A Field Guide to the FROGS OF BORNEO

,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
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. 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: [email protected] Website: www.nhpborneo.colll

1 ~1t>1;()7· X )

Tcxt copyright (Q 2005 Natural HislOry Publications (Borneo) Sdll. Bhd. Photogmphs and illustrations copyright
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 Key to the Frogs of Borneo. .. ............ . A Checklist of Anuran Amphibians ........ .....

.............. II .15 ..... 2...

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

...... -+ I

family Bombinatoridae ............ . Family Megophryidae ..

. ...................... ........................... 43 . ............ .... ............... ........ ................. 45

Famil y Bufonidae (Tme Toad:\)

........................... 67

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

. .... 93

Family Ranidae ........ ...... . ................. .

. ... 111

Family Rhacophoridae Further Reading .............. . I\cknowlcdgcmcnb Index .......

............ .

...... 159 ..19-4

............. 195 196

cf>rejace

10

th e § irst edition

I has been j ust under ten years since a discussion on the veranda of (he 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.

I

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' 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

I

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\\ 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 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.

E

1 There

j,

one "pedes of frog on the Indian ~\llx:ontin~nt that i, reponed to con~ume a~ part of ib diel.

aquatic \cgetatioll

Fig. I (opposite). A ~mall :.Irc
GENERAL BIOLOGY OF FROGS

parotoid

heel

extra joint

Fig. 2. Outline ()ffrog showing anatomical Fui Lian)

feature,~

referred

to

in text. (Drawn by Tan

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

GENERAL BIOLOGY OF FROGS

Fig. J.

Slagc~

of dc\ .:Iopmcnl of lad poles. (Drawn by Phillip Yong after Tan i-'ui Li:tn )

3

GENERAL BIOLOGY OF FROGS

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 onl~

lTIa)

require

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

GENERAL BIOLOGY OF FROGS

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

GENERAL BIOLOGY OF FROGS

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 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~

D

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
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. 10

~.

Plate of trog" ~

Icg ~

fried in batter.

c{)istribulion and Classification

of fProgs of C/3orneo bout 150 species of frogs occur in Borneo. The exact number is uncertain because new species are discovered every year. Thinyseven 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 ('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.

A

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. Fonyfive 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 Roughbacked 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 ............. . Webbing between lingers thin. membranous

................ Pefvphr\"l1e l

............ 3

3.

Colour black with irregular. small light spotS or lines. ............. . .............. ................. .. Staurois Imopa/lllallI S 80dy not black ........ ................. ................................ -J

-1-.

Top of head and Not a~ above

5.

Body green .................................................. . Body not green ......... .... . ................ .

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 Wehbing not red

9.

Forearm with flap of skin along ouler edge . Forearm with no skin nap ... .. ....... .

~ides

extremely rough and wany .. .... Thdol/erllla horridum ................ .... 5 ...... 6 ......................... ...... . S

... .. .. ... ...... 9 10 . RhacoplwTUS partillliJ ...... 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 ................................ 14 Sides and thigh without blue spots .....

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-~

Not

yellowish with black spots. abow

a~

. ................ N/wcophol"ll.\' allgu/ilV.lfriJ

.............................

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 IR area. as large a~ or larger than the eye No large gland behind eye. ................... ...................................... 2-118. Top o f head between Without ~uch ridges

e)e~

with a pair of narrow. raised ridges. ................ .

19

....... 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 .

.___ 26

25. Skin of back wi th one Of more nal"l·ow. elongme ridgcs . 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:':;"'.. M(,~i!l'hIT.\·

27 . Conical proJ('ctioll of , kin from tipof snout No projection of \kin frolll tip of ~no Lit .

28. Eardrum \-i,ihil: (rig. 21. L:ardrum not \'j"ibk . 2Y.

.1U.

.... 39

29

Lnder~ide oj foot with ('nlarged. ra ised spade- li ke proj('ction . Lndersid(, of foot not as aboye .

Tip~ offing(,I'~ ('xp~mdcd into hlunt pm_h. Tips of tillf'(,fS not expanded

J I . To(', wchhed . _ To('s \\ithout wehhing

1l1lSIIIil

MegopirlTs kobdl"llSirii

._.i 2

...... . ............ .

~I

. CuI/wi/a' ..... .... ..... Koloula hrile(l(o .. ..... .. ....... 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

Snout nOI ~o long.

/;O/"ll{'CIII;1

......... 33

~ 1,1I"::"::11 fnunJ '-"l t~ on \l nunt ~I u lu "'lei "3 t"II~ -I~g~~d

«I

'I'.:"~'.

3 1 Sp(~":JC< Ll J Cdlll,'111' .Ir~ hurn'''~'' .n,d '''rd _ ' t:II~(lLJnl~r~d Ji<<."ll\ .,~d III

Di'"m:liull> .lIllLln~ I h~"l

arc'

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 Toes not fully webbed

...................... ...... ... ..... ... .. ... ... Lilllll()/leetes kuhlii .................................... .. 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

blui~h or whitish ............................ ........... KlI/ophrYllus lieterocliifll.\· Spot(S) black. ... .................................... ....... Ka/oplirYlllo p/ellfusriKlI/li

4 1. Spots

-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
~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 le..\1.

18

d~'L'nplion,

in

KEY TO T HE FROGS OF BOR.!"lEO

-+..j.. Tpp of lirst two t\JL:~ while or hlu.:." . Toe" nOl as ahove .

-+.i

..... -+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 grey 10 black . Coloura!ion otherwi ."c

;1

thin pall' blue or II hili~h linc. sides

....... . ...... .. ............... . Rlllw /1I("tllo.\(/

.... .. .. . -+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. Body dark bro\\'n or black. will1

.. ....... .. ........ .. .......... .... ..... ... 50 ycllowi~h

(lr n."ddish spots,

lighl edge to upper eyelid . Cnlouratioll (ltherwi~e

al\\"ay~

II ith ... .. 5 1

.. 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 ..... .... ,. .

Throat without dark central

.. Hoplobmmclw s ruguloslIs

~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 ................ . To.::., at least partially wcbbnJ .................. . 60.

60 ........................................ fi5

Tip~ of lingers and toes distinctl~ pointcd Tips of fingers and toes rounded or blunt

61. I lead plu~ body less thim 2.5 em . Head plus bod) more than 2.5 CIll 62. Ikad not \Iider than body I-kad wider than body ..

.. .. 61 .. 62

....................... Leprahmchclla !>pp J LeprohmchillllllliXmps

.............................

......................................... Lepra/a{a.l spp.s .................................................... 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.: mlcro,copt". 5.:.: ut',cnptlons 1Il1<:.~t . S) Ta.~{)nol11ish :If(· ~tilltr)ing to ,on out the ,pt"(ic~ of thi, fcnu., anu ~pccic, an.· t-cing ui'ico'cr~d 't("auily. Sec (k~cripllon' IIltcxt.

20

JnOlh~[

III

"ithout J

the procc"

IlC\\

KI:.Y TU THE FROGS

6~ .

Bell~

or RORNI:.O

light ill colour. 'pco.:I-.kd Ilith tine bbd, doh . . -...... . Lt'{lfOhmrl/lwlI Ilo1
R(' II ~ gr~'::

found abolc 1000 m :litltLld.;'_

.... Lt'pro/1nlc/rilllll

6:' . .--\n II1\lT[ed \·-,I].lp~·d ridgc on hud, b.;'ll\.;'enllw ~(l ,ud] ridg~'

~ll\Iuldcl"

11/"11/(11/11111

.

6(, .. __ ...... _ 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', Tip~ oj' toe ~ not II ilk-ncd inti) di,tinct pa(\, . 68. ,--\ .,mall. bla,'I-. X in o.:l'ntrc of b'ld. 1\0 'Ilch Illarklll!,' (,l).

77 . (,~

...... Lt'/'loplr'TJlf

/J(!/'h"I/1((/

6 t)

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

Tip, (>ffinger, 1101 II ider th:m ]a,[ joint _

,\ II hltc Ill' lT~all1·cololired 'lrcah. rrul1l

c~c

to armpit . . .--1l1villia

73 lhll'd ~,nJ fifth t\IL'~ \Ichhcd to till> Third und lii'th lO~" Itot 11,'bbed [,> liP'

/luidiKd' 7"Y.

al/JOlllti,

Itlliftl

.... ..11/10111<11:111"1'1 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. A few rounded rough area" or back smooth

... 75 .............................. 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\\

No ;.uch ridge

-edged bony ridge above the enrdrtllll . ................ .. PO(lpedares mi/op/lIIs .. ......... .............. .............. ....... .. 79

79. Eardrum covered b) dark bnnd extending along shoulder to armpit , ....... ...... ......................................... .......................... Pot.\pcdare~ lI1acmris ........... .... .. 80 Pallcm olben. ise ... ... ..... ... ......... ....... .......... . 80. Outer t(k: \\ ebbed to ba;.c of pnd Outer toe not II ebbed to pad ..

81 ... R3

81. Third toe webbed to base or pad on bOlh ~ides ............... . ............... 82 , Phi/millis hosii Third toe not 'Webbed tu pad on both sides . .................. . ... .. ..... Phihmllo il/gerill 82.. Baek and sides bright green with dark ~p()h Hack and sides brO\~ n or blaek , ....... "....

...... JH£risrngelly.l· killahallll'llsi.1

, 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
22

KEY TO THE FROGS OF BORNEO

lB. Tips of finger~ not much widcr than next joint Tips of lingers abouttwiec width (lfncxtjoim . ........ .... . 8 ~ . Sidc~

with enlarged ~kin

gland~.

producing: a rough or pebbled

............... ........ . .. ...........

Skin of ,ide not as above

... .... ........... 84 ····· · ........... ... . 86 ~urt'acc

. Raila g/andll/oul

....................

... ... ........... 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. Legs rather short: skin rough

................. . Legs long: skin smooth ......... ....... ......... ... .

88. Snout sharply pointed Snout nm poimed .

............... .. .. ......... 87 ....... . lngl'l"(//w haflll'luiJ

.......... 88 .............. Pulypedmes cof/c Hi ... 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('
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'
A. A. A. A.

14

jilligillea (~·ltX'quard ) * fJ llihei Inger* IWllit.l'chi Inger'"

/witlil'ca Inger'"

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. L. L. L.

keneplIien.\·i.\· (lnger)*

kuhlii (Tschudi) laticcps (Bou1cngcr) 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. H.. H.. H.. R.

erythmea (S<"hlegel) g/ulldlliom Boulenger

hosii Bou!enger larerinll/culata Barbour & Noble 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* 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 I~

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 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

T

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

.l!

fnfC'i

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 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'. theG iant Ril er Frog (LlllllllllltTle.1 /t'I'om/llI) 1110'-1 frc4u('ntl~

FiA. 111. ro.UTl ne'" (It tn:.: frog er-hanging rhlllo II aUOII

III

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~ \
ECOLOGY OF BORNEAN J--ROGS

"llCh dead

a~

the

~tiek~ frog~ tKalophr\'lIl1sL ~pI."nd mo~t

of the time under the 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. lIl\\~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 lean~ ..

of the

fore~t

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 Topeeies \\ 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.!, 'Ille 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
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 ('Oll~lnne~ ~pidef~,

ladr()k~ ,

bt't'tlt's, I-..m;.did\ and other onhoplefan~ , but aJ1t~ arc mort' than ."We ofih pn:~ . The generali~b tend to eat larger prl'~ :md to inge~t fewer pr('~ p("r meal. For "",\
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 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
D

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 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.)

T

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

T

he Bornean

specie~ u~ed

to bl!

c\a~~itil'd a~ mcmbcr~

of the Family

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"

T

he genu .. LeproIJul( Iwl/a Include;; the .. rnallest adult frogs in the

fatluh r-Iegophr)ld.lt:. ,mo 1.., \...no\\11 onlv from Borneo and the BllnQ:ar.1ll l ..omellmes c.!Iled Natutl
ar~' ~IX

spcl'les

in Borneo and three. Ll'{JlObrachl'lJ{/ II/jobergi (Sanl\\ak). L. bafllellsis (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~ \\
:" 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 Sabah, northc11l S,lra\\ ak. and nonhea<;tern Kalimantan.

\'.e~tern

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.~:

In

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 150 500 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 Slrir~'d

,~eras allae

Dring

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 ott the CO:l'l of BUIl1t'o.

"til

MEGOPHRYIDAE : LEPTOBRACI-IIUM

Genus Leptobrachiul1I Large-eyed Liner Frogs the 1l10~t \~ idely di~tribll1ed ~pecie~ of Lcprob/"(/chiu/II 111 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\ .

P

re\"iou~l~.

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
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,·/\. UJO(."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 :

Fig. 17.

LEPTOHRI\CIIIU~\l1

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 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'
A

~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 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 ba~e

Fig_ 211. L"I110/
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. 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(ieide 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~

T

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. 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. r-.lale~:

remale~:

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