
occu r" on Sumat ra.
Genus Rlwcophorus
11 the Borneall ~ r)(.'d~'.), of Ihis genu, o f tree fro~ ... live in fO~ ... I!>.
A either pri ll1ar~ or old .,ecol1dar~. 0l0... 1 HI]O\\ ch,.'\:ltions though th ree
are in montane c ll\" ironmel1l .... We do nm \...no\.\ how high they climb
abme the [or..::.1 noOl". but we h;.l.\c ..,een tree frog ... al le:I<;[ 'ic\cn melre<; abO\c
the ground and hal'c heard calll> from indi' idual~ in the canup). T he Bornean
"'pl'cie::. fall il1lo ,1.'\ eral group~. Fir ... \. there are [ho,>c that breed in flowing
\\alt:r. including R. ollgulirmrril, R. C,WlII{)PIIII{"f(/IIiS and R. SWllli. and an:
relmi\'el) ... mall. All the othl'r BorneaTl ,pcde~ breed in !>landing \\~uer.
though t\\O ~maJl ' lX'ci,,>, R. OI'IIl'lIllif"lIlof/IS :md R. kaj{/u. !>eelll to U'i! ver)
~hallo\\ water. such a<:. !>coeps and shallU\\ !>wamp'. The remaining ~pecies are
"JXlmr' breed('r~. often using \\alluw!> made 0) forc!>t pigs or rhinos. ,\hl)
induded in lhi~ genus are at il:a!>t three Bornl'an frogs thaI tan glide; il i~
possible thaI more ha\e thi!> capacity. The known g.liding s pecic~ ba\e
extensi\c\) \\ebbed lingl'f!> and toe,. \\hkh greatl) ;m:rease their ~\II"face
area hel'au~e they <;preml the tingcr~ and toe" inltllCdiatei) when Ihey jump.
It i~ liJ.-eJy that tht· gliding frogo; u .. ~, thi~ abilit~ 10 C!>r.::lpe enemies :lnd a.<; a
mean!> of de,cending from the l·:mOp) of the rorc~t 10 their hr\!eding o;itco; in
10\\ vegetat ion arou nd pond ...
OesCI'iption- adults: Small to large frogs with e\l~'n!> i \"e l )' \\r.:bbcd toe.;;
and atlca~1 !>ome \\ebbing between the tingers. nlerc b some rOml of fringe
or Ibp!> or soft pmjcctions from the edge of the limbs and at the heel. All
sp~'de!> have tip .. u f fing.ers and loc!> e;o;:panded into wide, round o r oval palb.
The hind limb .. aTe long and ,lender. The e~Lf(.l nllll is \i~ible. T he e)l:!> are
large and the pupil cOlllraCh·into a hori7.0ntal ~tit . Colour.llion \'aric,> widely.
from hrigbl green 10 or~)ngc 10 yello\\ to brown and from unifonll 10 barred
or moulcd. T he ~J.- i n oflhe back j .. u<,uall) fineJ~ p!..'bbled, but nla) have .. m;11I
bump), or proj('ction ...
Ta dpole : Mo~t !>pecies hatt: reb1l\el~ large. f:l1 tadpole!> with relathel~
dl'~'p fim. but in ~Oll1e specit'" they are o;mall. !>lender. ami have naITO\\ t:lil~ .
All u~u<lll~ 11;1\1.:' ~ome marl..ing ...
175
RH ACOPHORIDAE : RHACOPI-I ORUS
RlllIcoplturw; ullgllliroslris Ah!

\ l a~\...ed Tr~'1: Frog (Fig. 1(0)
Descriptiun : A ~rnall frog wJ!h a \\ id(' head and a ~hon pointed ~noUi
\\ jth ,harp edges. The to\:'. C\ccpt for the fourth. aTC webbed to thl: b:!\c of
the patb. The two outer finger' :Ire fully \\ebbed to the pad;;. The ~\...jn i~
~m()oth 0111he head and bac\... and granular on the che~1 and 11('11). The frogs
arc pale £~~ -gn:cll tn ;;and) bro\\ n. Sume Indi\ idua[s ha\e darl-. ntl"har; on
the hac\... and "hite ~pot~ on the ... ide of thl' head or hody. The :,id"'!iof thl'
bod) and the inner surf:u:c\ uf the legs :lre lIsll:tll~ ~I:llt)\\ \\llh \e\emllargl'
bl:l~·\... \P<lh. 11lc chest and bell) arc ~arly white.
\ Iale: 3 [-JJ 1111ll.
Fema[es: 45-51 1111ll.
Tndpnll': The bod) i ... mal and ~tig htl} llalll'n("d aho\e. The ~Iender tail
i., 1\\ ice a, tong a~ the bod). The tipS around the mouth arc "ide and form a
cup-til-.c ,truclure. The body i., darl-.. almost black on lOp :md nn lhe ~ide".
The tailmu ...I..:1c has large dar\... ~pOb and lhe upper fin has smalkr on~". Tota[
length reathc,> J6 mm.
l-"ig. 100. Hlw, ""ho(/f\ lIIH!ufirow,." \hl.
176
RH ACOPII ORJDAE: RI-IACOPIIOR US
BahilS and habilal: This i!> a frog of primary ron:,;!, ~roratlil'alh

common along clear, rod) !>!r~ams above 1000 mctn:!>. Calhng male .. perch
on !>mall trees mcrhanging ~tream~ u<,uu.lI) several metre" abO\l' the water.
Tht' wdpolc<; li,·c in g ravel riffle<;.
Call : A !>harp chirp.
DiSlribuliun: So rar know n only rrom Ihe mountain!> of \\l'!>tem Sabah.
Rlwcopllofll.1i llppemliclI/(I(US (G unther)

Frilled Tree Frog (Fig. 101)
J)esc ription: This sma iliree frog has a Iriilllgular head \\j lh an almo!>!
conical lip 10 Ihe snout in m;lies. In female!> the snout has an enlarged conit-al
proj~(· ! ion . giving them an odd ··boal-nosed·· appearance. The iOes are ahout
Ihree-fourth, \\ ebbed. wilh the l'nds of ~e,reral IOI!S projecling beyond the
Fig. 101. RhaWl,lulI"l1l (/1'1'1'lIdicllf(l/1/, (Glimh ... r) .
177
RH ACOPIIOR I DAE : RH ACOPIIORUS
\\ebbing. The 1\\0 outer finger~ are pUrliall) webbed. but thc others are
un\\~bbed. The ~k.in 01" the upper ~urra<.:e~ i" emered \\'ith 111UI1) ilTegular
~l11all bump~. \\hich tend to be larger on the side". The oUier edges of the
fort"ann and the leg haw a narrO\\ \\a\'y~t"dgt"d fringe of \\..111 . i\ nan'O\\' tlap
of ,,\..in mn~ acro"" the bod) jmt belo\\ the lent. The upper ~urfw.:e~ arc g.rey
gnx!l 10 brown. \\ilh \aTiable dark l11arking~. The under"ides of the lwad and
bod) arc \\hiti"h. \\ilh a \!.."ry "light yello\\i~h ting!.." . About half of the
indi\'idllah h:J.\·e a pinkish linge on the front of the thigh .
.\-Iales: 30-37 111m.
Females: -11-50 111111.
Tadpole : Bod) 0\ al and rounded. The rather di-TP tail I" about twiee the
length of the body alld Il:l~ a ural\ 11 OUI tip. The body is dark grc) ish bra\\ n
\\"ith darker nos<;hands at the reur e.\ tending on 10 the b:lse of the tail. Total
It''ngth reache" .10 mm.
Habits a nd habita t: Thi<; "pecies lhe .. in prilllaq or old secombr)
tor6t at 101\ elevation!>. 11 ha .. hL'"en found in peat ~WUlllP\ and in \\ell-drained
f"or!.."q,. IIlales form large calling group" arollnd low mar~h) aT!.."::', or at slight
depn:\"ion, in tht" fore~t noor bcf"ore any r.:lin waler has acculllulak'd. The
tllale~ perch on t\\igs and Jea\'e~ 01" .. mall trees Ollt" to three melre~ ahml.·
ground. Th~' tadpoles develop in these ~hall()\\ oadit"s of standing \\ :l1er.
C all : A ~hort to long . . t"ries of soft did-Jng n()te~.
Distribution: found throughout Horneo in ilppropriate en\"ironll1ent~.
Thl" ~pecies also occur~ In Peninsular ,\1alaysia. SUlllatm and the Philippine
hlands .
RhacopllOr/lS baillel/sis Inger

Kinabalu Tree Frog (fig. 1m )
Description : A medium-~ilcd Iree frog with a ~hort. pointed "IlOUt. Tht"

t06 arc wehht"d to the ba~e of the pud~. The ollter fingers are half-\H.:bhed
and the inl1er ol1es a lillie less ~O. The ~J..in of lht" top of the head and back i..
~llloOlh or finel) pehbled. The bell) i~ ~·our~l·l~ pehhkd. Thert" is a ~moolh
ridge of ~kin running from along the outer linger along the edge of the
fort"arm to the eJbO\\ and another olle from along the IIUler toe along the
IIl\\er kg: 10 the het"!. Tht, hed ihell' has a distinct pointed .. J..in nap. This
"peei~'" i.. \ariahk in colour:uilln and it i" possible thal:111 indi\idualllla)
change colour depending on lemp..-rature and light. The head and hod~ are
...andy b~'lge 10 dark chocolate brov.ll. \\ llh either dark tran~\ er~c l1larJ..ing~ or
178
RHACOPJ-IOR[DAE: RHACOPHO RUS
irregular light blotches. There i~ a weak ridgc of white bumps ahmc the anus.
Ma[es: 50-55 mm.
Females: 60-65 mrn.
Tadpo[e : Thi~ i~ a large tadpole \\ith an o\a~. ~[ighll) flattened hod). The
tail i~ shaped like an elongate kaL \\ith a blunt tip. and j, about twice the
body length. The colour i~ medium brown . slightly [ightl.:f below. Total length
reaches 75 mm.
Habits and habitat: This frog [i\l~~ in prim:II)' and disturhed rore~t~ at
[000- 1500 metres ahme ~ea lew!. Males call in groups around ~mall ponds .
oflen at the edgc of rore~t~. All tadpoles have heen found in these small
ponds.
Call : A deep rallle.
Distrihution : Kno\\11 onl) from the 1ll001l11ains of \\e~le1ll Sabah and
northeastern SanJ\\ al\.
Fig. 102. Rhacoplw/"lts /)a/fleIlS;,\ Inger.
179
RI-IACOPIIORIDAE : RIIACOPHORUS
Rhacoplwrus cyallop"llctat"!J' Manthey & Steiof

Blue-"potted Tn:e rrog (rig. 1031
Description : A ~m<ll! tree frog with a large eye and a pure white _~pot on
the lip bclo\\ the C) ('. ( Rlwcopho/"l/\ glilmi al<;o has a \\hite SpOl at that point.)
The \\ebbing n:achc, the bas(' or th~' pad~ on all toe" ncepl the rourth , The
two outer finger~ are about half 10 lwo-third~ webbed and Ihe inner lingers
less webbed. The eye is aboul as large as the ~noUl. The skin is ~mooth abow
and coar<;ely pebbled belo\\". There is a narro\\" fringe along the edges of the
outer linger and toc. Some frogs ha\"(: ,('\cral hump, along the edge or the
for~'arm, The colour i~ medium 10 durl br(m n on the head and back with a
dark ... r bar between the e}e~ and usually Olle or t\\O crossbars on the back.
The sides are black with a number of bright. light blue spots: usually the lOp
of the inner tOe'S and webbing hase the same pattern.
!\-Jales: 2H-.1:"i nun.
Fcmak~ : 3-+---+3 mm.
Tadpole : Unk.nO\\ n.
Habits and habitat: Thi~ species has been seen only in primary forests
allow elevations. It has been found in flat and hilly fores!. Although !adpole~
have not been identified with certainty. they probably develop in riftlc~ 01
Fig. 103. RluU"o{JI/(J/"III nfllllllfllllh'lafll\ I\bnlh.:~ & Sicioi".
ISO
RHACOPHORIDAE : RHACO PIIORUS
cll'ar ~trealll<; ,incc that is the erl' ironml.!nl where adulb have alwa> sheen
l·ollected.
Di stri bution : KnO\~n from Kalirnarnan. Sabah. and SnrawnL It
proh;lbl) \\ ill be found in Brunei. Else\~ hl'rl.! it occur" in ...o uthern Thaihmd
and Peninsular 1\lal:t> ... ia.
Rllllcopltorus dlllitellsis Boulengcr

JUlk Tree Frog (Fig. 10.+)
Description : A sm .. lI tree frog \~ it h a l>harpl) poin ted l>nout and a s]endl'r

body. The tOl'l> ;Ire \\ebbed to the b"l~e of the pad.... The finge ..... are almol>1
full) \\ l'bbed. The skin orlhe bad i... smooth. There is a narro\\. \\ hite-edged
ridge of lokin along [he outer edge or the forearm and another along [he lower
kg. The heel has a '>mull t'onical or round flap of ~"in. A lran<;\'er,e ridge of
lolin project~ al1o\ c the anus. The entire upper <;url~Lt'e of t hi~ frog ilo pule jade
green \\jth or \\ithollt tin~ \\hite dots. The upper eyelid and a mlrro\\ line
along the edge of the "lout are reddi <;h bro\\ n to purple. The top of lhe \\ eb
rig. !U~. Rful("opht,nH '/IIIi/('/lI/1 HllulL-nger
1X1
KH ACOPHORI J)AE : RII ACOPH ORUS
b~t\\cen the out~r toe" i" red. One (lIthe odditie~ of thi" "pccie~ i" that the
limh hone . . are lurquoi'e gre<;,n and \ l"ihle through Ih~ llc..,h from Ihe
umkr"lrJe.
~Ialc, : JJ- HI mill .
....emale . . : -19- 50 mm.
'IiHlllulc : A largL' tadpole, \\Ith 1K"~t\). m31 bod~. Thl' tail 1'" moderate!)
d~.:p and :I little le~ .. Ih311 t\\ ice the bl}d~ length. T he boLl) ;lnd rront half of
thl,tail an:- pa le gre~i~h bro\~l1." \\:1\). blad.,. \>:nicalline cro~"'e~ the tail al
mid-kngth, ht'"yolld \\ hieh th..: t:1I1 b da rker. Total length reaches 50 mill.
Habib and hahitat: T hi~ b a Ir(lg of primar~ fore!'lt in !lat or hl ll ~ countr)
III the lowland". Link is kntm n of it.... hahll". It form., large br":L'thng groups
around fore!'.t pond .. at unpredict;tble inh.:nah. apr;]rt'ntl~ ..,t imulah:rJ h)
"uddell heavy rain:>. T h..: tadpole" Jt\e 1Il the:>e pond"
C all : ,\ ~hort, \\eaL eric\.,et-1Jkc trill.
Dii)lribulion : Kno\\1l from S:thah. Snr'l\\aJ.. alld rwnh centra l
K;llim;ll1tal1. It aho occur.. in SUlllatra.
RIWl"opltortl 'l el'ereUi B uulengt'r

~ I o"") Trc~ Frog (l-ig. lOS )
Description : A !'.mall "'I~K:J..~ frog. \\]{h slender. long leg ... and a shon.
blunt .. nout. T he webbing on the til": ... doe~ not quite rea1.:h thc ha..,e of the
patk The lingL'rs are webbed on I) atth~ ha..,l's. T he skin oftbe upper ,urtaccs
i~ rough, \\ ilh r11a ll ~ tillY pointed proj ~'o.:tio ll ' m cr the head and bad. T here
i\ a "emi·\.·i rl·k of .., pi k ~ projection!> running o\ cr Ih .. shol1luer~ between the
l'arurunh. ;lI1d l1uml'rou.., "'111111ar, though ~maller. ... pin} hlllllP' ~cal1ered o\a
the ha~k. The foreann and 100\('r leg h;\\e a 1"0\\ 01 "'l1wl1. cone-<;haped
proJcctioll ... along the outl'r ..:dgl'. nlC<;'e frog!> are mOille.! in colour... from
light gret'll to dark bro\\ ll. T he lOp of the head i:> darJ... \\ ith :l light har
bel\\l;'l'n thl' eyes. T he bat· J.. i" a mixture of patches of tan and bru\\n. T he
upper thigh, are lIsuall~ greeni..,h ) t'llu\\ \\ lIh hro\\ n ... pot.... The Ullllcr .. idc 01
the leg~ i ~ grl.'l'n.
i\ tale<. 30 -J2 1Il111 .
r:":lllale~; -1549 mill.
Tndpole : L'nkno\\ 11.
Ilahil'i and IlHbilat : Thi" fwg li\ 1.''' In primaJ;. ~lIbmU1l!an.: ;lIld montane
fore..,b from 1100 to 1800 1llt.'lrc" ;lhO\c ,ea ]e\·el. It i~ often found in rn(l" ... ~
area~. a ... un tallen log!> near !>trcam'. Lillie is ktlO\\tl of it.... h;lbih.
RHACOPHORIDAE: RH ACOPH ORUS
Call : A series llf sholl. har<;h no!!:!)..

Dis lrihulion : ;"'Iounlain<; of \Ie:.lern Sahah and northea:.tern Sarall:lk.
Also knoll n from P:llawan in the Phil ippine Islands.
FiA. 10:'. Rlwn'flllIlI"IlS ('I'nell; Bouknl,'cr.
RhacopllOrWi galilli (Inger)

Shon- nosed Tree Fro£. (Fig. 1(6 )
Desniption : A small slender-bodied !re'\.' frog with a Icry ~hon ~nout

and prominent. bulging eyes. All toe, except the fourth are wehhed to the
ba:.e of the pad!>. The outer lingers arc a!lllo~t fu!!~ II ebbed. The upper
qlJ"faees of the bod~ arc "l1ooth, though there usua1!~ i, a ~mall. ,harpl~
pointed. ICI1ieal projel·tion near the edge of the upp.:r eyelid. Then' is :11<'0 a
"mall JXlimed projection:1\ the hed ami three or four other<; hct\\ccn tho: heel
and the base of the outer tOt'". The bad, I" lI"uall) ught tail Ilith !>cIl'r:tllarge
dark ~pot<; . Therl' i, a con"pK'UoU~ while Spilt belO\\ the e) L' , The hidden
~urfaee:. of the thigh~ are a rich golden yeIlOl\.
183
RH ACO PH ORIDAE : RH ACO PHQRUS
r-. l alo;!'~: :1f)....JO mm.

Fernak,: 35- JR mill.
TlIclpolr : Til!, ~IJl<!JJ !adpolt' 1J.1.~ nil 0\,11 botl} di'lincIJ) J1nuened helU\\.
and a broadl) rounded ~nolil. Tit.: lail j<; ,lender. wp ... ring 10 a rounded lip.
and i ~ al mO'II\\ it:e the hod~ lengt h. T hl' m(lUlh i ... su rrounded b) \\jdt' lip<;
Ihal hang down [omling 11 cup-like ~l rU(' lUro;!'. The beak ... art" black. Ihid. amI
V-~ha[X'd. The bod) I~ } e llowi<;h hrowll. The tail mU~t'k' ha'i hrO\\ n or black
"pOh. Totallenglh reaches 25 mm .
Habits and h a hitat : T hi", ~p"'t'k~ Ii\c, in primary fo re~t in hill} WUlllf}
lrom near ~ell k\(,:l LO 750 metre~. It li\ l'~ in vegetation overhangi ng lhl'
rime" of clear. rot:k) "'Iream". Calling m ale~ foml too,e group... t--:e m a l e~
all:.u.:h their white [0:1111 nC~1 LO ka\c<; o\'e rhanginJ:! th~' ~tn·a m . The "mal t.
"tender laupo1e .. move inlO rillle ... ;md live In the cre\ice~ among the gr~l\cl
and J'(x:b of Lhl' hOllom.
Cull : A ~inglc ~harp note. a high-pitdlCd t:hirp.
Dist r ibutio n: KllI.l\\n from Sabnh .llld SarawaJ...
IR-l
RHACO PI-IORIDAE: RI-I ACO PH ORCS
Rlwcophorm' /wrrissolli Inger & Ha ile

Bro\\nTree ~rog\Fig. 107 1
OCSlTilltion: A mediulll-"i/cJ In.:~ frog with :l pointed. ~h arp-edgcd

... nout. 1\ 11 of [hl' 10('''' an: broadl~ \\ebbed [0 thl' ha"c or the pad ... Thl' thrcl'
outer tingcr~ ar~ :Jbo \\ebbed to [hl' pad .... The :-.~in of the had, h. line!),
pebbled or ~moo[h. The belly i" coar~ely pl..'bbled . There arl' no ridge~ 01' flap"
of ,,~in on the limh,. The he;ld ;md ba(:k of thi~ ratlwr plain frog :lre reddi,h
bro\\ n or clay orO\\ n. The !>ide uf the head ~OIllCI i1l1t.'~ ha~ while Th~ "po"'.
sides and thc hidden ~urfaces of the leg" arc gn..'I..'nbh yellow. The \\~'bbing of
the linger!> and loe!'i is dark hrO\\ n nr bl;ld ...
jvl;lle,,: 50-56 mm.
Fen1:lIe ... : 60-70 mill.
Tadpole: Th~ !xxI) b O\al. gelll'ra l l~ rounded aOO\'e and bdo\\ . The tai l
i ... aboul 1\\ ice the body length . Til..- hod} and !ail are dark grc~. Total length
rl'al..·hl;'~ -+ I mill.
Habits Hlld hahital : Thi ) !>pec i e~ ha~ been found in primary and old
~econdal) forc~h in hiU~ <.:auntry at to\\ e\c\ alkm". Lillte is known ahtlUl th~
usual forag ing hl'ight of th is 'pecie;;. bUI it dl')t'I:nm to within a fc\\ metre~
of t h~ ground to breed. All tadpole ... I..n ~m n have been oh"~n cd in water
ViI!. 107. RII(/('OII/Jllntl IlfIr"'~Wlli Inger ~I... Hade.
I X:;
RIIACOPIIORIOAE : RHACOPHORL'S
1\ ilhlll Irl'L" "holes" 1--+ metre~ abo\"e ground. Foam ne~t~ ha\e b.:en ~el'n
atlached 10 hark jU<;1 ahm·e a Iree hole. Some tadpole ~ile~ I\cre hole~ in Iree
!run"'.;: olher\ Wl'n~ formed h~ the fLl~ing of trunk bllttre~se~ making rather
large ianb.
Call : UnkIlO\\n.
Distrihution : KnO\\ n from Sabah and Sarawak.
Rhacophorus kajau Dring

While-cared Tree' Frog (Fig. IllX)
Dl'scription : 1\ \L"I") ~rnall IreL" frog. \\ ith OLller fingL"rs onl~ ahoulone-
third \\.:hhed. T(l':\ arl' ,Lhout hall-\\C'hhed. The .. hill i.. lind) rehhkd nil th.:
back and coal"!>d~ pebbled on the hdl~. A narl"ll\\, \\a\~ ridge of !>kin ruth
along the outer finger and aliter edge of Ihe forearm and a ~imilar ridge run"
along the Ollier toe and edge of the lower leg. There is a narrow tralh'·er~e
ridge belm\ Ihe anll<;. The colollr i~ leaf~ green 011 all urrer <;urf..'lC"C". ll~uall)
1\ ilh J11111UtC \1 hill' \POh \catlerL"d {)\ er the hac\... head and e\po.,ed \Url"<Kl'\
or Ihe limh\, The eardrum and the inner linger~ are \\hih:, The ]o\, ... r ~id ... :.
and bt'll) an: without pigment. The webbing i~ black and the hidden ~urfaces
of the thigh han." an orang ... ling ....
Fig, 10K Nlwn'p/ro/"lll J.."llW Dring
186
RHACOPIIORIDAE: RHACOPIIORUS
Male,: 18-20 nUll.

Femaln: L'nkno\ln.
Tadpole : The hod:; i~ O\al and rounded. The tail is drawn out into a
narrow tip and i~ about twice the body length. A large hrown blotch (O\'ers
moq ofthl"' top and sides of the bod). The tail inu,cle ha~ brown spot~. but i\
mo~tl) pale. Total length reat·he:. 17 mm.
Hahits and habitat: This is a poorl) knSAIl species that has been \<..'en
only a fcw limes. It lives. so far a~ i~ knmJil. only 111 primary fore~t at
elevatioll!> belO\I 750 metres. :-.-tak!> hale been found on ]eal'es of ~hrub~ and
!>lllalJ trees overhanging ~mall. slowly flO\\ing intermittent ,I reams and
marshes.
Call: A '(rit'~ of single or douhk dit'b made at intcna\s.
Distribution : Sabah and Sarall uk.
Rhacop/lOrtlS lIigropa/matlls Boulenger

Wallace', FJ:;.ing Frog (Fig. 1()9)
Description : A ler~ large tfee frog II ith an almo,! rounded snout and
very large hamh and feet. Fingers and to'"':' are fully \\"ehhed. The skin of the
body is vcry line!) pebbled. bUl ha~ no projections or ridge!>. The forearm ha,
a \1 ide. \mooth-edged flap of "kin thut extends 10 11K- t'dge oflhe OUicr linger.
The heel ha~ a wide. rounded nap of skin Ihat continues On!O Ihe outer edge
of the foot. All upper ~lIrraees are emerald green. sometime~ with a few
glo~s~ Ilhite ~pots on the b:lek. The underside i!> white . becoming )ellowish
on the chest and helly. The sides of the hody and legs are deep yellow. The
webbing of Ihe feet i~ black. fading into )ello\', to\\"ard~ the lips of Ihe toes.
Males: 79-89 mm.
Femak~: 89-100 mm.
Tadpole: The bod) i~ lj\'al and rounded. The lail is a bit k~~ than twice
the body length and i~ rather deep. The hody i~ pale gre) ahove and white
hl"lO\I. The tail ha~ hlack ~pots 011 the IHll,clc and fim . \1 ilh a dllrk edge to
Ihe upper fin. Total kngth reache~ 50 mm .
Hahits and habitat: Thi, i, Ihe original "Flying Frog of Borneo"". It is a
resident oj primar) fore,! al km de\ atiOlb. Frogs rome dO\I n from the foresl
canopy to breed at turhid p(lnl~ of \\ ;nero and foml ~mall group'" around wild
pig or rhino \1 allows. Lurg~' fO:Hll ni!Sh aft' attached either to Illud banks or
leg<:tation olerhanging: the pools. Large number, of ladpole~ hal e often
been I"llund inexlrernl"l) mudd). Illrbid \1·;1.tcr.
1S7
RH ACOPHORIO!\E : RH ACOI'H ORUS
C all : ;\!UI tle~o..:rih~'d.

Dbtribulion : K n()\~n I'TOm Kalimantan. Sabah and Sara\\aL II
undoubtedly oe(:ur~ in Bnl11..:i. The "pccie~ ha~ al~o been found in :,olllheftl
Thailand. Peninsular l\ tahl) ~ia and SUlllaira.
RhacopJlOrlis pardafi"i GUnlher

Harlequin Tr..:e f rog {Fig. 110)
Dt'sniplion : This i~ ~\ ~!llall to mL'dlll!n - ~lZed Iree frog . \~ith a rounded

"lWUI. The toe~ and Ihe Ihree oul..:r ling..:rs ar..: tu ll ~ \\ t'hbt'd l0 the pad~. The
,kin or the hack i, ~m(lo[h. \\hilc Ihe bel1~ i., · (Jar.,cl~ pL'"hhlt'd . A wide
~m oolh - l'llg..:d Ilap of ~kin run, alo ng the Oliler edge or the t\m~anl1 and hand.
The h ...(') has a rounded flap of :--kin. The upp ... r ~urf,!~''':' ,If": tan In reJJi~h
bro\\n. \\ilh Jark markings often forming an X in Ih..: llliddk uithe hack.
,,,
RHACOPHORIDAE : RHACOPHORUS
il. l o~t indi\'idual~ h:l\l' ~e\eral \Ihite Spot,> on the back ':lnd ~01lle hal'e yellO\\
or blue "poh on lh~' ba~'k and upper ,urf;I~:C~ of the leg'. The ~ide, and beJJ~
are ~ ello\\ i~h. II jth black 'POI', on the "ide~ and :1TI orange nelworh. on the
bl.'ll). The I\ebhing on thl' hand .. and fect i, orange-red.
i\la\e'>: 39-)5 mill.
f emale,; 55-71 mm.
Tadpule : The hod~ 1, Ola l and rather deep. The tail is slight!) tel>~ than
1\\ ke the hod~ length and tapers to a naITOII tip. The hod) b pale light
brown. Some ladpolt'~ have "eleral blal.'k ~pob on the body. while Other~
or
ha\'e Ju~t om: ~pot un the ,ide thl.' lwad. Total kngth n.:aehe, 45 111m. The~e
t:l(lpoh:~ m:l~ be confu,..:d II ith tllO~e of R(wa c(wfcOIIOf(/ tx:cau~..: of their
'polled pattern. hut onl) the latter hille II hite glandular pa t ~·he .. o n the
under,idc.
Fig. 110. RIrIlt'UphoI'1I1 (),mlall \ Gumh..:r.
IX9
RJ-I ACOP J-IOJ{Jl)AE: RH ACOPIIORLS
Hahits and habita t: Thi') Img:. \\ hich IS a proficient glider. i~ found in a

\1 ilk \ aricty of forested habilat~ al 1(1\\ ck\ation~. It form~ small breeding
group~ around rnar~hes. pond~ and quiet ~idc pooh of qreams. It is relati\"ely
common along logging roads where ~trellm~ are ortcn blo{:kcd and form
pook The nonmll l"oragl11g hahitat is unknown . but is presumabl) in the
forest canopy. The tadpole:. d<.:\elop in ~tallding waler.
Call : A hrief ra~py chuckle.
Distrihution : Known from Kalimatllan . Sabah and Sara\\ak. It \1 ill
probably bl." found in Brunei. [t also occurs in Sumatra and thl." southern
Philippine hland~.
RlwcopllOrus reil/wardti (Schlegell

R~'inwardt"~ Flying hog (rig. 5 & 111)
Description : A medium~siLed \0 large tnoT frog \1 ilh a hroad head. The

10(,S and three outer fingers are fully webbed to the pad~. The ~kin i~ granular.
more coarsely all the sides and belly. The olller edge of the forearm ha~ a
\1 ide. ~m()oth-cdgcd Ilap of skin. The heel has a wide . rounded flap .
(.."ontinuing 'IS a nl.l1TOI\ wip along the edge of the lower leg. The upper
surf<l(:e:. are a rich. Jark gr<.:en ~callcrcd 1\ ith dark spots. fading into golden
yellow to\lard~ the ~irJe~. Th<.: ~irJcs ha\c a hluck hand SCI \\ith wrquoise hlue
spots. I n some frogs. the blaek i~ limitd 10 small area ... ~unounding the hlue
spots. The webbing is black \\ilh golden )ellO\\ and lighl blu\.' :,poh. The
Ihroat. eheq and belly are pure white.
Maks: -1-6--55 mill.
Female ... : 56- 65 mill.
Tadpole: The tadpole i~ large. \\ ilh an o\"al. ~hghtly tlattened body.
typit'al of lhi~ genu .... The tail i~ le~~ lhan twice Ihe hody length. The ~hape of
the wil is leaf-like. \Iilh it~ deplh about one-third ih length. Tntal lenglh
reaches 60 mm. The colouration is greyish bro\1 n \\ ilh St· altered blac k doL~.
Habits and hl.lbihlt: Thi:. handsome frog ha~ been found onl) in prilllar~
fnreqs at 1m\" eie\·'lIions. It descends from the canopy 10 breed around sl."llli-
permancnt pools all the forest floor. As this species has been found only a fe\\
timc) in BOl1lco. little i ... knOl\ n of its hahits. Adults probably feed on canop)
insects.
Cull : A 1Ll\\ cr'K"kling t·huckle .
Distribution : KnO\\n from Sabah and Sara\\ak. It aho ()n::ur~ 111 Ja\a.
Sumatra and Penilbular r-.lalay ... ia.
190
RHACOPHORIDA E : RH ACOPHOR US
RIWCOIJllOrll S rttJipes Inger

Red-legged Tree Frog (Fig. 11 2: )
Dl'scription : Thb b a ~Illa ll tree frog \\ ith triangular. JXlinted head. The
toe~ :lre full) \\,.'bbed. The ou ter fi nger.. are aho eXlcn ... i\ d) webbed . but the
web doe ... nol 4ui1e reach the pad~. The ~ki n is <;J}1omh un the upper <;urfaces.
but coar..,d ) granular on the hcll~. There are no fri nge::. or nap<; of <;kin on the
li mhs. The colour of the upper ~ urf.aees i, [an 10 reddi,h hro\\ n \\ i[h a few
bro\\ n ~pot~ or c ro<;~IKlI1lh.. The webbing of tht' hands and fccl and [hl' hidden
s urr:Jl"e~ of the leg ... a f l' bright o range-red . The under<;ide of the bod~ i<; pale
)'cllo\\ i~h \\ ithout the llrange network ..een in H.. 1)(II"{{alh.
~ I ale .. : 3~-~tJ mm.
Female,: -'8-50 Illlll.
Tadpole: Unkno\\ n.
Habi LS and hahita t: I.ittle i~ known about [hh ~pecie,. 11 h;l, heen found
unl) in primal) fore,1 at 10\\ e!e\·ation,. II form<; hreeding group, all
\egetalion \\i1l1ln 1\\0 mClre) of the ground. it ~eelll<; to hreed in ,hurt.
191
RHACOPHORI DA.J:: : THE LODER:vIA
unpredictable burst~. ~ubsequentl~ di~appcaring:, \"othing i;; I-.nO\ln of its

normal foraging habitat.
C all : Unknown.
Oi stri/Jution : KnO\\ n ~o far only from Sabah and Sarawak.
Fig. 112. RlwC/)p/roru .1 mfipe,\ Ing('r.
Tlieloderlllff /lOrridlllll ( Boulenger)

Rough TrL'c Frog (Fig. I 13)
Description : A small tree frog. with the head as wide as the hod). The
tips 01 the finger~ are expanded into I'er) wide rad~. that of the fir~L (inner!
linger a<, II ide a~ the eardrum. II hieh i.. \ i.. ihlc. Till.' outer finger, arc II ehbed
almo .. t Lo the ba .. e or the pad~ <.lnd th,,: loe~ an.' compklel) lIebbed to thaI
point . The skin or all upper surface, is l'v\ered lIith lllan), small \\hitish
~pines cluslered onto round II art~ on lhe top and side of the head. sides of the
body. and upper sides of lhe limbs. The ,mall .. pines of lhe back are arranged
into irregular lines and clu~ters. The upper surfaces are dark browlI. with
large hlack area, on the side'>, The under,ide IS ·Iarl-. grL'~ II lIh hold II hite
m:trbling.
~Iales: LnklH)\\ J1.
r-emalc .. : ..J.0--+9 ll1m .
192
RHACOPHORIDAE : TH ELOOl:. Rl\ IA
Ta dpole: The !adpole'~ body j" O\aL ... Iightl) nal1ened. and \'er~ dark on
the upper w rface and !>i de~ . The tai l lapcr~ to a l>harp point. l\ la_xHllulll length
i~:12 mill.
Hahits and huhilal : The on I} :.peci men," of [hi" "pcde~ e, C'r discmered
in Bo nll'O \\ erc in 10\\ land rain fore ... t. The tadpole.'> were found in a \\ater·
contai ning tree hole about J.5 lll\.'trc~ abU\c Ihe ground in the ~allle for<:~t.
Dc"pile the scarcil), of re~·onb. ' . . e ~U~peCI thai thb 'pecie~ j" not rare.
bcl·ilU"C the rough "kin and dark colour pro\-idc C\l"ellcnt carnou tlage for a
frog cling ing to the trunk of a trec.
Dis tribution: The onl y Bome:m rCl'urdl> for Ihi" frog :Ire from ea.'>lern
Sabah. It is also knO\\n from . . outhern Thailand and Pl·nin ... ular :-' I:11u).'>ia.
Fi ~. 11 3. /Iwloderma horridllll1 1B\)ulengcr,.
193
J u rth pr l\eadill bo·
Berr). P.Y. (1975). The Alllphibian FnUll1l of Peninsular Mullly.lill. Tropical

Press. Kuala Lumpur. x + 130 pp.
Campbell. EJ.F. (1l)Y4). A IItdJ.: rfmJllgh (IU' LOIrfand Roill Fores! or

Sabol!.
\ 'atural lIi~tor~ Publicaliom momen). KOla Kin:lbalu. \"iii + 8.~ pp.
Inger. R.E {1966 J. The SnfClIllllics and Zoogeography of tlu' II lliphibin of

Horneo. Fieldialla: Zoo/mi.l 52: 1---4()2. (Reprint. 1990)
Inger. R.F. and R,B. SILlching fl989). Frogs or St/buh. Sahah Parb
Puhlication no. 10. Sabah Park.\. TflIslee~ . Kota Kinabalu. i\ + IJJ pp.
Inger. R.F. and R.B. Stuebing (1992). The Montane Amphibian Fauna of
Nonhwestern Borneo. Al afayan Nature }ollmal .+6: I-51.
Inger. H. .F.. R.B. Smelling and Tan Fui Lian (1995). New species and n('\\'
records of Anurans from Borneo. The Hames Bulle/ill of Zoology
(Singapore!-L\(I): 115-131.
Inger. R.F.. R.B. Stuebing and Tan Flli Lian ([996). Frog:~ and Tomb. In:
Wong. K.;\I. and A. Phillipps (eds.) . Kinahulll - SlIlIlllli! of Borneo (2nd
Edilion ). The Sabah Society. KOla Kinahalu. Pp. J5J- J67.
Inger. R.E and Tan Fui Lian (1996). Thl? Narl/rat His/ory of Alllphihians mui
Rel1lile.1 ill Sahah. Nalllral H islOr~ Publications (Borneo). Kota
Kinabalu. xx + 101 pp.
Wong. Anna ( \99-1). Popll\mioll Ecology of Arnphihian\ in dilfncnt altitudes

of Ki naballl P:lrk. Sabol! .\1 USl'1I111 lmll1w{ I (2J: 29- .18.
Wong. K.r-.1. and c.L. Chan ( 1997). ;\'1011111 Kinal!atl/- Bomeo's MaKie
MOl/nwin . N.'llurai II j'-loT:; P ub1ication~ (Borneo ). KOla Kinabaiu. \'iil +
Y5 pp.
19-1
,J Ick n o I r-{pd coO'eill eni s
Support for our field \\or}.. in Borneo has COllle from the f-ic!d t-. l useum of
N:uural I-li~lory (Chicago), Ihe International Tropical Timber Organi/alion.
lilt: r\ation:li Science Foundation (CSA). :IllU the John D. and Catherine T.
MacArthur Foundation. WI,' r~l·t'i\ed much logistical ,uppOrt from Dmuk
Lamri All. Director. and ;-' lr Francis Lie\\, former DCPU1) Director. Sabah
Park,.; Prof. Ghazally hlll"i!. formerly of UniH:r~ill Keb;mgsaan ~ l ala)'ia
iSab;th Campus): lJatu Cheong H Choull and Mr James Dawos 1'. lamit urlhe
Fore"'lr~ Department. Sarawal..: Dr Charle .. Leh. Sara\\ a" ;o. lur"cum: and staff
of Sahah ForC,>t [ndu~lri~,. We llmllk D:nuk CL. Chan. Dr Anhur Y.c.
Chung. Dr Indraneil Das. Dr I\tauricc KOIlc]al. Bjom Lardner. Kehin Lim.
John I\ l urph~. Tee Kim Ling and w.~ 1. Poon rur u:.e uflheif phOlogrnph~ and
Tan Fui Li:lIl for use of her dra\\ ing~. Our work in Ihe field \\ null! nOI hu\e
been pm~ible or plca~urahlc \\ ithout the help and :ls,nciation of Tan Fui
Llan. Ahlll Biull. Paul Yumbull. D,lIl~on Kandong. Frcdd) Paulu:.. Frederic~
Franci,. P;lIr;c~ Franr.:i~. Dr Anna Wong. Alhcrt Lo. Shahbudclin Hj Sahk).
Saifuddm Sen:l\\i. Tmb Jainudin. ,\nung IIcma\\:m. Setiawan. Tum Frist.
R~ an RiLl. G:lUn Sureng. Aba~. Jawu. BJJok. Puulu~. ~ I amo. and Illan~ other
Ikld a'::.blants. Our \\-ork III Ihe !.\hmatory \\;1:. greally :ls~iqcd b) ;\ir Alan
Re~elar and ~ 1 <; Ca<;~andra Redhed We are \"el") grmeful w all of these
per~ons and orgamlalinn,.
195
ll/de.\"
C
"
.I"wlllu. 21. -'8. 67-8
1I111</lIId",lm" Inger. 21. 2-1. MI-9. 6!i
(""/11,,,/1(1, 17.
ImXJb,
J~. 93
( Buult.·ng~·TI. ::!5. 9.1--1
ml()ll! Inger. Tan & Yambun. 1-1 JlllIa Kic\\. 25
Jrdi~IIl<'11 4\ 1'1I:4UJrJ). 2-1, 69 JIIl/I!!1 (Barbour &. ~otJkl. 25. ',14 _~.
Inger. .21. 2-1. 70
gllil'l'i
{1II11111£111
I(l/idi~rll
Ing~r.:!l. 2-1. 70 I. 71
Inger. :! I. 2-1
"
C/1II1'l'mW f!H('{I i\kll.:4u •.lfJ. I Y. 2_~. ,10,
_
' -'. Ijj·o. 95
ftp/UI"" (Gilluht'n. ::.~. ,~. (,7. hi-\, 2-1-8
Chc..:\":Ii~t.
7'1. 3, 72 Cla"ilic;JllOn. 15 1.1
"J1!1(fillgllrl II1gU. 25. 7:'-.4. 7J.
IIWIII7" Inger. :!.~. 7-1 D
pl(//\It)/Iw lngC'r. 21. 15 lJi,nihulion . I I 1-1
'pllm/iler I ;"hl<,:quarJI. :! I. ,25 . .'5. 75.
~ E
/01.,.,,111/1 Dring. 25
Anuril. II
F
II !-ejCl1unu
R(/rbtl/lfllili. 1.1 ('(IIla/Lom IGr:lIenhur-I).\). 21.16.
k"lim<lIlfWlf'lIt/1 h)..;md3f. 2-1, -I.~. ~ 29.311. IIJ I-I. IIJ
Biol(Jg~. 1-6 hllllloclUlr;1 !Hoic). 22. 26. 21).
Bombin:uoriuJc:. II. 1.'_ 2-1. -I, 11-1---1:•. liS
Bulo,7f1 FoIl-tal..... X-I,/
fI)/'uGr3\enhl}r-!. 17. 2_~. 7ft 7.77, Fr,'g'
711. 110 Bush. JJ. 161
di"'~"11I r..-!cr'. Ir.. :::5. _~5. J.7. Arollll.167-9. 167
79-80. 79. III Golden-It:ggt:u. 161 -'. 162
jrutaV"',·lngcr. 17. 2:'. n. 76. 77-'). Gret:ll. 1(,2. 16J
71'l. 80 H o~·s. I 65-{). HiS
mdmrl"li<l1l1 S.:hn'·ldcr. 16.25.29. R<"d-e~ei.l. 16-' -1. 16-1
~(). 76. 80. tU Sharp-,nOUlcJ. 166-167. 1M
f/lwdnpUllflllll I3nulengcr. 10.25.38. Cinnamon. l,'iiJ---t>lL 100
HI 2. 112 Creek
l3ulmlldal'. 11.2-1. r.7 Kuhl"' 3~. 12-1-5. 115
Ilullfroc:,. II em·kel.19. 151 2. 151
·\Il1Cri~·3n. 10 O\\;.Jrl
Bam.leu. 11.2'). 10-1 i. IUS Inj1ds. IIS-19. 119
Bro"n.21J. 10_....... 1IIJ 01.1 atf Litter. 12. -15 6
196
INDEX
Kin<lh<llu. ~6 Ri\ukt. 1~5--6. 125

MJobcrg·s. ~6--7. ~7 Rock. 31
Rough-~il.kd. ~B Black-'polled. 30. 156--7. 157
Striped. 48 Green-_pollcd. 1:'i7-R. 158
\\eb-fooll:d. ~7-S ·Poi'onous. 1-1R-<.). 149
rbt -headed. ~3. -1-1 Rough-backed. 120
Flying Rough-sided. JR. 147 8. 147
Rdn\\ardt\. I~. 14. 33. [90. Saffron-b... llied . .'10..,3. 95-0. \)5
191 S,-'cp. 139. 140-1. 1-10
Wallace·s. 11 . .:1.'.187-8. ISS Slender Li ller. .'5. 5 ~-5
Gra".29.114-- 15. 1I5 Dring·s. 55--6. 55
(;r~en Padd~. ~9. 14f> 7. U6 /I,·jolllane.58
Guardian . .'':1. 36 Painted. 58-9. 59
Rough. [3. 37. 38. )~0--1. 121 Sar<lwak. 56-7. 56
Smooth. -'7. [~O. 1~9-130_ 130 \\'hilc-bcJlied.5 7 8. 57
Holc-Ill-I he -Head. 117 18. 1i7 Spadefool. 3-1. 93
Horned. ]::'. 35. 60--1 Blolch-sided . 94 5. 9-1
Ilorne~n. 8. 3h. 64 5. 65 Brook.,.. 9-'--4
Dring',. fl} Stick). 35. 97
Kinabalu. 61. 62 illlt":mltXliat.... 99-100. 100
/l.lolllane. M Kinahalu. 97-8. 97
Rough. 63--4. 63 Lesser. 102. 1U2
Lilt..-!". 32 ..1.5 Rulou' -~Ided . J6. 37. 100-1.
Ahboll·'.38 1"1
Black-eyed. 53. 54 Vari ahle. 98. 99
Large-e)~d. 49-50 Stream
Lowland. 50-I. 50 Spotted. 31. 3 _~. [52--4. IS]
/l.iontane. 5~. 5J Slriped. 152--4. 153
SpOll~d. :; [. 52 S\\;Jmp
Long·,nomed.90 Greater. 30. J-I. JR. 123----1. 123
/l.iahogal1). 9. J.'. 1-19-50. 150 L ... ,,,Cf. 131-1. 131
.\1angrole. 9. 29. 3S. I U 1-1. 113 Pe;)!. 38. 11X-9. 129
t\!al',h Tai\\ane,e. 10.19. 3,\;. 115-17. 116
Brown, 1-I3--t 1-13 Torrell!. 74. 131----1
!\<lTrO\\-moulhed. II. 9.'1. 107 Brown. U7-8. 137
Berdlllorl'·'. 107-8. 107 MOnlane. 13~ ', _ 1.'5
,5.
B(lme~n . . lOS 9. II)\) Xorthern. 1.'5----6. 1.'6
Least. III. III Whitehead·s. 13X. 139
PUlholc. 109-10. I]() Tree
l' lldtllc. 1.'19 Blue-'fX)\fed. 180-1. 180
Yello"-hcllied . .\0. 141 ~. 142 flro"ll. -'3. IX5 0. 185
R.i\er Cinnamon. J3. J~
Giant. 13.30..'14. _~5. 3S. 3'). Co!lcn\. 170-1. 170
1~7-8. 127 Dar[...-t":ared. In _'- 173
l{ough -bad.. etl. 13. 1::'2-3. 122 hk -e.Jr... d. -'-1.174--5. 174
197
INDEX
r(lur-lin ~d. ~Y. 171 2. 171 \IIhl<'I,..'I/I·i.1 Inger. 25. HJ2. 102
Frilkd. -'~. 177-8. 177 K allll,{iI. ~
Hariequill. 12. 3~. _'5. ISX-<)(). balml<l (.\lli[lerl. 17. 25. 29. U'-'----l.
1H3
'"
Jade. 12. ISI-2. 18 1 !Jld(I!I'<IIGra~ I.~. 17. 26. 29.114-5.
Kinotb:llu. 178-". 17') IU5
.\1.1,,!..eu. 175. 176--7. 176
\lo",,~ . 18::!· .' . IS3 I.
Ori<'!ltal. II l .,pIO/)ru{'hdl<l. 20. ~:'-6
R~tl - kg:g:etl. Ilil ~. 192 hllilu'lI.lil' Smith, 2-t. 4:'i. 46, 4 S
Rough. 192- 3. 193 hrt'I'iCf'IIl Dn ng. 2~. 45
Shorl-n""'.· d ..':'i. IS.' --1-. 183 l!I}ohagl Smith, 12. 2-t. -t.~. -to 7. 47
\\·hJtc-eared. 186---7. 11'16 pll/mala Inger & Stuebin£. ]4. ~5.
Tree Hole. 3.'1, 105---6, iOn -t7-R
\\·hnc-lippcd. J~ . .'1:'i. I--W-:'. 145 plln'a Dring. 2~, ~:'i. ~H
\\ 'rinl..leu. 1.32 ;,ua,wwe Dring:. 24. ~S
LCl'lohmcililllll. ,2 ..'5. 50 ~9
C (lbbolll Cochmn. 20. 2-1. 38. ~9-50.
(i1l.,lrIIl,h,TIWiil<,., 50
h()/"11,·"" .\i~ (lloukngcrl. 17. 2.'i. 96 fi;1II11111fi;/"lji,\ r.. 1 ~[kIllU" 2-t. 52
/l<'lI<lrld:'.lfIIli Ta:- lor. ~ I. 2--1. 4Y. 51.
H 52
H"l'lohWI"tU'illl'< /l/UIIIIIIIIIIII IFi,dl€'f). 21. 2 ~ . ~9. 5 2.
rugu/rllll' tWlq:mannJ. [0.20.26. 53

29.30 . .'1t;. 115-17. 116 1Ii.~rcl/" Berry & Hcndrickson. ~O. ~~.
~9. 53. 54
Cwill/!/I"!III!m d301lIeng<'f!. 18.20. l.l'pl<l/a/m. 2(L .'15. ) +..5
117IS. 11 7.13~ (lran!l .\\at,ui. 2~
(1rill~i DlIhoi,. 2 ~ . :'i.~ 6. 55
f!f(loili< IGiinthcrl. 2 ~ . :'i6-7. :;6. 58
/umli<h Mahui. ] ~ . 57-f( 57
Illgualla
(]loulcngcr). :'..'1. 26.
halll"lI~i.<
maul'Q I nga. fl. 1aldarin. Aliln.l.:
118-19. 119 Y:ullbun. ~-I. 58
prum .\J.tll..mu" ~ ~ . 57. 5S Y. 5,)
'-"pl()l'linll(' hOl"hollic(f IT~chudi I. ~ [. ~:'.
K
82- 3. 8.'
Kalo/,llf"I11J. 18. J:'. 38. 97
LllllIlo/li'('{('~, III) 20
/W/II<'1I.I1.1 Ki<,\,. 25.1)7 S. 97
MJWI'Cllo linger. Boc~dl & Tauii!.. J.
t;O~ Dot .. &. Ila:L.. , 25
::6. I]()
hnenJt'iIlIHI Boulenger. IS. 25. <JS.
hll-Ihii. 1~S
'J')
(ill<'hi (]rtgerl. 12. 1-'.]1. ~fl. 3.~, -,fl,
irllt'I1I1Uilll,1 Ingt'f. 2:'. 99-101J. 100
.H JK. 1211-1. 111. 130
!whimla Dring. 2.~
li>mWI'/(1II linger!. 9.13.20.26.
1'/<'ufV.lliglll(IITsehudd. 18. 2:' . .'16.
12~-J. Ill. 12~
37.98. l()(J---I. iOl
III,<,/"iIK inl I. Y. ~(l. ~6,~O ..' ~ . ,'~.
PIIIICI<lf/,-, Pda,. 2.~
12 ,~----l . 123
198
INDEX
kr/1('pai,>IHi< (llIger). 26 h",(/mm'l'i ml ~thl. IX . .:!6. 107 8. 107

!./lillii 1Dllllle:ril &. Hibron). IS. 211. II(I"h'Cllli~ I'arler. IS. 26. J5, .1,6.
-'0. 12-1 :'i. 125. 1-11 lOS 4. I()\)
!lIIi"p~ rBoulenger). IS. 211. 12.~ 6. maw/irail Inger. 26
125 1"'lpun'lI Inger <'I: Frognt:r. 18.211.
Itponllll,l L-I.nJer"unl. 1.\). 10. 1,1. 109.111. 111
20.26 . .10. 34. 35. 38. 39. 7.'. pCl/'I,!!I'1lfI In);cr & Frogncr. 18. 211.
12l 12-1. 12.~. 127 X. 127 109-10. J 10. III
1II0/,><il1l11'; II-\IC\\). 9. 22. 26. J8.
128-<). 129
/>(II(II'(III"llIi< (Rouicngc.n. 12.21. 26. N
35.36. 120. L!<) un. Uti
,'lI/"llIIlUrmd(1II (lngcr). 22. 26. -'8. I,ieflll (l-'t'lt'r\). IX, 27. 3.'1. 3-1.
131-2. iJ l 159---60. 160
r/w('()illl< (In,l:cr, Ilocadi 8: Tauliki.
1~1-,2 0
On·ido:,\~IJ. [J9--IO
~I I"dllulli~ 11~"Ulcn);c-r). 18. ~6. I ~').
~lcgoph0iJ'le. 11. 2·t -I,' 1-10-1. 1-10
If,'goJphn<. ,11. 35. 60-1 /U<'I'i.1 (Gimlhl'n. 1t\. 16. 30. 1.'19.
b(/I!l..u~1) (Houlen)!erl. 17.2-1.11 1. (,2 !-t1-1. 1-I2
drimd Ingl·T. Sluchmg 8.: Tan. 17.2-1.
"(,.'
nlortllllilllll' Ingt:T. 17.2-1. u). (,J---L
[>
Ihlrmi/w<. S-I
"''''1'1/1 IHoukngcr). 1;'i
J..oilu\(II/1II ~i'll)..nlU' &. ,\ lahul. 11. I1mi! (Bnukngcrl. 16. 25. 32. 34. 35.
2-1.6-1
7J. 8'+-5. 8!"
11<1\1111'1 (Schkgc-II. 8.17.2-1, 3K. Nt
1~l\l'xluard I. 25
III(/Wltll/i.\
6-1-5. 115
rIIcM/lllngcr. I~. 16.15. X5-6. 86
,I"·ril'tll~rlln.
22. -'5. 38 . .'9. 7-1 . Lt2---l Pclobati,jae. -15
(lmorupa/wlIIl.1 \1l iJt'lliJ. 211. 13-1
Pdop/wl'lll'. 15.36.87
jerboll ((;imlhnl. 26. U4
Opt Dring. 25
kill(1bo/llt'11:>Il ling,.n. 22. 26. U-I 5.
',(lIemhai ~B()lllc·ngcr). ~5
135 mifem (,\hx·quard). 15. ~7. 8S
/!tlltmphr!ra/lIIl1s (~ I msuil. 26. IJ-I
'/IOJ!uJ!hililll I ng~r l~ StUdlHtg. :!5.
,,'p/lII/)(IIC'1II1.1 (i\1,lhuit 211. I,~ fl.
1.'6 ",
<iCItIllIl (8ouJengt'TJ. 25. S9-9O. tW
flh(u'(lIIIt''''~ linger & Gnti,!. 211. n-l. }'III/(IIIII/." 2J. 1-. .'18. 161
137-1';' 137 <.1('/11111 Dring. 27
,,,w, iitlf (Ingcr &. Gritb). 26. I_q Il/IlOellll\ Smilh. ~7. 169
"lIihll<'aili IBuuk·ngal. 211. I-,t;. I.'W <IIl/wuiuIII Inger. 27. 161-3. 1f,2
.I/"l<Il'ilnlldla
Imllilll~ Inger. Stud' llIg ,\; mn.27.
fI//Idol//ltI'",t,.r,J. 19.211. J,'. I(J:; fl. 162. 16.
106 diler,'elll Ingcr, 17
,Iii, n,/r\,/u. 1(17
199
INDEX
t'nrIIlVI,hllwlllllt) Stu.;bing & \\'(lng. ~i!!lllIlill (ju nl h cri. 19. ~7 ..11. 7.'.
'27. 16 .~---4. 104 15'2---4. 15.'
Iw,ii IRnulcng~rl. n. n. Itl!. IbmJat". 11.26. II.,
165---6. 165 Rhncophoridae.II.17.159
/II~l'I'i Dl'lng. 2'2. '27. lli6 167 . If)() RIIlI('oph()nI~. 12 . .iX. 170. 175
keram,lIe Oring. 27 ""'(IIIi/WII"!:> Ah!. 12. 16.27.175.

Ir)ll~i"/'II\ IRll uicn gat. 27. 16l) 17tl 7. 176
IIIjoiJn-).!i Smith. n.
168, 169 aplit'lltii(lIitilll' IG(lnl her). 27. 3-1.
p,'leni (B(Jukngeri. n. lli7 '.l. 167 175. 177 X. 177
re/il,(ii Inger & Slllcbing. n.
169 /la/wll!>i., Inger. 1:;.27.178- 9. 179
\(Iueri r.,·lalkrnu, & RieJt". '27 chmol>lmCWlII.I" I\ l anlht"~ & Skiol'.
1I'I/ilS Dring. '27 12.16.27.175. 180---1. I~O
umhrtl Oring-. '27. 169 dulil<'/l.lil H()ult"ng~r. 1'2. 2X. lXI'
Pol.l'JIf'dilln. l(il.)-70
c·o//eIIlIHoult"ngt"rJ. 23. '27.170-1. '"
""1'1'<'111 Hou lenger. 16. 2X. IX'2-.1.
17H liB
lell('/l/llI;It/\ IGraH'nhor.,tJ. 2 ... 27. (<1.1<//1110 Bou lt"ngt"f. 16. '2il
2lJ. 1m. 171 2. 171. 113 gmllli1 Ingn). 12. 16. 2:t3.~. 175.
111{/(1tJ1I:J IBou1cl)g~rJ. 2'2. '27. 171-3. 180.18_'-----1. 18]
173 harl'ls.I(lIIi 1111'er& I l ail~. 16. 2X. -'.'.
()/iiopillH I RoulcngCr). 2~. n.
J 4. 185---6, I ~5
174- 5. 174 /..lIjauOring. 15. 2X. 175. 1:-;6 7. IS6
fl(,lIlio/mjo lIi~mf!a"!lIlIll\ l1(lulcngo:r. 11. 15. LS.
\IIb(/~l'aTschudL '2U. '2:;. 90---1. 90 .'3. 36. 187-8. 188
/'<I,.,li~li\ Gimlht"r. 12. 15. ~S. -'2. 35.
R \88- 90, 1~1j
Rmw.3:-;.I ·B
H'immrdll ISdlkgt"ll. 12. I ... I 'i. 2K
/)(Imlllica RncTtg.;r. 23. 26. 1·-13---4.
.'-'.36. 190. 191
14.1
ru{rpn Inger. 15 .28.191-2. 192
hll'lhii. 12X
Roek Skir~r. 13 ..,--1.1 5:) Ii. 155
c"a/co/ioluISchkgdl. 19. '26. "''2. 35.
144-5. 145. IS!) s
l'ITllil'<1l'n (S<:hlcgdl. 19.27.29. SI<Illmil. 154 5
146---7. 146 /(I/(l/M/III(I//iS IRoulcngef). Ll. [5. ~7.
f!lalldlllo.lu Bouicng-a. 23. 27. J:-;. 3-1. 154. 155---6. 155
[+to 147-8. 147 n(l/af()1' ICiumhert. 19.27.30. 15--1.
IIO.lil HouknglO'r. 1'1. 27. 31. 14:-;-'1. 156---7. 157
141) IiIh<'l'ilil1~!111 Houkngt"f. 19. n. 15-l.
11II1'rllII<I('II"I/<I Bnrbour 8.: 1'.oblt". 23. 157-8. 15~
27. 14-l
/mlmJ:>1I Welef'l. 19. n. 3-'. 1-l9-50. T
ISH Thdodf'I'III(I
lIicob.u·ifllli) IStolic7bl. 22. 27. 19. IlOrrldI4lIlIHOllk 11g~rl. 15. 2S. 1'l'2.1.
151-2. 151 II)]
l'i('lImlla BOlukng-t"f. l'l. 27 ..,1 . .,5. Toad,

1:;1-3. 15 ,~ Blaek. .'~
200
TND EX
Blad..-"poued. :!<) Gll ibc's.70

Common Sunda. tlO. S l KinalXlh!. 70 1. 71
Crested. 35. _n. 7<)- 80. 79 Long-fingered. 7J---I. 73
Cnh'. X2 -'. ,11;\ ~Iontant' . o()
O\\arf.87 . Spin). 35. 75. 7S
Climhing. 88-9 \\ hite-liplktL 6X---9. 6S
Kinabalu. 87. 88 Swamp. 38. 81-2. 82
Lov.1amL 89-90. S9 Aq uatic. ')0 I. ')U
Rh er. 76--7. 77 Tree
GianI. J 7. 76. 77-9. 7S tho" n. :'2. _~ ... J5. 8..- 5. 8S
Slender. 67-8 l"Ibrblcd. I:! . 85--6. 86
f3 rown. 35. 7::!- J. 72 True. 07. 70
OllarI'. 7.. Tradition. 7- 10
201

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,f1bout th e 13001\

A ,"' jcld G uide 10 the f rogs or Borneo

Front cover: RhacQpllOrtls reiml'Urrltl. one of

2000 - 3000 m Balldar

Robert F. Inger and Robert B. Stuebing

Natural History Publications (Borneo)

Naturalllistory Pllblicatiol/s (RomfO) Sd". Rhd. 1 ~1t>1;()7· X )

Tcxt copyright (Q 2005 Natural HislOry Publications (Borneo) Sdll. Bhd.

AI! rigins rescrvcd. No part of this publication lIlay be reproduced. ,tored in

First publishcd 1997.

A Fi{'"ld Guide to the Fro!!;s of Romeo (S{'"cond Edition)

Fmnti,picce: Map of Borneo.

Preface ............ . . ... vi

General Biology of Frogs ....... "'" .•.•.......................•....... 1

Frogs in Human Tradition and Culture .......................... ..... ........... 7

Di stribution and Classification of Frogs of Borneo .............. II

Ecology of Bamean Frogs .................................................................... 29

family Bombinatoridae ............ . . ...................... ........................... 43

Family Megophryidae .. . ............ .... ............... ........ ................. 45

Famil y Bufonidae (Tme Toad:\) ........................... 67

Fam ily Microhylidac (Narrow-mouthed Frogs) .......................... . .... 93

Family Ranidae ........ ...... . ................. . . ... 111

Family Rhacophoridae ...... 159

Further Reading .............. . ..19-4

I Danum Valley Rest House resulted in the publication of Frogs of Sabal!.

descriptions for most of the genera.

I cOlllinue to be discovered in Borneo. Study of this part of the rich fauna

"eryone can recogni,l;I.' a frog. which is a good indication of ho\\

E distinct and relatively unilom1 these animals arc. Their distinctive

Another distinctive characteristic of frogs is their complex life cycle.

frog (Kalol,hrynlls pfellrtNiXlIla) produces grcal quantities of annoyingly

e~pilc a divcr~ilY of about 150 specie,s in Bomeo. frogs do not play

D a partit'u!arly prominent role in the folklore and traditions of the

Llemon~trated by (he wa) frogs nee. panic-stricken. from a ~n akc . A

"MallY years agu. a man lil'illg ill a I'Wage IU'{I/' MOl/III

chicke n". In SlIbah, an exot ic species culled ··bongkok Taiwan·· or Taiwanese

Fig. ~. Plate of trog" ~ Icg ~ fried in batter.

bout 150 species of frogs occur in Borneo. The exact number is

A uncertain because new species are discovered every year. Thiny-

If we examine the anatomy and behaviour of adults and tadpoles in !>omc

Ncverthek~~. Borneo's fauna does show links !O adjacent areas. Fony-

Mount Lumaku in Sahah, and 011 Mount Mulu in northern Sarawak.

FiA. 5. Recently emerged frog let of Reinwardt"s Flying Frog (RhacophofllJ

Key to the Frogs of Borneo

2. Webbing bem een linger;, thid.. lleshy ............. . ................ Pefvphr\"l1e l

3. Colour black with irregular. small light spotS or lines. ............. .

5. Body green .................................................. . ...... 6

8. Webbing red ... .. .. ... ...... 9

9. Forearm with flap of skin along ouler edge . . RhacoplwTUS partillliJ

13. Pak blue spot.~ on \ide~.

15. Sidc-~ yellowish with black spots. . ................ N/wcophol"ll.\' allgu/ilV.lfriJ

19. Parotoid gland n:lrro\\ and elongated: colour often orange.

20. Tips of fingcr\ natlencd and squari;.h in outline ....................................... 21

25. Skin of back wi th one Of more nal"l·ow. elongme ridgcs . .___ 26

26. Projection on C) did an m 'el'hanging triangular nap of ~kin . ... 21

27 . Conical proJ('ctioll of , kin from tipof snout .. M(,~i!l'hIT.\· 1l1lSIIIil

28. Eardrum \-i,ihil: (rig. 21. .... 39

2Y. Lnder~ide oj foot with ('nlarged. ra ised spade- li ke proj('ction .

.1U. Tip~ offing(,I'~ ('xp~mdcd into hlunt pm_h. ...... . ............ . ~I

J I . To(', wchhed . _ ..... .... ..... Koloula hrile(l(o

3-l. Fir~! finger missing .............. ...................................... Micruhy/a petrigt'llu

40 Spots present on side of body in from of hind leg ... 4.1

4 1. Spots blui~h or whitish ............................ ........... KlI/ophrYllus lieterocliifll.\·