(r--lorth Borneo)
Rana df!corala
Y!IX')UARU,
1890
Cd/amaria /aff!rali.. M()( '()I '/\RD, I B90
Pd/lll'OKonia CPphdllllr/ MOC<)I.ARD, 1890
AMPHIBIANS f:I REPTILES of MOUNT KINABALU (North Borneo) Rudolf Malkmus, Ulrich Manthey, Gernot Vogel. Peter Hoffmann & Joachim Kosuch
with 384 photographs
by
c. Bruhl, c.L. Chan. I. Das. W. Denzer, T. Eitz, R. Gerhardt, A. GOtzke. W Grossmann, T. Hiltida, W. Hoot D. Knowles, U. Kuch. M. Kunkel. B. Leideritz. K.E. Linsenmair. W. Malkmus. S. Manthey, M. Maronde. M. Matsui, J.e. Murphy. A. N611ert. M. Pfeiffer, H. Sauer, M. Schroth, R.B. Stuebing. T. Ulber. M. Veith. S. Wilson. and the authors
A.R.G. Ganmer Verlag K.G. 2002
Frontcover:
Backcover:
Phi/aufus auran/iulIl © R. Malkmus
Trimeresurus cf. SlimatrallLlS © K.E. Linsenmair
Address of tbe authors Rudolf Malkmlls, Schllistr. 4. D-97859 Wiesthal , Germany (Introduction. Mount Kinabalu - the su rrounding environment, Amphibia (with exception orlhe Hlmily Ranidae» Dip\. lng. Ulrich Manthey, Kindelbergweg 15. 0-12249 Berlin, Germany; Emai l:
[email protected] (Sauria (lizards) and Te
tudine~
(turtles, tortoises, and terrapins)
Dr Gcrnod Vogel, 1m Sand 3. D-69115 Heidelberg, Germany: Email:
[email protected] (Scrpentcs (snakes» Dip\. Bio\. Peter Hoffmann, 1m Oberdorr41, D-79292 rratTenweiler, Gennany: Emai l: dorin.hotTmann@t-
Translation from the German manuscript (with some exceptions) by Herprint International CC, P.O.Box 14241 , Bredell 1623, South Africa
rSBN 3-904144-83-9 © 2002 A .R.G . Gantner Verlag Kommanditgesellschaft, FL 9491 Ruggell Distributed by Koeltz Scientific Books, Herrnwaldstr. 6, D-61462 Konigstcin I Germany Fax: (+49) 6174 937240 Email: kocltz@ t-ollline.de Phone: (+ 49) 617493720 Internet: http: //www.koe ltz.com Layout by Sibylle Manthey, Berlin / Germany Design by Ulrich Manthey. Berlin I Germany Printed by Fuldaer VerlagsanstaIt , D-36037 Fulda / Germany
ontcnts
Contents ~---
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Preface ........................................................................................................ ............................ ...... Acknowledgements ...... ... .... ....... ................................ .... ... .... ................... ... .......................... .... How to use this book ............................................................................................................... . Map of Borneo ...... ................. ........ .................. .................. .................. ............... ...................... Introduction ............. ... ... .... ..... ... ... ....... ... ............ ... .... .............. ....... ... .... ................ ....................
7 8
9 11 12
Mount Kinabalu - the surrounding environment ......................................................... 13 Topography, Geology, Geomorphology.. .................. ....... .. ............... ...... .............................. Zooge graphy ....... .............. ....... ............... ...... ............... ............. ................... ........ ....... ......... lilnate .............. ....... .... .. .. ........ ............................ ..... ....................... .... ......... ... ... :. .. .... ... .. ...... Vegetation ......................... ... .......... ...... ............................................... .. ... ... ...................... ..... Tropical lowl and vegetation zone (to 900 III a.s. l. ) .............................................................. Tropical lower montane vegetation zone (900-1800 m a.s.!.) ...... .......... .......... .................... Tropica l upper montane vegetation zone (1800-3000 m a.s.1.) ............... ............................. Tropical subalpine vegetation zone (3000-3700 m a,s. !.) .................................................... Fauna ... ......... ....... .. .......... ....... ............ ............ .. .. ... ...... ..... ...... .. ...... ..................... .................. Herpetofauna .... ....... ....... .......... ....... .......... ....... ........ ... ... ....... .... .. . .... ... ..... .. ......... ..... ...... ....... Tluman settlements in the vicinity of Mount Kinabalu ........................................................ A retrospective glance on herpetological research in Borneo and the area ofMt. Kinabalu .....
13 15 16 20 21 22 23 25 28 32 38 40
Amphibia (amphibians) ........ ................................ ................................................................. 46 Anura (taiJless amphibians or frogs) .................................................................................. ImpOltant traits for the identilication of frogs and their larvae .................... ........................ Morphology. ........ ... ....... ... ...... ....... .... ... .... ... ...... ..................... ... .... .... ... ...... .............. ............. Habitats of tadpoles .... ......... .......... .... .................. .............. ............. ........ .................. ...... .. ..... Larval feeding types .............................................................................................................. Bioacoustics ....... ....... ....... .. ... ,............. ,..... .. .. .. ............ .............................. ,. ...... ............. .... .... Reproductive strategies ......... ............. ............. ...... ........ .. ...... .. ........ ......... ... ... .. ..... ....... .... .. ... Key to the families of Bornean frogs .................................................. :................................. Preliminary key to the larvae of Borncan frog fam ili es .... ............ ............. ........ .......... ........ Bufonidae ........... ...... .. ...... .... .................. .. ............ .. ............................ ............................... Megophryidae ......... .. . .. . .. .......... .. ... ... ....... ... . .. ..... ......... ... ............ .. .... ...... .. ... ............ ......... Microhylidae .......... .. ........ ..... ......... .... ...... ....... ....... .......... .... ............ ...... .......... ..... ....... ..... Ranidae ....................................... .............. ......................................................................... Rhacophoridae ................................. .. ................ ............. .. ...... ...... ...... ... ........ .. .... .... ... ...... Gym nophiona .... .... ............ .... ............ ...................................................................................... lchthyophi idae .... ... .. ... ... .. ..... .. .. ......... ,.. ............ .......................... ............... ........................
46 48 49 53 54 56 58 60 61 62 88 115 133 175 218 2 18
5
Contents
Reptilia (reptiles) ...................................................................................................................... 220 Squamata (scaly rcptilcs) ... .. .......................................................................................... ..... .. 220 Sauria (lizards) ... .................. .... ... ........... .... ...... ....... ......................... ..... ... ................... .. ..... . 22 1 Distinguishi ng features in lizards ...... ... .. .. ... .................. .. ... .. ......... ..... ... ......... .. .... .. ... ..... .. 222 Key to the famil ies or Bomean lizards .................. ... .... ... .. .... ........ ... .... .... .... ..................... 223 Agalu idae .... ................. ... ................................... .... .. ........... ........ ....................... ........... 224 Anguidae .......... ......... ...... .. ..... ............. ... ... .. ....................... .. ..... ....... .. ..... .. ......... ... .... ... . 250 Gekkonidac ................ ...... .............. ............. ............. ......... ....... .. .. ... .. .. ..... ... .................. 251 Scincidae ......................... .................................................... ...... ...... .... ...... .... .... ....... ..... 269 Varanidae .............. .............................. .......... ................................................................ 293 Serpcntes (snakes) ................ .... ... ......... ........ .... ...... .. .. .. .. ..................................................... 297 Distingui rung features in snakes ....... ...... ... ... ............... .. ........................................ ......... 298 Key to the rami lies of Borncan snakes ......... ...... .............................................................. 299 olubridae ..... .. .......... .. ...... ...... ... .. ... .... ............ .. .. ..... ...................... ............................... 300 Crotalidae ....... ..... .. ..................................... .. ................................................................. 373 Cylindrophiidae .. .. ...... ... .......... ... .. .... ........................ .. ................... ........ ... .... ......... ..... .. 380 Elapi dae ................... ...... ............. ........... ... ..... ... ................................. .. .... .. ...... ... .. .. ...... 382 Pythonidae .................................................................................................................... 389 Typhlopidae ................................................................................................ ... .... ........... 392 Xenopeltidae .............. ... ........ ... ......... ....... ...................... .................. .. ........................... 393 Testudioes (turtles, tortoises, and terrapins) .. .................................................................. 395 Distinguishing fea tures in turtles ............... .. ........... .............................................................. 395 Key to the fami lies of Bornean turtles ......... .. .... ............. ... ............ ......... .......... ... ... ....... ... .... 395 Bataguridac .... ..... ..... ...... ..... ...... ..... .... .............. .. .. ................................................ ............. 396 Trionychidae ........... .... ............ .. .......... ..... .... .... ........ .. ........... ... ............. .... ..... .... ... .. .... ... .... 399
Glossary .......................................................................................................... ............................. 401 Bibliography .......................................................... ... .......................................................... ....... 403
Index ....... ...................................................................................................................................... 417
6
Preface
Preface ompared with previous times, the substantially improved possibilities for travelling have enabled an increasingly large number of people to reach other countries or even distant continents. Amongst these are a number who, be it at home or outdoors, spend their spare time wiLh intense studies of certain group of animals and/or plants. mall by comparison are the numbers of those who are able to study as a profession subjects such as behaviour, systematics and evolution, or the geographical distribution and eco logy of animal and plants. For many of these, and other nature lovers, journeys to poorly known zoogeographical zones will only be ensiblc if they are able to recognise at lea t a portion of the biological species living there. nabl ing this is one ofthc main goa ls of this book. Thc author have dealt in the main chapters of their treati e with all species of amphibian and rcptiles currently known to exist on Mount Kinabalu. Rising to 4,095 m Mount Kinabalu i the highest mountain between the Himalayas and the alpine mountain ranges of ew Guinea. It i situated in the north of the tropical island of Borneo, in the Malayan state of Sabah, and is marked by a variety of ecologica l conditions which, in turn, have created an extraordinary diversity of species. The visitor from central Europe will be su rprised to learn that the Kinabalu massif is home to 77 species of amphibians and J 12 species of reptiles, which equa ls the species diversity of tile entire European continent. The prescnt book is by no mean a compilation of existing litcrature publishcd by other author . Rather all authors except one have becn to the Kinabalu region themselves and have partly worked there for severa l months. They have therefore been able to combine the results of their own studies with an in-depth analysis of the literature to create a treati e original in character. The more than 30 pages of introductory chapters contain valuable information that may be interesting not only to the herpetologist. The attentive reader will be particularly pleased to find :
• The thorough analysis of the relevant primary literature with synonymy lists of scientific names,
•
something which was, unfortunately, insufficiently attended to in many books of a simi lar nature published during the past decades; A wealth of information , not yet otherw ise published, on the behaviour and eco logy of the individual pecies; Identification key for all herpetological taxa known to occur in the Kinabalu region; Excellent colour photographs of live representatives of almost all species; o cillograms and audiospcctrograms of the calls of many arnphibians .
In order to make the book acees ibJe to an international readership, it has not been published in the original language of the authors, but in Engli sh. "Amph ibians and Reptiles of Mount Kinabalu" places a tool in the hands of amateur re earchers as well as professional herpetologists which will not only eliminatc many identification problems in the field, but which can also be used for the identification of photographs and preserved material or that ofrecorded amphibian calls. I am convinced that, de pite the recent publication of a considerable number of checklists and some books more orientated towards the general public on the amphibians and rcptilcs of this region, this book will fwd ajustificd place in the libraries of many nature lovers. Those interested in the herpetofauna of Southcast Asia will receive it with particular appreciation.
Berlin, January 2002 Dr. sc. Rainer Gunther Musewn rur Naturkunde der Humboldt-Universitiit zu Berlin
7
Acknowledgements
Acknowledgements ------~---
We wi h to acknowledge the belp ofa number of people during the preparatory tages of this book. Many people contributed infonnation, technical assistance and organisatorial help. Many thanks to all of you: Markus Arning, John H. Beaman, Wallcr Brodt, JuJian Dring, Helmut Fleil3ner, Uwe Fritz, Klaus Kittel. Klaus Kuhn, Petcr Lenk, Nicola Lutzmann. Bcmhard Malkmus. Mark-Oliver Rodel , Andreas Schliiter, and Barbara Sein. We arc grateful to Kraig Adler. Carsten BrUhl, han Chew Lun, Tndraneil Das. Wolfgang Denzer, Thomas Eltz, Ronald Gerhardt, Armin Gotzke, Wolfgang Grossmann, Tsutomu Hikida, Walter Hodl , David Knowles Ulrich Kuch, Michael Kunkcl, Bernd Leideritz, Karl E. Linsenmair, Walter Malkmus, Sibylle Manthey, Manfred Maronde, Masafumi Matsui, John . Murphy, Andreas Nollert, Martin Pfeiffer, Horst Sauer, Martin Schroth, Robert B. Stuebing, Thomas Ulber, Michael Veith, and Steve Wilson for the permission to use tbeir photographs. Several of ollr colleagues reviewed one or morc sections of this book. In palticular, we wish to thank Rafe Brown, lndraneil Das, Patrick David, Wolfgang Denzer, and Jimmy McGuire. We also acknowledge the help of Patrick David. Robert F. Inger and Vall Wallach regarding taxonomical problems. To cllratorial staff of the following institutions we owe a debt for loaning us specimens under their care: Museum fur Naturkunde der Humboldt-Universitat zu Berlin (ZMB), Berlin (Rainer Gunther and DeUef Langer); British Muscum of National History (BM), London (E. icholas Arnold and Colin McCarthy); Field Museum of atural History (FMNH), Chicago (Harold K. Vori and Alan Resetar); Zoologisches Forschungsinstitut und Museum A. Koenig (ZFMK). Bonn (Wolfgang Bohme and Wolfgang Bischof1); Forschungsinstitut Senckenberg (SMF), Frankfurt/M (Gunther Kohler); Instituut voor Taxonomisehe Zoologie (ZMA), Amstcrdam (L. van Tuijl); Museum alional d'Histoire Naturelle (MNH ), Paris (Annemarie Ohler); The Raffles Museum of Biodiversity Research, National University of Singapore (USDZ/ZRC), Singapore (Kclvin Lim, Peter Ng and C.M. Yang); Kyoto University Zoological Collection (KUZ), Kyoto (Masafumi Matsui. Tom lJikida and Hidetoshi Ota (Okinawa»; Nationaal Natuurhi torisch Museum (RM H), Lciden (MarinLis S. Hoogmoed and RA.J. in den Bosch). For the preparation of drawings we would like to thank Vera Heinrich, Jan Ilnizdo, and Petr Kliment. Maklalin Lakim, francis Liew. Klaus Riede, Anna Wong, and Paul Yambun made our stay at Kinabalu Park very pleasant and provided help in many ways. We arc indebted to Rainer Giinther (ZMB) for presenting the preface. Finally our heartfelt thanks to our families for their patience and support in many ways. Last but not the least, we would like to thank Sven Koelt7, our publi her, for keeping a positIve attitude towards tbis publication during all stages of it production.
8
•
How to use this book
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How to use this book
This book is a comprehensive guide to the repti les and amphibians occurring on Mt. Kinabalu treating all species rccorded from the area and including all related taxa known to occur on Borneo. Tnclusion of all taxa serves the purpose to identiry also species, which are not yet recordcd from Mt. Kinabalu. EBENI-IARD & SJOOREN (1984) Ii led three species of snakes for Mt. Kinabalu, omitted here, viz.: Calamaria lowi BOULENOER, 1887, Ptyas carinatLls (GONTIIER, 1858) aJld Cerberus Iynchops (SCHNEIDER, 1799). Since no sources were provided for these records, the three species have been ignored in the present work. DAS & [ MAIL (2001) listed additional lizard species [or Mt. Kinabalu that have not been included here as the compilation is still under revision (pel's. corum. DAS). After a general introduction to the orders of amphibians (Anura, Gymnophiona) and reptiles (Squamata Testudines) occurring on Mt. Kinabalu supported by numerous illustrations, explanation of terms, formulae, and analyses individual families are discussed in detail. Taxa are introduced alphabetically irrespective of any taxonomical or phylogenetical relationship. With regard to illustrations preference was given to eolOlu' photographs that were taken on- ite in the area of Me Kinabalu, even if this meant to accept an inferior photographic quality. While the photographs primarily serve to illustratc the general appearance of a species, additional drawings are provided for specific features (e.g. oral fields in tadpoles, details of pholidosis in reptiles). Genera arc differentiated via dichotome keys based solely on morphological criteria. Additional data useful for unambiguous identification can be found in the species account. Discussion of an individual genu includes a table of distribution for all Bornean species and a key to these spec ies. AJ I taxa recorded from Mt. Kinabalu so far arc given in boldface. Species accounts are standardized providing the following information: thc valid scientific name, reference ofthe original description including the locus typicus (type locality; origin ofthe type specimen), a chresonymy list (as of end of 2000), a diagnosis, description of extcrnal features as well as ecological and ethological observations, and distribution . Scientific denomination follows trinomial classification. The basic unit of zoo logical classification is the species. Its scientific name is composed of two names (binomia1), that of a subspecies of three name (trinomial). Thc first name is the generic name, the second the species name, and the possible third the subspecies name. The taxonomic concept of subspecies is subject to controversial debates and we have thercfore linlited information on subspecies to unambiguous characters.
Synonymy/Chresonymy lists are limited to those references, which refer to an occurrence of the pecies on Mt. Kinabalu. Species munes and relevant synonymies are given with the respective authors' names, year of the publication, and page number(s); oompletc references can be found in the bibliography. Diagnosis is a compilation of generally valid data and serves to distinguish a taxon from other taxa on generic, specific or sub specific level. In case of a monotypical genus, the relevant taxon is differentiated from similar Bornean taxa. Description is based on own examinations and/or the standard publications for the region, e. g. DE RoolJ ( 1915 , 1917), INGER (1966), and MANTfl EY & GROSSMANN (1997). A descriptive summary of adult extemal features is accompanied by data on ontogenetic developmcnt or developmental stages where appropriate (c. g. amphibian larvae) . Measurements relate to standardized methods described in thc introductory chapters and are given in rnillimetrcs.
9
How to u e this book
Ecology/Ethology summarizes data on general behaviour, habitat preferences, vertical distribution. population density, feeding, predators and antipredator behaviour. It hould be noted that there is a painful lack of ecological and ethological information on many of the specie . Distribution compi les the entire geographical distribution of a species as well as its distribution on Borneo (for localities see map on page 11). Distribution maps are limited to those species for which a larger number of loeaJjties are known. Locality data relate to Our own studies, those taken from the literature are marked differently. Mt.lGnabalu contains listing of all known localities on Mt. Kinabalu (> 500 m a.s.l.) where a taxon has been recorded. Apart from a few unidentified placcs, all localities are shown on the maps on page 200, 203 and 204. Their spellings are very much inconsi tent in commonly u ed maps, and we largely follow the pellings u ed on a draft map kindly made avai lab le to us by BEAMAN. It is unfortunate that older record u ually do not include altitudinal information. 1n many ca e the altitude of the actual collecting site i likely to vary substantially from that of the stated locality as only thc nearest known location is given in publications. pecie detennination of specimen hou ed in lhe col/eclion ofKinabalu Park (HQ-Co/leclion) has been accepted without nlrther verification. Literature u ed in tbe introductory chapters and further reading are marked by an asterisk (*) in the bibliography section of this book. Errata: Fig. 94 on page 101 does not depict a member of the genus Lepto/a/ax but rather a ranid. The below given descriptions were received too late for inclusion in the text of this book.
Phi/aulliS erYlhrophthalm u STUEBING & WO~G, 2000, type locality Mt. Muruk Mio, 1550 m asl , Sipitang District, Sabah (4° 22.715' , 115° 50. 125 ' £) is a specie of the vermiculatlls-group, resembling P. aurantium. It differs from that pecies in having no dark pots neither dorsally nor ventrally and reduced webbing reaching only the mid- ubarticular tubercle of the fourth toe. An onia anotis I NGER, TAN & YAMS '.2001, type locality g. Wario (880 m asl), Sayap Station, Kinabalu Park, Kota Belud District, abah (6° 14'N/ I1 6°32'E) is the only Bomean species of Ansonia without a vi ible tympanum. Conical tubercle on back almost homogeneou in size; first finger do not reach wollen tip of econd; tips of outer fingers expanded into distinct, patulate discs; web extending beyond distal subarticular tubercles of third and fifth toes. SVL of one female 52 mm, of one male 37 mm. Sphenonlorphus aesclileticoia u..GER, T LAKJM & Y AMBtN, 2001, type locality Mt. Lamaku (1350 m), 14.6 km from Mendolong, Sipitang District, Sabah (4°52'NIl15°3 E) is a short- legged, small ( VL < 45 nun) species with 4 supraoculars, 6-10 subdigitals (fourth toc) and u ually 28-30 scales around midbody; prefrontals do not meet in the midline; with dark lateral band. pecimens from Mesilau wa provisionally referred to this species. Sphenomolplltls crassa INGER, TAN, LAKlM & YAMBUN, 200 I, type locality Mendolong (670 m as I), ipitang Di trict Sabah (4°54'N/ I15°42'£) is a short-legged, large (holotype VL 82 mm) species with 4 upraoculars, 18 subdigitals (fourth toe) and 32 scales around mid-body; prefrontals widel eparated; no dark lateral band. Sphenomorphlls tanahtinggi INGER, TA , LAKJM & YAMBUN, 2001, type locality Mt. Lamaku (11 0 m asl) ipitang District, Sabab (4°52' 1115°38'E) is a long-legged peeies (SVL max. 64 mm) with 5 supraoculars. 16-17 subdigita ls (fourth toe) and 40-42 scales around mid-body' prefrontals narrowly in contact or eparated: with dark lateral band.
10
rntroducti on
Introduction Covering an area of739 204 square ki lometres, the biggest of the Sunda ls lands, Borneo lies in the central tropics. As far as size is concerned, it ranks tbird among the largest islands of the world, after Greenland and New Guinea. It has more than a thousand years of varied history, during which Indian , Chinese and - from the 16th century onwards - European tradesmen and colonial powers strived for territory, resources, and commodities. It was not before the end of World War IJ tllat the territories of Borneo became independent.
.. J11 1949, the formerly D utcb part of Borneo become Indonesian under the name of Kalimantan (535 834 km2 ) • .. In J963, tho formerly British northern part of Borneo joined the Malaysian Federaey; the provinces Sabah (73 620 kml) and Sarawak 123 985 km') were established. >I< In 1984, the forolerly British protectorate Brunei (5 765 km') gained independence. The economic dominance Brunei exerted in the region lead Europcan to using its name in gradual linguistic distortiolls (Brunei ~ Barllney~ BOnloo) to Signify the whole island. This name is still in corrunon uso nowadays. Gunung (= MOLlnt) Kinabalu, 4095 m, is the highest sUl11mjt of the entire Malay Archipe lago and is sitllated in the far north of Borneo, in the province of Sa bah. Apart from the summit of Makital Kal imantan (2987 m) it towers more than 1500 m above all the other peaks of the island. In former times indigenous people regarded the mOllntain as a place where they expected the soul of their deceased to be gathered in a village of the dead (Aki Nabalu), which was strictly off limits. From the mid-19th century onwards, the mowltain has been radiating a virtually magic attraction for botanists and zoologists due to its unrivalled diversity of species. ALmost instantly after having been incorporated into the Malaysian territory, it gruned the status of a protected Park, and even the capital ofSabab, tbe former Jessclton, derive its name from the great peak: Kota Kinabalu. For tourists it bas been an unrivalled attraction from the very beginning:
12
during the first decade after its first ascent, only 53 people climbed the mountain, whilst nowadays tcns of thousand annually follow their tracks. In pite of this mass rush, the mountain has not 10 t a bit of its aura. obody manages to elude the magic and majesty radiated by the granite giant emerging from the rain forest. The elevated language used in films and leaflets to entice the tourist to abah, the land of tbe " Sacred Mountain" , and to evoke the paradisaical wonders which await the visitor shows quite unmistakingly: the tonne!" rea lm of the dead has undergone a m tamorphosis to a national sanctuary. in biological terms, Mt. Kmabalu has remained what it Llsed to be in the last century: a great centre of biologicaJ diversity which is tillparalleled - not only in the Sunda area, but also worldwide.
Mount Kinabalu - the surrounding environment
Mount Kinabalu the surrounding environment In order to understand the distribution of specific species at any place n the earth, it is necessary to bave a clear idea about the fundamental abiotic and biotic conditiollS of the respective area under examination. The valid criteria are: * the geological and geomorphological condition s >I< the zoogeographical circumstances >I< the reccnt climatic conditions >I< the trucmre of the vegetation cover * the anthropogcncous impact on the environment The following chapters try to delineate an overview of thcse basic conditions for the area under inspection .
Topography, Geology, Geomorphology Mount Kinabalu, with 4095 m the highest mountain between the Himalayas and ew Guinea, is located at the northern tip of Borneo (5°57'-6° 18' , 11 6°28'-
I 16°45 'E). The central ~inabalu massif forms the shape of a horseshoe open to tbe nortb. Tbe uninterrupted rocky cJjJf facing south sends two ridges to tbe north separated by the J 500 m deep Low's Gully (the western ridge already descends rapidly after 2 kilometres, while the eastern one stretcbes far to tbe north, reaches once more all altitude of2576 m in the summit of Tambuyukon and descends in the range of Mt. Templar-Mandalon) and continues to the east down to Poring jointed into variolls lower ridge to the north and to the south. The massif is one ofthe most impoltant water catchments in abab and the source of many streams (e.g. Sg. Kinaram and 1\mran flow to the north, Wariu , Panataran, and Kadamaian to the west, Liwago, Mesiiau, Luidan, Langanan, Kinapassan, and Kapuakan to the east). Together with their in part temporary, tributaries and crossing through all altitudes they represent just about every type of habitat one may expect ill tropical montane running waters.
fig.] ML Kin
13
Mount Kinabalu - the surrounding enviromnent
Fig. 2 Rock arca polished by ice during thc last glacial, with nunataks in the background {northern plaLeau of Mt. Kinabalu .
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The area examined comprises in principle the entire mountain massif above 500 m above sea-level, but, for reasons of inaccessibility, i largely restricted to localities around Poring, Headquarters (inclusive of the Summit Trail), Mesilau, ayap, Kiau, and Bundu Tuhan. About 80% of the area lie within the boundaries of the Kinabalu Park. The crystalline basemcnt of northern Borneo is covered with a thick sequence of marine sediments which developed in the Tertiary pcriod. During the middle of the late Miocene period (somc 15 million years ago), these layers of shalc, siltstone, and sandstone were folded and uplifted to the mountain rangcs of the rockers and Trusmadi. During the Pliocene period (some 10 to 5 million years ago), a ubvolcanic intrus ion was exposed uplifting the overlying layers at the northern end of the rockers. The central swnmit plateau - today called Mount Kioabalu - consist of a granitic ceotre (hornblende adamellite), covering a surface of 150 km2, and an elevation of2000 and 3000 ill, surrounded with a ring of porphyritic and ultrabasic rocks (serpentine). These are followed
14
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by tertiary edimcnls at lower elevations. On the so utheastcrn slopes (Pinosuk Plateau) up to 140 L11 thick Pleistocene layers of gravels are depo ited. During the last Pleistocene period, the swnmit region was covered by an icc-cap, about 5 km' in extent, reaching down to about 3000 m. Tllis glaciation has left distinctive landfonns not only in the summit area (polished and plucked su rfaces by the erosional action of the ice 00 thc plateau, nunataks, U- haped gullies, moraine material with subangular boulders at the end of such gullies and near Pakka Cave, cirques on the summ it), but the geomorphology of the lower lopes, situated in the periglacial zone, was tamped likewise (deposits by blockstream and mudflows on Pinosuk, forms of valleys and canyon ). The agents of the receol, chiefly chemical, weathering under tropical conditions (high temperatures and humidity) can be studied on the denuded summit area (exfoliation of slabs of rocks) and especially in regions deforested by man (partially up to 1800 m) with considerable soil erosion.
Mount Kinabalu - the surrounding environment
Zoogeography Together with the islands of Palawan, Sumatra, Java, and the Malay Peninsula, Borneo forms part of the so-called Sunda Plate, a southeastern spur of the Asian continental shelf. Between Borneo and mainland Southeast Asia, large parts of the Sunda Plate were exposed for much ofthe Tertiary. It was not before the Pliocene period that a shallow shelf sea separated the continent from the Sunda Islands. The Quaternary then was a period of repeated alternation of colder (glacial) and warmer (interglacial) spaces oftime. During the glacials a substantial portion of the water was bound terrestrially as glacial ice causing eustatic sea level changes worldwide (> 100 m max.). Under these conditions large areas of land of the Sunda Shelf were re-exposed and connected Borneo with the Asian continent. Distinct indications of this intermittent Quaternary falling-dry are given by submarine channels. They indicate the courses of former rivcrs at times when the platfonn was above sea level. Corresponding to the alternation of colder 311d wanner periods, these land connections were formed and broken several times. Fig. m The area of Sundaland (surrounded by a broken line) exposed at times of lowest sea Icvel (dark shade) during the last glacial with ancient river system (submarine channels).
Situated east of the unda platform, and reaching depths of several thousand metres in places, the Makassar Straits separates Borneo and Sulawesi. The latest connection between these two islands existed in the middle Eocene, more than 40 million years ago. During the Tertiary, arcs of volcanic islands were important as potential land bridges and island hopping routes for organisms to and from the Philippines, Borneo, and Sulawesi . The climate of the Sunda region during the glacials was probably 2-4°C cooler and more arid than it i today wIllie the interglacials had a wann and humid climate. These paleo-climatogeological conditions manifcst themselves in the existing zoogeographical situations. They help us to explain many disjunct plant and animal distribution patterns and the composition of species in the different geographical regions. II is significant that a major zoogeographical bOLmdary scparates Borneo and Sulawesi: the Wallace's Line. The islands west of this boundary - thus including Bomeo - form part of the Oriental Region, those to the east 'of it belong to the WalLacea, a transitional region between Orientalis and Australis. The relationsillp of the herpetofauna of Bomeo is signi ficantly closer to the Asian mainland and other Sunda islands than to Sulawesi despite its shorter geographical distance to the latter. This is proven by facts such as 70% of the snake and 43% of thc lizard species of Bomeo also occurring on the Malay Peninsula whereas only 25 and 19%, respectively, exist also east of the Wallace's Line. Nearly all non-endemic species of Bornean frogs (ca. 35%) are also found on thc mainland , but only 7% also on Sulawesi (of which 70% are fTOgS settling in cultivated areas which are very likcly to have been introduced by man). The large diversity of species of amphibians and repti les on Borneo in general and on Mt. Kinabalu in particular can be based for a large part on the following factors: • Most of the species of animals and plants show a tendency to increase their diversity the nearer to the equator they OCClLr. Borneo is situatcd in the central tropics within the species-rich Oriental Region . The amphibians and reptiles native to this region furthermore include numerous genera which are known for their rapid
15
Mount Kinabalu - the sun-ounding environment
speciation (e.g. Ansonia, Meristogenys, Phi/auILlS, Calamaria, Sphenomorphus). • Tbe repeated connection and separation of Borneo to, respectively from, the mainland during the Quaternary favoured an exchange of faunae in one situation and speciation processes in the other resulting in ca. 25% of the snakes, 45% of the lizards, and 67% of the frogs being endemic to the island. If the hypothe is presented by BRAl'
16
The geological age ofMt. Kinabalu is high enougb to enable many genera to have reached considerable degrees of differentiation through adaptive radiations manifesting themselves in the f01111 of localized endemites which bave adapted to certain ecological niches within certain altitudinal zones (e.g . Ansonia gllibei, Ansonia ju/iginea, Kalophrynus baluel1sis, Leptobrachium gUllungense, Phi/au/us saueri, Lepto/a/ax maurus, Calamaria schmid/i. etc.). Recognizing these species as localized endemites, however, requires a considerable amount or caution duc to the fact that our knowledge of the distribution of the animals is still very limited. As the average temperatures were several degrees lower than today during the Pleistocene, some oftbe forms classified as localized endemites may turn out to be in fact glacial relict which previously inhabited also lower e.levations, but were then forced to retrea1 to higher altitudes when the temperatures rose . Today they occur only in montane isolates scattered widely over the entire islands.
Climate
. -------
- 111C MfII"ei Pclt'ei spur ofrered a fil1e pO>ili0l1 for a Scll1Jlarilltll, at any height belween 4000 ft <mel 000 ft. Our letll WdS pitched at IIbout 4700 ft, ,111(1 we found thell the lhennom ley IYlftrked 75' In Ule micl·delY sh.lde, 56 ilL "IX
rencer Sl John , l.ire ill the I'orests ortlle l'elY Fil.,t. 1862
Borneo maintains a tropical rain forest climate up to altitudes of above 1000 metres. It is characterized by an annual precipitation in excess 0[2000 mm, no dry periods for several months, day temperature of +27°C, +22-23°C at night. and a relative bumidity ofusual\y above 90%, and thus lies in the range of the monsoon weather pattern of tropical A ia. he constantly pre ent equatorial low pres urc area dominating the Indo-Pacific region, known as the rnter-Tropical Convergence Zone, shi fts by following the position of the sun to the south in winter and to the north in summer, triggering the north-
Mount Kinabalu - the surrounding environment
1--
I
[mm)
700 -
----------
II 600 n
1987
200
L
- - 1988
I~ I I- 400 I 300 I 500
-~--------
1989
l
1990
1 I
I I
J
I 1
February January March 1Fig.
rv
April
June May
July
August October December September November
Monthly precipitation at the Headquarters of Mt. Kinabalu in the years 1987-1990 (after
east monsoon (November/ Deeember through March) and the southwe t mon oon (June through eptember), respectively, with their heavy zenith rains. Substantial precipitation is, however, observed throughout the year in all parts of Borneo o that the monsoon merely cau e it to intensify in certain seasons. Notwithstanding this situation, extended dry periods are observed aperiodically with monthly precipitations of less than 30 mm. yclic changes in the atmospheric circulation (amplification of the southeast passats) in the form of the so-called EI ino Southcrn Oscillation often result in monthslong dry periods at irregular intervals (e .g. in 1977, 1979, 1982/83, 1992, 1997/98) exposing the ecosystem rain forest to sub tantia! stress and favouring catastrophic forest fires . For example, in 1972173 a prolonged drought distllrbed all woody plants of the upper montane forest in orne areas with shallow soils on Mt. Kinabalu.
Mi\I~
J996b) ]
The general climate may be altered significantly in places due to the individual topography. Outside prolonged rainy or dry period , a very characteristic daily weather pattern can be ob erved on Mt. Kinabalu : The first hours of daylight are marked by a clear or minorly overcast kyo Depending on the intensity of sunlight and wind, humidity drop from 90-95% to 40-70% out ide the fore t and in the crown of the trees, while it oscillates very little within the forest. Air saturatcd with moisture ascends the slopes of the mountain following its relief. As the temperature drop by an average of 0.55 to 0.65°C per 100 m incrcased altitude, the cooling watcr vapour condenses. The condensation level during the morning hours nonnally lies between 2000 and 3000 metres. This results in the formation of a ring of fog and cloud up to 3000 m altitude, covering the cntire mountain down to about [500 m during the course of the afiernoon. Heavy rain showers,
17
Mount Kinabalu - the surrounding environment
slopes so that insolation and wind exposition are often in the fonn of thunder stonns, usually et in highly variable factors. For example, entirely disfrom between 2.00 and 4.00 p.m. often lasting up simi lar climatic conditions may be found when into the early night hours (i .e. 7.00 to 9.00 p.m.). While the annua l pattern of the temperatw'es is relatively balanced, pronounced oscillaFig. V Oscillations of temperature at various altitude on tions are discernib le during the cow:se of a Mt. Kinabalu (measured in August/September 1986). day. They are least apparent in the cloudy modified after K ORS liN ER moss forest and most distinct above the tree 1990 line. Corresponding oscillations are found in Co I I I the relative humidity. The summit region is I l'-,\ known to experience occasional snow falls ,.",/ \ 30 and frost at night. ! \-50m II • The visitor from central Europe rinds a I I SepilokiSandakan rather agreeable climate between 1400 and SOO m! 1600 m which was appraised already by 20 the pioneer explorers of Mt. Kinabalu in the 19th century (see quote at the beginI ning of this chapter). This climatic ituaI I KamborangQh 2500 m I tion is subjeet to numerous localized aber10 rations . lsotherms and isohyets show no parallel alignment anywhere on the mountain as a result of the very differentiated structure of the relief (in the form of gorges, canyons, outcrops, edges, and ridges) and 12 14 16 18 20 22 24 2 4 6 8 10 12 14 16 18 hrs the varying exposition and inclines of the
- -0--:---11(;
l8
;--0---
Mount Kinabalu - the surrounding environment
comparing an exposed ridge with a stream that flows in a protected gorge, even if both are situated at the same elevation and only a few hundred metres apart. It is signific31lt that it is exactly these gorges which represent the upper altitudinal distribution limit of many spccies offTOgS. The topograpbically induced network ofmicroclimatic niches experiences another fine-tuning of its differentiation by the vegetation cover. Below 3200 m there is only one natural type of plant [ormation on Mt. Kinabalu, i.c. forest. Depending On the heigbt of its crown canopy, significant aberrations of the individual climatic components may be observed within vertical distances of as little as 10 to 60 metres . The climate immediately above the leaf litter up to about 2 m is marked by being very constant as far as temperatures are concerned (daily variation usually less than SoC). Due to the ineffectivcness of air currents, the ground level is constantly covered by ai.r which is saturated with moisture; the relative humidity thus measu.res more than 95 % most of the day, with a minimum of 80-90% between 12.00 a.m. and 3.00 p.m . It is only when the dry winds accompanying the EI Nino weather phenomenon take cffect that the hu.rnidity may decrease to less than 60% for prolonged periods of time. The partly many-tiered erown canopy blocks out the sun so that merely I to 3% ofthe sunJight reach the forest floor although the intensity may vary
locally during the cou.rse of the day due to the shifting of sun-exposed patches. Entirely different climatic conditions are found at the level of tbe crowns si nce they are highly exposed to wind and insolation. The result is that individual climatic components are subject to substantial oscillations during the course ofa day, with temperatures varying by more th311 lOoC, and humidity by 50 to 60%. Between the floor and the crown level, the climatic components experience alterations of all kinds, so that it is just to speak of a vertical tiering of climates according to the tiering of the rain forest itself, in other words a "storey climate". Climatic peculiarities, lastly, result ii-om tcmporary gaps in the forest caused by wind storms and slides of the soil, and in the immediate vicinity ofstreruns. With amphibians and reptiles being poikilothermic aninlals. their activity is limited to a relatively narrow bandwidth of climatic preconditions. The knowledge oftbc many-faceted climatic conditions in their biotopes is therefore an essential tool for the understanding of their often insular niche segregation in a larger biosphere, their activity rhythms, and numerous other adaptive strategies. Figure VI demonstrates this principle clearly by showing calling activities offTOgS on the same site during various weather conditions, while fig. VIIT on page 26 shows the influence of the vertical distribution.
Fig. VJ Calling activity of frog species on Sg. Liwago (1450 m) in August 1990. A - afler one week of drought (reI. humidity < 60%). B - after two hours of lbundersloml between 3.00 and 5.00 p.m. (reI. humidity > 95%). no. of species
no. of species
9 8
9
A
8
7
7
6
6 5
5 4
B
4 3 2
3 2
1
3.00
1•
4 .00
5 .00
stream -associates
6.00
7.00 •
8 .00 p.m.
3.00
shrub and tree dwellers off from streams
4.00
5.00
6.00
7.00
8.00p.m
ground dwellers off from streams
19
Mount Kinabalu - the surrounding environment
Vegetation 'The r 'ople of abab po~se on this famous mOlmlain what I believe i richesl aJ'ld most remark.lblc assembldge ofpldl'lts in the world." LJ.H Corner. 1978
nw
With the exception of the slopes ofLbe Kadamaian valley, the fianks ofMt. Kinabalu above 500 mused to be covered by virgin rain forest wltil as recently a the early 20th century. The e montane forests are home to the largest variety of plant species in the entire Malcsian region as the perma-l11oist Southeast Asian tropics are referred to in the cience of phytogeography. Besides hundreds of upper montane, subalpine and alpine endemisms (notably neoendemi ms in the vegetation facies on ultramafic ubstrate), it includes truly spectacular plants such as Ra.Ulesia which parasites on vines (Tetrastigma), sixteen species of carnivorous pitcher plants (Nepenth es), twenty-five specie of Rhododendron. and not less than 610 species of ferns, 350 of mosses, and 750 of orchids. BEAMA I\ & B F,AMAI\ (1990) estimated some 4000 species of va cular plants to occur on Mt. Kinabalu .
Fig. 4 Flower of RaJJlesia pricei. Mamllt Valley. On Mt. Kinabalu Ra,/jlesia is common round the Mamllt Valley at 900-1500 m a.s.1.
20
Every visitor to Lbe mountain cannot but note how the physiognomy of !.hc fore t change the higher he ascends. The mountain i virtually covered by stacked rings of vegetation zones with each having a specific composition of species. These altitudinal vegetation zone are the re ponse by the flora to the altitudinally changing climatic and edaphie conditions. otwithstanding this there is no strict demarcation between one zone and the subsequent one, but a gradual transition. While Indo-Malayan taxa dominate the lower elevations, the moderate altitudes are home to mainly East Asian ones, and AustralAntarctic floral elements reign above 2800 m.
Fig. 5 Coelogyne ragio/erens. a beautiful orchid .
Mount Kinabalu - the surroundjng environment
Tropical lowland vegetation zone (to 900 m a.s.l.)
Climate: Levelled, moist-warm ; average annual temperature +26-27°C; precipitation throughout the year (2000-3000 mm). Soil: Yellow latosol with a thin layer of humus in the form of mould or mould-like compost as a result of a rapid rotting and mineralization of leaf litter. Vegetation: Multi-layered dipterocarp rain forest Upper storey: 30-50 m high tree giants (Shorea. DiplerOCatpus. Hopea, Vatica), moderately evident epiphytism. Middle storey: Dense, closed crown canopy 20 m above the ground (Annonaceae Meliaceae Myrtaeeae, Sapotaeeae), bamboo, lianas. Bottom storey: Lacunate, 8-10 m high underbrush; vines, bamboo and rattan thickets; plank roots; the floor is covered with a closed layer of lea flitter.
Water: Mountain streams with moderate to trong currents and occasional cascades; pronounced tidal changes relative to the weather. Structures: stream bed sediments dominated by coarse gravel, forming an extensive scattered pattern (hyporheic interstitial area); sand and gravel banks; rocks 3Jld fallen logs which protnlde above the water surface; pernlanently or temporarily connected rocky pools with fine sediment and deposits of leaf litter. Accounting for the strong CWTent and shade (usually only 2-5% of the sun light reach the water surface), no higher aquatic plants and fewalgae can develop. The water of these mountain streams is in general very rich in oxygen, poor in dissociating salts, alkalines, and acids as a result of the filter perfoLlnance of the fW1gai plexus which is associated with the root systems of tbe trees; pH 6 to 7; water temperatures rather stable (+ 17 to 20°e). Ephemeral rain pools or puddles and phytothelmes (water bodies fit for vegetation): limited to cavities in trunk or limbs oftrees, tree buttresses and )janas, as well as bamboo internodes. fUllnels in plank roots, and in the axjls ofleaves (e.g. Pandanus).
21
Mount Kinabalu - the surrounding environment
Tropical lower montane vegetation zone (900-1800 m a.s.l.)
Climate: Levelled, moderately moist-waml; average annual temperature + 18-24 ° ; precipitation throughout the year (> 2500 mm).
22
Soil: Micropodzolic soils with a layer of raw humus who e thickne increases with increa ing altitude . Vegetation 900-1300 m: Tran ilion from ubmontanc dipterocarp forest to lower mixed Fagacea forest (30-35 m). 1300- 1800 m: Mixed Fagacca fore t (oak-chestnut forest) . Reduction of the crown height to 15-18 m; very large variety of species (Lauraceae, Theaceae, Myrtaceae, Elaeocarpaceae), first conifcrs (Da cry dium , Podocarpus, Agathis); the reduced height of the canopy permits a higher inten ity of light on the floor resulting in a thicker underbrush. Phoropbyte (carrier trees for epi phytes) are thickly covered with mosses, ferns, and orchid ; the first cutancou ferns (Hymellophyl/um) and terrestrial mosses appear pointing towards the presence of cli matic element typical fo r mos y forests . Water: Usually fas tflowing, ca cadeous. cold (+ 14 to l7 0 C) mountain tream with the structural traits of submon tane stream . The banking vegetation includ es thickets of shrubs (Gunnera, impatiens. Begonia, Pito patha) and Rattan (Ca/af11u • Daemonorop ). Phytothc)mc : Dead pitchers of Nepen thes and leafaxils, while cavities in trees and Iianas are rare.
Mount Kinabalu - the surrounding environment
r Fig. VU
Daily oscillations of air and = l P e r = the confluence of Sg. Silatt-Silau and Sg. Liwago (ca. 1400 m a.s.l.) in August 1994 (after MALKM S 1996e).
Iem p (OC)
22
-
Airtemperalure along - - - Sg. Silau-Sllau. 1 m above the ground
\
\ 21
\20 19
[ Airtemperature. 50 m away - - - from Sg. Silau-Silau. 1 m above the ground Watertemperalure of Sg . Silau-Silau. - - - 20 em under watersurface '---
1
[ 18 17
[ 16
15
114 13
L
0
2
4
6
8
10
Tropical upper montane vegetation zone (1800-3000 m a.s.!.) Climate: Levelled, cool-moist; ave rage annual temperature + 10-\3° ; precipitation tllroughout the year (> 4000 mm); daily, long-persistent cloud layers; extremely high atmospheric humidity. Soil: Very acid brown earths, podzols and peaty gley soils. Vegetation: Cloudy moss forest. An 8-12 m high forest of conifers, Ericaceae, Myrtaccae, Fagaceae, tree fcrns (Cy athea) and Bambusa. with dense underbrush . The trees arc usually stunted and deeply packed with cpiphytes such as mosses (es-
12
14
16
18
20
22 lime hrs
J
peciaJly Bazzanis. Plagiochila. Mastigophora), liverworts (Lepidozia), lichens (Usnea), orchids (e.g. Dendrochilum, Coe/og)me, Bulbophyl/um), and club mosses (Lycopodiatae). Many specie absorb water from the fog ("fog-stripping"). Since thc plaster of mosses rcleases water only very lowly, the coating of the phorophytes never dries up completely. Wafting mists tum the stunted forest with its heavy load of epiphytes into a bizarre, spooky environment which accounts for its name "elfin forest" . While the mossy fore t rises up to 3000 m altitude on the southeas!em side, it is replaced along the Summit Trail between 2600 and 3000 m by ultrabasic rock ( erpentine) with highly demineralized, variously moist soil which can'ies a small-crown forest « 8 m)
23
Mount Kinabalu - the surrounding environment
Fig. 8 Usnea lichens C S. Manthey
Fig. 10 Nepenth es villosa M, Maronde
ci
o !!.g~1 ~ ~ossy
forest. along, the ummit trail.
with colonies of few icadjng pecie (Podocarpus. Daclydium, Myrica, Leptospermum) which is marked by a substantially reduced epiphytism. Water: Almost exclusively very rapidly flowing to
24
torrential streams with many cascades which often full dry during prolonged dry periods above 2500 m. Phytotheimes: Pitcher of Nepenthes arc very common; cavities in trees are rare.
Mount Kinabalu - the surrounding environment
Tropical subalpine vegetation zone (3000-3700 m a.s.l.) Climate: Distinct lluctuations in the temperatllre during the course of the day (often > 10° ); cool; average annual temperat1.1re +8-1 O°C; precipitation throughout the year (no data avai lable); extreme winds with chil ling and dtying effeet common; relative humidity valying between 20 and 100%. Soil: 3000-3200 m: A layer of acid brown podzolic soils and peaty podzois. 3200-3700 m: Accumulations ofpodzolic soil limited to eroded grooves and rifts between large granite boulders (skeleton soi ls). Vegetation: Low-growing « 5 m), very dense co lonies of hard-leafed plants (Leplospermum, SchifllG, Rhododendron, Myrica, Daphlliphyllum. Phylloclades, Podocarpus) which form the tree-line already ben.veen 3200 and 3400 m for edaphic reaons (the potential tree-line in the Indo-Australian region lies at 3700-3800 m, Mt. Wilhelm on New Guinea being an examp le). Species ofthe same gen-
era colon ize crevices and rock fissllres which are filled with humus up to the summit zone (4100 111 a.s.I.), scattered over an alpine rock desert (tropical alpine dwarf-scrub). Water: The water levels of the treams are directly related to the amount of precipitation; water-filled rock basi ns arc occasionally found. Phytothelmcs: Pitchers of Nepenthes villosa below 3200 111.
25
Mount Kinabalu - the urrounding environment
750
sSOO
,
~_'
10 8 6 4 2
altitude (m)
12&0
__-='~=
1750
altitude (m)
2250
~
2750
1750
750
2~___ ~~__
S5(\Q
2250
1..,..,_,~-=.=-..-. 2OQO
2750
~
15 Rhacophoridae
Bufonidae
10 5
0
o
10 8 6 4 2 0
3 2
Megophryidae
-
lchthyophiidae
o 50 40
7
Amphibians
30
Microhylidae
5
20
3
10 25 20 15 10 5 0
o (/)
Ranidae
Q)
endemic species to Mt. Kinabalu
1
·u Q)
c. (/)
Fig. VIII umber of amphibian species in the di fferent altitudinal vegetation zones on Mt. Kinabalu,
t (OC)
(/)
Q)
·u Q)
c. (/)
30
rain (mm) >5000 >5 OQ.. .......
25.8
6000
0
Q; .c
20
E ::J
c:
12
10 2075
500
2788
750
10
I
4000 9
2000
1000 1250 1_ 1750 2000 2250 2500 2750 Tropical vegetation zones on Mt. Ki nabalu
500-900 m lowland
900-1800m
1800-3000m
lower montane
upper montane
subalpine
Fig. VII I While average temperatures decrease continually with increasing altitude, frequency and quantity of precipitation incrcase. The altitudinal distribution of amphibians and reptiles is influenced to a significant extent by both factors: the diversity of species of repti les decreases dramatically with increasing altitude; although numbering fewer in species, amphibians are better adapted to lower temperatures and moisture and therefore extend their ranges a 1000 m higher into the montane regions.
26
'0
Q; .c
E ~
Mount Kinabalu - the surrounding environment
a ltitude (m)
2750
750 ~500
'%JlQI~_""'~_
2500
3000
1250 1750 2250 2750 1500 2000 2500
12
Agamidae
8 4
12
Gekkonidae
8
5
4
4
12
2
Scincidae
4
1 4
2
Varanidae
2
4 2
35
25 20 10
65 --~----------~~~r-~~--~~
60 Reptilia
50
40
•
Tropical vegetation zones on Mt. Kinabalu
500 - 900 m
lowland lower montane
30
upper montane
20 10
subalpine
(J)
Q)
'0 Q)
a. (J)
-
• endemic species to Mt. Kinabalu
o
Qj
.a
E ::l
c:
Fig. IX Nu mbcr of rept ile species in the different a lti tud ina l vegetation zones on Mt. KinabaJ u.
27
Mount Kinabalu - the surrounding ellvironment
Fa una "The firsl il'l1pres ' iol1 which one per ceives in the Lropical foresls is '-hfll almost no dl1imat life cal1 be foul1d: A.R. Wallace The Malay Al"chipelago, 1869
-.:
~ ~.
Froln top to base Fig. 14 Traglllll.l'jovolliclIs. the Smaller Mouse Deer and Calloscillrtl. preFo Iii (Fig. 15), the White-striped Tree Squirrel, appear to be common in the lowland zone. The Palm Civet ParadoxlIrliS hermaphrodillls (Fig. 16) was also ob erved up to 1800 In and higher.
28
The diversity of faunal species of the Kinabalu massifis in no way inferior to that of its flora. It is a pure forestfauna of the East Oriental region with a high content of endemisms. On the other hand tropical rain forests are characterized by a quantitatively unproportional ratio between pJant and animal biornasses with an average of 1000 tons of plant mass to only 200 kilograms of animal mass. Tbe fact that the number of animal pecies is notwith tanding substantially higher than that of plants ' is explained by the large diversity of invertebrates which amount to little in weight. Most of the species are furthermore rather rare so that a high species diversity (richness) goes along with a low population density (evenness). The spatial niche segregation ofthe individual species - a large percentage of the latter is confined to the canopy - is so perfectly adapted to the habitat that every observer who enters a tropical rain forest for the first time (and who would not do so with the expectation to encounter an unlimited species richness?) will sureJy be disappointed and agree with WALI. ACE'S statement quoted above. The following is an inventory of what has been recorded to date from Mt. Kinabalu: 110' species of mammals, e.g. Orang Utan (Pongo pygmaeus), Red Leaf Monkey (Presbyfis I'lIbicunda), Grey Gibbon (Rylobates moloch), Slow Lori (Nyclicebus coueang), Tarsier (Tarsius bafleGnus), Flying Lemur (Cynocephalus variegatus), Pangolin (Mallis javanica), Porcupine (Trichys lascieLllata), Malay Bear (llelarefos malayanus), Leopard at (Felis bengalensis), Mou e Deer (Tragulus sp.), Barking Deer (Munliacli sp.), 22 species of bat (Chiroptera), 26 squirre ls ($c iuridae), 7 tree hrews (T\.Ipaiidae), and 7 civets (Vivcrridae). . 65% of the mammal species endemic to BOJ11eo occur on Mt. Kinabalu.
Mount Kinabalu - the surrounding environment
306 species of birds, e.g. Great Argus Pheasant (Argusianus argus), 6 species of hornbill s (Bucerotidae), 11 woodpeckers (Picidae), 19 cockoos (Cuculidae),3 pittas (pittidae), 17 bulbuls (pycnonotidae). 8 broadbills (Eurylaimidae), 13 sunbirds and spiderhunters (Nectarinidae), 5 drongos (DicruFrom top to base Fig. J7 Argusial1l.1s argus. a common lowland residcnt is morc heard than seen . The Rhinocero Hornbill, Buceros rhinoceros (Fig. 18) is rearly een so near. 11 was observed on Mt. Kinabalu up to 1750 m.
>
'"
..
-5 c: ::E
::i
o
From top to base Fig. 19 An unknown spider. ~~. 20 PJ!!ebopyll~rJ!. }!!!3!s a fro~~eJ:leadquarte~
ridae), 29 laughing thrushes (Timaliidae), and 9 barbels (Capitonidae). 48% of bird species endemic to Borneo have been recorded from Ml. Kinabalu. 40 species of fish; predominantly spccies which have adapted to fast-flowing waters by the development of adhe ive organs, e.g. catfishes (S isoridae, Bagridae) and gastromyzontids (Gastromyzontidae). Only a low percentage of the legions of arthropods has as yet been described: crustaceans (Crustacea), milliped (Oiplopoda), centiped (Ch ilopoda), spidcrs (Arachnida) and espacially insects (Insecta). Insects arc represented in thousands of spec ies in many orders: brilliant odonates (Odonata), butterfli es (Le pidoptera; e.g . Papi lion idae , Pieridae,
29
Mount Kinabalu - the surrounding environment
..
~ iii
::< ~
~~_IOOWi_ O
From top to base Fig. 23 Pseudomyagrus waterhousei Fig. 24 Brahmidia hearseyi F~. 25 G.!!J:!!.~ j;ypse.!! _ _ _ _ _ __
From top to base Fig.2 1 Pyrops illtricata !:ig.22 A~ew ly~merged~cad!:. _ _ _ _ _ _ _ _
30
Danaidae Lycaenidae; phingida , Gcometridae, Drepauidae) and beetle (Coleoptera; e.g. Cerambycidae, Lucanidae, Bupre tidae , Elateridae, Passalidae, Curculionidae), bizarre stick insects (Phasmida), crickets (Orthoptera) and bugs (Hemiptera); very diverse, in many case colonial and
Mount Kinabalu - the surrounding environment
therefore contributing more individuals than any other order of insects are termites (Jsoptera), ants and bees tHymenoptera). Most species occupy certain ecological niches defined as that part of the environment which a species actually exploits - and partly repre ent highly specialized morphological and ethological adaptation as a result of complex co-evolutionary developments of animals and plants, or of different animal species. They are fixed components of the rain forest and cannot survive without it. The highly advanced degree of differentiation of the adaptations also provides a clear indication of the old age of the Malayan rain forests as a stable ecoystem . As far as amphibians and reptiles are concerned, other animals are of vitaJ importance as predators and prey. Unfortunately that the prey spectra ofthe specie occurring on Mt. Kinabalu are as poorly known as their predators. The major part of the food of metamorphosed amphibians is composed of arthropods and their larvae, worms, and molluscs. Larger species, such as Limnonectes leporina. may also prey upon smaller vetiebrates, including member of their own species, and crabs. Most frogs are broad-spectmm carnivores (dietary generalist ); dietary pecialists are rare. M icrohylids, for example, feed mainly on ants and temlites. Predators of amphibians are particularly numerous among small mammals (e.g. civets, rats, otters) birds (owls, hawks, thrushes), and snakes (Amphiesmaflavifrons. Amphiesma sara·wacense. Dendrelaphis pictus, Rhabdophis chrysargos, Xenochrophis triangu/igerus) , but also include large spiders, centipedes, and crabs. The carnivorous larvae of dragonflies (Anisoptera) and aquatic beetles, hygrophilous snakes, and fish prey upon tadpoles living in the mountain streams and some may also ingest frog eggs. Nothing is known, on the other hand, about the predator of anuran larvae which live in stagnant pools . Potential dangers might come in the fonn of terrestrial crabs, larvae of small dragonflies, pond skaters (Baptista sp.), the larvae of mosquitos (Toxorhynchites sp.) and young, aquatic snakes. Due to their substantial sizes and physical powers, snakes (e.g. species of the genera, Python and Ptyas) play impor-
Fig. 26 Dendrelaphis pic/us eats a frog of the family Ran ida.=. (photog~pJ:. ~~not taken from Bomeo_).__
tant roles as predators among vertebrates. Many of the climbing species (e.g. Boiga, Dendrelaphis, Gonyosoma, Trimeresurus) prey mainly upon birds and plunder nests. Smaller arboreal (e.g. geckos, agamids, scincids), and fossorial species (e.g. Calamaria, Typhlops, Sphe/Jomo.rphus) feed mainly on arthropods and worm; the snail-eating snakes of the genera Pareas and internatus are largely specialized in preying upon molluscs and Gongylosoma upon spiders. Predators of repti les are predominantly found among predatory small mammals, wild boars, and birds (e.g. hawks, cockoos, thrushes, pittas). Lizards are the favourite food of many snakes, the smaJler ones being often captured by large hunting spiders. Besides the entirely unresearcbed parasites, the numerous predators arc a major controlling factor of the populations of amphibian and reptiles. Being carnivorous, amphibians and reptiles are secondary consumers of otber secondary and primary (vegetarians) conswners with amphibians usually occupying a midfield position in the food chains as second, third, and fourth level consumers. This is aJso true for many reptiles, although larger pecics may also be top predators "In many of food chains, within the food web.
31
Mount Kinabalu - the surrounding environment
Herpetofauna As of 2000, 77 species of amphibians and I 12 of reptiles are known from Mt. Kinabalu . Be ides some species settling in cultivated areas which occur in tbe marginal zone (e.g. Kaloula plilchra, Polypedates leucomystax. Rona et~vthraea, Raila nicobariensis, Fejervaty a limnocharis, Gehy ra Inutilata, Hemidactylus frenatus , Hemidactylus garnotii), the herpetofauna of the Kinabalu massif is composed mainly of forms particular to submontane and montane forests . These inhabit either trees at various storeys, the leaf-I ittered forest floor tbey may live subterreslrially, or have adapted to habitats dominated by mountain streams. Many species spend their entire life in the branchwork of trees and may only descend to. or near, the ground for a short period of time to reproduce; others are ground-dwellers, which occasionally can be fOlmd in the lower vegetation stratmn. While some species (e.g. Chaperinajusca, Rhacophorus nigropalmatus) undertake cxtensive journeys through the fore ts others inhabit rather small niches which they do not leave for years. The vertical distribution of some may stretch over more than -1300 m in altitude (e.g. Ansonia juliginea, Ansonia /ongidigita, Pelophryne misera, Cyrtodactylus ball/ens is, Phoxophrys cephalum. Amphiesma sarawacense), while others are only known from certain. well-delimited localities (e.g. An 'onia guibei. Leptola/ax maurus) . The adaptations to a life on the trees are particularly manifold. Arboreal reptiles have developed specific organs which aid climbing, such as scansory pads (Gekkonidae). strong, curved, and pointed claws (Gekkonidae, Agamidae), keels between the ventral and lateral scales ( erpentes: e. g. Dendre/aphis. Chrysopelea) . or prehensile tail s (Serpentes: Trimeresurus) . Frogs, on the other band, may have adhesive, glandular surfaces on their ventral sides and the pad of their finger and toe tips (Rhacophorids possess a cartilaginous joint between the subterminal and terminal joints which allows the pad to bc placed onto thc surfacc at a right angle). To be able to bridge larger distances between branches and trce trunks, frogs employ various leap-
32
ing techniques which are aided by particularly long hind limbs. Snakes havc solved this problem by employing a longitudinal muscle contraction which results in a stiffening of the body axis. Some species have acquired highly specialized adaptations, such as the ability to reduce the sinking speed of a frce fall by paragliding as a result of an active enlargement of the body urface thus enabling them to bridge larger distances in an oblique, controlled gliding flight. This may be achieved by a spreading of the ribs (flying snake: Chrysopelea), which may also be extremely enlarged and interconnected by skin membranes (patagia) (flying dragons: Draco), by employing appendicular lateral skin fringes like sails (flying geckos: Pfychozoon), or unproportionally large digital webbings (flying frogs : Rhacophorus). WALLACE (1869) was the first to describe and illustrate a "flying" frog (Rhacophorlls Iligropalmatlls) from Sarawak, a species also present on Mt. Kinabalu . Rain forests are marked by a distinct stratification of animal cOITIllHmities ill relation to its zonation. The various genera of scincid lizards (Scincidae) may serve as an illustration of this fact, with Sphenomorphus and Mabuya living mainly on the forest Door, Aptelygodon on tree trunks , and Lamprolepis in the canopy. In the case of agamid lizards, the circumference of the tree trunks leads to another type of segregation. with Draco distinctly favouring the mighty trunks of dipterocarps ,
Fig. X Rhacophorus nigropalmaflls alter WALLACE 1869.
Mount Kinabalu - the surrounding enviIonment
Gonocephalus trunks with a smaller diameter (usually < 30 cm in diameter), and Phoxophrys and Aphaniotij,· slim trunks and bushes. Frogs of the genus Megophrys spend their lives on the ground, PhilaUius amoenus prefers levels between ]0 and 200 em above the ground in the mo sy forest • and Phi/autus bUllitus occurs at 3 metres plus on trees. Due to a lack of research we know next to nothing about the herpetofauna of the canopy. In contrast to the arboreal species, numerous ground-dwelling inhabitants cxhibit particular adaptations to a shady, moi t environment. Roots, fallen logs in all stages of rot, rock flakes , and an all-covering layer of leaf litter offer a highly differentiated range ofnicbes. Besides cacci1ians (i .e. Ichthyophiidae) and terrestrial frogs (Lepto brachium, Megophrys, Ka/ophrynus), the leaf lilter community includes numcrous fossoria! to semifossorial snakes (e.g. Bungarus, Caiamaria, Cylindrophis, Cal/iophis, Pseudorabdion, Oreocalamus, Typhlops), forest skinks (Sphenomorphlls). worm skinks (Brachymeles) , and glass lizards (Ophi·aurus). Blind snakes (Typhlopidae) and dwarf snakes (Ca/amaria) exhibit typical adaptations to a burrowing-boring way of life in the forest floor with the head being reinforced and compact with a reduced scutellation and reduced eyes (the blind snakes have eyes covered by scales), no distinct slimming of the neck, a short, spine- like, pointed tail suitable for anchoring, and a very smooth seutellation of the body. Mabuya and Sphenomorphus actively utilize shifting sun patches on the forest floor for their thermoregulation . Frogs of the genera Ansonia, Hllia, Limnonecles, Meristogenys, Rana, Staurois, and keeled skinks (Tropidophorus) strongly depend on mountain streams; some snakes (e.g. Amphiesmaflavifrons, X enochrophis trianguligerus) and the turtle Dogania subplana are hygrophilous. The larvae of 71 % of the anuran species of Mt. Kinabalu are known, 56% of which are inhabitants of mountain streams , the remainder develop in ponds and phytothelms. The species of the genus Phi/autus, which are known for their eggs to develop into froglets within a shell of jelly, deposit theiI eggs terrestrially at sufficiently moist sites (cushions of moss, rock crevices, beneath bark, and in dead pitchers of Nepenthes). Some ran ids (Limnonectes
Fig. 27 Semiadult Rhacophorus everetti /Ilacrosce/is, a~ e~amp l eJ:.or lich!n~esi~ _ _ _ _ _ __
finchi, L. /eporina, L. palavanensis) engage themse lves in parental care for tbeiI developing offspring. Tadpoles living in mountain treams are usually adapted to a life in certain ecological niches with some posse ing highly specialized adhesive organs to withstand strong currents. A remarkable feature in the appearances of amphibians and reptiles is the high content of shape and colour elements which correspond with plant structures (phytomimesis). 10 this respect some representatives of the families Megophryidae, Bufonidae, and Microhylidae imitate dead leaves (inclusive of the leaf's ribs and imperfections caused by phyllophagous insects), while lichen and bark structures are found in the body coloration of various frogs (Rhacophorus everetti, Ansonia hanitschi) and reptiles (Dra co, Phoxophrys, Ptychozoon), while again other resemble twigs and Iianas (Ahaelulla, Dendrelaphis. Pareas, Psammodynastes, Trimeresurus). By their shapes and colour patterns, frogs of the genera Ansonia, Meristogenys, and Staurois blend perfectly into their surroundings of rocks in the plashing zone of small waterfalls whieh are partly overgrown with algae. Some Amphibians and reptiles further enhance these cryptic effects by remaining motionlessly attached to the surface when they feel they might have been discovered. Many species (e .g. ofthe genera Phi/aulus, Rhacophon/s, Bronchocela, Phoxophrys, Draco) furthermore possess the ability to rapidly alter theiI colorations. This physiological colour change is hormone-driven and may be triggered by changes in
33
Mount Kinabalu - the surrounding environment
Fig. XI N umbers of species in an uran families and genera present on Borneo and Ml. Kinaba lu.
0 Bombinatoridae Barbouru/a Megophryidae Leptobrachella Leptobrachium Lepto/a/ax Megophrys Bufonidae Ansonia Bufo Leptophryne Pedostibes Pe/ophryne Pseudobufo Microhylidae Cafluella Chaperina Gastrophrynoides Kalophrynus Ka/oula Metaphrynella Microhyla Ranidae Fejervarya Hoplobatrachus Huia Ingerana Limnonectes Meristogenys Occidozyga Rana Staurois Rhacophoridae Nyctixa/us Philautus Polypedates Rhacophorus Theloderma
22
9
20
30
40
13
4
2
2~
5 1
2
7
8
20
f
8
3 ~ 1
~
2 1
2 21
J
22
39
[2
~
~
i1
10
4 4 2
- 35
~----
22 1
15
14
2 3
4
8
0
10
8
10
14
20
endemic species to Borneo
34
10
30 species on Borneo
40 endemic species to Mt. Kinabalu
l
sPElcies on Ml. Kinabalu
Mount Kinabalu - the surrounding environment
Fig. XU Numbers of specie in saurian fami lies and genera present on Borneo and Mt. Kinabalu .
Agamidae Aphaniotls Bronchocela Complicilus Draco Gonocephalus Harpesaurus Hypsicalotes Phoxophrys Pseudocalotes Anguidae Ophisaurus Dlbamidae Dibamus Gekkonldae Aeluroscalabotes Cnemaspis Cosymbolus Cyrtodactylus Gehyra Gekko Hemidaclylus Hemiphyllodaclylus Lepidodaclylus Luperosaurus Ptychozoon Lacertidae Takydromus Lanthanotidae Lanthanotus Scincidae Apterygodon Brachymelis Dasia Emoia Lamprolepis Lipinia Mabuya Riopa Sphenomorphus Tropidophorus Varanidae Varanus
27
11
9
6
2- - -
-.------i--- ---
2
o
29
4
1
1
2 1
11
2 2
3_--t _ _ _
22
I
~
=--±-l~= -
+---=-- - I~I
- - - - _ -___--::40
16
4 4
4 1
2
1
14 -!-~
o
-+-----
10
20
endemic species to Borneo
30 species on Borneo
6
JL 2 2J 40 endemic species to Mt. Kinabalu
species on Mt. Klnabalu
35
Mount Kinabalu - the surrounding environment
r Fig. XIII
umbers of species in snake families and genera present on Borneo and Ml. KinabaJ u.
o Acrocordidae 2 Anomochilidae 112 Colubrldae Ahae/ul/a 2 24 Amphiesma Ap/opel/ura 1 Bolga 15 Calamaria 11 21 Can/aria 1 1 Cerberus Chrysopelea 2 Dendrelaphis 4 Dryoca/amus 2 Dryophiops 1 Eiaphe 3. Bapoidls '11 E:nhydrls l Si Fordonia 11 J2 Gongy/osoma 11 Gonyophis 11 Gonyosoma 1 Homa/apsis 2 Hydrab/abes Intema/us 2 Lepturophis 1 1 Liopeltis Lycodon 3 Macrophlstodon 2 29 Oligodon Opisthotropis 1 1 Oreoca/amus Pareas 2 2 Psammodynasles Pseudorabdion 14 Pseudoxenodon 1 Ptyes Rhabdophis Sibynophis Stegonotus Stoliczkia Xenelaphis Xenochrophis Xenodennus Crotalidae Ovophis Trimeresurus Tropido/aemus Cylindrophiidae Elapidae Bungarus Ca/liophis Neja Ophlophagus Pythonidae Typhlopidae Ramphotyphlops Typhlops Xenopeltidae Xenophldionidae
o
20
30
40
J
2 1 2 4 1
11
12 2
2
1 11 1
1 1
4 1 1 1
1 1 1 3
Ii
2
5
- --- ---
10 20 30 40 50 60 70 80 90 100 endemic species to Borneo
36
06
10
species on Borneo
endemic species to Mt. Kinabalu
1
species on Ml Kinabalu
50
Mount Kinabalu - the surrowlding environment
the surrounding temperatures, the intensity oflight, the level of humidity, but also by excitement. These colour changes do, however, not only serve the purpose of camouflage, but also play roles in commwucation and themloregulation. A perfect example for the function mentioned last i Phiiautlls aurantium which pos esse ' guanophores in its kin which retlect sun light. It enables the frog to avoid overheating by turning almost white in response to being exposed to sunlight while resting on a leaf during the day. Aposematic, brightly contrasting warning colours, as known from eotropical anurans (e.g. Atelopus. Dendrobates) are almost entirely absent in the species of Southeast Asia. The cbange of climate-induced ecological actualities, particularly between altitudes of 1200 and 1400 m, and between 1700 and 1900 m, respectively, corresponds not only with changes in the composition of the flora, but also with those of the fauna. umerous species of amphibians and reptiles reach their maximum, respectively minimum, vertical di stribution limits at these two altitudinal levels. The overlapping existence of highland and lowland forms in the area between 1200 and 1400 m causes the slight increase of number of species, though tendentious decreasing with increasing altitude. The oftenobserved congruences of these distributional relations in species of various genera underlines tbe distributional ecological importance of these boundaries. A drastic reduction of the specific diversity becomes evident at elevations above 1800 m, in the transitory zone to the mossy forests, so that the statement by I NGER (1978), "One of the disappointing things about Kinabalu's frog fauna is that it show so little altitudinal stratification.", clearly demonstrates the need for fundamental revisions. This tatement was very obviously based on an insufficiently large pool of information on the distribution of the amphibians in this area at that time. While our knowledge of the vertical distribution of reptiles on Mt. Kinabalu is grossly insufficient (only a handful, partly doubtful records exist for altogether five species from above 1800 m of which the records of Rhabdophis murudensis and Cyrtodactylus baluensis from 2500 m currently represent the maximum of the vertical distribution of reptiles from Borneo), at least some of the more vocal amphibians provide us with more acceptable in-
fomlation on their altitudinal distribution. Many genera (e.g. Leptolaiax, Leptobrachium, Megophrys, Kalophrynus, Ansonia, Philautus) appear on Mt. Kinabalu in the fonn of species groups, with the distribution ofthe indivi<;lual species being limited to certain elevations. It requires only an ascent up the mountain at night and the personal experience of just how sharply the individual vocalizations supersede one another and can be beard sympatrically only in narrow overlapping zones, to eliminate any doubts as to the existence of such abrupt vertical distribution boundaries. Figures VllI and IX (p. 26 and p. 27) shows clearly (with the exception of the area between 1200 and 1500 m) that the nwnber of species diminishes gradually with increasing altitude. Variou reasons are accountable for this. A particularly important factor - especially as far as reptiles are concerned is of course the climate whose upportive qualities for poikilothennic species of tropical areas deteriorate rapidly with increasing altiulde (lower temperatures, increasing precipitation, frequency of cloud cover). With increasing altitude the following changes are observed: • a reduction of the variety of ecological nichcs which is largest in the dipterocarp forests; • a reduction of the availability of food with the specific diversity and density of potential prey (mainly invertebrates) being subject to substantial limitation with increasing altitude; • a reduction of potential spawning sites. As a result of the topography, stagnant water bodies are absent and the streams above 2000 m usually have steep gradients. Their water levels often fluctuate between drying up completely and extreme high water. With the exception of Ansonia juliginea, whose larvae are unknown, none of those specie of frogs are found above 2200/2300 m which deposit their spawn in streams. For reasons already discu sse d in chapter "Zoogeography", the occurrence of endemic amphibians and reptiles on Mt. Kil1abalu correlates positively with the altitudinal level between 1400 and 1900 metres. It must be said, however, that their stati as endemites can at best be considered tentative as the totality of the species inhabiting the mountains of Borneo is not sufficiently known. In fact, many species previously considered endemites of
37
Mount Kinabalu - the urrounding environment
Mt. Kinabalu bave since also been recorded from otber mountains of the island, examp les being Leprobrachella balLiensis, Megophrys baluensis (?), Occidozyga halliensis, Sphenomofphorus kin abaluensis, Meristogenys kinabalue/1sis. Pelophryne misera, Cyrtodactylus baluensis, Oreocalamus hanilschi, elc. At present. 13% of the amphi bian species of Mt. Kinabalu are considered endemic to the mountain, i.e. Leptolalax a raya i. Leptolalax maw'us,
Leptobrachium gU/1ungense. Megophrys kobayashii, Ansonia j'uliginea, Ansonia guibei. Kalophrynus balLiensis, PhilaLilus amoenus, and Phi/autus saueri. AU of them have as to date only been found above 140011500 metres. 7% of thc reptiles of Mt KinabaJu arc currently regarded endemic, i.e. the lizards Hypsica/otes kinabaluensis,
Complicitus nigrigu/aris, Ptychozoon r/Jacophorus, Sphenomorphus tenuieulus, and the snakes, Calamaria schmidti, Hydrablabes pral!fi'onlalis, and TrimereSllnlS malcolmi. Since some of these species descend as far down as to Poring, their stati as endemites must be considered doubtful.
Human settlements in the vicinity of Mount Kinabalu "Enough has been discovered for il to be realized that fo1' many dl'limdls the rainforest habitat is es. ential and that they cannot Uvc in secondary forest: Whitmore, 1988 'The loss of biological diversity is the great tragedy of our age - J COn ider it to be U1C single most important and lasting of our specie on this plal1et.' Davis B. Wake, 1994 Discovelies in the Niall Caves, Sarawak, prove that there has been human life on Borneo for at least 40000 years. The population density, however, was at such alow Level until the last century, that around 1900 more than 90% of the island were still covered with a complete and coherent primary forest.
...'"
---------------------------------Fig. 28 Vegetable plantations between Bundu Tuhan and Kundasang .
38
Mount Kinabalu - the surrounding environment
Although SH ELFORD (1916) had already complained abollt an extensive destruct iOIl of the forests in the vicinity of settlements, the real decline of the rain forests did not start until the end the World War II . By 1957 the island had already lost 25 % of its forest. In 1953, Sabah, whcre every citizen could claim land as long as he wa willing to clear and cultivate it, was still covered with fore ts to an extent of 86%. [n 1983 there were only 63% left, in 1990 only 4 L%. The time bas passed in which one could still write like MJOBERG (1928): "The gigantic island of Borneo is clothed from head to foot, as it were, in green virgin forests ." How did this dramatieal decline come about? The most important reasons are: * Drastic increase of population. This leads to a continually growing need for new areas to settle in, whicb automatically implics infrastructural measures and woodland clearances. * Slash-and-burn as well as shifting cultivation by farmers; during some years with prolonged droughts in relation to EI Nino Southern Oscil-
lalion events between 1982 and 1987 as well as in 1997/98 trus kind of agriculturc caused fires of devastating extent; a total area of about 5-6 million ba. bad to be sacrificed to these flIes in the course of just the last ten years. * Highly develop-ed industrial nations (especially Japan) bought perrrussions for extensive logging for a period stretching far beyond the turn of the millennium. * Agro-industrially organised, large concerns promote the monocultural cultivation of oil palm trees and mbber plants on the cleared soil. These developments did not spare the forests ofMt. Kinabalu, as well. The surroundings of the mountain are inhabited by the Dusun people. Traditionally, they practise shifting cultivation (economy of changing land based on slash-and-bum cultivation: after a period of a 1-2 years' cultivation of a soil extremely lacking nutrients there follow fallow periods of various lengths), but restricted their economic actions to the glens of Kadmaian and the vicinity of Ranau, for the mountain itself was a
39
Mount Killabalu - lhe urrounding environment
religious taboo. It was not before the middle of the 20th century, when the belief in the mountain gods had deteriorated to a mere myth, that extensive forested areas facing south were cleared - up to an altitude of L400 metres. Between Kundasang and Ranau extensive vegetable plantations were estabUsbed; in L958 a jeep track was built between the west coast and Ranau which was extended in 1972 to the so-called Transsabah Higbway connecting Kota Kinabalu and Sandakan. In 1964, 71 I kml of the Kinabalu massif were declared a protected Park under the administration of Sabab Parks; mountain lodges (in between 3200 and 3800 01) and a visitors ' centre (Headquarters, at L524 m) were established, followed by additional ones at Poring (500 m), close to "Hot Springs", in 1967, and in Mesilau (1860 m) in 1998. Eventually, 25.5 Ion 2 were sacrificed to enable the establishment of a copper mine in the vicinity of the Sungei Mamut source. In order to compensate for the loss and to silence critical voices, the park was enlarged to the north into the Templer region. This region covers an area of93 km2, but large parts of it consist of substituted forest on logged-over land. In 1983, one of the big fires raged in the eastern foothills, and it was only due to sheer coincidence that it did not reach tbe buttresses of the mountain itself. The omnipresent tampering with fire continues to be common practice around the mountain and is not liable to any restrictions. In 1984, the park boundaries were redefined and the protected area was reduced by 26 Ion 2 . Two areas of a particularly rich biodiversity were sacrificed up to an altitude of about 1800 m during these reductions, the rain forests of Bukit Hampuan and of the Pinosuk plain. Temperate fruit and flower farms, a dairy farm , and a golf course were established on the deforested aeras. Whell in 1989 first steps were taken to reintegrate Bukit Hampuan into the park, a fire destroyed the major part of the remaining forests. Within the boundaries of the park as well , there are deficiencies hard to understand, e.g. the introduction of a sewage into g . Silau Silau, water supply taken trom the upper parts of Sg. Silau Silau, Sg. Liwago and its tributaries. During persistent periods of drought this leads to a partial running dry of entire stretches of the rivers and brooks, the overdimensional new road at Mesilau with ditches
40
for rainwater (40 cm deep) running alongside without the slightest support for animals to crawl back up. These ditches fonndeadly pits for many of them (in March 1998 we fOUlld 34 withered Ansonia and Merislogenys and three Sphenomorphus within a stretch of about 100 metres). TravelUng from the Headquarters down to Ranau, respectively frOlD Sayap down the Wariu valley, is an experience of a devastatingly harsh scenic contrast. Mountain forests still untouched and pure a second ago abruptly give way to a chaotic view without any gradual transition ; the traveller is confronted with the entire range of unthought-of methods of overfelling and overexploitation: all over the plaxe randomly dispersed buts of corrugated iron, slopes eroded by mud avalanches, bare ridges, residues of secondary forests crumbled to charcoal , simmering rubbish dumps polluted brooks ~~th an increased water runoff and increased sedimentation rates, and the churned-up areas of copper mines. Ignorance, Lack of organisation, unlimited pursuit of profit, and the bunger for land amongst an incessantly growing population have contributed to the gradual shrinkage of the belt of a protecting buffer zone around the mountain . But: the time of patchwork tailoring is drawing to an end: it is not possible anymore to bum away forest in the south and add its territory in the north, for one is running out of wood resources there as well. In the meantime, the mountain's substance has become totally exposed ; its overboarding biological treasures are forced to recede to a highly fragile and extremely jeopardised insular territory.
A retrospective glance on herpetological research in Borneo and the area of Mt. Kinabalu "The collector is far too prone to kill at o;ight ('very animal h cdptures; he i ' usually a bird of passage.1ll1d ha tlot the tune to devote to the patient and difficult observation of dn ru'liY1'1a\'s behaViour and habits of life." R.w. helford, A otUfdli t in Borneo, 1916
Mount Ki nabal u - the surrounding environment
Unti l thc beginning ofthc 19th century the Netherlands d id not have a fum grip 011 more than a couple of forts along thc coast. Vcntures into tbe interior of the islands, into the rcalm or cannibals, were looked upon as su icidaL Not before the time between 1820 and 1850 did the "Natuurkundige omissie" organize research excur ions, which invaded the central part of the island (e.g. by Borner, Gallion, Muller, Schwaner). The first zoologically important collection (or mammal and birds) was established on behalf ofthe Rijks Museum van atuurlijke Historie at Leiden. The herpctofauna did not attract any attention for quite a long time. The great snake atlas by SCHLEGEL (1837) does not mention more than one species as far as Borneo is concerned, Hydrophis pelamis. SClILEGEL (184854), GRAY (1845, 1859), and GONTHER (1859, 1864) then described for thc first time the amphibians and reptiles of Borneo stored ill museum collections at Leiden and London. Shorter herpetological works were published by BLEEKER between 1857 and 1860, and EOEUNG in 1865. fn 1885, FISCHER described a small collection made by Grabowsky, in 1892, Mo QUARD classified one by Chaper. The first check list of the amphibians and reptiles of Borneo (GONTH£R 1872) comprised already l57 species. Since by now more than a hundred years no significant progress has been accomplished as to the knowledge about the distribution and ecology of thc herpetofauna of Kalimantan, which suffcrs from a lack of research an)"vay. The scicntific research in the nowadays Malaysian part of Borneo has undergone a totally different development. Its beginning is inextricably intertwi.ned with the reign of the " White Radjas" of the Brooke dynasty, who resided at Kuching/ Sarawak. Scientifically interested Sir James Brooke was of paramount importance in this respect. He established his own collection, founded the Sarawak Museum at Kuching and roused interest in undertaking trips into the interior amongst naturalists and civil servants of his own administration by providing generous s upport. Without doubt, A.R. Wallace, who toured Sarawak in 1856, was the most well-known amongst them. During tbis time the first publication dealing with local herpetological data (MOTLEY & DILLWYN 1855) occurred.
Undcr the ru le of James Brooke, the "Sacred Mountain" became an object of scientific research. His co lonial minister Hugh Low reached the summit plateau for the first timc on the L1tJl of March 1851 , and he repeated the ascent together with Spencer t. John in 1858. The latter vividly depicts the expedition in his book, " Life in the Forests of the Far East (1862)". The trial s and tribulations, which lasted for weeks, did not mainly consist of the physical challenge of the ascent itself, but rather ofthe problems with establishing one el fat the base camp (the Dusun village Kiau) and with the recruitment of porters, who had to be convinced to invade the sacred aura of the mountain. The topograpby only aUows one route of ascent, wh ich was also taken by all subsequent expeditions. Still today the Summit Trail sticks to this route from 1800 m onwards, i.e. from Kiau (914 m) passing Lumu Lumu (I 676 m), Kamborangoh (2195 m)andPakka cave (3109 ill), up to the region of the summit. New route up Mount Kinabalu MesiJau trail promises more challenges Source: The tar, Friday, January 8, 1999 KOTA KINABALU
Trekkers intending to make their way up Mount Kinabalu, whether for the first or umpteenth lime, can now try out a new route which is also morc chal lenging than the existing trail whieh has been used more than 30 years. The new route, called the Mesilau Summit Trail, is located on the lesser-kno\vu eastern face of the mountain. rt starts at the recently-opened Mesilau Nature Resort ..... near a restaurant at the resort. The new 6 km route ends at the Layang Layang hut and hikers will have to trek for another 2 km before reaching Laban Rata ...... the new trail will trek through a montane forest and pass the Mesilau waterfall. ..... ''After 2 Ian at an altitude of 3000 to 4000 m, the trail goes downhill through a valJey ... , hikers can see a huge Agathis damara tree with its unique branches and roots," .. .. After passing through a number of streams and about 2.5 km into the journey, a 45 m-high waterfall can be seen. Then for another 2 km, the path snakes up a steep and narrow hill, the toughest part of the route .... after this section, tbe trail levels for 1.5 km and pitcher plants abound on both sides of the path .... the trai l then takes hikers tJlrough a rocky slope before they reach the Layang Layang hut at an altitude of 2702 m .
41
Mount Kinabalu - the surrounding environment
OUTH CHINA SEA
Rovi.--f'o-lIeW4d
'rltll.
Pl.'" _ _
... ~
'I '1OdOl' .... ., ... 11. . 1'3.4.1!.0 " i} 'It, P.,_I\.. 'J . ~ It , \.. 1..0.,..'. hell" 13,45-' /II. AI.,..t'(Gr.. Pull. 13";'3!r1
\1' .
40",,',
~. kl'r•• {d .... rd" ....." ....
G_ MII':I''-O __ ' ••
p"~.
't _
13,401
It has to be pointed out already at this stage that these altitudes, which were applied to later material, should be given a closer look before being taken over. This is especially vaJid as to "Kamborangoh, 2195 m". In pite of intensive research , many species (e.g. Ansonia longidigita. KalophfJ'nils pleUlvstigma, Cyrtodactylus baluensis. Sphenom01phus kinabaluensis) which were collected by Chasen and Pendlebury in 1929 and attributed by SMITH (1931) to thi s location could not be found at this altitude. Presumably they were collected somewhere alongside the track between Lumu Lumu and Kamborangoh . While the first attempts to climb the summit plateau are to be treated as rather sportive events without any scientific fruits worth mentioning, the ornithologist J. Whitehead did not only manage to climb the highest point ofthe plateau (Low's Peak, 1888) for the first lime, but also to earn scientific
42
It
'''',3''''' tto
n\Qtlilled
aMr
P~"Dl.fllQ"" & CHAI'EN 1932
merits; to Whitehead "belongs the credit of the first extensive exploration of the mountain and to him we owe our first real insight into the zoologica l treasures" (MO ULTON 1915). In 1893, WHlTEHEAD'S brilliant work "The Exploration of Kinabalu North Bomeo" was published. It was a decisive contribution to the formation of a new aura, the aura of an unfathomable biological trea sure. During the following decades Mt. Kinabalu became the place of pilgrimages for botani ts and zoologists alike, but never did a herpetologist occur amongst them. The herpetological section of Whitehead's collection was not edited by him himse lf but by B OULENGER (1887) and MocQuA RD (1890). Their list of species presents the first scientifically recorded amphibians and reptiles in the Kioabalu region. DUling the following 40 years civil servants of the Sarawak and Raffles Museums brought minor
Mount Kinabalu - the slUTounding environment
collections from Mt. Kinabalu every now and again, which were described by herpetologists who were not acquainted with the countIy by own experience; examples are the systematic descriptions of the collections of A.H. Everett by G.A. BOULENGER (1893, 1896), of R. Hanitsch by G.A. Bo LflNGER (1 99), of 1.A. Griswold by A . LOVERIDGE (\938) and of the biggest one, comprising 600 specimens, of EN . Chasen and H.M. Pendlebury by M.A. SMITH (1931). Due to the habit of considering the collector when creating the names of new species, we frequently encounter in them the names of scientists and of civil servants working as curators who dealt with fields of research totally different from herpetology (e.g. Trop;dophorus beccari;, Ovophis chasen;,
Pedostibes everelli. Oreocalamus hanitschi, Calamaria griswoldi, Philautus mjobergi, Meristogenys whiteheadi. etc.). Since the mid-fifties of the 20th century the research into the herpetofauna of Sarawak. and Sabah is very much intertwined with the personality ofR.F. Inger, now the Emeritus Curator of the Amphibians and Reptiles at the Museum of Natural History, Chicago. His classical work "The Systematics and Zoogeography of the Amphibia of Borneo (1966)" comprises the entire knowledge about the amphibians of the island available at that time. It also shows, however, how fragmentary the knowledge was back then. While already a high percentage of the descriptions of species is based on very old ma-
terial , data about ecology and ethology can be found only in fragments or are totally missing . This is the result of a research methodology predominant during more than a hundred years, which is characterized quite concisely by Shelford, as quoted in the heading of this chapter. As to the data concerning distribution, INGER himself remarked: "Analysis of the distribution of amphibians within Bomeo at this time is more likely to reflect the distribution of collectors than of frogs.". During the subsequent years INGER wrote (partly in co-operation with other authors like BACON, DRING, EMERSO , FROGN ER GRln , HAlLE, HOSMER, KING, LLOYD, TAN, VORIS, and WALKER), a number oftreatises on population dynamics, the bio logy of reproduction, and the settling in ecological n.iches by various species for the fust time. Apart from that he published many taxonomical papers describing numerous new species; he wrote particularly recommendable works about the systematics and ecology of tadpoles. Further descriptions of new amphibians were published by ISKANDAR (1978), DRING (1983a, L983b, 1984), MALKMus (1992b, 1996d), MATSUI (L986 1997), MALKMUS & RJEOE (1996a), MALKM US & MAT La (1997), and MANTHEY & STEIOF (1998). Due to these new insights and data the nwnber OfknOWll and described species has increased from 93 (L"IGER 1966) to L52 (Amua from 9\ to 146, lchthyophiidae from 2 to 6) amongst them Barbourula kalimallta-
Fig.31 M.A. Smith
-:Fig. 32 R.F. Inger
Arcbiv: Forschungsfustitut Senckenberg
43
Mount Kinabalu - the surrounding envirolUnent
nensis by ISKA"lDAR, 1978, a member of a family not yet verified for Borneo until then. Comments 00 the ecomorphology, ethology, activity rhythms. and the bioacoustics of specific species were published by DENZER (1994b), DRING (1983a, b, 1987), EMERSON (1981), HARDING (1982), HOFFMANN (1995c, 1998c, 2000), LAKIM, YAM BUN & AIRAMIl (l999), MA LKMUS (1989, 1992a, 1993, 1994a, 1994b, 1995a, 1996a, b, e, 1997, 1998), MALKMUS & RIEDE ] 996a, b), MATSUI (1979a, 1996), MATSUI, Wu & YorG (1993), MATSUI, YO:-JG, ARAYA & IlAMto (1996), and WONG (1994). Research on rePti le\~tilllingers on a. rather low level compared with that n amphibians. The latest total survey is provided by e work ofN. OEROOIJ (1915, 1917), which eocompas es the whole Indo-Malayan territory. MANTI·my & GROSSMANN ( 1997) published a volume, which comprises the whole variety of all known genera of the Southeast Asian herpetofawla; its impressive galJery ofbri II iant colour photographs adds to its cientific merits. Amongst the more recent revisions of the description of families, genera and species, INGER'S works (partly in co-operation with GREENBERG, HOS(l..fER, LEVITON, and MARx) occupy once again a dominant position, bcsides those of BACON (1967), HrKIOA (1982, 1990), GONTIlER & MANTHEY (1995), MANTHEY (1991) MANTHEY &
GROSSMANi\ (1997), MAI\THEY & DENZER (2000), MUSTERS (1983), 0tA & HIKJ[)A(1991) and WAI.LACH & GONTllER (1998) . HAILE (1958) and STUEBI"G (1991) published check lists of the snakes of Borneo, INGER & TAN (1996) did the same with regard to frogs, the lizards of Sa bah were listed by TAN (1993), the amphibians and reptiles of Brunci by DAS (1995), the amphibian and reptiles of Mt. Kinabalu by MANTHEY (1983) and MALKMU (l994b), the amphibians of Kinabalu by INGER, STUElliNG & T AK (1996) and INGER, Tt,"" & y ;\.\4BUN (2000). It is pleasant and satisfYing to note that recently five books about Bomcan amphibians and reptiles, written in a popular scientific vein, were published (JNGhR & TUEBING 1989, 1997, INGER & TA 1996, S, UEBING & INGER 1999, LIM & DAS 1999). Around the middle of the 20th century Mt. Kinabalu lost its formerly dominant position among the research territories of Borneo. This was not exclusively due to the f.:'Ict that one was tempted to believe that after a whole generation of scientists' intensity of work its zoology and vegetation were sufficiently explored; another important role played the fact tbat an exacerbated infrastructure and the use and deployment of helicopters made the remotest areas accessible (especially the central highland of Sarawak, the mountainous regions between Gunung
I'Fig. XV Graphic illustration of new records of amphibian and reptile species in the Mt. Kinabalu regiO~ (> 500 m a.s.l.) during the individual decades since the beginning of observations.
50
I
47
40
30 20 6
7
1971~1980
1891-1900 6 1931-1940 0 1951-1960 1887·1890 1901·1930 1941-1950 1961-1970
L-____
44
27
19
10
o
j
I
Reptiles
199~ 12000
1981-1990
• Amphibians ___________
Mount Kinaba lu - the surrounding environment
Mulu - Murud - Lumaku and rocker, the Danum Valley in the eastern part of Sa bah, etc.). The expansion of the examination area brought about the insight that many species, which were believed to be endemic, turned out to inhabit other areas of the island as well (cf INGER 1978, lNGI:;.R & TUU3INCi 1992). This was, however, noth ing new to ornilhologi ts (M/OBERG 1928). Surprisingly, investigations undertaken since 1984 lead to the discovery of numerous new species of amphibians and reptiles, all of which still have to be treated as endemic species. This demonstrates that the investigation of the hcrpetofauna ofMt. Kinabalu is far from approaching a final conclusion . Figure XV demonstrate the historical development of coming to know the range of species at Mount Kinabalu . Scientists doing field work regularly brought back collected material from their travels. Most of the herpetological material is looked after in the following collections in museums throughout the world: The British Muscum of Natural History, London (BM); Field Museum of Natural History, Chicago (FMNH); Kyoto University Zoological Collection (KUZ); Museum of Comparative Zoology, CambridgefMass. (MCZ); Museum National d' Histoire Naturelle, Paris (MHNP); Naturhistorisches Museum Wien (NHMW); aturhistorisches MusemTI Basel • (NMB); Osaka Museum of Natural History, Osaka (OMNH); Sabah Parks-Field Museum, Headquarterl Kinabalu Park; Sarawak Muscum, Kuching (SMK); Forschungsinstitut Senckenberg, Frankfurt/ Main (SMF); Raffies MusemTI of Biodiversity Research, National University of Singapore (USDZ/ZRC); Sabah State Museum, Kota Kinabalu (SSM); Coli . Universiti Kebangsaan Malaysia, Sabah Univ. Campus (UKMS); United States national Museum, Smithsonian Inst. , Washington (USNM); Zoologiscbcs Forschungsinstitut und Museum A. Koenig, Bonn (ZFMK); Muscum fur aturkunde der HumboldtUniversitat zu Berlin (ZMB). Onc could still agree with Ingcr's comment on amphibians, dating from 1978, althougb a moderate modification secms recommendable: "Almost none of the information on natural histOlY comes from information madc at Kinabalu. There is much to be done."
This quotation does not on ly delineate a deficiency, but also functions as an appellation to draft a programme of future research activitics. It is also an appellation to get going. Especially three field s should form the centre of investigation :
*
*
*
Systematics: a whole variety of systematic problems arc to be tackled in this context. TI1is should not be attempted on a merely morphological basis, but also by including and considering ecological, ethological, and molecular data. Naturally, orne uncertaintie will not yield to clarification, as tbe products of a systematic approach are artificial and - like WILSON (1997) adequately pointed out: "scienoe to a bigh degree, but a bit of art as weU". Distribution: nearly all the data about places where animals were found is restricted to the close environment of Poring, the Headquarters, Mesilau, Sayap, Bundu Tuhan, and Kiau; apart from the summit trail there are no data conceming the areas above 2000 metres. All the data refer to the lower stratum of the forest (below 5 metres). Future investigations should include areas outside this very limited scope of observation, which barely covers approximately five percent of the total space. Almost nothing is known about the equitability (relative density and distribution) of the various species. atural History: we barely know anything about the natural history of most of the species (including the larvae). Tbcre is a lamentable lack of studies rescarch dealing with ecology and ethology (e.g. reproduction, acoustics, population dynamics, activity rhythms, settlement in niches, ctc.). As a crucial step forward in investigating the herpetofauna of Mt. Kinabalu, Sabal1 Parks havc initiated a program in 1991. Thcy work together with INGER, who monitors the population dynamics ofsclected species at three sites at different levels of elevation.
This book attempts to sum up the present state of knowledge about the amphibians and reptiles of Mount Kinabalu. It soon will become evident that the number of unsolved problems is vast. No doubt: "There is much to be done" !
45
higher taxa of anurans Suborder Archaeobatrachia uperfamily DiscogJossoidea family Bombinatoridae Genu Barbotinila .. uperfamily Pelobatoidea Family Megophryidae Genera Leptobrache/la Leptobrachium Leptolalax Megophrys
uperfamily Bufoooidea Family Bufonidae Genera Ansonia Bufo Leptophryne Pedo tibes Pelophryne
Superfamily Microhyloidea Family MicrobyJidae Genera Calluelia Chaper;na GasLrophrynoides Kalophrynu.'i Kaloula Metaphrynella Microhy/a
Superfamily Ranoidea Family Raoidae Genera Fejervarya Hop/ohalraclws Huia Ingerana Limnonectes Meristogeny ' Occidozyga Rana Staurois
Family Rhacophoridae Genera Nyctixalus Phi/autus
Anura
Important traits for the identification of frogs and their larvae According to MAYR (1975) thc members of a species form a genetic and ccological unit and a reproductive community. The phcnotype, which presents itself in the form of never absolutely identical individua ls, is a summary of traits which reults from both endogenous (genotype) and exogenous (biotic and abiotic influences of the environment) factors. The selective pressures exerted by the latter may have favourable, eliminating, or varying effect on the development of individual traits. Such traits, or groups of traits show a more or less pronounced consistency specific to individual taxa and are therefore of importance for the classification of an animal. •
48
The species as a genetic unit: the ana lysis of the genotype is thc task of tbe laboratory biologist. As far as the frogs of Southeast Asia are concerned relevant work bas just begun. The spec ies as an ecological unit: it manifests itse lf in many-faceted interactions with the animated and unanimated environment, in particular by behavioural patterns typica l for the spec ies or genus, and by physiologica l adaptations. It requircs only a single apomorph ic feature to prevent populations from producing fertile progeny. Such differences may, for examp le, be variations in the aposematic behaviour, communjcation, activity rhythms , social structure, choice of ecological niche and calling sites, ctc . As far as Mt. Kinabalu is concerned, there are sta tionary bank-dwellers along the streams (e.g. species of the gencra Lepto/a/ax, Ansonia, Me rislogenys, Huia , Staurois, Rana ), ground-dwe ll ers of the forests which never (e.g. Pelophry ne, Kaloph rynus), or only for the purpose of reproduction visit streams (e.g. Leptobrachium , Megophrys ), or arborcal species which never (e .g. M etaphrynella, Nyctixa/us, Phi/aulus), or also on ly for the purpose of reproduction visit streams (e .g . Pedostib es, Po{y pedates , Rh acophorus).
•
The species as a reproductive community: behavioural pattems which can be linked to reproduction are to a large extent ritualized and particular specific to a species . This servcs to ensure a reproductive scparation among closely related, sympatric species. Ritualized behavioural parameters can be found in all interactions related to reproduction. Oviposition and volwne and appcarance of clutches may as well provide information as to which species or genus we are dealing with. Diumal, small frogs, for example, which show a "flagging" bebaviour with stretched-out hind limbs are certain to belong to the gcnus Staurois; if we encounter a free-hanging foam nest or one which floats on the water surface, we can be sure it were Polypedates or Rhacophorus which deposited tbemthere. The advertisement caUs often offer the key for distinguishing specics which cannot be separated, or are djfficult to diffcrentiate, by morphological traits (e.g. species of Phi/au Ius or Leptobrachium montanum vs. Lep lobrachium gunungense).
As valuable as biochemical and ethological data are for the identification of species, they usually require the emp loyment of costly instrumentation, large amounts of time, or depend on the coincidental concurrence of certain factors. The herpetologist in the field will therefore make use of the immediately availab lc traits of the pbenotype (external morphology) for the identification of a species in most of the cases. Many species of Bornean frogs can be identified without doubt on the basis of morphological traits only. It is on ly for highly variab le and phenotypically very similar forms (c.g. in genera suc h as Ansonia, Leptobrachium, Meristogenys, Microfly /a, Pe/ophryne, Philautus) that even the experienced systematist is faced with ditIiculties. As a consequence the identification keys for these genera are unsatisfactory. It is important for the user of this book to know the meaning of the tcrms used in the species accounts and how the metric details were taken, and the same appJie to the definition of the call characteristics. Only by this knowledge will it be possible to verify and compare them with own data.
Anura
Body shape and size of a frog u ually ugge t an affiliation to the family, and often point to a certain genu already. In order to ub equently determine the re pective genus and specie , a erie of traits are of great importance, such a the hape of the nout and pupil, the hapc, relative ize, and po ition of the tympanum, and certain kin tructures (skin surface smooth, granular, rippled, warty; size, hape, and arrangement of glandular raise , skin ridges and appendages; meristically highly variable groups of keratinized pikes on the lower jaw, chest, and finger ). Metric ratios and comparative data of two body dimension are often much more important than indications of linear size ( VL of adult specimens) as they are not only relevant for the identification of a pecie, but also for that of the repective sex. Here, we restrict ourselve largely to the following indices: • tibia lengthlsnout-vent-Iength (TU VL); • comparison head length head width; • distance of the anterior eye margin from nostril ;
• distance of the posterior eye margin to tympanum; • horizontal diameter of eye horizontal diameter of tympanum; • inter-ocular di tance - inter-narial distance; • length of finger and toe ; and the p ition of the tibio-tar al articulation when the hind leg is plied forward to the body, etc. The presence, re pectively absencc, of mandibular and vomerine teeth, presence or ab ence of nuptial pad in male , and the morphology of the tongue may al 0 be of importance. The arrangement and hape of the tubercles on the lower side of the hands and feet are often of great taxonomic relevance, and 0 are the hape of the finger and toe tips and the extent of the interdigital webbing. To de cribe the latter we will u e a formula in accordance with GLAW & VE~CE (1994): "Finger and toes are numbered. ach toe or finger is followed by the number of the phalanges, which are not webbed (in parenthese ). Only the visible phalanges, which are bordered by the subarticular tubercle , arc considered. 'i' mean inner side, 'e' means outer (exterior) ide of a finger or toe. 0, '4i(l)' mean 'at the inner side of the fourth toe there is one unwebbed phalange'."
Fig. XV I Place of measurements in anurans.
Fig. XVII Anuran palate.
Morphology
snout-vent length internal naris aperture (choana)
vomerine teeth
49
Anura
supratympanic fold
truncate or spatulate or enlarged triangular
canthus rostral is crest tympanum
disk
semicircular
3
spinose warts parotoid gland
slightly enlarged metacarpal tubercle
skin granular
not enlarged (sometimes pointed)
thenar tubercle
Fig. XIX U nderside of the hand ofa frog. femur knee
vent tibio-tarsal articulation (heel)
4
pointed projection supernumerary tubercle
dermal fringe tarsal-metatarsal articulation
cutaneous border
5
Fig. XVlIl External morphological characters of frogs and toad s. intercalary cartilage
" subarticular tubercle - metatarsal tubercle
terminal phalange glandular
Inner metatarsal tubercle
region circumarginal groove
outer metatarsal tubercle tarsal fold
Fig. XX A sec tion through the tip of the digit of a treefrog.
Fig. XXI U nderside of the foot ofa frog.
4
e 2
5
Webbing formula : Fully webbed; 4i(O) 4e(O) i=illller side e=cx(erior side of phalanges
Webblll£ fomlUl a' 41(2) 4e{ I); 4i(2) : 2 phalanges of (he inner side of the fourth (OC are un webbed 4e( I): I phalange of the ex terior side of the fourth toe is unwebbcd
Fig. XX II Webbing fonnula (after
50
GLAW
&
V ENC'ES
Full formu la; I (O . ~) 2i( I )2e(0.5) 3i( I )3e( I) 4i(2}4e(2) 5(0) we usc a simpli fied version: 1(0.5) 2(l i O.5) 3(1 / 1) 4(2! 2} 5(0)
1994) for describing the extension of webbing.
Anura
Fig. XXIII Shape of snout of Bomean frogs.
Fig. XXIV Pectoral girdles (cart ilaginous areas arc stippled).
truncate, blunt (e.g. species of Allsol/ia. Pedo.lliiJe.l)
round (e.g. species of Leplo/a/ax. l\1erislUgel/)'s. Occidu::),ga , Rana)
sloping (e.g. species of PO(lpedales, RaHa )
A fi r mistern a l pectoral girdle (epicoracoid cartilages fused in the midline) have specie of tile familie~ Microhylidae. Ranidae and Rhacophoridae.
protruding (e.g. species of AI/sollia , Bllfo, Ka/ophrl'lllls)
a ngular (e.g. species of Leplll!Jrachilllll.
RI/(/cophol"l/,~)
In some cases tip of snout with a denna l projection. e.g. triangular projection (Megop hrys lIa.wla). con ica l tip (females of Phi/a/lll/s pelen-i) .
Next to the habitus, it i often the colouration and the colour pattern which give a pecie it typical appearance (e.g. in Leptolalax piclu , Pedostibes everetti, Chaperina fusca, Rana signata, or yclixalus piC/liS). The pigmentation of the iri i unmi takable in orne form : Leptobrachillm monla1111111 ha , for example, a white, and Leplobra Ilium abbotti a blui h eye background, Rhacophorus angulirostris po cs cs a blue upper margin of the iri , Leptolalax arayai how a coloured egment covering ca. 120 0 of the iris, while a network of blood ve e l cover the eye of Leptolalax piclus and Philaulus bUllilus. Some pecies exhibit a marked exua l dichromati m (e .g. Phi/aullls
amoellllS, Rhacophonl . angu/iroslri , Rhacophorus everelli). Referring anatom ic re earch of frog, detail of the kull , the shoulder (pectora l) girdle (arciferou or
An arciferal pectoral girdle have species of the fillni lies l3ufonidae and Megophryidae.
firmi temal), and details of the vertebrae are of great importance for determining a specimen' sytematic po ition. The larvae of anurans (tadpole) can al 0 be identified by species- pecifie externa l criteria. Major morphological trait include the po ition of the piracle, the ana l tube, the eye, no tril ,the hape and extent of the caudal fin, morphometric relation ,colouration and colour patterns. However, the most important group of character for identification oftadpole are tho e a ociated with the oral field (shape and extent of marginal oral papillae, the number of row of labial teeth, and that of the keratinized beak). The rows of labial teeth are again described in a formula which wa originally propo ed by AloTlG (1970). It mu t be pointed out, though, that the proportion of the body of a tadpole may vary with both the indi idual and age a
51
Anura
Fig. XXV Embryonic and larval development of a frog (substantially simplified after stages 1
2 ·13 cell division
fertilized egg cell
14
15 16 rotation neural tube
O '_ .
17 tail bud
18 muscular response
19 heart beat
GOSNER
1969),
20 21 22 gill cornea tail fin circulation transparent circulation short outer gills
neural folds
23 - 25
26
30
mouthdevelopment ~~ ~, ,,'_. ~ long outer gills gill reduction
,,~.,,;"
_
31 - 38
limp bud toe development development
,subarticular tubercles visible
Leg bud visible
40
42
41
43-44
45
46
,., metamorphomouth & snout ; ", £\ }~ ~~ '7z.._G ~ . sis complete transformation ~~
tail reduction
.
arms visible under skin
arms appear
mouth slit clearly visible
-
tail only as bud
Fig. XXVI Lateral view of a tadpole, caudal muscle
lateral (sinistral) spiracle or spiraculum
leg
anal tube
Fig. XXVII Structures of the oral disc of a tadpole, labial teeth rows
beak
Teeth rows undivided
Teeth rows divided
2
3 4 5
2 3 papillae
52
5(3-5) / 3 5(2-5) /3(1-2) Teeth row formula after AmG 1970,
Anura
l
Habitats of tadpoles
Fig. XXVIII Tadpole of LeptobrachiulIl glln ungense [rom g. Tibabar (1850 m), showing system of lalcralI ine organs.
The type of a water body and its altitudinal location can also assist in the attempt to detennine tadpoles as the larvac of the individual anuran species occupy in general very specific niches. Thc origin of a tadpole is therefore valuable information for the taxonomist. I NGER (1985) distinguished 9 of those microhabitats ("rn ' in species accounts) for mnning waters and 7 for stagnant ones.
can the structures of the oral field (e.g. in Leplobrachium, Meristogenys) . If visible, the arrangement of the lateral organ, and the position of glands, may be of importance for the identification of a species. The significance of larval morphology for the systematics can be demonstrated by a look at the Amolops-group where the genus Amolops was split up into the gcnera Amolops, Huia, and Meristogenys on the basis of morphological differences in their tadpoles (YANG 1991) and larval characters were crucial for detcrmining the evolutionary relationships among the taxa.
1. Streams a. Torrent: surface of water broken, occasionally foaming, bed of rocks (diameter > 5 cm) and boulders, current strong (e .g. Leptobrachella, Meristogenys ) b. Rifflc: surface of water in shallow waves but never foaming, bed of gravel, current moderate (e.g. Leplo/a/ax. Ansonia, Rizacophorus cyanopunctatus) c. Shingle area: surfacc of watcr smooth, bed of flattened rocks (3- 10 cm), current weak to moderate (e.g. Rana ehalconota, Megop/zrys nasllla) d. Leaf drift: areas with deep accumulations of dead leaves, drifts up to 1.5 m deep and 10 m long,
Fig. XXIX Microhabitats in a mountain brook (in part. aller
M~LKMUS
1999).
typical microhabitat for larvae with ventral sucker disk like e.g. Meristogenys
~-_'7""J:!:i~r:r;:::",L
rocks, exposed to the current }: :"f·~ ·-,·t~...~~~.. _.
• ~
interstices of gravel leaf drift barrier of leaf drift, caused by rocks or branches and trunks / _________ fallen into the stream ~
I
typical microhabitat for larvae with cup· llke expanded lips like of and funnel-shaped lips like of Megophrys _
L ____~ ___
-=-___
LePtola/:x:---:_::!!!!~_____
------
53
Anura
current weak (e.g. Leptohrachium montanum, Rana signata, Rllacophorus gauni) e. Open pool: ponded parts of streams, bottom silt to sand and gravel (e.g. Leprobraehium monlanum. lvJegophrys nasufa) f. Side pool: an embayment off the main channel, partially or completely cut off from the current, bottom fine t.o coarse sand, usuaUy covered by dcad leaves. current weak or absent (e.g. RanG signata. Leptobrachilln1 montanum. Bt!fo divergens) g. Pothole on rock bank: water-filled depression on rocky bank, bottom bed rock, often with dead leaves (e.g. Rana signata. Rana eha/conota) h. Seepage area: trickle of water less than 2 cm deep, usually emerging from a steep clay bank (e.g. Limnonectes lruh/i) i. Pool of intermittent stream: up to 0.5 m in maximum surface dimension, depth up to 5 cm, bOItom fine silt, sand, and pea-gravel, current only after moderate to heavy rain (e.g. Limnonectes kuhli, Raila cha/conota)
2. Microhabitats away from streams a. Seepage area: film of water less than I cm deep emerging from a slope of clay and small rocks, current. negligible (e.g. Linlllonecfes ktlllli). b. Rain pool : rain-filled depression of variable size and depth, bottom usually clay, often with dead leaves (e.g. Kalophrynus pleurostigma. Rona linchi. Polypedates otilophus. Microhyla). c. Animal wallow: rain-pool modified by the action of mammals, bottom fine silt, length more tban L Ln, depth up to 15 em (e.g. like in 2b) d. Log cavity: mainly open on upper surface or side of a log, surface up to 15 x 20 em, depth up to 10 cm (c.g. Chaperina fusca. Nyctixalus pictus. Microhyla borneensis). e. Hole in living surface root: small holes up to L8 cm in diameter and LO cm in depth. height above ground 5-16 cm (e.g. Nyctixalus pic/us). f. Buttress tank: cavity fonncd by auastomo is of two or more buttresses, surface lip to 30 x 40 cm, depth 30 cm, height above grouod up to 4 m. g. Hole in trunk or large branch: surface area up to 20 x 30 cm depth 30 cm, height above ground > 0.5 111 (e.g. Chaperil1a fusca, Metaphrynella sllndana, Nyctixalus pictus).
54
h. Water-filled phytothelmous cavities in bamboo internodes, leafaxils (Pandanus), natural cups in litter (fruit husks, palmtree spathe, fallen leaves) and dead Nepenthes pitchers: very reduced water content (e.g. Leplophryne borbo/liea, Metaphrynella sundana. Phi/aUiLls saueri)
Larval feeding types As adaptations to their individual habitats and partly in order to be able to make use of certain food resources, anuran larvae have often developed substantial modifications to their body stmc!Ures (e.g. shape oftbe body, development ofeaudal musculature and tins, position and stmctures of the mouth area). In principle two categories are distinguished, i.e. endotrophie and exotrophic larvae. Endotrophic larvae gain nutrition for their development from parental sources (yolk): these froglets may either hatch from egg capsllies (Phitau Ius) or are free-swimming, nidicolous, nonfeeding tadpoles (Kalophrynus. Pelophryne). All the other species of Bornean genera have tadpoles of the exotrophic type. i.e. they are free-swimming, feeding larvae. (J 986) distinguished 5 larval feeding types: I) Bottom suspension feeders: in microhabitats away from streams and in stream areas with weak currents where fine sllspension and leaf dri fl cover the bottom. TadpoLes of this type (generalized pond-type) show no morphological specialization (e.g. Bufa [fig. 39J. Rana, Po/ypedales [fig. 40]). 2) Mid-water suspension feeders: in microhabit.ats away from streams (raiJl pools, pig wallows, potholes); larvae with large branchial baskets with dense ruffles, lacking beaks and denlicles (e.g. Chaperina [fig. 41], Microhyla [fig. 42]). 3) Obligate benthic feeders: in streams with moderate to strong currents; tadpoles feed on the epilithic film on rocks to which they cling. They have cup-like oral disks (e.g. Rhacophorus), sucker-type oral disks (e.g. Ansonia [fig. 43]), or abdominal suckers (e.g. Huia [fig. 44]. Meristogenys); larvae are rheophilous, with elongated, depressed bodies, long muscular tails, low fins, and small, non-bulging eyes.
INGER
Anura
4) Macrophagous feeders: larvae inhabit a wide range of microhabitats in streams; they are common in leaf dri fts and in interstices of gravel on the stream bottom (e.g. Leptobrachella, Leptobrachium [fig. 45] , Lepto/a/ax [fig. 46]); larvae with heavy, sbarp beaks. 5) Surface film feeders : larvae dwell in the entire range of stream microhabitats from riffles to potholes; they suspend particles of floating material from the surface film by means of their upturned funnel mouths (e.g. MegoplllYs [fig. 47]).
I
Larval feeding type 2
I
J
il :J
0 6-______~~~__~~~~~__=_~~~~
I
Larval feeding type 2
J
il ________________ ~ L
From top to base: Tadpoles of the genera Fig. 39 BI!(O Fig. 40 Po/ypedares Fig. 41 ChaperillCl Fig. 42 Microhy /a.
From top to base: Tadpoles of the genera Fig. 43 Ansonia Fig. 44 Hllia Fig. 45 Leptobrachillm Fig. 46 Lepto/a/ax: Fig. 47 Meg ophlYs.
55
Anura
All these types of larvae are represented on Mt. Kinabalu although the bottom suspension feeders, which are typical elements of lowland waters, feature much less prominent than the rheophilous benthic feeder. Due to geomorphological and hydrological reasons, the nwnber of water bodies which would allow a successful development of amphibian larvae is very limited above 2000 m on Mt. Kinabalu; here, endotrophic fonns dominate (e.g. Philautus species). The realization of an identification key to the larvae is impossible at present since only 50-60% of the amphibians of Borneo are known by their tadpoles (comp. T}J GE R 1985 , 1992a, b, TNG ER & WAS ERSUG 1990). We must therefore be content with a key to the families here.
Fig. 48 A calling male of Lepto/a/ax pictus from Wariu, Say1p, 1000 m .
Bioacoustics On reaching maturity the males of most anuran species are able to vocalize. Most species develop - partly in both sexes - several types ofvocalizations (advertisement, release, distress calls, etc.). Among these, the advertisement calls of the males are of exceptional importance with regard to biological and systematic sciences.
I
Listeni.ng to nightly concerts of frog voices is certainly one ofthe most memorable experiences during a visit to a rainforest. The volume of thesc concerts is again and again surprising. It is mainly due to the resonant properties of the vocal sacs (sac-like extrusions of the bottom of the oral cavity) which vary in ize and position, but are gen-
Fig. XXX Circactian calling activity of some anurans on Mt. Kinabalu (after
Megophrys ba/uensis
-
Megophrys kobayashH Leptobrachium montanum Lepto/a/ax arayai Lepto/a/ax pic/us Kalophrynus ba/uensis Metaphrynella sundana
----
Staurois /uberllinguis
Phi/autus mjobergi
----
-------
..............................................
..............................................
Phi/au/us aurantium
L 56
I
o
I 996b).
.... .... ., - . ..
Ansonia hanitschl
Rhacophorus angullrostris
MA l KM1;
I
I
I
I
I
I
I
3
6
9
12
15
18
21
maximum calling activity
limited calling activity
Anura
--.----
,-------
I~
i__
II~:~ P
---
- - - - - - -------,
Which part of an acou tic signal must be considered the "caU" is often a qucstion of the definition choscn . 1n a series of signals largely made up of about equal pulse , each signal considered a note and the entire signal series a call. (e.g. Phi/ali/lIS ollrontiuIII).
call
• note means a pulse group (n) • note duration means the time from the beginning of the first to the end of the last pulse in a note • intcmote interval means the time from the end of one note to the beginning of the next note (i) • note repetiiion rate means the number of notes per second • p: pulse duration (ms) • ip: interpulse interval (ms) • pulse rate means the munber of pulses per second
ip
Fig. XXXI Physical call characteristics.
the localization of the caller, and the timulation of the sexual partner. thus serving a reproductive isolatjon mechanism. ]n conjunction with aggression calls the advertisement call simultaneously functions as an acoustic marker for an occupied territory and the maintenance of the same, thus regulating population densities. In order to prevent confusion clo ely related species which occur
erally positioned in the throat region . The sounds are actually produced by the channelling of air back and forth between the lungs and the oral cavity passing past vocal chords which oscillate and produce sounds which are then amplified by the vocal sac. The biological significance of the advertisement call lies in the identification of the own species,
------ 1I as an example.
Fig. XXXll Graphic illustration of acoustic signals using Phi/autus mjobergi
I
«» j
I I o
1
2
3
20
140
40
I
160
time [ms]
timers]
1 Sonagram (graphic illustration of an acoustic signal illustrating its course of time, its composition oftrequencie • and the relative intensity of the individual frequency components. The horizontal axis indicates time, the \ vertical one frequencies) . 2 Oscillogram (serves to interpret structural traits recognizable in a sonagram). 3 Enlarged section of an oscillogTam showing a high resolution of structural clements in order to perfoml de\ tailed analysis of the call/note structure.
-- --
---
------- -------
I
I I
57
Anura
sympatrically have developed more distinct call structures than those with an allopatric distribution (e.g. in species of the genera LepLObrachium and Philaurus). But it is not always the male which actively attempts to attract a sexual partner. For example, female Polypedale.l' leucomystax ready to mate advertise by producing single croaking sounds and a rhythmical dnunming with their toes (seismic communication) for minutes on end . On Mt. Kinabalu we are met with spatially finl1ly established calling colonies (e.g. of Megophrys baluensis, Metaphrynella sun dana, Staurois tuberilinguis. Ansonia longidigitata. Philawl/s aurantium) , those with calling densities which are unequally distributed over larger areas (e.g. Leptobracilium montanum, Kalophrynus ball/ellsis, Phi/autus bunittts and P picfus), and specics whieh produce singular calls at large intervals (e.g. Merisfogenys orphocnemis, Rhacophorus angulirostris). There are di- and nocturnal callers. Some voices give evidence of an adaptation to the acoustic background of their individual habitats. The dwellers of mountain streams for example produce sharp clicking, whip-cracking, or chirping signals in order to rise above the sound level of their biotope (rushing of water) and facilitate communication. The fact that advertisement calls are species-specific is of immense value for the field herpetologist since especially morphologically very similar species are typically distinctly differentiated with regard to their bioacollstics. It also helps to gain an overview of the local distribution patterns of the individual species. Until about 40 years ago, the description of advertisement calls was Limited to onomatopoetic descriptions such as a short, whip-cracking "cheep" -sound for Meristogenys or a burring "emrr..." for Phi/autus petersi, or by using musical note . Obviously thcse methods of illustration are very subjective and little informative. In order to record the physical call parameter (call length, fi'equency structure, intensity repetition rate of the impulses. length of intervals. etc.) objectively, the calls arc recorded on tape and, with the aid of computers, displayed in the form of sonagrams and oscillograms thus rendering them fit for comparisons. Sonagrams provide a basis to analyze sound levels, relative volumes, and the timing of acou tic signals.
58
When comparing analyses of voices it must be taken into consideration that factors such as humidity, temperature, light, and stimulation by the given acoustic environment, but also the volume produced by the individual, may have substantial effects on certain caJl parameters (particularly volume, frequency, speed of the pulse rate). To facilitate acoustic communication within the acoustically heavily polluted rain forest the individual species choose certain frequency bands. Just like radio stations use various carrier frequencies, frogs occupy various acoustic "chatmels", in other words acoustic niches within the overall backgrowld noise of their habitat.
Reproductive strategies In a worldwide survey, D UELLMAN & TRUEB (1986) distinguished altogetl1er 29 different reproductive modes which differ in the choice of oviposition sites and type of development of the larvae. 0 less than 10 of these modes have been recorded for frogs on Mt. Kinabalu:
I. Eggs aquatic A. Eggs deposited in water 01. Eggs and feeding tadpoles in lentic water (e.g. Chaperina. Kalollla, Occidozyga, Rana, Bufo) 02. Eggs and feeding tadpoles in lotic water (e.g. Lepfobrachella, Lepto/alax, Ansonia, Meristogenys, Huia, Staurois, Rana) 03. Eggs and feeding tadpole in water in tree holes or aerial plants (e.g. Metaphrynella) 04. Eggs and non-feeding tadpoles in waterfilled depressions (e.g. Kalophrynlls) 05. Eggs and non-feeding tadpoles in water in tree holes or aerial plants (e.g. KalopfllYnus, Pelophryne. Phi/autus) B. Eggs in foam nest 06. Foam nest on pond; feeding tadpoles in pond (e.g. Polyp edales, Rhacophorlls)
n. Eggs terrestrial or arboreal
C. ~ggs on ground or in burrows 07. Eggs hatch into feeding tadpoles that are carried to water by an adult (e.g. Limnonectes jinchi) 08 . Eggs batch into froglels (e.g. Phi/alilus)
Anura
Above from top to base Fig. 49 A typical foam nest of Polypeda/es leucomystax. Fig. 50 Eggs ofa Philallllls species in a depression ora dry fern. Left from top to base: Fig. 51 Tadpoles of Chaperina /usca in a bambusintemodium. Fig. 52 In waterfi lled tree holes live tadpoles of some species (e.g. Met~phrYllel!a).
D. Eggs arboreal 09 . Eggs hatch into tadpo les that drop into water-filled cavities in trees (e.g. Nycfixa/us pictus)
10. Eggs hatch into froglets (e.g. Phi/aufus) E. Eggs in foam nest II. Nest arboreal; hatching tadpoles drop into ponds or streams (e.g. Po/ypedafes, Rhacophorus) ; foam nests were found not overbanging water, but the tadpoles would hatch during rain showers and then be washed to a water body.
Like tbe choice of oviposition, the details of the individual clutches (e.g. arrangement, number, size, and pigmentation of the eggs) are indi cative of genera and species. Unfortunately, only a relatively small perccntage of species are known by their clutches. Species dwelling the upper montane zone tend to produce smaller cl utches with larger eggs (e.g. Lepto/a/ax, Pe{opluyne, Philaulus). Whether all or at least some of the anurans at the various altitudinal levels are reproductive ly active throughout the year, is still llllknOwn . A far as Borneo is concerned, respective information is only availa ble for lowland rain forests (I NGER & GREENBERG 1963,lNGER & BA 'ON 1968). They sbow a circannual reproduction for individual species.
59
Anura
Key to the families of Bornean frogs (in parts after
MANTHEY
& GROSSMANN 1997)
Teeth present in upper jaw (A) .......... ......... ... .. . 2 • Teeth absent in upper jaw ......... ................ ........ 7 2 An intercalary cartilage between the 2 terminal phalanges (8); 1" finger opposed to remaining fingers ............ ........ .... ....... ...... .. Rhacophoridae
D
c
• No intercalary cartilage; first finger not opposed to remaining fingers .... ..... ........... .. ..... ................ 3 3 Pupil vertical (C) ................ ........ Megophryidae • Pupil horizontal (D) or round ......................... .. 4 4 Vomerine teeth present (E) ..... .. .. ....... ............... 5 • Vomerine teeth absen!.. .. Ranidae (Occidoz.yga)
E
5 lnner metatarsal tubercle shovel-like (F) ...... ..... . .... ...... ........... ........ ..... Microbylidae (CaJluella) • Inner metatarsal tubercle not shovel-like or absent ....... ........ ........... .. .. ......... ............. ...... .. ........ 6 6 Fingers fully webbed (G) ... ...... Bombinatoridae • Fingers without webbing or rudimentary webbing ............. .............. Ranidae (except Occidoz.yga) 7 Skin dry, warty; parotoids (H) usually present ... ................... ................ ............ ..... ... .... Bufonidae • Skin moist, smooth or rough, not warty; parotoids absent.. ... ....... Microbylidae (except Ca/luella)
60
H
Anura
Preliminary key to the larvae of Bornean frog families (in parts after
MANTHEY
&
GROSSMANN
1997)
A
Free-swimming larvae ....................................... 2 • Direct development within the egg .................... . .... .... ..... ... .......... .... Rbacopboridae (Philaulus) 2 Spiracle lateral (A) or ventral (8) ..................... 3 • Spiracle not visible, only I row of tccth (C) ...... . .... ..... ....... ... ................. Bufonidae (Pelophryne)
B
~~ c
3 Spiracle lateral (A) ........................................ .... 4 • Spiracle ventral (8), no keratinized jaws, toothless ... ..... ...... ....... .............. ... .... ..... Microhylidae 4 Oral disc directed ventral (D) ........................... 6 • Oral disc directed terminal (E), without rows of teeth ... ......... ..................... ..... ............................. 5 5 Lips funnel-shaped (F) ........................................ . ..................... ......... Megopbryidae (Megophrys) • Lower lip horseshoe-shaped (G) ...... ... Ranidae (OccidotJIga)
G
@)
6 Marginal papillae of the mouth surround the entire oral disc (H) .......... Megopbryidae (with teeth: Leptobrachium ; without teeth : Leptobrachella: with or without teeth: Leptolalax) • Upper lip without or only with lateral marginal papillae (I) ......................................................... 7 7 Anal tube medial (K) ........... ............................. 8 • Anal tube lateral ..... Ranidae (Hopiobatraclllls, Linlllonectes), Rhacophoridae (Nyctixaius, Rhacophorus, Theloderma) 8 Abdominal sucker (L), more than 2 rows of teeth on the upper lip ... .... ......... .......................... ........ .. ......................... Ranidae (Huia, Meristogenys) • With or without oral sucker (M), 2 rows of teeth on the upper lip ................................................. 9 9 3 rows of teeth on the lower lip ......... Bufonidae (keratinized upper jaw ab ent or divided ( ): Ansonia; keratinized upper jaw continuous (0): BII/O, Leptophryne, Pedostibes) • At least 6 rows of teeth on the lower lip ............ . .......... .................. ..... ... ........ Ranidae (Stallrois)
N
o
61
Bufonidae
Bufonidae Bufonid are co mopolitan in temperate and tropical regions (except for Madagascar and the Au traloPapuan region) . Oriental bufonids have an arciferous pectoral girdle, the sacrum has di lated diapophy es, an omostemum is absent, a bony temum pre ent. There are 5-8 holochorda l, procoelou pre acral vertebrae. The maxillae and premaxillae are edentate. Most species have dry, thick, glandular skins, often with pustular wart . The pupil i horizontal. Mating takes place in an axillary amplexus. Most bufonid produce egg clusters in tring form (eggs normally pigmented) and deposit them in water. Their tadpoles are free-swimming, aquatic, have beaks and denticles: the piracle i inistral, the anal tube medial (except in PelopIIlYlle ). orne genera have torrent-adapted, oro-adhe ive tadpole with large, expanded oral disks and suckers (Ansonia) . 111e specie ofPelophty ne have non-feeding larvae which survive entirely on yolk. 80mean bufonids occur in various habitats, some are terrestrial (Bufo) , others arboreal (Pedosfibes) or both, living partly on the ground and partly on low vegetation (Ansonia, Pe/ophfJl/le); Pseudobufo is aquatic. Nearlyall pecie are distributed in primary and secondary forests, only a few appear in areas modified by man. On Mt. Kinabalu, bufonids can be found over a wide altitudinal range. Of the three frog pecies known to exceed an elevation of3000 m a.s.l., two arc bufonids (Ansolliafuliginea. Pelopllly ne misera) . The fami ly is represented on Borneo by 6 genera and 29 species, of which 5 genera and 12 species occur on Mt. Kinabalu.
Key to the Bornean genera of Bufonidae (after INGER 1966, MANTHEY & 1 Tympanum visible (A) .. ... .............................. ... 2
• Tympanum not visible (in part) ....... Pedostibes 2 Parotoids present (8) ....... ...................... ........... 4 • No parotoid glands .............................. .............. 3
3 Toes fully webbed (C), body and limbs robust; adults > 70 mm ......... ..... .... .... ... ...... Pseudobufo • Toes not ful ly webbed, body and limbs slender; adults < 65 mm ..... .... ..... ....... ................ Ansollia 4 Atmo tone phalange offirst finger projecting from fleshy web (D); adults < 40 mm ....... Peiophrylle • two phalanges of first finger projecting free of palm or web, hands without fleshy web .......... 5 5 Finger tips rounded, not dilated ...... ................. 6 • Finger tips dilated into truncate disks (E) .......... . ...................... ......... .... ......... (in part) Pedostibes 6 Underside of foot with a large elongated tubercle near bases of each toe; body and limbs slender; length offemur and tibia > SVL (F) .. .... .... Leptoplrrylle
•
62
o such tubercles; body and limbs robust; length of femur and tibia < VL (G) .. ... .......... .... . Blllo
E
GROSSMANN 1997)
Bufonidae
Genus Ansonia
STOUCZKA,
1870
Members of thc genus Ansonia are known from southern India, Thailand , the Malay Peninsula, Borneo, and Mindanao (Philippines). Bornean species are mall to medium- ized. brown to black toads (SVL < 65 mm), with a dorsolaterally weakly flattened . elongate body with lender. long legs. Tympanw11 distinct (except in A. anofis, see p . to) ; in most specie rostrwn projects over the mouth ; nostrils above symphysis, large eyes (diameter ~ snout length), pupil horizontal. Tips of fingers and toes mostly rounded, some species with truncate disks. Finger lengths 3>4>2> 1, toe lengths 4>5>3>2> 1; fingers webbed at bases, webbing of toes varies with the spccies. Skin rough, usually warty ; parotoid glands absent. Adult males with subgular vocal sac (except in A. guibei), nuptial pads and mandibular spines present. Ova rather large and unpigmented. Tadpoles small (usually < 20 mm) and adapted to a benthic life in flowing water with a streamlined body (except in A. leptopus), and a cup-like, sucker-type oral disk; two upper and three lower rows of labial teeth, upper beak divided or absent, lower beak divided only in a few species (e.g. A. guibei). One species (Ansonia an otis) with an abdominal sucker. Body coloration usually black. These toads live in forest streams with strong currents (except A. /eptopus). Bornean species inhabit primary and old secondary forests in hilly to montane areas. They can be found on the forest
floor and on leaves in the lower stratum of vegetation. On Mt. Kinabalu, this genus has a distinct altitudinal stratification with A./uliginea reaching the maximum vertical distribution (3480 m) of all amphibians of the Sunda Islands . Some species are local endemics, restricted to very limited areas (A . Juligin ea, A. guibei). E leven species are known from Borneo, six of them occur on Mt. Kinabalu.
Remarks: There is some doubt about the occurrence of A. albomaculata and A. /eptopus on Mt. Kinabalu. INGER (1985) described larval Ansonia from Sg. Kipungit which he originally believed to beA. albomaculata. ln a later paper (L'IGER 1992b), however, he distanced himself from this view by leaving the specific identity of these larvae open and referring to them as Ansonia 'crucifonn" due to their dorsal pattem. On the other hand, A. albomaculata is listed again to occur on Sg. Kipungit in the check list of the amphibians ofMt. Kinabalu (INGER, STUE.BING & TAI\ 1996), while the works by INGER & TAN (1996a) and INGER & STUEBING(1997) do not mention it for Sabah, but only for Brunei, Sarawak, and Kalimantan . Another larva, which could as yet not be assigned to a species, was described by INGER {I 992b) from East Mesilau (1460 m) and from Sarawak. Like A. guibei, it has both jaws divided and is therefore referred to as Ansonia "beaks divided". In a new checklist of INGER et al. (2000) A. albomaculata, A. /eptopus and even A. minuta are mentioned for Kinabalu Park.
Tab.] Di stribution of Ansollia -specics on Borneo. Sabah
Ansonia albomacu/ata (:-;GER, 1960 AI/sonia l 'uIiJ.:iflea (MOCQUARD, 1890) A1Isol/ia guibei [ NGER, (966 A"so"ia Iratritselli i NGER, 1960 Ansonia /afidisca LNGE R, 1960 Ansonia /eJ)/o{lLis (G ONTHER, 1872) A I/sol/ia /OlllfidiJ!itli h '-GJ;R, 1960 AllSollia minuta i >lGER, 1960 Anso"ill p/atysOlt/11 h '-GF R, 1960 AI/sotria Spilill/jfer (MOCQUARD, 1890) Ansonia forrenfis DRING, 1983
• • • • • • •
JZinabalu ?
• • • ?
•? • •
Brunei
•
Sanlwak
•
• •
• •
• • • • • •
Kalimantan
•
• • •
63
Bufonidae
Preliminary key to Bornean species of Ansonia and 51b toe webbed to tips; skin flap on vent present ... ...... ....... ... ....... .......... ................ . guibei
6 Disk of 3"' finger subequal to diameter of tympanum ..... ...... .......... .................................. latidisca
• 3111 and 5111 toe not webbed to tips; no skin flap on vent .............................................. ..................... 2
• Disk ofJ,d finger much narrower than tympanum
2 I" flDger does not reach disk of 2nd when fingers adpressed .................. ........................................ 3
7 Distance tympanum - mouth half or less than half the distance nostril - mouth: flattened habitus .. ............... .... ......................................... piat)'soma
)'d
• I" flDger reaches disk of 2nd when fingers adpressed .................................................................... 9
3 Sharp tarsal ridge present ................................ 4 • No tarsal ridge .................... ............................. 5
......................................................................... :. 7
• Distance tympanum _ mouth more than half distance nostril - mouth .......................... hanitschi
8 Snout projecting, oblique in profile .... torrentis • Snout vertical in profile ........ ..... ........ juJiginea
4 A wlllte spot below eye; a whitish band from eye to arm ... ........... .... .... ..... ...... ......... albomaculala
9 Dorsal warts low, rounded .................... leptopus
• Without such pattern ............................. minuta
• Dorsal warts elevated, spinose ...................... 10
5 Tips of outer fingers dilated into spatulate disks
10 No interscapular spot ...................... IOllgidigita
........................................................................... 6 • Tips of outer fingers rounded .......................... 8
Ansonia /uligillea (MocQuARD, 1890) Blljo /liliginellS MOCQUARD, t 890: 158 - Type locality: North Borneo Neclophlyne alliludinis S~lInl. 193 I : 14. pI. I : fig. I - Type locality: " Pakka. alt. 10,200 feet". Mount Kina Balli, abab (syn . after I NGER 1966 : [02 ; ["GER & T/\ 1996a: 554) P~DlEBl:RY & CHASEN 1932: 21 Pedo.l'libes altitlldiflis, BARBOUR [938 : 192 Ansonia altitlldinis. INGER & Tl.iEBING 1992: 4[-51 Allsalliajitiigineu, b :GER 1966: 102, 1978: 312: MATSUI 1979: 314; MAstlf(Y 1983: 21; MATSlI, HIKIDA& NAMBU 1985: 152; MAll-.~I\JS 1988c: 176; INGER & DRI"C 1988: 463; h ."-It & Sn ~flIN<; 1992: 41-51; M.\LK~IU' I 992a: 106; "C.I· R, llEBI'<J & TAN 1996: 362; IN(;fR & TAN 1996a: 554; INGER el al. 2000: 16
• An oval interscapular spot present; tips of fingers dilated ........................................ spilllliijer harp; lores oblique, a little concave; tympanum half diameter of eye; tips of fmgers and toes slightly wollen; finger webbed at their base; webbing of toes 1(0.5),2(1 /0.5), 3(2/ 1.5),4(3/3),5(2); a weak outer metatarsal tubercle; mandibular spines b lack, arranged in a double row; skin dorsally and laterally with low, rowlded warts, sometimes with melanic spinules; lower side of body granular. Brown above, eyelids and snout darker; flanks and lower sides black; legs light brown below; abdomen with yellow or whitish spot; iris blacki h with tiny yellow pots.
EcologyfEthology Diagnosis A medium-sized, moderately stocky pecies, with a vertically truncate snout-tip in profilc; lip of I" fingcr not reaching base of wollen tip of 2 nd ; nuptial pad composed of a few keratinized spines: restricted to the high mountain region.
Description Male up to 36 mm, fcmales up to 44 mOl; TU VL 0.38-0.41: head as wide as long; canthus rostralis
64
Nearly nothing is known on the life history of this toad . It i re tricted to the high mountain region (> 1500 m a .. 1.), to the upper and subalpine forest zone and beyond the tore 1 line. We observed a specimen a tive at night on the fore t floor in the rocky region with very dense vegetation near Pakka Cave (3050 m). At the sanle site PE.'1DLEBURY & CHASEN (1932) had found a toad, "the stomach of this specimen contained fragments of beetles and ma ll crustacea".
Bufonidae
Fig. 53 AnsoniaJi.diginea from Pakka , 3050
~.
Distribution Ansonia fuliginea is only known from Mt. Kinabalu: Marei Parei; Kamborangob ; Layang Layang; Pakka; Panar Laban (1500-3480 m); Sg. Mesilau (East); Mentaki Ridge (1800-2000 m).
Ansonia gllibei
INGER,
1966
Ansollia gllibei INU I'R.. 1966: 104 - Type locality: Mesilau Cave camp, 1800 m, Mount Kina Baiu, Sabah Amonta guibei, L"OER 1978: 312; MANTHEY 1983: 21 ; INOER & DRING 1988: 463; MALKl.IUS 1988c: 176; INGER & S11JEBING1992: 41 -51 ; MALKMUS 1994b: 221 , 1994c: 87; INGER, STUEBr.-'G & T AN 1996: 362: lNGER & TA)~ 1996a: 554 ; LAKI~t eta!. 1999: 33: M AL KM I,;S & Kosuc1I2000: 121-124 ; INGER et at. 2000: 16
Diagnosis A medium-sized sp ecies of Ansonia with long limbs; snoul vertical in profile; first finger reaching base oftip of 2 nd finger; toe webbing reaching
tips of 1"', 2nd , 3m, and 5th toe; flaps of skin flanking vent; no vocal sac. Restricted to high montane regions. Description Males up to 32 mm, females up to 34 mm; TLiSVL 0.47-0.56; head as wide as long' canthus rostralis sharp; lores weakly concave; tympanum one third of eye diameter; tips of fingers and toes swollen; fingers free of webs; a large, round pabnar tubercle; toe webbing 1(0), 2(010), 3(0/0), 4(2/2), 5(0) ; small inner and outer metatarsal tubercles; skin above with many round warts, sometimes with dark spinules; flanks and ventrum granu lar; mandibular spines black, arranged in one row. Dorsally and laterally dark brown, sometimes with lighter regions; ycllowish brown crossbars on hind limbs; knees and upper arms lightened; belly blacki h with whitish granules; chest and throat greyish
65
Bufonidae
'!'l
~:;; Ii
o !.ig. 54 _A pair of Ansonia guibei fron: Sg. Mesilau East, _1850 m.
rosy ; toe webbing greyish brown; nuptial pads black; iris golden brown with fine black network. Tadpoles with very broadly rounded snout; oral disk wider than widest part of body; upper and lower beaks divided; labial teeth dense, fom1Ula 2/3' ta il tapering to a rounded tip in the last third; tail length 64% of total length (23.8 mm at st 29); head and
Fig.
xxxnl
Lateral and dorsal view of the tadpole of
body uniformly black dorsally, c ireul111aterally lighter; caudal muscle dark brown to black; fins transparent with dark margins. Ecology/Ethology Ansonia guibei is resl:Iicted to the high montane primary forest of the Mesilau valley. Here it lives on boulders protruding from the water and areas near the banks of the rapidl y flowing mountain stream. It is noetumal. When threatened it may also leap into
Ansonia guibei.
1 -
Fig. 55 Tadpole of Ansollia gllibei.
1 mm
R. Malkmus
66
Bufonidae
velY rapidly flowing water, even into cascades, dive, and hidc in rock cavities below the water surface or near the bank. In mid-August we observed a fema le which carried a male, attached to it in axillar amp lexus, to the stream. The male produced frequent erie of impulses with high-pitched, metallic clicking and cracking sounds. Large numbers of larval A. guibei were found in the Sg. Mesi lau ( 1800-1950 m) in the month of March where they preferably resided on large boulders in moderate to strong currents (m: la, b, c). They were at about the same level of deve lopment (Sf 25-30) indicating that reproduction follows an annual rhythm. The p rey pectrum of A. guibei is wide with a large variety of arthropods being accepted: cicadas, pentastornids, small grasshoppers, plantlice, aphids, beetles, small butterflies, spiders, mites, hairless caterpill.ars, and beetle larvae.
Ansonia itanilschi INOlOR., 1960b: 484 - Type loca lity: Mount Kina Balu, Sabah 811/0 (Ansonia) penangensis. (not AI/sonia pe/langensis TOLlCZKA, 1870) MocQ AIUl 1890: 160 Blifo penangensis, (not Ansonia penOflgensis S TOll 7KA. 1870) HANITSCIi 1900a: 74; VAN KAMPio.'I 1923: 75 BII/o leploptts, (not Btt/o leplopus GON11lI!R, 1872) SMl111 1931 : 13 (part); 1NGBR 1956: 395 (part) Ansonia mimI/a, (not Ansonia minllla INGFR, 1960) MANTHEY & D IiZER 1982: 12; ~1 A'/TIIEY 1983: 2 1 Ansonia albomaclIlC//a, (nol Allsonia albomaClllala IN OER, 1960) MALKMIiS 1991 a: 28 Ansonia cf. albomtlcl1lala, MALK.~IUS 1992a: 108, I994b: 222 Amonia "allilsehi. INGf.R 1966: 106, 1978: 312; M AiS!';1 1979: 314; M ANTIIEY 1983: 21; MALKMUS 1985 : 7, 1987: 278, 1988c: 176; I:;OER & DRI ~G 1988: 463 ; MALK.\1\JS 1989: 184. 1991 : 28, 1992a: 106; INGER & STUESING 1992: 41-51 ; WO~G 1994: 25-37; M ALKMUS I 994b: 221, 1994c: 89; ilOFFMANN 1995a: 9; ING~R. STUEBING & TAN 1996: 362; INGER & TA.' 1996a: 554; INGER & TA~ I996b: 73; MALKVl US 1996b: 20-26, 1996c: 281; M Ar-TIlEY & GRaS MANN 1997: 28; H OFnlA.'1\ I 998b: 85; LAKIM et al. 1999: 33; lNGERet al. 2000: 16
Distribution The spec ies is known only from the eastern and western Sg. Mesi lau, between 1800 and 1950 rn a.s.L, on Mt. Kinabalu.
Diagnosis A mediwn -sized, moderately slender highland species with long legs; nout oblique, loping in profi le; I" finger not reaching disk of 2nd •
Ansonia hanitsclti I NGER, 1960
Fig. 56 Allsonia hanitschi from Sg. Silau-Si lau.
67
Bufonidae
From top to base Fig. 57 Dorsal pattern of a female of Ansonia hani/selli from Sg. Silau-Silau. Fig. 58 Belly of two females of Ansonia hanitsehi.
Description Males up to 32 mm, female up to 37 mm; TLiSVL 0.45-0.57; head as broad as long; canthus rostralis sharp; Jores veltical; tympanum half eye
I ::< Ii @
68
Bufonidae
males were only encountered during the day (between 6.00 a.m. and 6.00 p.m.) perched on the banks of streams, rock formations, sand and gravel banks, and on emerged rocks in sprayzones. ln contrast to most of the other amuans their calling activity is hardly affected even by prolonged period of drought. Although territories are marked acou licalJy, aggressive behaviour may be witne sed occasionally. In tills regard it was obscrved that one of two males, which were perched opposite each other at a distance of only 1 cm and called antiphonically, suddenly raised on its lUnd limbs, grabbed the head of its opponent and pressed to its chest for about 30 seconds. A captive pair produced Fig. 59 Male of Ansonia hanilschi. strings comprising 45-73 eggs on rocks submerged in water during the day (HERRMANN & ULBER 1992). gitlPoring (500 m), INGER, STUEBING & TAN (! 996) Tadpoles live in rocky streams with moderate to for " Layang Layang" (2600 m), but both these strong currents (m: la, b , c). records are doubtful, because of its normal altitudinal A. hanitschi can often be found resting or foraging range. on the leaves oflow (up to 2 m) shntbs near streams during the night. Captive specimens accepted ants, flies, pentastomids, earthworms, spiders, and the larvae Fig. XXXlV Sonagram and oscillograms of Ansonia hanitschi (Sg. Liwago, 1500 m: 3.00 p.m. +2 10c) of beetles and butterflies. Call: (+2 10c) A frequen tl y repeated note series which is very much reminiscent of the chirping of a cricket. Each series takes 6- 10 s and is composed of equal notes with a note repetiti on rate of 6 per sec. Each note (duration 140-\50 ms) comprises a group of 3 pulsed single notes (duration 21-50 ms) which are repeated at intervals of 25-35 ms; frequency 5.4 to 6 kHz.
2.40
2.60
2.80
3.00
3.20
sec
Distribution A. hanitschi is only known from severa l mountain ranges in western Sabah and northern Sarawak. Mt. Kinaba)u: The species is very common in the Silau-SilaufLiwago region ( 1400- 1850 l!l); Kadamaian Falls ( 1600 m); Mesilau Cave; Sayap; Sg. Kinateki; HANITSCH (1900a) recorded it from "2 100 and 4200 ft."; WONG (1994) mentioned it for Sg. Kipun-
69
Bufonidae
Aflsonia lOflgidigita INGER 1960
slender; tip of I" finger reaching base of swollen tip of 2nd finger.
Ansonia /ollgidigita /ongidigita lNG ER, 1960b: 480 - Type locality: "at 4,200 feet on Mount Kina Ball1, North Borneo" Bufa leptaplis. (not Bufo /eptopus G 01\.IlER, 1872) B OULENGER 1887b: 95; MucQuARD 1890: 159; WHITEHEAD 1893 : 121 ; BARTLElT 1894: 204: HANrrSCH 1900a: 74: VA • KAMPEN 1923 : 75: SMITH 1931: 13 (part); Ansonia /eplopIL1. (nol Hufo /eptoplIs Gm.'flER, 1872) LN GER 1956: 395 (part) Ansonia /ollgidigila /ongidigira. INGER 1966: I I l. 1978: 312 Ansonia /oflgi(Jigita. MA:
\LK.."IUS I 996c: 281 ; HorFMA~ 1995a: 9: INGER & TAN 1996a: 554; HOFrMAN:\' 1998b: 85: LAKlM et al. 1999: 32; INGER el a!. 2000: 16 Ansol/ia sp., l:-:llER 1985 : 20
Description . Males up to 50 mm, females up to 70 mm; TLfSL 0.53-0.61; head slightly wider than long; canthus rostralis sharply angular; lores vertical; tympanum two fifths of eye diameter; tips of fingers and toes swollen; fmgers without webs; a large round palmar tubercle; toe webbing 1(0), 2(0/0) 3(1.5/ 1-1.5), 4(2 .5-3/2.5-3),5(0.5-1); 2 low, round to oval metatarsal tubercles; limbs with small, elevated, spinose warts dorsally and dorsolaterally; throat finely granular, other ventral surfaces coarsely tubercular; 2-4 rows of brown to black mandibular spines. Dark brown above, with small irregular Iight spots and singlc brick-red warts; obscure light crossbars on limbs; throat brownish, chest and belly yellowish with brown or black spots; nuptial pads
Diagnosis A moderately large to large, slender-bodied Ansonia with an oblique, projected snout; fmgers long and
-----
-
-
Female of Ansonia /ongidigitafrom Sayap, Sg. Lurnotok Kecil, 1100 m.
g
tn: o '--_~"-'-"
70
----
Bufonidae.
.c
~
u Ul
'Ic5
e
E
0~ '---_
0.:
_
Fig. 61 Tadpole of Ansonia longidigita from Sg. Kcmantis.
o
Fig. 62 Male of Ansonia /ongidigita from Kundasang, Sg . .tg~sep~.'- 150~ ~ _ _ _ _ _ _ _
(duration 36-44 ms) consisting of2-3 pulses (durayeJlowish brown; iris light golden brown, with a tion 7-33 rns). The internote interval takes 31-36 ms, fine black network. Tadpoles teardrop-shaped; snout broadly roun- the dominant frequency is 3-4 kHz with a harmonic at 10.8 kHz. ded; eyes dorsolateral, oral disk ventral; labial teeth rows formula 2/3; upper beak divided by a large gap; tail tapering Fig. XXXV Sonagram and oscillograms of Ansonia IOllgidigita to a pointed tip; body and cauda l (Sg. Langanan, 800 m; 7.30 p.m. + 22°C) muscle dark blackish brown ; total length 20-25 mm.
EcologylEthology This toad is distributed over a wide altitudinal range (150 to 2100 m a.s.1.) in primary and old secondary forests . It is active at night, and by day stays hidden beneath logs, leaves, and in crevices of rocks. Its breeding sites are clear, swift, rocky streams. The males call at night from spray-moistened rocks, gravel bars, and from the leaves and twigs oflow vegetation (up to 50 cm) forming calling chomses. Tad pol es I ive on the bottom of streams with moderate to strong currents in primary forests (m: la, b, c).
~~
t-I ~ tit H 1.40
1,60
1,60
2.00
2.20 sec
sec
Call: (+22°C) The long note series (0.9-10 .5 s) is composed of long, high-pitched trills consi ting of 4-135 notes at a note repetition rate of J2.5-15.9 per second, each note
71
Bufonidae
Distribution Ansonia /ongidigita is known from abah, Brunei, and Sarawak. Mt. Kinabalu: On Mt. Kinabalu, the species has an extens ive alti tud inal range with a recorded maximum of 21 00 m, but it i les common above 1000 m a.s.1. Sg. Panataran, 4800 ft. (WHITEHEAD 1893); Kenokok, Kiau, Sg. Kadamaian; Sg. Liwago/Silau-SilauJ Tibabar, Kamborangoh (SMITH 1931 , as A. /eplopus, 2100 m); Sg. Mesilau' Sayap: Sg. Kemanlis, Sg. Wariu, Sg. LumotokKecil; Sg. Luidan, near Bundu Tuhan (1100 m); Poring: Sg. Langanan (very common between 650 and 850 m a.s.I.).
Ansonia platysoma
1960
Ansonia pla/ysoma I 'lGhR, I 960b: 487 - Type localit"y: LlIid!lll River near Bundu Tuhan at 3,3 00 feel on MOllot Kina Balli , N o rth Borneo
Ansonia pla tysoma. IISGER 1966: 102, 1978 : 312 : M ANTHFY 1983 : 21 ; h GEH & DRlKG 1988 : 463; M ALKM US 1988e: 176: i NGER & ST~b Al NG 1992: 25·37; M ANTHEY I994b: 222: I NGER, STUhl,l l 'lG & TA~ 1996: 3 62 ; INGER & T A" 1996a: 554 ; M "'IKMUS I 996c: 281 ; H o r FMANN 1995a: 9
Diagnosis A small , very lender species with a distinctly depressed body and head ; snout strongly projected,
Fig. 63 Female of Ansonia cf. plalysoma from Sayap, Sg. Wariu.
72
I NGER,
Bufonidae
sloping in profile; I" finger very short, not reaching disk of2nd ; tips of outer fingers spatulate; mandibular spines absent.
Description Males and females 20-25 mm ; TLlSVL 0.51-0.58; head longer than wide; canthus rostral is sharp; lores vertical; tympanum very close to the eye, halrthe latter's diameter; tingers slender with a rudimentary web between first 2 fingers; a low, round outer palmar tubercle, an inner one at base of 1st finger; tips oftoes swollen; toe webbing 1(0), 2(0/0.5-1), 3(lIJ), 4(3/3), 5(l.5); 2 low metatarsal tubercles; skin above and laterally with low warts, sp inose on limbs ; belly and chest coarse ly granular, tlu'oat finely so; males without mandibular spines. Dark brown above and laterally, sometimes with ye llowish dots; limbs with light crossbars; below blackish, throat and belly with small whitish spots; nuptial pads blackish brown.
c c
~ ..... :z: o r
..:
...o:I""'-=;""""-MOoa..-.,....;._ _;a...;,jI;;_ @
Distribution Ecology/Ethology Oulya few sites are known for this species from Sabah Little is known of this species. It was observed in and Sarawak. Mt. Kinabalu: Sayap: Sg. Kemantis, primary forest (150- 1375 m a.s. 1.). Males call Sg. Wariu, Sg. Lnrnotok Besar, Sg. Minodtuhan; Sg. throughout the day with considerable persistence Luidan, near Bundu Tuhan; Poring: Sg. Kipungit from rocks which may be sprayed by water in, or (600 Ill), Sg. Langanan (900-950 m). from the banks of, forest streams, usually in the vicinity of small cascades. At night, A. platysoma can be found on the leaves of low vegetation Fig. XXXVl Sonagram and oscillogram of Ansonia platysoma (up to 50 cm high), sometimes (Sg. Wariu, lOOO m; 2.00 p.m . :+24°C) also at some distance from the streams. CaU: A series of high-pitched tri lls, similar to a clicket song. It usually consists of more than 100 weakly pulsed notes (duration 10-15 illS, intemote interval 100-150 ms ; the interval between the last two notes of a note series is distinctly longer; note repetition rate 5-7 per sec) with a frequency of 7.8 to 8.2 kHz.
kHz
8.89 sec
10 r-----------------------------------------~
~ T~ I 1111 j Iii I " i , :11 i
II
j 1,1 II 't'lil
111111, 'I 1'1111 IH II i I ,
5
73
Bufonidae
Ansonia spinulifer (MocQUARD, 1890) Bufo spillll/!fer MOC(JUARJ), 1890: 160 - Type locality: Kina
dorsolateral warts large, elevated, juxtaposed, with strong spines; characteristic light oval patch between shoulders.
Balu
Description Males up to 41 mm, females up to 50 mm; TL/SVL 0.48-0.53 ; head broader than long; canthus rostralis sharp; lorcs vertical; tympanum two fifths of eye diameter; tips of fmgers dilated into small rOlmd di ks, tips of toes a little smaller; fingers free of webbings; a large palmar tubercle present; toe webbing limited to the bases rudimentary; 2 metatarDiagnosis sal tubercles ; dorsal skin see Diagnosi s above, beA medium-sized lowland species; snout project- low granular; males with 2-3 rows of dark maning, oblique in profile; Jst finger reaching base of dibular spines. tip of2"d; toes almost free of webbings; dorsal and Dark brown to black above, with a light oval or diamond-shaped spot __ _ _ __ _ _ _ _ _ _ between shoulders; a Fig. 65 Allsonia spifllllijer from Poring, 550 m. light streak on groin, and light crossbars on limbs; the light areas sometimes intermixed with brick-red warts; side of llcad light brown; below blackish with white dots, sometimes forming a network; nuptial pads dark brown ; iris golden brown. Tad poles teardropshaped; snout very broadly rounded; body widest in plane before eyes; eyes dorsal; oral djskas wide as body; labial teeth rows foonula 2/3; upper beak divided by a large gap; tail t.apering in the posterior third to a rounded tip; uniformly black; total length lip to 20 mOl.
Ansollia feplopus, (not Bufo /eplOpus
GOl',cTlU:R, 1872) INGER 1966: 110, 1978: 212; MALKM US 1988d: 7,1989: 282 Allsonia spinlllifer, BOULENGI:.R 1894a: 81; VAJol KAMPE.'I 1923 : 79; INGER & D~t""' 1988: 463; INGER 1992b: 229: INGER & STUESL'IG 1992: 41-51; WONG 1994: 25-37; HOFFMM").I 1995a: 9; L'IOER, SWEBIl\G & T AN 1996: 362; INGER & TAN 1996a: 555; MAi.KMus 1996c: 282; LAKIM el al. 1999: 32; INGER et aI. (2000): 16
Ecology/Ethology A. spinul~fe,. occurs in primary forest up to 1000 m altitude. It is
74
Bufonidae
after INGER 199Zb
after INGER 1992b
L __
5mm
10 mm
--- ---
Fig. XXXVU Tadpole of Ansonia spimdifer.
nocturnal and has been observed on the forest floor and in the neighbourhood of streams. Males call from the top of rocks or from low vegetation, but do not form calling aggregations. Tadpoles live in forest streams with swift currents (m: I b, c), clinging to smooth rocks and feeding on the organic film (algae) there. Occasionally, tad-
Genus Bufo
LAURE TI,
1768
This cosmopolitan genus is the only Bornean toad genus containing species outside the Oriental tropics. None of the five Bomean speeics of Bufo is restricted to this island, three of them also occur on the Malay Peninsula. The Bornean pecies are medium-sized to large toads, with rathcr short limb , tocky shapes, thick warty skins, and large parotoid glands behind the eyes. Tips offingcrs and toes rounded, not dilated; no webbing between fingers , all Bomean species with distinct tympanae . The male with a median
-----~
Fig. XXXVIl] Oral disc of the tadpole of Ansonia spiTlu/ifer.
poles can be found ascending vertical, watersp lashed cliffs up to heights of 5 cm. Distribution
Known only from a few sites in Sabah and arawak. Mt. J(jnabalu: Sayap: Sg. Kemantis (1000 m); Por-
ing: Sg. Langanan, Sg. Kipungit, Bat Caves.
subgular vocal sac and nuptial pads; mating in axillary amplexus. The species of Bufo lay egg strings with numerous, pigmented ova in stagnant or slowly flowing water. Their larvae are of a generali ed type, with spheroidal bodies. keratinized beaks, and deep fins . These toads are terrestrial, dwelling in both primary and secondary forests as well as clearing and plantations; Bufo jltxtasper can be found in agricultural land, B. melanosfi cfus in villages and towns. On Mt. Kinabalu, this genus is rarely encountered at elevations above 1000 ill a.s.1. Five species are known from Borneo, two of them occur on Mt. Kinabalu.
-
Tab. 2 Distribution of Blifo -species on Borneo. Saban
Buro asper G RAVEN HORST, 1829 BII/o diverg(!ns P ETF.RS, 1871 Buro jllxtasper [NGER, 1964 Bu/o me/anostic/us S nIl'EIDER, 1799 Bu[o quadriporca/lls B OULEI\GER 1887
• • • •
Kinabalu
• •
Bntnei
• • •
Sarawak
• • • • •
Kalimantan
• • • • • 75
Bufonidae
Key to the Bornean species of Bufo
A pajr of crests between eyes and parietal region .. ............ .. ..... .. .... ....... .. .............. ..... ............. .... .. 2 • If crests between eyes present, they do not continue back onto the parietal region ... .......... 3 2 Parotoids long, sharply raised, their maximum widths at most 1/5 of their length ..... ............ .... . ....................................... ........... ... quadriporcatu.~
(based on INC;ER 1966)
3 A bony ridge curving around the eye to the parotoid gland .. ... ..... ....... ..... ........ meianoslicfus • o such ridge .. ......... ........ ... .............. ............ .... 4 4 Parotoid gland elongated, much longer than eye in diameter ........................................... juxtasper • Parotoid gland round or triangular, about the size of the eye ....... ...... ........ ... ........... ............... .. asper
• Parotoids triangular or oval in shape, their maximum widths at least 2/5 their lengths ........ . ................................... ............ .. ........... divergens
Bufo divergens
PETERS,
1871
811/0 diverge" , P El FR~, 1871 : 579 Type locaUly: Sarawak BI10 biporcows. (not Bldo hiporca/tls GRA VE'IHORST, (829) M OCQUA RO 1890: 158; VAN KAMPE'I 1923: 88; S\,III H 1925b: 30, 1931 : 30 811/0 biporcorlls biporcoll1s. IN(ibR 1956: 394 Bulo biporcaflls di vergem . II\'OER 1966: 60, 1978: 3 12; MA I\T IIEY 1983: 20 811/0 dil'ergeTls. l'IorR, S llJHIING & TA'I 1996: 362; lliG CR & T" 1996: 555
Diagno is A relatively mall- ized, stocky toad; nout with a small median bulge at the tip, projecting, and oblique in profile; a pair of narrow bony crests between the eyes; toes webbed over half their lengths; 1" finger longer than 2nd.
Description Male up to 45 mm, females up to 65 mm ; T LlSVL 0 .32 -0.48 ' head wider than long ; canthus rostralis harp, lores vertical; tympanum about 3/ 5 the diameter of the eye; tips of fingers and toes blunt, not swollen; supernumerary metacarpal tubercles
76
Bufonidae
present; toe lengths 4>3>5>2> 1; toe webbing J(l), 2(1/1), 3(1.5/ 1.5), 4(3/3-3.5), 5(1.5); inner metatarsal tubercles oval; a smaller, rounded outer one; skin above and laterally with many small, conical sp iny warts; a row of enlarged warts between eye and groin; a row of spinose tubercles on inner edge of tarsus; venter coarsely granular; parotoids triangular to oval. Reddish brown above, often with dark pots; dark interorbital marking; limb with dark crossbars; nuptial pads black; below yellow or light brown , with or without dark mottling; iris golden brown with a dark network. Tadpoles: Head-body portion of tadpoles oval, slightly flattened above; eyes and nostril dorsal, oral disk ventral, subtermina l; no expanded lips (in contrast to the larvae of B. juxta:,per); labial teeth rows formula 2/3. Head-body portion and caudal muscle dark brown to black; venter without pigmentation; dorsal fin dusky, ventral fin only along the base so; total length 15-17 mm . Ecology/Ethology Tllis toad Lives in forest floor litter and is confined to lower elevations « 1000 m a.s.I.). It can be found in both primary and secondary forests, plantations, and village clearings. B. divergells is mainly active at night, although it was also observed hunting during daylight hours. It feed on a wide variety of arthropods, favouring ants and termites. The males call at night, forming calling communities along the edges of spawning sites. Tadpoles occur in the shallow areas of stagnant or slowly flowing water in small streams (m: Ie, t). They are bottom suspension feeders (ING ER 1985, L986). Call: A series of short, raspy trills. Sonagram not published. Distribution Bulo divergens i known from Borneo, Sumatra, and the Natuna Islands. Mt. KjnabaJu: This species was mentioned for Mt. Kinabalu only once, i.e. in a check list by MITH (1931), as "known to inhabit Mt. Kina Balu above 3000 feet", which refers presLUnab ly to the Kiau region. Recently, tadpoles were found near Poring.
Bufo juxtasper ING ER, 1964 I NG~It. 1964: 154 - Type loca lity: Sungei Tawan, Kalabakan. Tawau District. Sabah Bu/a usper. (not BII/v asper GRAVENIIORST, 1829) M OCQUA RD 1890: 158; VA' KA~ IP EN 1923 : 82; S~11'I H 193 1: 30 Bllji) jllXlasper, I "GE R 1966: 69; MATSU I 1979: 313; MA "'T11I',Y 1983: 21; M AI"~I US 1985: 13. 1987: 277. 1988b: 9, 1988c: 176. 1992a: 106; I 'IOER & STUE!}1 '0 t992 : 41-51 ; YlALKMUS 1994b: 220; WO,G 1994: 25-37; HOfFMA.\; 1995a: 9; I NGER, STUIillING & T AN 1996: 362; I NGER & T A.'i 1996a: 555; L AKIM et al. 1999: 32: INGER et a1. (2000): 16
Bufo jlL'(laspel'
Diagnosis A very large, stocky toad, with a broad, blunt head; ear-openings rather small ; a thick supraorbital, but no parietal crest; toes fully webbed, except for the 4'h toe; I " finger equal in length to 2nd. Description Males up to 125 mm, females up to 215 mm; TUSVL 0.42-0.48; head wider than long; snout pointed; canthus rostralis rounded, lores vertica l; tympanum smaller than one third of the eye diameter; tips of fingers and toes swo ll en; subarticular tubercles large; supernumerary metacarpal tubercles present; toe lengths 4>3>5>2=1 ; toe webbing 1(0),2(0/0) 3(0/0), 4( 1.5/ \.5), 5(0); metatarsal tubercles large; a sharp tarsal ridge; skin above and on tbe sides covered with small to large, fOlmd warts; parotoids large, as long as their distance from the tip of the snout; ventral sides coarsely granular, each granule tipped with a melanic spinule. Grey to earth-brown above, Lighter on tbe sides; venter greyish white with dark mottling and spots; nuptial pads black; iris yellowish golden with a fine black network. Tadpoles: Head-body of tadpoles oval; eye and no trils dorsal, oral disk ventral; labial teeth rows formu la 2/3(1-2) . Head-body, and caudal muscle blackish brown ; disk region greyish; venter golden yellowish; total length up to 20 !TUll . Ecology/Ethology Bt10 juxtasper lives in primary forests and disturbed habitats as ociated with the activities of man (secondary forests, agricultural regions, villages) . The altitudinal distribution range is moderately wide, and it is usually encountered below
77
Bufonidae
78
Bufonidae
::i @~~~~~~~~~~~~KW~~~~~~~~
."-]g. ~9 T~pol ~O~Blifo juxtasP3r from A~ehl- Sumatra.
Call: Tlte call consists of a single, resonant, hoarse cbirp. The Dusun name for B. juxtasper imitates the sound of two calling toads as "buangkut" . There are two types of calls, a shorter and a longer one . The fonner lasts 0.07 to 0.16 s and is repeated 2 to 2.8 times per second; dominant frequency 540-730 Hz; hannonics are not evident. The longer call is a typica l trill in which a shorter note (0.05 s; 6 pulses) is foUowed by a long one (> I s; 130 pulses), dominant frequencies slightly higher than in the shorter call (MATSUI 1982a) .
Distribution 1000 m a.s.l. This species is active by day and night. Borneo and Sumatra. Mt. Kinabalu: Common in Toads of this species were found on gravel banks, disturbed habitats along the boundaries of the Park (vall ey of Sg. Kadamaian near Kiau, Kebeyau; large boulders, and fallen logs on the bank and in the middle of the lower Sg. Langanan and Sg. KadaPoring; lower Sg. Langanan, Sg. Mantukungan; maian, but also far away from water on the leaf Mamut Copper Mi.ne), but rare in primary forest (Sg. Kipungit, Sg. Sasapan; Sayap; lower Sg. Silaulitter of the forest floor. When captured, this toad Si lau, 1500 m a.s.I.). produces a large quantity of a fou l smelling, white secretion. Its diet includes a wide variety Fig. 70 Oral disc of Bu/n juxta,spe,. from Aceh, SumaTra. of spiders, worms and insects, particularly ants and termites . In the Poring region it was observed along ant paths formed by the large ant, Camponotus gigas, consuming considerable quantities of these insects. The males call from along slreanl banks. The species name is onomatopoeic in being reminiscent of the call of the species. The strings of gelatinous, brownish grey eggs are attached to aquatic plants and rocks in spring basins and ca lm sections of streams and rivers (m: I e, 1).
79
Bufonidae
Genus Leptophrylle FITZI
GER,
1843
The genus Leptophfyne compri cs two species: L. cruentata TSCHUDI, 1838. of Java, and L. borbonica ( K UHL & VAN HAS ELT, 1827), of the Malay Peninsula Sumatra, Java and Borneo. It is the only genus of Indo-Malayan bufon ids w ith an arciferofi nn istemaJ pectoral girdle.
Lepfophryfle borboflica
(K UHL &
VA
HAsSELT, 1827)
Hy laplesia borbonica K [;IIL & VAN HA SSELl in SOI1.4(iP-L, 1827: 294 - Type locality: "East Indies", restricted to ".lava" by INGER (1966) OCOplll:Vlle borbo/lica. I I\(,ER 1966: 75, 1978: 312; rYwnlF Y 1983: 21 Lep/op/"Jme borbonica, INGER & STUEBC\'O 1992 : 41-5 I; i KGER.. STIIEBlNO& T AN 1996: 362; INGER & TAN 1996a: 554
Diagnosis A sma U, slender bufonid with very long, slender limbs; bead without bony crests; snout short pro-
80
jecting slightly over the moulh; tympanw11 distinct fingers and toes long, with small. round disks on the tips; no webbing on the fingers, toes webbed over half their lengths. Description Males up to 31 mm. females up to 50 DlI11; TLlSVL 0.56-0.70; head as wide as long; canthus rostralis sharp, lore vetiical tt'aig ht; tympanwn oval, one third the eye diameter ; fi nger lengths 3>4>2>1; toe lengths 4>5>3>2> 1; subartic ular tubercles large ; toe webbing ](1),2(1/1-2),3(23/2-3), 4(3-3.5/3-3 .5), 5(2-3); inner metatarsa l tubercle oval. outer one rou nd; nuptia l pa ds present; skin above and on the sides with numerous warts; paroto ids weakly developed; a row of dorsolateral glands; lower side granular; a row of conjcal tubercles on the oUler ide of the lower ann and tarsus; males with a median su bgular vocal sac. G rey ish to brown above and on the sides, with darker and lighter markings, usually forming an X in the centre of the back; limbs with dark crossbars; ventrum yellowish, mOltled with brown; throat
Bufonidae
,--------
after B ERRV 1972
Fig. IXL Tadpole of Leptop!u"t..lle
b~·bonica .
edged with black; head-body and caudal muscle black; fins blackish grey; total length up to II rom.
~
s::;
E ~ :;
:i I)
Fig. 72 Leptophryne borbonica (male) from North Swnatra,
nc~r
Berastagi, 1450 m.
greyish to brown; undersides of the legs grey to yellow with brown spots; iris golden brown. Tadpoles: Head-body portion of tadpoles oval, slightly depressed; eyes and nostrils dorsolateral , oral disk ventral, subtennina1; lower lip bordered by papillae all around; labial teeth rows formula 2(2)/3 or 2(1-2)/3; beaks finely serrated, white, narrowly
Genus Pedostibes G "
THER,
1875
The species of the genus Pedostibes are native to the Malay Peninsula, SLUnatra, and Borneo. This genus contains medium-sized to large, moderately stout toads ~tb long legs. Parotoids present or absent. Fingers long dilated at tips into truncate disks; tips of toes round and swollen. Tympanum present or absent. Males with median subgular vocal sac. The species produce small, pigmented eggs
Ecology/Ethology early nothing is known on the natural hi tory of thi specie other than it is terrestrial and occurs in primary forest where it favours the proximity of forest streams, especially eepage areas, from sea level to 1500 m altitude. The eggs arc not arranged in strings, but deposited in clumps of 200-300 in calm sections of streams (m: If). The tadpoles are benthic feeders . Distribution Malay Peninsula, umatra, Java, and Borneo. Mt. Kinabalu: The onJy hint towards an occurrence on Mt. Kinabalu is found in INGER (1966) as "Kota Belud District, Mount Kina Balu", which presumab ly refcrs to the Sg. Kadamaian region.
in trings, which are deposited in calm si de pools of small forest streams. The onJy tadpoles known are those of P hosii; they are benthic feeders and simi lar to those of the genus Bufa. Pedastibes are arboreal and occur in primary and old econdary forests at altitudes ranging from sea level to 1000 m; nearly nothing is known with regard to the vertical distribution on Mt. Kinabalu. Three or four species (see remark) are found 011 Borneo, two of them also occur Oil Mt. Kinabalu.
81
Bufonidae
Key to the Borneau species of Pedostibes (based on INGER
Tarsal ridge present ..... ............. ................... ...... 2 • No tarsal ridge .......... ............................ .. .......... . 3 2 Parotoid oval or circular; lateral and medial borders convex ............................ .............. .. . rugostls
• Parotoid triangu lar; lateral border straight or concave, medial border convex ................ .. hosii
3 Parotoid glands present; tympanum visible ........ ..................................................... .... ..... . everett;
• No parotoid glands, tympanum invisible .... ...... .. .. ........... .. .. ..... .. .. ........ ..... .... .. ............
maculatlls
1966)
Remark: [NGER (1958c) described the species P rogostlS which, however, is hardly distinct morphologically from P e veretli, stating: "The main differences being the absence of a tarsal ridge, the smaller digital disks and the greater separation ofparotoid and eyclid in everetti" ( ING ER 1966). Further consideration of variation in P rogosus demonstrated (MANTH EY & GROSSMANN, 1997), that none of his diagnostic characters were distinct from character states exhibited by P. everetti.
Tab. 3 Distribution or Pedostibes-species on Borneo.
-1I - _ I +- I 1 '- - I - -I _- _ _-_ \- -C-r---Sabah
Pedost;bes everett; ( B OULENGER, 1896) Pedostibes hosii (BOUUi NGbR, 1892) Pedost;bes maclliatlls (MOCQUARO, 1890) PedoSlibes rugoslLV l NGER, 1958
I
Pedostibes everetti
( B OULE GER,
Neclophry ne everelli BOL'Lh~GER 1896b: 450 Mount Kinabalu, Sabah Neclophry ne everelli, VAN K .....\\PfN 1923 : 68;
1896)
'I)'pe locality: MI1 H
1925b: 31.
1931: 30 Pedoslibes everelli. BAlUIOUR 1938: 192; I l'GER 1966: 90, 1978: 312; M i\);THEY & D ENZE R 1982: 17: MAl\THEY 1983: 2 1; MAU:'MUS 1988c: 176; I 'GER & STUE8I NG 1992: 41-51; [ "GFR. S T U Ii 81 ~G
&
T AN
MANTHEY & GROS
1996: 362; I NG I' R & 1997: 38
rAN
1996a: 5 5;
\\ANN
Diagnosis A medium-sized to large, moderately stout Pedo tibes with egg-shaped parotoids; kin of the upper surfaces very warty ; no tarsal ridge; above green with large brown pots. Description Up to 87 mm; head broader than long; no cranial ere ts: tympanum one thjrd the eye diameler; fingers webbed on the bases: finger length 3>4>2> I:
82
-
I
Kinabalu
I
Brunei
-
I Sarawak
i
-
I Kalimautan
-
toe length 4>3>5>2> J; toe webbing J (0), 2(0/0, 3(0/0) , 4(2/2) , 5(0); two metatarsal tubercles. Green above and on the sides, with small to large, irregularly scattered , dark brown to reddish brown spots; limbs green with brown crossbars; lower sides cream-coloured, immacu late or mottled with brown . Ecology/Ethology The only report on field observations is found in MANTffEY & GROSSMA."IN (1997) reading in translation: "On Mt. Kinabalu, in the environs of Poring, we found nearly 200 juvenile toads on tbe bank of a partially torrential stream on a clearing in the rainfore t 650 m a.s. l.) in Augu t. Here, the stream was blocked by wedged drift wood which had caused it to form a pool. ... The juveniles measured 10-15 mm and presented themselves openly on the sparseJy vegetated ground with only little leaf littcr." omc spccimen were raised in terraria. While
Bufonidae
Above
Fig. 73 Pedostibes everetti, nearly 20 years old from Sg. Kipungit D, near Poring. Left
Fig. 74 Pedostibes everetti, juvenile from the same place as above.
Pedostibes maculatus (Mo they were terrestrial and diurnal during the initial 4-6 weeks, they then began to climb and shifted their activity to the hours of dusk and night. After only one year they had grown to 60-82 mm SVL.
Distribution Tills species is only known from "Mowlt Kinabalu, Sabah" (BOULENGER 1896, rnGER 1966), presumably from the Kiau region. A more recent record stems from the Sg. Kipungit II region (650 m a .s. l.) (MANTHEY L983).
Q ~U),
1890)
Neclophryne maculala MocQuARD, 1890: 162 - Type locality: Kina Balu NeclOph,yne maculara, VAN KAMPEN 1923: 71 PelopiJryne macu/ara. BARBOUR 1938: 193 Pedostibes macu/afUS, INGER 1966: 92, 1978: 3 12; MANTHEY 1983 : 2 1; I NGER & STUEB ING 1992: 41-51 ; INGER, STLIillING & TAN 1996: 362; ING6R & TAN 1996a: 556
Diagnosis A small, slender Pedostibes with long limbs; fingers long, tips with trwlcate disks; head w ithout bony crests; tympanum invisible; parotoid glands absent; DO tarsal ridge.
83
Bufonidae
Description Males up to 42 mm, females up to 51 mOl; TLlSVL 0.54-0.60; snout truncate, projecting in profile; finger lengths 3>4>2> 1; subarticular tubercles moderate large· fingers webbed at the bases; tips of toes not dilated; toe lengths 4>3>5>2> 1; toe webbing 1(1), 2(1 / t) , 3(1 /1.5), 4(3/3),5(1- \.5) ; an oval inner and a round outer metatarsal tubercle; numerous round warts with black spines above and on the sides. Brown with irregular dark spots above and on the sides; li~bs with dark crossbars; greyish below. EcoiogyfEthology There is no rnformation on the life history of this tree toad. Distribution Distributional data is limited to "Mt. Kinabalu, Sabah".
Genus PelQphrylle
BARBOUR,
1938
The genus Pe/ophryne is distributed on the Malay Peninsula, Sumatra, Borneo, and the Philippines. The dwarf toads represent the most distinctive genus of Oriental bufonids: Very small toads (SVL < 31 mm) with a moderately stout body and slender limbs; hands and feet with flesby webbings, partly reduced phalanges; tympanum visible , parotoid glands absent. Coccyx fused to sacral vertebra and expanded dorsally; only seven presacral vertebrac. Males with subgular vocal sac; nuptial pads 1md mandibular spines may be present or not.
I I I I I I I I
- --- - - - - - - - --
afle r MOCQU'AD 1690
I- - -- - - - - F ig. XL P~ostibes maculatus
The females produce large, non-pigmented, whitish ova rn smaU clutches (usually les than 20 eggs). Suitable spawning sites are rainwater-filled depressions and phytothelmes (e .g. leafaxjis and tree holes). Larvae small, subterminal, circular oral disk with reduced labial teeth and weak beaks; piracle not visible. They are non-feeding and subsist entirely on yolk. Pe/ophtyne are terrestrial and can be found in low vegetation in primary and old secondary forests. The altitudinal distribution of tills genus is wide-ranging, stretching from sea level up to 3100 m a.s.1. Eight species are known from Borneo, but only one occurs on Mt. Kinabalu.
-
Ta b. 4 Distribution of PelophrYfl e-species on Borneo. Sabah Peloplrryne api DRING [983 Pe/ophryne brevipes (P ETE RS. 1867) Pelopillyne exiglla BOI:.ITGIlR, 1901 Pelophryne g lletllheri (BOL'Ll'NGER, 1882) Pe/ophryfle macroli.. (BOULf.NGFR, 1895) Pel ophryJle misern (MOCQUARD 1890) Pe/ophrYll c rhopoplrilillS INGER & STUEIlI~G 1996 Pe/ophrvne si$:flala (BOULENGER 1894)
84
Kinabalu
Bruoei
• •
•
&,rawak
•• ••• • ••
Kalimantan
•
• •
Bufonidae
Preliminary key to the Bornean species of Pe/ophryne Tips of fingers dilated ....................................... 3 • Tips of fingers not dilated ................................. 2 2 Dorsum black, without extensive light areas ... api • Dorsum brown, with lighter areas.. ........ misera 3 Tympanum at least four fifths of the eye diameter ................................................................ macrotis • Tympanum at most three fifths ofthe eye diameter ...................................... ..... ............ ......... ........... 4
4 5'h toe webbed up to the disk .............. guentheri • 5dl toe not webbed up to the disk .................. .... 5 5 A yellowish white stripe between eye and tympanum ....................... ............................ . brevipes
• No such light stripe below the eye ... rhopophilius
Pe/ophryne misera (Mo
QUARD,
1890)
Nectopilly ne misera MOCQuARD, 1890: 161 - Type locality:
"Nord de Borneo". restricted here to "Kadamaian River, Mt. Kinabalu" (based on data provided by WlIITEH~AD 1893) Neclophryne misera. WHITEHEAD 1893: 177; BOULb'GER I894a: 81; S~lInt 193 I: 14 Nectophryne gllentheri. (not Nectophryne gllelllheri BOLLENGER, 1882) BARl LETI 1894: 204 Pelopilly ne misera. BARBOUR 1938: 193; J GER 1954a: 250, 1966: 88, 1978: 312; MANTIIEY J 983: 21; MAlK.\lu$ 1985: 12, 1987: 277, 19 Sc: 176: (" ,FR & STLEBING 1992: 41-51 ; IN(WR, SlUFBING & TAN 1996: 362: MALI:.\lUS I 996c: 280; I"GER & T"" 1996a: 556; INGER & Sn.EBING 1997: 87 PeloplllYlle brevipes, (not Hylaplesia brevipe~' PETFRS, I R6 7) MALKMLS 1989: 183, 1991a: 27, I992a: 110, 1994b: 221. 1996a: 129; INGER c l aL 2000: 17
Remark: Roux (1906) placed Pe/ophryne exigua in the synonymy of P. guentheri. INGER (1966) referred it "in the absence of distinguishing characters" to the synonymy of P. brevipes. In the latest check list of the frogs of Borneo (INGER & TAN I 996a), P. exigLla appears again as a valid species . P. signata was likewise synonymised with P. brevipes by INGER (1966), does consequently not appear in said check list (I 'GER & TAN 1996a), but is treated as a valid species by INGER & STUEBIl'G (1997) without an explanation. For the purpose of the present book, P. exigua and P. signata have been included in the species list with reservations, but excluded from the key as the latter is based on distinguishing morphological traits.
0.5-1.), 4(2/2 -2.5), 5(t)· subarticular and metatarsal tubercle hjdden or absent; skin above with irregularly cattercd, mall , conical warts, forming fragments of dorsolateral rows; sides and venter coarsely granular; males with or without mandibular spine and yellow to brown nuptial pads. Upper ide blackish brown, reddish brown to ochre; dorsum occasionally with 1-3 large, dark spots of variou shapes and an interorbital , trapezoid-shaped marking; hind limbs with dark cro sbars; dark flanks often harply set off from li ghter dorsum; a light supra labial spot between eye and tympanum; lower sides yellowish white, with large, irregularly
Diagnosis
A mall montane spccies with a modcratcly tout habitu and slender limbs; snout truncatc, vertical in profile; tips of fingers and toes not wider than bases of phalanges; fingers with fleshy webbings. Description Male up to 21 mm, female up to 23 mm; TUS VL 0.34-0.41 ; head as wide a body; canthus rostralis sharp, concave; lores vertical; tympanwn half to two thirds of eye diameter; finger webbing 1(0), 2( I1I), 3(2/2),4( 1); subarticular and palmar tubercles hidden ; toe webbing thick, 1(0.5), 2(0.5/0.5), 3(0.5-11
85
Bufonidae
i :::;
a: @
Fig.:...76
Pelop~ryne
misera from Liwago tTail, 1550 m.
scattered brown spots which may fuse to fonn a crude and incomplete reticulation; iris golden bro"vn, with dense black marbling. Newly metamorpbosed toadlets with highly contrasting colour pattern: blackish brown laterally, a white dorsolateral line between tip of snout and insertion of hind limb; upper side whiti h grey with large blacki h brown marking; upper side of hind limbs black with narrow white crossbars. Tadpoles: Head-body of tadpoles ovoid; eyes dorsolateral, oral disk ventral, subterminal; labial teeth rows formula I/O; bcaks weak; spiracle not tubular; body and caudal muscle blacki sh brown, fins lighter; total length 13 mm ; fre hly metamorphosed toadler 5.8 mm.
86
Ecology/Ethology Pelophryne misera is locally abundant in tbe montane forests of Mt. Kinabalu (1450 to 3100 m a.s.I.). While the toad was found active, foraging for food, and calling during the day at subalpine elevation it could never be discovered outside its hiding places (beneatb leaf litter, in rock crevices, holes in the ground, dead pitchers of Nepenthes) before the on et of dusk at altitudes below 2000 m. Its locomotion is a slow striding galt with the body rai cd high. Moving around like this, a foraging toad will plunge its head deeply into mosses, lichens, and leaf Iitter every now and again . [ts prey (arthropods < 5 mm; ants, spiders, hymenoptera, wood lice, etc.) is stalked, visually fixed, and then
Bufonidae
"! ~
'iii
::;;
a' @
Fig.: 77... EglE' of PeI9phrylJ ~m!!e!:a
!rom Sg. liwag~ . Fig. 79 Pelophly ne misera, three days old ,
".--.._.......- - - ,
"shot down" with the tongue. Males usually call from the leaves of shrubs, 10-50 em above the ground. Until now, the clutch of only one female from Sg. Liwago has become known. It was discovered in mid-August, in the depression of a dead leaf, in which water had accumulated and consisted of 10 yellowish white eggs with diameters of2.8 mm on average. The larvae emerged after 16 days, one of which metamorphosed under temuillm conditions after another 44 days.
Call: So far, single notes and note series have been recorded. Single note (+ 19°C) repeated at intervals of4-10 s, duration 90-110 ms, frequency 3.54.5 kHz; the note is pulsed and has a short grace note. Note series (+ 19°C) consists of 7 -12 pulsed individual notes (40-50 ms) with internote intervals of 100-130 ms; ------Fig. XLl Sonagram and oscillogram of Pelophryne misera at 7-8 ,kHz and peaks up to 14 kHz (Sg. Liwago, 1550 m; 8.00 p.m . + 19°C) these calls belong to the highest tones produced by bllfonid anurans.
Distribution This species is known onJy from two localities on Borneo, i.e. Mt. Murud (Sarawak) and Mt. Kinabalu. Mt. Kinabalu: Marei Parei spur 0550 m) , slopes of Sg. Liwago valley (1450-1600 m), Kamborangoh (2200 m) , around Pakka Cave (3000-3100 m) .
kHz
20
43,56 sec
.----------------------------------------, 4
10
t
•
,.
c
• I
It • ,
II H·t'I!lHt 87
Megophryidae
Megopbryidae The Oriental megophryids are di tributed from Pakistan east to China, the unda Islands and the Philippines. The family is characteri ed by a arciferous pectoral girdle and a cartilaginous sternum and omostemum. The pre acral are not fu ed. The sacrum ha sub tantially expanded <,iiapophy es and is fused with the coccyx; the epicoracoids overlap at mjd-line. The parahyoid i absent, maxillary teeth are pre ent, vomerine teeth absent (except in the species of the genus MegophIJ's). The pupil is vertically elliptical , except of Opillyophryne. Mating take p lace in an inguinal amplexu . All Bornean species have aquatic larvae, most with beaks and labial teeth (except in the genera Leptobrachel/a and MegophfYS), and depre sed, spheroidal body shapes (except in the genus Megophrys). Megophrys have funnel-mouthed tadpoles. Bomean megophryids are terrestrial and live in leaf litter layer of primary rain forests, particularly in the vincinity of brooks. On Mt. Kinabalu they occur up to altitudes of 2200 m (Lepfobrachella baluensis. Leptobrachium gUflungense). Oil Borneo, the family is repre ented by 4 genera and 22 species, on Mt. Kinabalu by 4 genera and 13 specie .
Key to tbe Borneau genera of Megopbryidae
A
B
1 Denna) appendage on upper eyelid and/or snout (A) ........... ................ .... ............. ........ Megophrys • No such appendage(s) ........ ............................... 2 2 Head conspicuously wider than body (8) .......... . .............. .................................... Leptobrachium • Head not wider than body .................. .......... .. .. 3 3 Tips of digits swollen, with pointed disks (C); adults never larger than 25 mm SVL .. .. ........ .... .. ........... ... ..... ............................ ... Leptobrachella • Tips of digits not swollen (D); adults larger than 25 mm SVL ...... .... ..... .... ....... ...... .... ... Leptolalax
Genus Leptobrachel/a
SMITH,
1925
Representative of the genu Lepfobrachella are only known from Borneo and the small islands Pulau erasan and Pulau atuna Besar (the latter is home to the endemic L. nalllllae GCfl\THER, 1895) we t of Borneo.
88
°/-;· -:' ;
' .1
Thi genu includes the smallest forms (SVL < 23 mm) of megophryid fro g . Habitu moderately stocky to slender with mooth or shagreened skin on dorsum and venter and granular flanks . A small, conical pectoral gland pre ent medial to arm in ertion. Limbs slender, fingers and toes ending in pointed disks; large inner palmar tuberc le, long, low inner metatarsal tubercle; nuptia l pads absent.
Mego phryidae
Tab. 5 Di stribution of Leplobrachelfa-s pecies on Borneo. Leptobrachella baluellsis SMml, 193 1 Leptobrachella brevicrus D RING, 1983 Leptobrachella mjobergi S\1ITII, 1925 Leptobrachelfa palmata )"G[R & STlJfB II'G, 1991 Leptobrachellu parva D RING, 1983 Leptobrachella serasall ae D RI'IG, 1983
I
All species, with the exception of L. pa/mata. have rudimentary webbings confined to the bases of the toes. Maxillary teeth present, vomerine teeth absent; tympanum di tinct. Larvae have a single filter cavity on either side; they lack the usual fri lled gill flJter rows, and also have no labial teeth. The head-body section is extremely elongated. The larvae of all species may be very similar.
Preliminary key to the Bornean species of Leptobrachella (based on
INGER
&
S TUEBrNG
199 1)
3rd and Slh toes webbed up to the terminal disks . ................................................................ palmata
• Toes on ly with rudimentary webbing ............... 2
2 Dark lateral stripe between eye and groin continuous ......... .......................... ......... ......... serasanae • With or without small, isolated, dark lateral spots .......................................................................... 3 3 Belly dark with many small, irregular, light spots; a dark streak on the supratympanic fold ... baluensis • Belly light, with or without a faint dark network; no supratympanic streak ................................... 4 4 An aligned ventrolateral row of elongated, slightly separated whitish glands between axilla and groin .............................................................. mjobergi • Ventrolateral glands round, isolated, not aligned
..... ..................................................................... 5 5 Adu lt males IS-17.S mm SVL; no tarsal gland .. .................................................................. parva • Adult males 19-21 mm SVL; tarsal gland present ............. ......... ...................................... brevicrus
Sabab
Kinabalu
•
•
Brunei
Sarawak
•
• •
•
I
Kalimantan
• • • • •
•
Adult dwell in leaf litter and appear on the banks of mall , rocky, clear treams during the breeding season . Their larvae are macropbagous feeders adapted to a life in well-oxygenated water dweUing inter tices of gravel and rock crevice at the bottom of rime . Six species are known from Borneo, two from Mt. J(jnabalu.
Leptobracltella baluellsis
SMITH ,
1931
Leplobrachella buluclIsis S~m'H, 193 1: 12 - Type locality: Kambomngah Nesobia mjobergi, (nol Leplobrachella mj6bergi SMITH, 1925) I"GER 1966: 49 (pari), 1978 : J 12; M ANTflH 1983 : 20 (part) Leplobrachella baluellSis, DRl~G 1983: 96; INGER & Sn..EBlNG 1992: 41-51 ; INGER & TAN 1996a: 552. 1996b : 72; LAK IM et a!. 1999: 33; II'GEK et a!. 2000: 16 Leplobraclrella mjobergi, (not Leplobrachella mjObergi SMfTIl , 1925) HorFMANN 19953: 8
Diagnosis A highland specie with "chirp" call. Flank glands scattered or fused in an irregu lar, short, ventrolateral series.
Description A small terrestrial frog; S VL of males 15-19.6 mm , females up to 23 mm; T LlS VL 0 .S OO-0 .527; tips of finger s a nd toes sl ig htl y dilated , sharpl y pointed ; finger len g ths 3> I >2=4 ; toe len g th 4>3>5>2> 1; su barti e ul ar tubercle s weak, but prcsent; dorsum mooth; flanks s lightly granular with s mall pale glands, the lattcr elongated in th e po terior portion of the lower flank , forming ventro lateral eries; males with a median subgular vocal ac.
89
Megophryidae
Di tribution There are only a few records [Tom Sabah and Stlrawak. Mt. Kinabalu: Sg. Lumotok Kecil/Sayap (900 m); Kamborangoh (2200 m).
Leptobrachella parva DRJl\G, 1983 I.eplobrachella parva DRING, 1983 : 89 Type locality: Gunung Mulu , Fourth Division. Sarawak Leptobrachella pan 'a. MALKMls I 994b: 219
Diagnosis A lowland spccies with a " buzz" call ; flank glands scattered and not in ventrolateral series I ike in L. baluensis .
.:: c:
I
,,: 0 """,_"""""
F ig. 80 Male of Lepfobrachella balllellsis from Sayap (900 m).
Brown above with a dark interorbital bar and a capular W-shaped marking; a distinct, dark supratympanic spot or streak; tubercles on flanks pale, dark outlined; limbs with dark crossbars; throat and belly greyish, with diffuse dark pigment.
EcologylEthology The fcw records from Mt. Kinabalu so far all came from the levels between the oak-che mut forest and the mossy forcst (900-2200 m) where thi specie inhabits the leaf litter layer and rock rubble near streams. The maJes call in the late afternoon and at night from between rocks or are perched on the leaves of shrubs or in low bush vegetation usually in the vic inity of streams with strong currents.
Call: (+22°C) Call duration 0.14-0.15 s; each call begins with a soft cl ick, followed by a series of 14 pulses (0.06 s) at a constant rate of275-280 pulses per second; dominant frequency 12-14 kHz, with a strong fundame ntal below 2 kHz; frequency modulation absent, but amplitude mod ulation present; call interval 0.38-0.40 s (DRL~G 1983a).
90
Description Very similar to L. baluensis; males 1518 mm, a gravid female 17.8 mm; TLlSVL 0.497-0.548; in contrast to L. baluellsis there arc no subarticular tubercles; no dark tympanic mask or stripe on the flank, and flanks not potted with black, but with small dots and black areas around wh ite glands.
Ecology/Ethology Calling males were found in hidden place on pebble beaches and boulders at the edge of streams and on rocks protruding from the water in dipterocarp forest. L. par1la was observed on the leaves of low bushes (30-50 cm) along Sg. Langanan.
Ca ll: (+22.5°C) The ca ll is a buzz lasting 0.150.16 s, consisting of 24-28 pulse . The pulse rate increases during the call from an init ial 50-67 pulses per second to 300-330 pulses per second towards the end. Dominant frequency 11.2-12.8 kHz; no fundamental frequency ; frequency modulation and intensity modulation are not evident (DRlNG 1983a).
Distribution Only a few records exist from Sabah and Sarawak from between 150 and 850 m a.s.1. Mt. Kinabalu: Sg. Langanan (850 m).
Megopbryidae
Genus Leptobrachium
TSCHUDJ,
ubstantially larger than male, with relatively shorter tibiae and narrower heads . Their eggs are pigmented. The tadpoles of Bornean Leptobracl!iulII are mod-erately large to large (70-90 mm) with a long, leafhapcd tail. Head-body portion robust, oral disk ventral, subterminal. A continuous fringe of short papillae surrounds both lip s. Black beaks vshaped, coarsely serrated; at least four divided rows of labia l teeth on upper lip. Eyes dorsolateral, spiracle mid-lateral; anal tube dextral, united to ventral fin. These larvae are macropbagous feeders and occur in relatively calm reaches of treams with weak to moderate currents. Adults can be found among the forest floor leaf litter of primary rain forests and on clearings. Five species are known from Borneo, three of them from Mt. Kinabalu.
1838
The species of th is genus are widespread in Southeast Asia, ran jng from southern China to the Philippines, the Greater unda Islands and Bali. Its Bornean representatives are moderately large to large stocky frogs with a head broader than, or as wide as, the body; snout not projecting, tympanum visib le. Maxi llary teeth present, vomerine teeth absent. Tips of fingers and toe rounded (except in L. nigrops); inner palmar tubercle circular, not extending along the fir t metacarpal oval or round, flat axillary glands and small femoral glands; skin of body surface with a network of low ridge delimitating closed cells. No dermal palpebra l projections. Males lack nuptial pads and have a median subgular vocal sac. Females are
Tab. 6 Distribution of Leptohrachium-species on Borneo. Sabah LeplObrachillm abbott; (COCHRAN. 1926) Leptobrac1,illm gunllngense ~ALKM US. 1996 Lep/obrachiwn hendrick.",ni TA YLOR, 1962 !:!:E!!!!!rachillm mOlltallllm FlsClfrR, 1885 LeplObrachillm nigrops BERRY & HENDRI CKSO:-4, 1963
Preliminary key to the Bomean species of Leptobrachium Upper part of iris orange ......... ....... hendricksoni • Upper part of iris blackish ............................... 2 2 Tips of fi ngers sharply pointed; subarticular tubercles present ... ...... ..... ...... ..... ..... ...... nigrops • Tips of fingers rounded; no subarticular tubercles ....................... ... ................................................. 3 3 Lower side with white and black pattern; weakly bluish ring around the eye .......... ............ abbotti • Lower side with no such pattern; distinct white ring around the eye ............ ... ...... ..... montanum • Morphologically indistinguishable from L. monlanum, but with significantly different bioacoustics .... .... .... .... ..... ... .... .. ..... ... ....... ..... .. .. . gunungense
• • •
Kinabalu
• • •
Brunei
•
•
Leptobrachium abbotti
Sarawak
• • • •
Kalimalltan
(COCHRAN,
•
• 1926)
M egophrys abbolli eOCH RAJ' , 1926: 446 - Type loca lity: Ba1ikpapan Lep/obrachium hasselli, (no 1 Leptobmchillm hasselfii TSCfI UDI, 1838) INGER 1966: 29 (part), 1962/63 : 46, 1978: 312 (part); MATSLI 1979: 306 (pan); MANTHEY 1983: 20 (part) LeplObrachillln lI/ontal1um. (nol Lep/obrac/!iul/1 1/10nUIIlU/1I FISCIIER, 1885) j"GER & STUEBIN , 1992: 41-51 (part) Leptobrachiulll cf. mon/arlllln, (not Leptobrachilllll mOrl/anum FISCHER, 1885) MA LKMUS J994b: 218 (part) Leplobrachillm abbOt/i, ING ER, STlJEBIN Q & TAl' 1995: 123, 1996: 362; MALKM US I 996c: 278; INGER ct al. 2000: 16
Diagnosis A stocky, broad-headed frog with bulging eyes; short, lender hind limbs; in contrast to L. montanUn! a distinct ventral pattern of black and white; no white ring around the eye, but a small bluish arc in the distal portion of the eye.
91
Megophryidae
4(2.5/3), 5( 1.5); inncr metatarsal tubercle low, oval in shape; dorsum smooth, often with a network of low ridges; a supratympanic fold between eye and axilla: under ide weakly rugose or granular. Upper side dark brown, occasionally blackish , with an indistinct, even darker pattern between the eyes and on the back; upper sides of legs with dark bars; ventral side of body white with large, dark lead-grey to black '" ~" pots between the chest "iii :; and thc cloaca, which Ii become more prominent o Fig. 81 LeplobrachillllJ abbol/i from Sg. Kipungit l, 550 m. distally, peppered with white dots ; juveniles with a brick-red supratympanic fold, canthus rostralis, and lateral glanDescription dular protuberance . Male up to 75 mm, females up to 95 mm; TLI VL 0.33-0.39; head as wide as long; snout obtusely Tadpole see "Genus". Labial tceth rows formula pointed or truncate, not projecting; cantbus rostral is 6-7(2-6 or 2-7)/6(1-5): lateral inframarginal ridges on lower lip witll up to 16 groups ofsmall denticles sharp; tympanum distinct, half the eye diameter: a large, round inner palmar tubercle ; toe length on either side; tail lightly convex; tail length 4<3<5<2< 1, toe weakly webbed: 1(1),2(2/1),3(3/2), 1.5-1.9 times the head-body-Ienglh; total length 75-90 mm. Hatchling tadpoles light brown. They darken gradually towards earth-brown with obscure Fig. XLII Oral disc of LeplobrachiLlln abbo"i. dark spots on head, body and tail. Lower side of body and caudal muscle with irregular reticulation o'fthin black line.
R. Malkmus
92
J
Ecology/Ethology On Mt. Kinabalu, L. abbotti pre ent itself as an inhabitant of the dipterocarp forest between 500 and 900 m a.. 1. Here, it occurs along streams and rivers, a well as far awa from the e, on the leaf litter layer of tbe forest floor. It is nocturnal and feed on large insects (Orthoptera, Blatraria, moth ), their larvae, and spiders. The males do not form calling communitie . The eggs are cream-coloured and black in the entire animal hemisphcre. The larvae live in stream microhabitats of the types
Megophryidae
Ic, d, e, and f, sometimes in la or b. They prefer ca lm reaches and can be found exposed on rocks and grave l or hidden among dead leaves cove ring the bottom of stream pools. They feed on dead leaves or graze on algae which overgrow rocks. When di turbed they attempt to reach a sheltcr with great speed.
Fig. XLID Sonagraro and
Call: (+22°C) 1-3 loud, distinctly pulsed notes (duration 190-250 ms, of which the first note of a call is the longest; each note consists of 20-30 pulses), emitted at intervals ranging from a few seconds to some m inutes ; dominant frequency 1-2 kHz; pulse repetition rate 130-170. Distribution This species is probably present in all parts of Borneo. Mt. Kina balu: Sg. Kipungit (500-600 m) , Sg. Langanan (700-900 m), Sg. Kadamaian (below Kiau).
2.80
0
-
cillograms of Leptobrachium abbotti (Sg. KipungitI, 550m; 8.00 p.m. +22°C)
3.00
3.20
~ , ~z
3.40
L
r ,
3.60 sec
.I
.
5~~
10,------------------------------------------.
5
Leptobrclchium guntmgellSe MALKMUS, 1996 Leplobrachium gU1l1mge1lsis M ALK.Ml'S. 1996d: 297 - Type loca lity: Sg. arson (~ Sg. Tibabar) Leprobrachillln (LeplobrachitJm) gliflullgense. D LBOIS & OHl ~ ~ 1998: 23
Diagnosis Morphologically very similar to L. man/anum. Description Morphometric ratios almost identical to those in L. manlanum. Together with the latter, L. gunungense forms a pair of species which can hardly be differentiated morphologically, but which differ substantially with regard to their bioacoustics and their molecular genetics (KoSUCH, unpublished) .
Tadpoles very similar to those of L. man/anum; labiaJ teeth rows formula 6(2-6)/5(1-4, 1-5) or 6(1-6), only 4-7 short lateral rows of dentic les; total length (st 37) 63.9 mm; lateral side system often distinctly visible. In contrast to the larvae found in Sg. Tibabar, those of Sg. Mesilau have blackish lip margins. EcologyfEthology LepLObrachium gunungense inhabits the transitory zone between the upper oak-chestnut forest and the mossy forest (1750-2200 m a.s.l.) on Mt. Kinabalu. Here it displays a predilection for the slopes of creeks where it can sometimes be encountered in large populations. The males form incessantly calling choruses at the onset of dusk whjch reach their greatest intensity between 6.00 and 8.00 p.m. and are particularly noisy after rain showcrs. The maJes
93
Megophryidae
Fig. 83 Tadpole of Leptobrachillm gummgense from Sg. Tibabar, 1800 m.
.,
.,::;.§" a:'
o
call out of their hiding places (beneath leaf litter, from rock cavities and holes in the ground) and are Fig. 82 Leptobrachium gllnungense from Sg. Tibabar, very difficult to locate. 1800m. Up to an altitude of 2000 m the tadpoles of1. gunungense occur in large - - - numbers in interstices of the gravel Fig. XLlV Sonagram and oscillograms of Leptobrachium glllllmgense sediment and accumulations of leaf (Sg. Tibabar, 1800 m; 7.00 p.m. + 16°q litter in the side pools of streams (m: I a-f), above 2000 m the population density decreases rapidly. The water temperatures at these altitudes average only 13-15°C.
10.60
10.80
11 .00
11 .20
11.40 sec
kHZ ~_ _ _ _ __ _ _ _ _ __ _ _ _ _ _ _ _ _1-. 4.06 sec
10 ,
5
94
CaD: (+ l6°C) The relativel y stifled, nasal call consists of series of 4-16 individual notes (note duration 140160 ms; intemote interval 240-300 ms; dominant frequency J-1.4 kH2). A note is composed of 15-16 pulses (intcrpulse interval 3.6-ll Ins) with a distinctly longer inter-pulse interval (16-20 ms) between the ' first 5 and the subsequent pulses.
Distribution Leplobrachium gunungense is as yet only known from the tributaries of Sg. Kadamaian, Sg. Liwago, and Sg. Mesilau at altitudes above 1750 m a.s.1. With L. man/anum. which occurs below 1800 m, it shares only a narrow zone of sympatry.
Megophryidae
Leptobrachiu", ",ontanum
FISCHER,
1885
Leptobrachillll1 mOlllatltlm FISCHER, 1885: 44 - l)'pe locality; Pramassan-Alai Mountains, Southeast Borneo Megalophrys hasselti, (not LeplObmchium hasseltii TSClIUDI, 1838) VAN KAMPPN 1923: 13 Megophrys hasselli, (not Leplobrachium hassellii TSCHUDI, 1838) IN ",R 1966: 29 (part), 1978: 3 12 (part); MATsUI 1979: 306 (part); MA.WHIlY &; D ENZU 1982: 13; MANTHEY 1983: 20 (part); M'\LKMUS 1985: 8, 1987: 276. 1985c: 176 Leplobrachillm cf. montallllm. INGER & STUEBfNG 1992: 41-51; M,\LIC\IUS 1994b: 218 (part) Leplobrachilllll sp., M ALKMUS 1989: 181 Leplobrachium mOlllallum , M .... LKMI:S 1991a: 27, I 992a: 102; W ONG 1994: 29-37; H OPF\IAN"- 1995a: 7; I NGER, S TUFRING & TA ' 1995: 123, 1996: 362; INGER & TAN 1996b: 75; M ALKMU I 996b: 20-26, 1996c: 278,1999: 13-19; L AKIM et al. 1999: 33; I NGER et al. 2000: 16
Diagnosis Similar to L. abbotti, but ventral side not patterned in black and white; a conspicuous white arc in the upper portion of the eye; larger than 60 mm .
Description Males up to 62.7 mm, females up to 64.4 rom; TIJSVL 0.33-0.43; habitus and morphological details are very similar to those of L. abbotti. Head and upper side of body whitish brown to ochre, with a medial, highly variable, asymmetric, occasionally light-bordered spotted pattern, partly flanked by wavy bands ; dark spots also in the tympanic region and on the sides of the head. If the dorsal side is chocolate-brown, the spotted pattern may be hardly recognisable. Upper sides of limbs with dark crossbars; lower side grey to blackish, peppered with a multitude of white dots, lightened to violet in the gular region; axillary glands whitish . Juveniles show a brick-red supratympanic fold. Tadpoles similar to larvae of L. abbotti, but without fine network ofJines on lower side and caudal muscle; labial teeth rows formula 6-7 (2-5, 6, 7)/5-6 (14, 5) and 4-8 short inframargiJ131 ridges of denticles on lower lip; totallcngth up to 73 mm (st 41).
From top to base: Leplobrachiufll mOl/tanllm Fig. 84 from the Headquarters, 1500 m. Fig. 85 from the Headquarters, Sg. Silau-Silau, 1550 m. Fig. 86 from Sayap, 1100 m.
95
Megophryidae
From top to base: Tadpoles of Leptobrachiuni montanum Fig. 87 ventral view. Fig. 88 from Sg. ilau-Silau, 1550 m. Fig. 8,? from Sg. Liw! go.
'"
~"
~
0ci ...,_
_
Ecology/Ethology Leptobrachium montanum is one of the most common terrestrial frogs of Mt. Kinabalu (900-1750 In a.s.l.) and is encountered on tbe trails of the mountain forests almost regularly. It inhabits the leaf litter layer, even far away from running water.
96
A lthough some reports exist on its occasional diurnal ity (INGER L966, M ATS 1 1979) it is usually found outside its hiding places (beneath leaf litte r, roc ks, dead wood) on ly after th e onset of dusk, i.e . between 6.00 p.m. and 6.00 a.m. This is also the time when its call can be beard in the vicinity of streams, but also far away from these. Short duets an d vocal trios may arise, but these never escalate into choru ses like in L. gUn/lngense. L. man/anum is one of the few anuran species which continue calling to a considerable extent even during prolonged periods of drought. When disturbed, it either attempts to flee with short, low leaps, or closes its eyes and crouches to the ground with its limbs d rawn up to the body (ietisimulation) so that its phytolnimetic properties render it almost invisible. 1t s wim by using the hind limbs only while the arms are rigidly held forward at a right angle. Its cleansing behaviour includes numerous e lement of that of Kalophrynus baluensis (see the respective account). The ova of gravid females are cream-coloured with a sma ll area of dark pigment on the animal pole. The larvae of L. montanum are the most abundant tadpoles in the forest streams of their distribution
Megophryidae
area. They prefer calm reaches with moderate currents and are active at day and night. They often hide in leaf drifts and grave l interstices (m: I a-f). Adults feed on gra hoppers, moths. spider , roaches, etc.
Fig. XLV
C all: (+ 18 to 20°C) 1-3 (4) loud , vcry accentuated, pulsed notes (note duration 125-160 ms; internote interval 110-126 ms) which are emitted at intervals of a few seconds up to 5 minutes; dominant frequency 500-1100 Hz with harmonics up to 5000 Hz.
Distribution This species is probably present in all montanc zones of Borneo. Mt. Kinabalu: Sayap (Sg. Kemantis; Sg. Minodtuhan) ; Marei Parei; Bundu Tuhan; entire Headquarters region up to 1750 m altitude; East Ridge above Poring (1000 m).
Genus Lepto/aJax
DUBOI ,
1040
onagram and oscillograms of LeptobrachiwJl montanulIl (Headquarters 1550 m; 7.00 p.m. +20°C)
10.60
10.80
11.00
11 .20 sec
kHz 12.54 sec 10 . -- - - - - - - - - -- - - - - - - - - -- - - - - -- - - - - - - - - - - .
5
1980
Tllis genus is known from southern hina to Peninsular Malaysia and Borneo. Its Bornean species are small to moderately large, slender frogs (SVL > 25 mm). Width of head and body subequal; legs slender and relatively long. Vomerine teeth absent, tympanum visible. Tips of digits rounded, not swollen. Webbings between fingers absent, those between toe arc limited to the bases; large inner palmar tubercle not extending onto first metacarpal; inner metatarsal tubercle low, no outer one; subarticular tubercles on fingers and toes replaced by long, low, callous tis ue. Male withmedial subgular vocal sac, no nuptial pads. Females larger than males, with relatively horter tibiae and narrower head . The tadpoles of Lepta/a/ax are adapted to a life in streams, and have an elongated, slightly depre sed
shape. Small eyes, dorsolateral to dorsal , not visible from below; oral di k ventral and subtermina l, with cup-like expanded lips, at least one lip with short tooth rows; inframarginal ridges oflower lips lack mall denticles. Heavy beaks, serrated, and fully pigmented. Spiracle is sinistrally, mid-lateral. Tail lanceolate, 1.7 to 2.2 times the head-bodylength. Tail with strong muscle and low fins. Total length of larvae 60-65 mm. They inhabit clear treams in reaches with moderate to swift currents and occur yntopically with the larvae of Leptobrachella in the interstices of gravel bed . Adults can be found on the floors of primary and secondary forest where they prefer zones near streams with dense vegetation. The males call at night from branchcs of bushes flanking treams. Their calls consist of eries of put ating, highpitched notes. Six species are known to inhabit Borneo, four occur on Mt. Kinabalu.
97
Megophryidae
Key to the Bornean species of Leptolalax
•
Superficial pectoral gland present; uniformly dark brown on the lower side of the head ..... maurus o superficial pectoral gland; lower side of the head not dark brown ......................................... 2
4 Dorsum coarsely shagreened ................... aray a; • Dorsum smooth ................................................. 5
5 Dark, round markings on the snout ........ hamidi • No dark markings on the snout ................ pictus
2 Ventrum spotted .. ........................................ ...... 3 • Ventrum not spotted ................................ .......... 4 3 Forelimb bicoloured, elbow cream-co loured with-
out dark marking, contrasting with the dark lower ann ........ .......... ............... ......................... gracilis • Forelimb not bicoloured, elbow with the same dark marki ngs as the lower arm ........ ............... dr;ngi
Tab. 7 Distribution of Leplo/a/ax-species on Borneo.
Leptolalax arayai MATSl.I, 1997 LeptoLa/ax dr;" gi DUBOIS, 1987 Lepto/a/ax gracilis (GONTIIER, 1872) Lepto/a/ax hamidi M ATSUI, 1997 Leptolalax ",aurllS INGER. LAKI\I. BIU N & Lepto/a/ax pictll.~ M ALK\fUS, 1992
Klnabalu
•
• •
•
YA~IBU~,
1997
Lepto/aiax araya; MATSU I 1997 Leptolalax arayai M ATSUI, 1997: 163 - Ty pe locality: Headquarters, Kinabalu Park. abah LeptobrachiulII gracile. (not LeptobrachiulIl gracile GONTI~ 1872) BOULe; ,ER 1887: 9S; MocQuARD 1890: 163 MegaloplllYs gracilis. (not Leptobrachillm gracile GCNTHeR, 1872) BOULENGER 1908: 421 (part); VAN KMIPI'N 1923: IS (part) Megoplll)'s graCilis. (not Leptobrachillm gracile GIDn'IlER, 1872) Mml 1931 : 12 LeptobrachiulII gracilis. (no! Leplobrachilllll gracile GC'-ITHH, 1872) INGI?R 1966: 24 (part), 1978: 312 (part); MATSUI 1979: 305; MA'THbY 1983: 20 (part) LeplObrachel/a balueflsis, (not Leptobrachel/a baluensis SMITH. 1931) MALKMUS 1989: 182, 1991a: 27 Leptobracllella cr. balliet/sis. (oat Leptobrachella baillet/s;s SMITH, 1931) MAl KMUS 1992a: J03 Leptolalax gracilis. (not Leptobracllilllll gracile GC"THER, 1872) INGER & TUF81'10 1992: 41-SI (part); MALKM[;S & RIEDE 1993: 7; MALKMUS I 994b: 219; WO,G 1994; 29-37; INGER, TUfB",,'" & T.\.'1 J 996: 362 Lepto/alax sp., MALKMU' 1996c: 279 Leptolalax sp. "a". MALKMUS 1996b: 20-26; MANTHEY & GROSSMANN 1997: 7S. fig. 41 • Leptolalax arayai. MALKMUS & KOSIJ('H 1999: 161 ; MALIC~tUS 1999: 13-19; MALK.'I[;S 2000: 12
98
Saba,h
•
•
I
I
•
Brunei
Sarawak
Kalimantan
-
•
• • •
• •
-
•
Diagnosis A med ium-sized form of the gen us Lepta/a/a.x; . dorsum roughly shagrccned; chest and abdomen without pots; among the Bomean specie it is the one with the largest feet (> 50% SYL).
Description Male up to 36 mm, fcmales up to 38 mm; TLlSVL 0.60-0.62; bead slightly longer than wide; snout rounded, projecting beyond lowcr jaw; eye d iameter greater than snout length; lores sligh tl y concave; tympanum distinct, balfthc eye diameter; finger lengths 3>4=1=2; tibiotarsal articulation of adpre sed limb reache far beyond the tip of the nout; toe length 4>3>5>2>1; dorsum shagreened, flank with round tubercles; venter mooth ; a low supratympanic fold. pper ides light to dark brown, often with dark spots of various sizes; narrow, dark brown crossbar on upper sides of hind lim bs and lower arms; flanks light brown, distal third and inner sides of thighs orange red; tubercles completely or partly
Megophryidae
From top to base: Leptafalax arayai Fig. 90 female from Sg. Silau-Silau, 1550 m, and Fig. 91 m~e. - Fig. 92 male rrom Sg._Liwago.
From top to base: Tadpole of Lepta/a/ax arayai Fig. XLVI oral disc. Fig. 93 from Sg. Liwago.
framed with black; lower sides whitish, obscurely spotted with grey on the belly; anterior portion of arc of lower jaw blackish with light dots; iris whitish grey to light greyi h green, with a fine network of black veins; one segment (ca. '100-120°) in upper portion of iris bright reddish brown. Tadpolcs' shape as described under "Genus"; both lips notched in centres, lower one more deeply so;
both Lips covered with large, rounded papillae; labial teeth reduced to two small rows in centre of upper lip; lower lip without labial teeth; beaks heavy, serrated, black; tail l.8-2.0 times head-body-Iength; total length (st 28) 61 mm; head and body whitish blue with a metallic shine above and on sides; undersides lighter, not pigmented; caudal mu cle whitish blue with small white dots; fins dusky, not transparent.
99
Megophryidae
Ecology/Ethology Lepto/a/ax arayai is a very characteristic and common faunal element of the banks of streams crossing through montane oak-chestnut forests on Mt Kinabalu occurring at altitudes between 1400 and 1750 m a.s.1., occasionally up to 1900 m. It 1.00 1.20 1.40 1.60 1.80 sec finds shelter beneath leaf litter and bark, in holes in the ground, and cavities in dead wood. During prolonged periods of drought it often hides in the immediate vicinity of streams under rocks and wood on gravel banks, and leaves these only at the onset of dusk. Advertisement calls can be heard between 6.00 p.m. kHz 5.17 sec 10 and 5.30' a.m. (occasionally until 6.00 a.m.). The males call from the leaves and branches of shrubs and 5 bushes (20 to 200 em above the ground) on the banks of streams, very rarely also from boulders. The larvae are restricted to the in- Fig. XLVTI Sonagram and osciIlograms of Lepto/alax arayaj terstices of the gravel bottoms of __ _ _ _ _ J Sg. ~ il~u-Si~u!-1~5Q.mi 7.00 p.:-m :...+~°C) forest streams (m: 1b c, e). L. arayai is an "sit-a nd-wait" hunter which feeds primarily on spide rs, moths Leptolalax dringi D UBOIS, 1987 and hairless caterpi llars. A fema le which was startled in its hiding place feigned death with its body Lepto/alax dringi D UBOIS. 1987 (1986): 13 - Type locality: MOUJJt Mulu, Sarawak adpressed to the ground for 8 seconds, only to take Leploltl/ax dringi. INGER, SrU~61:'/G & TAN 1995: 119, 1996: to flight suddenly and escape with 50-80 em long 362; L AKIM et al. 1999: 32; JNGER ct a1. 2000: 16 leaps. Diagnosis Can: (+ 18 to 20°C) The 2- 15 second long call conA moderately stocky, large-eyed species with slensist of incessantly repeated, shrill notes . Each note der hind limbs; skin with rowld tu bercles above (duration 16·29 ms) i composed ofa double pulse and 00 the sides, most concentrated on the snout; chest and belly with small black spots. (3 .3 and 4 ms, respective ly; interpulse interval 89 ms) which is repeated at intervals of lOO-115 ms; domi nant frequency 5.7-5.9 kHz; frequency moduDescription lation weak; note repetition rate 9.0-9.3 per secMales up to 35 mm, females up to 48 mm ; TLlSVL 0.48-0.56; head width equal to body width behind ond. the anus; snout obtusely pointed, rounded in proDistribution file, not projecting beyond lower jaw; canthus Leptala/ax arayai is as yet known only from Mt. rostralis rounded , lores concave; eye diameter Kinabalu where it has been recorded from along greater than distancc between eye and snout; tymSg. Liwago and its tri butaries in the Headquarters panwn distinct, half the eye diameter; fi nger tips region. rounded; finger lengths 3>4=2= 1; pa lmar and
100
Megophryidae
subart icular tube rcles indistinct; toe lengths 4>3>5>2> 1; no webbing; subarticular tubercles elongated , obscure; inner metatarsal tubercle oval, no outer one; dorsal side with round tubercles, larger on the side ; venter smooth. Dorsum and head greyi h to reddish brown, usually with black markings; hind lim bs and lower arms brown with dark crossbars; e lbow often lighter; flanks grey with light tubercles; chest and belly whitish with dark spots. Ecology/Ethology Leptolalax dringi lives in primary and old secondary forests . Nothing elsc is known about its life history. Call: The call con ists of series (duration 1.8-3.5 s) of 8-10 pulsed note (duration 35-40 IDS; note interval 60 ms; note repetition rate 11 .0- L1.5 per second) ;
Fig. XLVLlI Sonagram and oscillograms of Leplolalax dringi (Gunung Mulu, 1800 m; 1.00 a.m.)
0.20 sec
11'.11 I1II
1:"'111111
IIIIIIII! I 11111 ~lIlf
I 5.68 sec
kHz
10 r-------------------------------------, 5
t", ••• , -tt""". I"""'"
,,""",,
LOI
Megophryidae
frequency modulation distinct; dominant frequency 7-7.5 kHz with a fundamental below 0.5 kHz.
Distribution Sarawak and Sabah. Mt. Kinabalu: Records have become known from Poring (600 m), Sayap and "Kinabalu", with the latter referring to a specimen in the British Museum said to be collected at 1800 m altitude.
Leptolalax maurus INGER, LAKIM, BI UN
&
YAM BUN,
1997
Lepto/a/ax mow'us 1 'GER. LAKlM, BIUN & Y A,\ffiCN, 1997: 48 Type locality: Mesilau Station, 1860 ro, Kinabalu Park.
Sabah Lepto/a/ax //laums. I NGER & STlJEBING 1997: 59; LAKIM et al. 1999: 33; MALKMUS 2000: 13; INGER et al. 2000: 16
Diagnosis A small, moderately stocky frog « 35 mrn); entire upper ide of the body dark; belly with a network of thin lines; a superficial, round pectoral gland on either &ide; back and sides covered with homogeneous, round tubercles . Description Males up to 26.1 mm, fema les up to 31.8 mm ; TLlSVL 0.50; widths of head and body subequal; snout obtusely pointed, rOlmded in profile; eye diameter slightly greater than length of snout; lores concave; tympanum more than half the eye diameter' tips of fmgers and toes rounded; finger lengths 3>4=2= 1; toe lengths 4>3>5>2> 1; back and flanks covered with round tubercles, smaller ones on limbs; ventral surface smooth; pectoral glands visible; a distinct, angular supratyrnpanic fold is present. Upper sides uniformly black, peppered with tiny white dots which concentrate on the flanks; hind
Fig. 95 Leptalalax maurus from western Sg. Mesilau, 1800 ID.
102
Megophryidae
L eptolalax pictus
MALKMUS,
1992
Lepto/a/ax piC/liS M ALKM s, I 992b: 3 - Typ e locality: Sg. Langanan, Kinabalu, abah Lepta/a/ax gracilis. (not Leptobrachium gracile GCNTlmR, J872) MALKMUS I 992 a: 104: INGER, S 'fUIlBING & TAN 1995: 122 (part) Lepta/a/ax pictus. MA LKM US 1992b: 3; MA LKML'S & RIEDE 1993 : 7; MALKMUS 1994b: 2; H OFFMAN'i 1995a: 7; I NGER, S TUEDlNO & TAN 1996: 362; 1l'GER & T AX 1996a : 553; MALKMUS 1996b: 20-26; MATSUI 1997: 164; LAKTht et al . 1999: 32; MA LKMUS 2000: 14; INGER el al. 2000; 16
Diagnosis A slender large species of Lepto[a/ax with a distinctive dorsal and lateral spotted pattern; skin smooth; lower side not spotted . .c
e
'fi (/)
::i
o Fig. 96 Ventral view of Lepta/a/ax mallrus from west__ _
e~ S~. ~esilau , l~OO~.
limbs slightly lighter above, with fOUI black crossbars between wWch are clusters of white dots; fore limbs blackish brown, upper side of the upper arm light brown; lower sides of the legs, and throat, chest, and beUy unifonnly blackish brown; belly with an irregular, fine, white network; iris reddish brown.
Description Males up to 36 mm, females up to 47 mm; TUSVL 0.55-0.61 ; head slightly longer thall wide; snout rounded, projecting beyond lower jaw; lores slightly concave; tympanum distinct half tile eye diameter;
Fig. 97 Dorsal pattern of Kipungit 11.
Leptola/a.~ piC/liS
from Sg.
EcologyfEthology Leptolalax maurus has as yet been found only at a few sites in oak-chestnut forests (1860-1900 m a.s .l.) and always among leaf litter. During an extended dry period (January through March 1998), a male was discovered around noon (1.00 p.m.) under a rock flake lying acros ' a spring stream (western Sg. Mesilau) . The eggs of L. maurus have the greatest diameter of all known species of Lepta/alax. Increased egg sizes are , however, also typical for the highmontane species of other genera (Philautus, Pelophryne, Ansonia). c: c:
Distribution This species bas as yet been found only in environs of the type locality, the region of eastern and western Sg. Mesilau.
~o
:r:
<>: @
103
Megophryidae
~
~
~ ex: Q
Fig. 9l!J~a!.e of LeplolalaxJi.£tll! !,:o~ ~g.:...Kj)Ungit]1.
finger lengths 3>4= 1=2; hind limbs long, tibiotar al articulation ofadpressed limb reaching beyond tip of snout; toe lengths 4>3>5>2> 1; dorsum and venter smooth; small tubercles on flanks and eyelids. Fig. 99 Calling male of Lepro/alax pictus from Sg. Langanan.
Dorsal side light brown with dark brown, irregularly scattered spots of ltighly variable size and sbape wltich themselves are bordered by fine, white partly interrupted outlines; an equally coloured, broad orbital band; flanks with numerous small to medium-sized, dark brown, white-bordered spots; 4-5 large and numerous small brown spots on the sides of the head between tympanum and tip of snout; upper side of lower arm and thigh with dark crossbars; ventral side of body white, with a dark tinge distally' iris greyish go lden, bilaterally with segments of 10-20% black and a reddish brown segment of about 120 0 in upper half; entire iris overlain with black reticulation that appears fragmented.
Ecology/Ethology Leptolalax pictus is a common appearance along streams in the upper dipterocarp forest and oakchestnut forest (500-1100 m a.s.l.) on Mt. Kinabalu. It activity patterns are identical with those of
104
Mcgophryidae
L. arayai. The males call from shrubs and leaves
and twigs of bushes in the vic inity of streams, occasionally from directly above the water.
Fig. L Sonagram and oscillograms of Lepto/a/ax pictlls (Sg. Kipungit 1. 600 m; 7.00 p.m. +20 0 )
Call: (+ 19 to 22°C) The call is shorter (1 .9-4.9 s) than in L. arayai. use a higher frequency band (6.87.2 kHz), and are more often paused (duration 4-5 s). Each note (duration 30-35 ms; internote interval 45 ms) is marked by an initial frequency drop from 7.2 to 6.8 kHz. The note repetition rate is I 1-13 per second.
1.25
1.30
1.35
1.40$8C
Distribution
Leptolalax pictus has as yet become kno\'m only from North Bomeo. Mt. Kinabalu: Poring: Sg. Kipungit, Sg. Mamut, Sg. Langanan; Sayap: Sg. Kemantis.
'111111111111111111111 11111111111111111 kHI ,--_ _--=~~~=~==.,____----....:., 5 .0Sec 20
10
Genus Megophrys KUHL & VAN lliSSELT, 1822 TIle genus MegoplllYs is native to continental Asia, Sumatra, Borneo, Java, and pacts of the Philippines. The Bomean species are medium-sized to large frogs with relatively short, slender hind limbs and large, wide heads. Most species have characteristic dermal palpebral projec60ns (particularly above the eyes and on the snout) and a sharply curved ridge of skin between the eye and the ann. Tympamml is visible, but often indistinct. Adults possess a conical axillary gland at the insertion of tbe ann . Tips of fingers and toes rounded, outer palmar tubercel and subar6eular tubercles absent or indistinct; inner palmar tubercle oval, extending along first metacarpal. Webbing reduced to basal rudiments. Males possess vocal sacs and nuptial pads. The Larvae are slender and have funneJmouths; beaks soft, serrated, lacking labial teeth, but with many short, low ridges; oral disk terminal, lips expanded into a horizontal, dorsally-oriented funnel; spiracle sinistral; anal tube med.ial, fused to the ventral fin or not. The tadpoLes dwell streams in sections
105
Megophryidae
with moderate currents. Thcy are adapted to feeding on plankton in the surface film. Adults are terrestrial and can be found in the layer of leaf litter in primary fore ts. The male of all three
pecies pre ent on Mt. Kinabalu form calling cheru e .
Five species arc Icnown from Borneo, four from Mt. Kinabalu.
Tab. 8 Distribution of Megophrys-spccics on Borneo. Snbah
Megopllrys baillells;s (BOULENGER, 1899) Me$!of)hrys drinJ!i INGER , S ruEBI:-4G & TAN, 1995 Megopllrys ed wardillae I:-4GER, 1989 Megopllrys kobayashii M ALKMUS & MAISUI, 1997 Megopllry.v lIaslita (SCHI.I!GI!L (858)
Key to the Bomean species of Megophrys 1 Head distinctly narrower than body ......... dring; • Head a little narrower or as wide as body ........ 2 2 Two continuous dermal lateral or dorsolateral fo lds; dermal rostral appendage present ........ .... lIasuta • Dermal fo lds not continuous; no rostral appendage
.................... ................................................ ........ 3 3 Large frog, VL > 90 mm; angle of jaw and end of supratympanic fo ld with large projections ..... ......... ....... .... .... .... .... .... .... ..... .... ... . .... . kobay ashii • Never larger than 70 mm; no such projections or only small ones ..................................... ............ 4 4 Vomerine teeth present .. .......... ..... ...... baluen sis • Vomerine teeth ab ent; no dorsolateral folds ..... . ... .... ....... .... ... ....... ................. ..... ..... . edwardinae
Megophrys baluensis (B Leplobrachium ballleT/se B Ol:LENG~R, I 99 : 453 - Type locality: Mount Kina Balu LeplobrachiulII baluellse, HA" rSCH 1900a: 74 Megalophrys baluellsis. BOl. LENGER 1908: 429; VAN K AMPE" 1923: 16 Megoplirys balueT/sis. SMml 1931 : 30; ~GER 1954a: 250, 1966. 46, 197 : 3 12; MATSUI 1979: 310; MAlffllEY 1983: 20; MALKM • 19 5: 9, 1987: 277, 1988c: 176, 1989: 181; INGER 19 9; 229; I OJ:R & TUEBINO 1989: 41 ; MALKMUS 1991 : 27 ; INGER & TLT.BI}.G 1992: 41 -5 1; MAuo.IUS 1994b: 220, 1994e: 86, 1995c: 8, 1996b: 20-26. 1996c: 280: INGER, TUEBL~ & T A.'1 1996: 362; lNGFR & TAN 1996a: 553; MAuo.1U & RIEDE 1996c: 152; LAKl/oI et a1. (1999): 32; MALKMUS 1999: 13-19; I NGER ct a1. 2000: 16 Megophrys cf. ba/uellsis. M ALK MUS 19923: 104
106
• • • •
Kinabalu
• • • •
Bnmei
?
•
Sarawak: ?
• • •
Kalimantan
•
Diagnosis A moderately large, stout tcrre trial frog. It is distinctly mailer (alway < 70 mm) than M. kobay ashii and M. nasl/ta. Snout very hort and without rostral appendage; dorsolateral, or lateral, glandu lar lump grouped in rows, but not forming continuou fo ld.
Description Males up to 45 mm, females up to 70 mm; TU VL in females 0.36-0.38, in males 0.40-0.46; head
Fig. 100 Semiadult Megophrys balLlellsis from
Tibabar, 1700 m.
g.
Megophryidae
Fig. 101 _Male o! "!..egophrys
baluensi!I!0~
.!he Headquarters.
wider than long; very short snout, vertical in profilc, without dermal appendages ; short, pointed deonal appendages above the eyes, followed by conical tubercles; tips of phalanges rounded ; no webbings between fingers; occasionally rudimentary we bbings between toes; finger lengths 3>2=4> I ; inner metatarsaJ tubercles low, oval, no outer one ; subarti cular tuberc le indistinct; toe Icngths 4> 3>4>2> 1; tympanum usually clearly visible, smaller than half the eye diameter; upper surface covered with irregularly scattered, usually elongate-oval glandu lar rises which form rows, and partly ridges, dorsolaterally and laterally ; a distinct, angled upratympanic fold between eye and ax iIla; maJes with nuptial pads on 1" and 2nd fingers .
Coloration and pattern highly variable and petfectJy adapted to pattern and coloration or leaf litter; upper sides dark bro.wn, reddish brown to ochre, lighter laterally and on limbs; lower side yellowish browll to dark flesh-red, with dark brown clouding; throat with dark or light median stripe; a triangular, blackish brown marking between occiput and mid-dorsum; between eye and margin of upper jaw alternating yellowish white and dark brown vertical bars; iris golden brown with black reticulation. A 3 cm long semi-adu lt specimen was light loamye llow with white ventrum, clouded with brick-red towards chest. Tadpoles sim ilar to larvae of M nasula. ostri ls elliptical, rim low, with very flat mid-dorsal projection; total length (st 3 J) 36.7 rom ; tai l length
107
Megophryidae
Fig. LI Oral funnel of M~g01!..h rys .!!alue!'sis.
tail length 2.09-2. 14 times head-body-Iength; tail tapering to pointed tip; body above and on sides dark brown with large loam-coloured spots; belly greyish brown , without pattern; caudal muscle brown with light spots, particularly on upper half; both beaks unpigmented with blackish band in the centre; tins dusky, with isolated dark flecks.
Ecology/Etbology On Mt. Kinabalu, M. ba/uensis occurs in the vicinity of streams cutting through oak-chestnut and
lower mossy forests (1280-1950 m a.s.I.). It prefers the banks of small running waters, particularly fields of rock rubble which are densely overgrown by shrubs (Begonia, Gunnera, Impatiens). Here, the frogs live in very stationary calling communities. Five of those colonies could be traced always in the same spots on tributaries of Sg. Liwago between 1984 and 1998. Choruses can be heard between dawn and dusk with maximum intensities between 5.30 and 6.00 a.m. and 6.00 and 7.00 p.m., respectively. These are started by one male beginning to call, triggering others to respond, and so building up to a choms whlch then lasts for 30-100 seconds. The frogs call from rock crevices, cavities among roots, and holes ill the ground. They leave their shelters at night and forage on the forest floor, occasionally c1imbillg on low bushes up to about 30 cm above the ground. Their tadpoles dwell mountain brooks with weak current (m: Id, e). M. baluensis shows rituaused cleansing motions, otherwise known from species of the geoeraLeptobrachium and Kalophrynus. Captured specimens show a pUlsating trembling shortly before they attempt to flee.
Fig. LIT Sooagram and oscillograms of Megophrys baluensis (Sg. Lanaga, 1500 m; 7.00 p.m. +l9°C)
2.10
2.20
2.30
2.40
~
': ' - -, -, - r[
108
'l.} -'~~""::
.,. F-·
2.50 see
" '~"'
Usee
._, ., ....... .. I
:~,-.'~ ~ ·<:( ·. '~.. ':l:
.
_
Call: (+ 17 to 19°C) The calls are emitted in series of 3-8 loud, metallic, sharp, and hard strikes at intervals of 550-700 ms. They consist, especially in their second third, of distinctly di fferentiated pulses and last 90-100 ms; carrier frequency 2.8 kHz.
Distribution Until recently M. baluensis used to be listed for the fauna of Sarawak, Sabah (INGER 1966, INGER et a1. 1985, I NGER & STUEBING 1992) and Brunei (DAS 1995).The latest check list of the frogs of Borneo (I NGER & TAN 1996a), however, lists this species only for Sabah, and INGER & STUEDING ( 1997) consider it to be endemic for Mount Kinabalu, stating: .oM. baluensis is so far known only from Mount Kinabalu". Mt. Kinabalu: Tributaries ofSg. Liwago (up to 1950 m on Sg. Tibabar); Bundu Tuhan; eastern Sg. Mesilau (1800 m).
Megophryidae
'800· '100m 1500 · 1800m
o •
M. baluensis M. kobayashii
Fig. Lm Distr~butioD 5!! Ml!..sophrys ba/ue11!!s_and M. kobay'-ashii in _the area of the !:Ie~_dquartcrs .
Megophrys edwardinae
INGER,
1989
MegoplllYs edwardinae ['IOcR . 1989 : 229 - Type locality: Nanga Tekalit, Kapit District, Sarawak ,IIJegophrys edwardillae. I NGER, STUEB ING & TA N 1996: 362; INGER ct al. 2000: 16
Diagnosis A medium-sized MegoplJrys; dorsum with scattered, round and elongate tubercles; a "hom" on the rough-sutfaeed upper eyelid; no continuous dorsolateral folds. Description Males up to 42 mm, females up to 82 rnm ; TL/SVL 0.43-0.50; head wider than long; nout obtusely pointed, rounded in profile; canthus rostralis sharp, curved; lores oblique, concave; eye diameter equal to eye-snout distance; tympanum obscure; vomerine teeth absent; fingertips rounded; palmar tubercles weak; finger lengths 3> 1>2=4; tips of toes rounded; rudimentary webbings on bases of toes; toe lengths 4>3>5>2> 1; subarticular tubercles obscure; inner metatarsal tubercle low, no outer one;
skin on dorsum and head with many irregular tubercles and short ridges; a large conical tubercle at rictus; upper eyelid with tubercles and ridges, central margin of eyelid with triangula r projection; sides of the head with ridges radiating from conical tubercles; a trong supra tympanic fold present; throat rugose; teat- like projection on chest near axi lla; belly smooth. Brown above and on sides; an orange patch on shoulder; two dark bars between eye and upper lip; ides light brown with irregular dark markings; ventrum pale brown with dark spots; lower lip barred; dorsal surfaces of limb with dark crossbars. Ecology/Ethology This species lives on the leaf-littered floor of primary forests. early nothing is known about its life history. It was found in low land and hilly regions at altitudes between 200 and 600 m a.s.!. Distribution Megophrys edwardinae is known from Sarawak and Sabah. Mt. KinabaJu: Only known from the Poring region.
109
Megophryidae
Megophrys kobayashii MALKMUS & M ATSUI, 1997 Megophrys kobayashii M ALK.\lU & M ATSUI, 1997: 3 1 - Type localiry: Headquarters. Mounl Kinabalu, Sabah Megophrys molltico/a, (not Megophrys molltana KUHL & VAN H ASSELT, 1822) SMITH 193 1: 12 MegoplllYs lIIonlitola nas l/ta. (not Ceratophryne nasuta SCHLEGEL, 1858) l NGER 1966: 41 (part), 1978: 3 12 ( p art); M AISUl 1979: 3 10; M ANTHFY 1983: 20 (part) Megophrys nasi/la. (not eratophryne nasuta CHLEGF.L. 1858) I NGIiR & SnJEBING 1992: 41 -5 I (pa rt ); I :-;GER, STlJ1j6ING & TAN 1996: 362 ( p art) Megop/trys sp., M ALKMUS 1995c: 7, 1996b: 20-26. 1996c: 279 Megophrys Iw bayashii. M ALKMUS 1999: 13- 19
Diagnosis A large fonn of MegoplllYs: in contrast to M. nasula with no long continuous dorsolateral folds; snout rounded without rostral appendage. M ko bayashii is most imilar tQ M. /igayae from the Philippines.
Description Females up to 109 nun ; TL/SVL 0.45-0.49; head and body stout, head wider than long; snout rounded in profile, projecting beyond lower jaw; no rostral appendage; snout somewhat shorter in length than eye diameter; upper eyelid dn;lwn out into a triangular projection; canthus rostralis sharp, lores concave; tympanum vi ible; supratympanic fold distinct; vomerine teeth present; tips of fingers rounded; ftnger lengths 3> 1>2=4; tips of toes blunt; toe lengths 4>3>5>2> 1; inner metatarsal tubercle ellipsoid; tibiotarsal articulation rcaching up to the eye when leg is extended; very short pairs of dorsolateral longitudinal folds; skin of head fused to skull; skin behind head ca1cifted~ eyelid with conical tubercles; lateral and ventral skin rugose or coarsely granular; sides with irregularly scattered, large, round and elongated tubercles; angle of jaws and end of supratympanic fold with irregular projections; arms and bind limbs with tubercles and short folds.
Fig. 102 Megophrys kobayashii from Sg. Silau-Silau, 1500 m.
LIO
Megopru-yidae
~ ~
:i g
Fig. 104 luvenile of Megoph,ys kobayashii from Sg. Silau-~il~u/.§ g..: ~w~g~, ~ 4~O ~ . _ _ _ _ _
Fig. 103 MegophlYs kobayashii from Silau-Silau trail, tEe_H<:.at!.q~arters.
ne~
Dorsum light brown; hvo conical , black tubercles on head ; large dark blotch below canthus rostralis and behind eye; lower lip dark brown, with narrow wltite bars; very large brown, white-bordered marking between posterior lower Iip and upper arm; lower arms with large brown spots; hind limbs light brown with darker fields around ridges; large dark Fig. Lrv Tadpole of Megophrys kobayashii.
R. Malkmus
spot on knee; lateral tubercles and ridges light brown with ilTegular dark spots ' throat brown, medially with darker spots; abdomen light brown with two parallel rows of large dark spots. Tadpoles similar to larvae of M. baluensis and M nasuta; rim of nostril mid-dorsally with pointed projection; total length (st 37) 41.8 mm ; tail length 2. 1. times head-body-Iength ; tail tip rOlmded; body and caudal muscle darker in coloration than in M baluensis and M. nasuta; with fewer lighter fleck; a light vertical bar on lower half of tail muscle near its root; fins dusky, marbled with. brown; centre of upper lip, margin of oral disk, the region lateral to beaks, and 'lateral tips of funnel black.
EcologyfEthology M. kobayashii dwells the layer of leaf litter on the floor of lower montane rain forests (1450-1675 m a.s.L.), preferring the neighbourhood of small streams, and hiding in holes in the ground, cavities among and in roots and among rocks . Its calling activity is limitcd to short periods of time in the early morning (6.00 to 6.30 a.m.) and in the late afternoon (6.00 to 6.30 p.m .), but calls can also be heard during rain showers (between 3.00 and 6.00 p.m.) when the daylight fades. It is usually one specimen which begins to call, triggering others to respond, and a full chorus is in place within 3-5 seconds. This species lives in pennancnt colonies. Larvae were found in small mountain brooks with weak current (m: Id,e). M. kobayashii appears to be rare.
III
Megophryidae
Call: (+2 1°C) a very powerful voice; call sequences last between 3 and 5 seconds and consist of 8-l3 notes with distinct pulse modulation; note duration varies from 135 to 188 ms(mean 163 ms); intemote interva1134182 ms (mean 150 ms); carrier frequency 1.3-1.9 kHz.
Distribution M. kobayashii is known only from the Headquarters region on Mt. Kinabalu, i.e. Sg. SilauSilau, Sg. Lanaga; Lumu Lumu (see also map on page 109).
Megophrys nasuta Cel'atopJllyne llasula
SCIILEOI:L,
kHz
10.15 sec
10 r-----------------------------------------~
5
F ig. LV Sonagram and oscillogram of Megophrys kobayashi _____ jI!e~~a~eE.S ,..!5~ O mi..6]O_p.m. +2 1°C)
(SCHLEGEL,
1858)
1858: 75 - T)'pc loca lity:
Su-
matra Megalophrys lllJ.$lJla, MOCQUARO 1890: 163 ; H ANITSCH 1900a:
74; VAN KAMP!;!\' 1923 : 10 Megopltrys monlieola nasllla, INGER 1954a: 250, 1956: 393, 1962/63 : 46, 1966: 41 (part). 1978: 310 (part); MANTHEY 1983: 20 (part); M ALKMt:S 1985: 9,1987: 277,198 c: 176 MegoplJrys nast/la. MITH 1931 : 9; INGER 1985: 13; INGER & SruEBING 1992: 41-51 (part); M,\LKMUS 1992a: 104; W ONG 1994: 29-37' HOFFMANN 1995a: 8; MALK.\tuS 1995c: 6, 1996c: 279; INGER, STUEDING & TAN 1996: 362 (part); LAKIM et al. 1999: 33; blGER ct al. 2000: 16
Diagnosis A large Megophrys; derma l rostra l appendage present, and upper eye lid drawn out into a triangular projection; two pairs of parallel, longitudinal, dorsolateral fo lds continuous between head and groin. Description Males up to lOS mm, females up to l25 mm; TLlSVL 0.34-0.46. A stocky frog with short, slender hind limbs; head wider than long; tympanum present, but usually bidden by skin; vomerine teeth present or absent; shapes and relative lengths of fingers and toes identical to those of M kobayashii; dorsum smooth with 1-4 large, conical tu bercles of black coloration; sides and ventmm mgose to coarsely
112
granular; 2 pairs of narrow dorsolateral folds; triangular dennal appendages on eyelids and rostmm are smaller in specimens from Mt. Kinabalu than in those from other parts of Borneo (INGER 1966). Specimens from Mt. Kinabalu are light to dark reddish brown above, sometimes with a dark-clouded central portion; a blackish brown, wide, V-shaped marking between eyes and central occiput; black bar behind and below eye; dark crossbars on upper side oflimbs; black spot on knee; throat brownish, grey to reddish, partly with small white spots; belly marbled light brown on a clirty whitish ground, lighter clistally; iris blackish brown. The structure of the skin, its body shape, and the colour pattern of this frog are an example for Fig. l OS Tadpole ofMegophrys nasuta from Sg. Kernantis, Sayap.
Megopbryidac
phytomimesis (dead leaves on the forest floor) developed to perfection. Tadpole: Head-body of tadpoles elongate, almost circular in cross section; eyes laterally, oral disk tenninal ; lips expanded to form a borizontal funnel ; open funnel as wide as body; lips without labial teeth, but with many short, low ridges; beaks thin, serrated; total length 42.2 mm (st 41); tail length 1.9-2.4 times tbe bead-body-length . Body brown dorsally and laterally; belly with or without dark spots; caudal muscle brown ; dorsal fin dusky ; ventral fin without pigmentation in its proximal two thirds, dusky near the tip.
rubble. The frogs spend the day bidden in cavities among the rocks, beneath dead wood, and in sub-terrestriallunnels, and emerge with the onset of dusk in order to forage. Their huge mouths eo-
Fig. 107 M egophrys flOsllta from Poring region.
Ecology/Ethology On Mt. Kinabalu, M. nasu!a is an inhabitant of the layer of leaf litter covering tbe floors of tbe submontane dipteroearp and lower oak -chestnut forest (500-1300 m a.s.I.). It is not only encountered in the vicinity of water bodies, but also far away from these, particularly on slopes of rock
113
Megophryidae
>-
£ ~ :;j
Q
Fig. 108 MegQphrys nasuta one day old lTom Pulau Pinang. Peninsu lar Malaysia. Fig. LVI Sonagram and oseillograms of Megophrys nasula (Poring region: 6.00 p.m. +20°C)
5.60
5.70
5.90
5.80
Call: A resonant " honk" note. The call consist of 9-34 of such honks. The very loud, in iti al notes of a call are often drawnout and resemble the song of cicadas. The notes last 160-] 70 ms; intcmote interval 150-160 ms; dominant frequency 2-2.5 kHz .
6.00 sec
,.
L.
able them to devour snails with shell diameters of up to 50 mm , large Orthoptera, moths, caterpillars, beetles, and even scorpions of up to 10 cm in length. Occasionally, a diumal specimen may be encountered. The ma les call from the vicinity of streams, particularly during the periods of dawn and dusk. They fonn very noisy choruses which, however, rarely last longer than 30 minutes. The white eggs (2 mm in diameter) are deposited on the underside of rocks and fallen 10 gs in those sections of streams where the current is moderate to weak (m: Ib-i ). The larvae prefer cal mer sections of forest streams, particularly accumulations of leaf litter along the edges of basins and on the bottom (m: I d, e, 1). They "suck up" floating particles from the water su rface by expanding their funnel-mouth. The latter does not serve as an adhesive organ (Bos HMA 1922, SCHMIDT 1976).
.....
Distribution Megophrys nasuta is native to
kHz 10
7.74 sec
r---------------------------------------,
5
.
114
-~
"'-~.
~
.-: ..
~
the Malay Peninsula, Sumatra, and Borneo . Mt. Kinabalu: Koung, Kiau; Sayap: Sg. Wariu, Sg. Kemantis, Bundu Tuhan ; Poring: Sg. Mantukungan , Sg. Mamut, Sg. Kipungit to Sg . Langanan (500-9S0 m).
Microhylidae
Microhylidae This famil y is rich in pecie and nearl y cosmopolitan in tropical and ubtropi cal latitudes. The Bornean specie are mall to medililll- ized frog with a roundi h shape and a mall head. They are characteri ed by a firmistemal pectoral girdle. Thc omosternum is ab ent, a cartilaginou ternum present. The vertebral coluITUl i di plas iocoelous, the sacrum is furnished with dilatcd diapophyses. Maxillae and premaxillae are edentate (except in Calluella) . The tips of thc fingcrs and toc mayor may not be expanded, and there are no intercalary pbalange . Tbe toes arc webbcd over half their lengths, or only on the bases (except in Microhyla berdmorei). A feebly rai ed skin fold i found between the snout and the groin. Amplexus i axillary. The 80mean species produce small , pigmented, aquatic egg from which free-swimming tadpoles
emerge. These po sess small terminal mouths without comified beak and dentic les. The tadpo les of KalophrYllus are nidicolous. They are characterised by a median spiracle, laterally positioned eyc , and the narcs do not appear unti l shortly before metamorpho is. Bornean microhylids include both terrestrial (Cal/uel/a, Chaperina, Gastrophrynoides, Kaloph ryn us, Kalollla, Microhy la) and arboreal forms (Metaphrynella) . They dwcll primary forests and econdary growth ; some pecie occur in distw'bed habitats associated with the activities of man . On Mt. Kinabalu, microhylids have a vertical distribution ranging up to 1800 m. Borncan microhylids comprises 20 species repreenting 7 genera. Mt. Kinabalu is home to 5 genera with 8 species.
Key to the Bornean genera of Microhylidae (after M Al\TII EY & GROSSMANN 1997) Ventral side with dark reticulated pattern (A) .... ...... ... .......... .... .... ..... ...... .. .... ... ..... .... '" Cllaperb,a • Ventral side with different colour pattern ... .... . 2
B.
--
2 Finger tips spatulate (8) ............... ......... ..... .. .... 3 • Finger tips with different shape ........... ........ ..... 4 3 Lower sides of outer fingers with double, enlarged basal tubercle (C) ... ...... .... .. ........ Metapllrynella • Lower sides of fingers without enlarged basal tubercle .... ..... ..... .......... .......... .... .... .. ...... Kaloliia
D
4 Tympanum invisib le ...................... ..... ..... .......... 5 • Tympanum distinctly visible (D) ... Kaloplrrynlls 5 Maxillary and vomerine teeth absent; inner metatarsal tubercle low .... ...... ... ....... .......... ......... ...... 6 • Maxillary and vomerine teeth present; inner metatarsal tubercle spatulate (E) .. ...... .... .. .. Calluella 6 Snout short « two times the eye diameter); toe tips not T-shaped .... .... ... .......... .. ... ... .. Microlryla • Snout long (> two time the eye diameter); toe tips T-shaped (F) ........... ..... ..... ..... Gaslrophrynoides
F
115
Microhylidae
Genu Chaper;na MocQuARD, 1892 Chaperina is a monotypical genus. Its only specie, haperina lusea, occur all over the island (Sabah, Sarawak, Brunei, and Kalimantan), in the southern part of the Philippines, and on the Malay Peninsu la.
Chaperina /llsca MocQu
RD,
1892
Chaperinajllsca M OCQUARD, 1892: 35 Type locality: Sintang, Borneo
Chaperillajllsca, SMITH 1931 : 2 1; INGER 1956: 398. 1966: 140, 1978: 312; MATSUI 1979: 322; MANTII EY 1983: 21: l"GER & STU~BING 1992: 41-51; R OZKOSNY & K OVAC 1994: 865; HOFFMANN 1995a: 10, 1995c: 27; r . •F.R. STU"BI~G & T AN 1996: 362; LAKIM ct a1. 1999: 32; l NGER et al. 2000: 16
Fig. 110 Chaperina ji/sea frolll Poring region, 650 Ill.
Diagnosis A tiny, long-limbed frog « 30 mm) with a very characteristic pattern on the ventral side, i.e. yellow spots separated by a dark brown network. uriously, the yellow colour will rub off onto buman fingers. There are small, flexible, spine-like derma l projections on the heels and elbows. Desc ription Ma les up to 21 mm, fema les up to 26 nun; TLlSVL 0.49-0.56' habitus slender to moderate ly stocky; snout rounded, projecting, longer than eye diam-
Fig. 109 Chaperina fusea frolll Sg. Wariu, Sayap.
~ ~
~ ~~~~~~£2~~~~~~~~ 116
eter; canthus not distinct, lores almost vertical; tympanum obscure; pupil horizontal; tips of outer three fingers dilated into round disks; tip of 1" finger not expanded; finger lengths 3>2=4> 1; subarticular tubercles sma ll ; tips of toes expanded into disks; webbing rudimentary, limited to bases of tocs; inner metatarsal tubercle oval; no outer metatarsal tubercle; skin above smooth or with low tubercles, sides with low, round tubercles; ventral surface smooth; supratympanic fold between eye and axilla; males with median subgular vocal sacs, nuptial pads absent. Dor urn and sides black, with or without small white, greenish, or bluish dots; limbs brown with dark bars; below a dark brovlIl network enclosing large yellow cells; iri dark brown. Head-body section of tadpoles oval, slightly flattened above, rounded below; snout tnmcate; tail tapering abruptly near end to a rounded tip; mouth terminal; lip not e panded; no labial teeth; no keratinized beaks or papillae; head-body-Iength (st 40) 21 mm; head-body portion black 3bove, lighter below; caudal muscles grey with black line at dorsal edge; caudal fins unpigmented; above iris a small, halfmoon-shaped, bright metallic golden spot. Freshly meta morpho ed juveniles are black below, peppered with fine white dots.
Microhylidae
c
I J:
a:
g
Fig. Ll2 Eggs of Chaperina/usca from Poring region, 650 m.
the walls of small water bodjcs or to submerged leaf litter (m : I g, 2b, d, e, g). We found adult and larvae at various stages of development in a rock pool ncxt to Sg. Lidian (surface area lO x 20 cm , dcpth 5-10 cm , filled with rotting mud and bamboo leaf litter). and another reproduction community in a phytothelmc in a buttress tank at ayap (surface area 20 x 30 cm, depth.25 cm , I m above thc ground). Here, we observed nightly territorial encounters between two males which lasted a couple of minutes. The opponents would raise against one another, chirp, and try to push each other away. Matings and oviposition take place at night. Fig. 1] 1 Ventral views of Chaperina/usca from Poring (1-5), photos : P. Hoffmann, and from Sg. Kemanti s, Sayap (6), photo: M. Schroth .
Fig. 113 Tadpole ofChaperinajilsca from Pulau Tioman, Peninsular Malay ia.
Ecology/Ethology On Mt. Kinabalu, Chaperinajusca is an inhabitant of the primary dipterocarp forest (500-1250 m) . It lives in the layer of leaf litter, occasionally climbing low vegetation . It i generally diurnal. The males often form choruses in and around small ephemeral water bodies with maximum depths of less than 25 cm rich in humic acids and infusoria (the tadpoles feed primarily on infusoria). A clutch comprises 6-10 grey eggs which are attached to
117
Microhylidae
& KOVAC (1994) found tadpoles in bamboo internodes (Gigantochloa levis) near Poring. The internodes lying on the surface and measured 10 em in diameter, were 30-40 cm long, and held rain water and fallen bamboo leaves. Other aquatic inhabitants were larvae of Coleoptera, Diptera, and Odonata. ROZKOS:NY
Call: (+ 220 ) Two squeaking, na al notes which are particularly distinct in their pulsing in the ir first and last thirds (note duration 400-550 ms; internote interval 400 ms), fo llowed by an interval of3-4 s and three short ( 100150 ms) postnotes; dominant freque ncy 2.5-3.5 .kHz; .£j-equency modulation and pu lse modulation present. Furthermore, long drawn-out sounds (up to 4 s in duration) with distinctly modulated pu lsi ng were noted (frequency 2-3 kHz). Distribution Chaperina Jusca is known from the Peninsular Ma laysia, Borneo, and the southem parts of the Philippines . Mt. Kina balu: Sayap: Gua Melayu, between Sg. Wariu and Sg. Kemantis ( 1050 m) ; Kiau ; Kenokok (1000 m) ; Sg. Lidian (1250 m); Sg. Kipungit (500-600 m); East Ridge abovc POling (1000 m). It is very likely that the spec ies occur also at greater altitudes on Mt. Kinabalu since it was recorded from 1800 m on Mt. Murud, Sarawak (SMITH 1925a).
0.60
1.00
1.40
1.80
2.20 sec
kHz ~------~------~--------~------~------~
5.09 sec
10
r------------------------------------------,
5
9.80
10.00
10.20
10.40 10.60
10.80
11 .00
11 .20
11.40 sec
-kHz
12.13 sec
10.------------------------------------------,
5
Fig. LVII Sonagrams and oscillograms of Cllaperilla Jusca (Sayap region, 1000 m; 8.00 p.m. + 20-22°C)
118
t
Microhylidae
Genu Ka/ophrYllus
TSCHUDI,
1838
Specie of the genus Kalophrynus occur in parts of Sou thea t Asia, on Sumatra, Java, Borneo, and the Philippines. The Bornean pecie are mall to medillm- ized frogs with pointed snouts, short hind limb , and relatively narrow heads. Tympanum present, but often indistinct; skin usually granular above and below. Tips of fingers and toes not dilated, 4,h finger and 51b toe very hOI1; webbings limited to less than half length of toes (except in K. pleurosfigma). MaxiUary and vomerine teeth absent. Males with median subglilar vocal sacs and nuptial pads.
These frogs deposit their eggs in aquatic microhabitats such as tree holes and bamboo internodes . Head-body of larvae oval, flattened above; nout broad ly rounded, mouth terminal. Lips not expanded; eyes laterally, visib le from below. Tail laneeolate. piracle mid-ventral , without frcc flap ; anal tube median, not open ; gut with only one or two coils. The larvae do not feed (no beaks, no labial teeth or papillae) and complete their development with the nutrition provided by their yolk. Adults are terrestrial, dwelling the leaf litter on the floors of primary forest . even species are known from Borneo, two of which appear on Mt. Killabalu.
Tab. 9 Di stribution of Kaloph'Yllus-species on Bomeo. SabJh
Kalophrynlls baillensis K.iEw 1984 Kalophry nus heterochirus B OU LEI\ GER, 1900 Kalophryllus intermedius I NGI:.R, 1966 Kalophry nus Ilubicola DRJ ~G, 1983 Kalopllrynlls plellrostil:m a T SCHI; D1 , 1838 Kalophryllus pUflclatus P ETERS, 1871 KalophrYllus subterrestris I NGER, 1966
• •
• •
Kinabalu
Brunei
•
• •
•
Sarawak
Kalimantan
• • • • • •
•
Key to the Bornean species of Kalophrynus (partly after ING ER Fleshy webbings present on palm .. ... .. nuhicola • No fleshy webbing on palm .. .......... ........ ....... ... 2 2 5"' toe projecting as far as or farther than 3,d toe ......... .. ........ ............. .. .. .... .... ..... .......... . punclatus • 511> toe not projecting as far as 3nJ ......... ............. 3 3 Portion of 4"' linger projecting from palm shorter than terminal phalanx of 3,d finger .. .... .......... ... 4 • Portion of 4"' linger projecting from palm longer than terminal phalanx of 3,d linger ..... ...... ... .... .. .. ...... .. .......... .............. .......... ............. pleurostigllla
• •
1966)
5 A light stripe or line from canthus to groin ..... 6 • No light stripe or line on snout or side .... .......... . .......... ..... ........ .... ......... .......... ........ ... intermedius 6 Bluish or whitish inguinal spots in a large dark area; adult females < 35 mm ; lowland species .. . ..... ........................... .... .. .. .. ............. helerochirus • Inguinal spots present or absent; adult females > 40 mm; montane species .................... baillensis
4 Only I tubercle under the 4 d, linger between the palmar tubercle and tip of the finger ..... .... .. .. .. .. .. ........ ........... .................. ... ....... ......... subterrestris • 2 tubercles under the 4'h finger .. .... ................... 5
119
Microbylidae
Fig. 114 KalopfllYflUS baluellsis from Sg. Silau-S ilau, 1500 m.
KalQphrynus baluensis I<1Ew, 1984 Kafopilly nus bofllensis Klll W. 1984: 151 Type locality: Kamborangob at 7,200 teel on Mount Kinabalu Kafophrynus pfellrostig mo. (not Kalophl)'lIus pfeurostigma Ts Hu m. 1838) SMITI~ 1931 : 21 (pan ); INGER I 954a: 250 Ka!ophry nus pleu/'Ostigma pleurostigmo. (not Kal ophry nlls pleurosligJIIG Ts °HL DI . 1838) M ATS!:I 1979: 320 KalopJzrynlls ball/ellsis, INGER & S n JEBlNG 1989: 59; MAL KMUS 199 1a: 28, I 992a: III ; INGER & SruE.BING 1992:41-51 ; M AI.KMUS 1994a: 9, 1994b: 224. 1996b: 20, I996c: 282; I NGER. STt..fBIN(; & TAN 1996: 362; INvE K & T AN 1996a: 557; M ·\ LKMUS & RIEDE 1996c: 152; INGFR & SnJEBI NG 1997: 96; I NGER etal. 2000: 16
Diagnosis A medium-sized brown frog; habitus stout; snout pointed , shorter than the eye di ameter; yellow patch(es) bordered with dark brown present or absent in the inguinal region; 5th toe with 1 subarticular tubercle.
120
Description A moderately large, stout microhylid; males up to 36, females up to 47mm; snout short, pointed; hind limbs short (TLlSVL 0.39-0.45); head wider than long. canthus rostralis di stinct; tympanum 1/2 to 3/4 of eye diameter; supratympan ic fold Dot recognisable; subarticular and metatarsal tubercles weakly developed ; tips of fi ngers and toes round, not dilated; finger lengths 3>2> 1>4; toe lengths 4>3>5>2> I; rudimentary webbings on bases of toes; skin finely tubercu lar, particularly on upper ide of body and limbs. Upper side medium to light brown, with individually highly variable, asymmetric, metiially oriented spotted pattcll1 ; hind limbs with dark crossbars; a light dorsolateral stripe betwccn rostrum and femur, paralleled below by a dark brown one; sides li ght brown, with partly densely arranged dark brown dot ; a dark reticu lated pattern in the inguinal
Microhylidae
region, pat11y with 1-2 large, yellow ocelli; lower side whitish grey witl1 smal l brown dots which become more numerous towards the throat; posterior hal f of belly sometimes with dark-bordered wh ite spots; iris pale golden with black dots. The colour pattern and body shape show distinct phytomimetic properties. Tadpoles probably similar to those of K. pleurosligma. Ecology/Ethology Ka/ophrynus baluensis inhabits the leaf litter layer of th e oakchestnut forests between 1400 and 1800 m a.s.l. Along larger streams with a high noise level, the species keeps a distance of at least 2030 metres from the water. It has also tormed a colony in the midst of the chalet settlement of the Headquar1ers and is locally common. The males call bet>veen 6.00 p.m. and 6.00 a.m., but can occasionall y also be heard during the day. The maximlm1 calling activity is reached between 6.00 and 9.00 p.m. after rain show rs. The frogs remain mute during proFig. 11 - ~alopl/fYllus baluensis ~~ ~e !i~d9uarte.:s . longed dry periods. The call is emitted from holes in the ground, crickets, hairless caterpiUars, and particularly spirock crevices, and cavities in roots and dead wood ders) in slow motion, fixes it visuall y, and then and is velY difficult to pinpoint. Depending on the state of excitement and the prevailing weather conseizes it with the tongue in a sudden fon-vard moditions, the calls are emitted at intervals of 3-10 s tion. Defence behaviour includes fleeing via a series of and often answered by other males leading to the quick, 20-30 em long leaps. When touched it exformation of a chorus. These do, however, not represent insular, geographically defined calling compels large amounts of a white, very sticky secretion from its dorsal glands which is probably efmunities which (like in Megophrys baluensis) are isolated from one another acoustically, but rather fective enough to deter even anurophagous snakes. variable concentrations of groups of individuals The frogs uses its hands to rub its chest, belly, and within a widespread population. Nothing is known flanks, while the hind Limbs take care of the distal portions of the body. It will then make a handabout its reproductive biology. Kaloplilynus baluensis forages for food at night, stand raising the posterior body off the ground. Without losing balance, the hind limbs rub over whereby it stalks its prey (ants, tennites, small
121
Microhylidae
the posterior two thirds of the body in circular motions. It is to be preumed that this serves to distribute glandular secretions over the skin. Call: (+20°C) The calls are emitted as euphollic, very weakly pulsed single notes at intervals of 6- lOs. Frequency 1.8 to 5 kHz; catTier frequency 2.6 kHz; harmonics at 7 kHz; duration 125 rus on average; frequency modulation present. Distribution Thi species is so far known only from Mt. Ki naba lu where records are as yet further limited to the Headquarters and Mesilau region. As the very characteristic voice of this microhylid has never been recorded from above 180'0 III altitude, it appears likely that the altitudinal information relating to the type locality (2200 m) is erroneous (SMITII ] 93 L) .
1.50
1.60
1 .70
~ 10 5
[
-----=-; ~~,
122
.,~
Fig. LV III Sonagram and oscillograms of Kaloph ryfllls baluensis (Headquariers, 1550 m; 7.00 p.m. +20°C)
1600·1700m
K. baluensis
~~
' - - - r
1500 -1600 m
o
1.90 sec
•
Fig. LlX Distribution of Kafophrynus baluensis in the area of the Headquarters.
z-
1.80
Microhylidae
KaloplJrynus pleurostigma TSCl-IUDI, 1838 Ka.loplllYlllis pietJrosligma Tsnlum, 1838: 86 - Type locality;
umalra Kalophl)'III1S plcllrDsligma plellrosligma. IN GER, 1966: 132 KaloplllYlIlIS plellrosligma. SMITH 1931 : 21; l"GER 1956: 396, 1978: 3 12; MANTHEY 1983: 21;MALKMt:S 1985: 8,1987: 279. 1988c : 176. 1988d: 9; IN C;ER & STl.BlING 1992: 41-51 : Il'iGER., STUEB[]\'G & T AN 1996: 362; M-\,\lTHEY & GI{OSSMA',N 1997: 54: LAKIM CI at. 1999: 32; INGER el at. 2000: 16
Diagnosis A medium-sized brown frog; habitus stout; head narrow; snout pointed , its length equal to eye diameter; a black ocellus at each groin' similar to K. baluensis, but larger.
Description
Tadpoles endotropllic; head-body portion ovaL snout broadly rounded; mouth terminal; lips not expanded; spiracle situated mid-way between eye and end of body, without free flap; tail lanceolate, tip bluntly rounded, 1.6-1.9 ti mes the headbody-length; total length of mature larvae up to 14 rum. Head-body grey to olive above, heavily dusted below; caudal muscle brown; fillS dusky or not. EcologylEthology K. pleurostigma inhabits the leaf littcr layer of dipterocarp primary forests between altitudes of 500 and 800 m on Ml. KinabaJu . Males call at night, to a considerably lesser extent also during the day, from holes in tbe ground, below leaf litter, or the edges of spawning sites. Tlus species chooses shallow temporary pools on the forest floor for breeding, such as holes in logs and road ruts filled with rainwater (m: 2b, d, e, b). The eggs are spherical, unpigmented, and deposited in clusters 2.5-3 mm in diameter. The development fro m gastrulation to metamorphosis takes 16 days at a water temperature of24-29°C. Newly metamorphosed froglets are
Males up to 50 mm, females up to 57 mm; hind limbs ShOlt, TLlSVL 0.36-0.46; head wider than long; nostril closer to tip of snout than to eye; loreal region almost vertical; tympanum usually distinct, 0.5-0.75 times eye diameter; finger tips round, not expanded; finger lengths 3>2> I >4; subarticular tubercles distinct; tips of toes like those of fingel's; toe lengths 4>3>5>2> 1; deFig. 116 KalophrYllus plellrostigma from Poring region. gree of toe webbing very variable, uSlI311y webbed over 1/3 to 3/4 of their lengths; dorsal skin granular (with keratinized sp ines in male specimens); belly coarsely granu lar; supratympanic fold present, but often obscure; vocal sac and nuptial pads present. Reddisb brown to loam-coloured or greyish brown above, usually with large dark blotches; dark crossbars on upper sides of hind limbs; flanks dark, sharply set off from dorsal coloration by a light line nmning fi'om tip of snout to groin; a black spot in front ofthe insertion of the hind limb; lower side yellowish whitc; throat usually dark brown, peppered with white dots; bel ly with dark spots; ilis bright reddish golden.
123
Microhylidae
cxcellent jumpers performing leaps of up to 60 em (LIM & G 1991). The defence, comfort, and hunting behaviours of the adult are very similar to those shown by K. baluellsis. Their diet consi t of spiders, mall caterpillar and beetles, but mainly of ants and termites. The stomach of a single individual contained up to 150 ants. When foraging the frog may also a cend into the lowest strata of shrubs up to 20 cm above the ground. a ll : The call consists ofa long series of un pulsed notes . The calls emitted by a frog from epilok (eastern abah) were irregular both in length
Genus Kaloula GRAY, 183 1 Specie of the genu Kaloula are present from Korea and northern China to the Le ser Sunda Islands, the Phi lippines, and ri Lanka. The Bomean species are medium-sized , emifo orial, stocky frogs with short thick hind limbs; head wider than long, snout broadly rounded, vertical in profile; canthus rostral is indi tinct; tympanum concea led by skin; tips of fingers not expanded, webbing rudimentary; large inner metatar al tubercle present. Males po e a median subgu lar vocal ac and coar ely granular skin on the throat. The frog depo it their egg in temporary pools. Tadpo le with oval body, flattened above; snout broad and truncate; cyes lateral, 6-7 times as widely separated from each other a the nostri ls; mouth terminal, no beak or labial teeth . The tadpoles of Kaloula di ffer from all other Bomean microhylid larvae by having the opening of the piracle at the end of the body above the anal tube. Anal tube medial, projecting over the lower caudal fin. Tail
(27-285 m ) and interval (138-348 m ), with a dominant frequency of 420-625 Hz at +27.5° . Frequency modulation present. Distribu tion Kalophrylllls pleurosfigma i found on the Malay Peninsula, umatra, Borneo, and the Philippines. TIle taxonomic tatus of the population from Java remains undetermined (MAT LI, CHANARD & ABIIITABHATA 1996). Mt. Kinabalu : 0 reliable records exist from loealitie above 800 m altitude (Poring - Sg. Langanan), with those from BWldu Tuhan, Lumu Lumu, Kamborangoh ( MITH 1931 , I~GE:.R 1966, MATsl;1 1979) being mo t likely referable to K. ba/uensis.
pointed, almost twice as long as head-body length. Larvae are uspen ion feeders. Adults are terrestrial , but may ascend bu hes and trees during period of heavy rainfall. They also occur in human settlements, agriculturally u ed land, and secondary crub; K. baleata i also found in primary fore ts. They are nocturna l and find suitable heltcrs for the day in holes in the ground , beneath pile of rocks, and fallen log. The genu comprises I I specie , 2 of which are widely distributed on Borneo. Their occurrence on Mt. Kinabalu is limited to cultivated lands in the marginal zones.
Key to the Bornean species of Kaloula Webbing reduced to basal fragments; a wide, pale brown stripe between eye and groin and from eye to arm .............. ....................................... pulchra • Webbing reac hes the middle subarticular tubercle on inner edge of 4th toe; pattern different from above ........... .... ............. .... ......... .... ......... baleata
Tab. 10 Distribution of Kaloula-species on Borneo. Kaloula baleata (MOLLER, 1836) Kalolllaplllcilra GRAY, 1831
124
I I
Sabah
Kinobalu
• •
• •
Brunei
Sarawak
Kalimanlan
• •
• •
Microhylidac
Kaloula haleata (MOLLER, 1836) Bombinotor baleaws MOLLE R, 1836: 96 - Type locality: Krawang, Java
Diagnosis A tocky frog with short, thick limbs and a short, broadly rounded snout; lores not concave; tips of fingers dilated into truncate disks; thc webbing of the toes reaches the middle articular tubercle on the inner edge of the 4th toe. Descr iptio n Males up to 61 mm, fema les up to 67 mm; TLlSVL 0.29-0.36; otherwise see Diagnosis. Finger lengths 3>4>2> 1; toe lengths 4>3>5> 2> 1; wcbbing 1(1.5), 2(1.5/ 1.5),3(2/2),4(3/3), 5(2); inner metatarsal tubercle oval, compressed, elevated; outcr one round ' skin granular above, or with low, rounded tubercles; smooth or rugose below' throat of males granular; often with a fo ld betvveen eye and insertion of forelimb. Dorsum dark or pale brown with a large dark spot in the centre; sides brown with dark spots; often with white spots in the groin region; pale brown below, with darker spotting, lighter towards the belly; iris golden brown. Tadpoles very similar to those of K. pulchra. E cology/Ethology K. baleata is a commensal species which finds refuges for the day under rocks, wood, and in the soil, but which has also been found in tree holes up to 2.5 m above the ground. Following heavy rainfalls, the males form noisy choruses along ponds, road ditches, and the margins of rice paddies. They inflate their bodies substantially during calling. The eggs are small and are deposited in clusters in puddles, road ditches, and small ponds (m: 2b). The food ofK. balaeta comprises small insects, pm1icularly ants.
Call: (+23 °C) A series of loud groaning honks in slow sequence. The notes are pulsed with a duration of280-300 ms, with a distinct decrease in intensity; dom inant frequency l-3 kHz.
Fig. LX Sonagram and oscillogram of Kalollia baleata
F rom top to base: Male of Kaloula baleata Fig. 11 7 from Poring, 600 m. Fig. 118 from Tam~n Negara ~P~Pc~insular ~al~ysia .
Distribution Peninsular Malaysia, the Greater Sunda Island, Palawan, and the Lesser Sunda [slands to F lores. Mt. KinabaJu: Rainwater puddles on the road side at Poring (600 m a.s.I.), edges of primary forest.
• kHz
': r
5.0 sec
j 125
Microhylidae
Kaloula pulchrll
GRAY,
183 1
Kaloultl pulchra GRAY, 1831: 38 - Type locality: China Kaloula pl/lehra, M...:rslJl 1979: 319: i KGER, S TUEBI:-JG & TAN
1996: 362
Diagnosis A very stocky, rottmd frog wi th short, thick hind limbs and a short, rounded snoul; lores weakly concave ' fingers long, their tips ex panded into truncate disks; toe webbing not extending beyond the level of the proximal subarticular tubercle of the 4th toe; a pair of prominent flanges on the sole of each foot between the heel and the base of the toes.
Description
Fig. 120 Ventral view of Kaloliia pllichra from Laos.
Males up to 70 rom, females up to 75 mm; TLlSVL 0.25-0.35; habitus similar to that of K. bafeata; finger lengths 3>4>2> 1; toe lengths 4>3> 5>2> 1; webbing rudimentary; inner melatarsal tubercle strongly raised, Quter one transversely oval; skin thick, finely pebbled or lubercular above; below smooth or faintly granular; lower back with widely scattered bump ; indistinct fold from eye to axi lla.
Dorsum brown w ith scattered dark pots; a w ide irregular, whitish to pale brown stripe from eye to f,rroin and from eye to insertion of arm; throat brown (to black in males); chest, belly, and undersides of legs yellowish with brown spots, sometimes form ing a network.
-
Fig. 119 Kalaufa pulchra fro m Laos.
126
Microhylidae
per second; dominant frequency 2500 Hz; no frequency or intensity modulations (HEYER 1971).
~
~ ~
=i
@ ~--~------------------------------~ Fig. .!21
!a~o l e _of Kalal/fa p l!.fchra ~om
Laos.
Tadpoles olive brown to black above, sometimes with go lden dots, and lighter beneath spotted with white ; fins transparent with small dark patche ; maximum head-body-Iength 14 mm , tail 26 mm . For further description sec under genus. Ecology/Ethology The bullfrog is a distinct commensal commonly encountered with in human settlements and anthropogenously disturbed habitats, in particular in land used for agriculture. It sUl"Vives periods of drougJ1t in underground hiding places which it excavates itself and which al so serve as shelters during the day. The males fonn choruses after rainfalls which may often include very many individuals (> 100) along temporalY stagnant water bodies. Here, the extremely inflated frogs can often be seen floating on the water surface . Females wh ich are attracted by the calling are immediately seized on arrival in an axillary amplexus. They dip their heads into the water at intervals, simu ltaneous ly raising the rear and releasing spawn . Any temporary pool may be uti lised for breeding purposes by this species (m : 2b, c) . The small eggs are laid in floating masses and develop at great speed. The frogs feed on small insects with a predilection for ants. Call: A loud groaning honk consisting of a single, partially puLsed note (0.56-0.60 s). Each note is composed of J 8-21 pulses at a rate of 32-35 pulses
Distribution The species is present throughout Southeast Asia. It has probably been introduced to the Sunda Islands and now occurs eastwards up to the Lesser Sunda Islands and Sulawesi. It is commonly presumed that it has reached Borneo only more recently. Mt. )(jnabalD: Only a single female specimen has become known from the Headquarters region , 1665 m (MATsUI 1979). This is likely to have been a translocated specimen as this spec ies is not normally a montane one and has never since been found on M.t. Kinabalu again.
Genus Metaphrynella
P ARKER,
1934
The genus Metaphrynella comprises only two species: the montane M. pollicaris (BOUl-ENGER, 1890) of the Malay Peninsula and the Bomean endemic M. sundana ( P ETERS, 1867) known from Sabah, Sarawak, Brunei, and Kalimantan .
Metaphrynella sundana
(PETERS,
1867) ,
Calohy la slIr/dana PETERS, 1867: 35 - Type locality: Pontianak (Borneo) Melaphry nellaslJndalla, M.~.KMt:S 1988e: 176, 1988d: 7, 1989: 185, 1992a: J 10; II\"GER & STUEBING 1992: 41-51 ; MALKMUS 1994b: 224, 1995b: 35; H OFF\lANN 1995a: 10,2000: 37; M ALKMUS I996b: 20·26, 1996c: 282; INGER, STlJEBING & T AN 1996: 362; H OffMANN 2000:37; L AKIM et al. 1999: 32; I NGER et al. 2000: 17
Diagnosis A tiny, stocky, grey to black, arboreal microhylid with a pointed snout and moderately short limbs. The underside of each finger with enonnously enlarged tubercles at the base which form accessory adhesive organs. Tips of the fingers dilated, tips of the toes not expanded. Description
Males up to 23 111m, females up to 25 mm; TLlSVL 0.42-0.47; head as wide as long; snout projecting,
127
Microhylidae
128
Microhylidae
Fig. 124 Hand of Metaphrynefla sundana from Poring, about 600 m.
"c '"
a little longer than eye diameter; lores almost vertical; tympanum visible, but somewhat concealed, 315 of eye diameter; finge rs w ith rudimentary, fleshy webbings; finger lengths 3>4> 2> I; toe lengths 4>3>5>2> 1; toes webbed for 1/2 to 3/4; a weak, elongated inner, but no outer metatarsal tubercle; tibio-tarsal articulation reaching tympanic region; skin above and below smooth or feebly granular, sid cs with larger tubercles; a curved supratympanic fold between eye and axilla; males with a subgular vocal sac, but no nuptial pads. Upper sides grey to brown, with irregularly scattercd black spots which are pcppered with small white dots ; hind limbs with or without. black crosslines; elbows often yellowish brown; lower sides dark grey to blacki h brown with lighter clouding, occasionally yell owish; iris reddish golden with fine black dorting.
ic.: ~
Fig. 125 Tree hole with foam and eggs of Metaphrynella SUI/dana near Poring. 550 m.
continue throughout the night until about 5.30 a.m . The largest calling community was discovered in a bamboo forest bordering the Sg. Langanan. The frequency and timbre of a call is often substantia lly altered by the size and shape of the treehole. Calling communities are always farther than J00 m
Fig. 126 Eggs of Metaphrynella sUl1dana from Poring. 1/
1
~.
Ecology/Ethology This microhylid can be found in tree holes (often in bam boo internodes) tilled with rainwater in dipterocarp primary forests, on Mt. Kinabalu at 500-850 m a.s.1. The males call at night from treeholes 0.3 to 10m above the ground with the openi ngs of these cavities not exceeding 10- 12 cm in diameter. Call ing activity was noted at Poring to begin as early as 4.00 p.m. in some instances, but normally commences from 5.00 p.m. These calls
129
M icrohyl idae
away from mountain streams if the latter create a high background noise. A male was observed emitting advertising calls during amplexus. [n February in Poring region a female was observed depositing 1300 eggs (egg diameter I mm) in ajelly string (5-6 rom diameter) in a water filled tree hole, 1.8 m above ground. The water surface was covered with a light whitish foam (HOFFMAN"N 2000). A clutch covered in foam was found in a bamboo interoode (m: 2g) at Poring. Call: (+23 to 25°C) The caU consists of a single, uopulsed piping note (80-150 ms) emitted at intervals of mostly > 3 s; dominant frequency 8001500 Hz; frequency modulation present, with peaks up to 21000 Hz. Distribution Metaphtynella sundana is endemic to Borneo. Mt. Kinabalu: As yet, this species bas on ly been recorded from the region between Poring and Sg. Langanan.
Genus Microhyla
TSCHUDI,
1838
The genus Microhyla is rustributed from China to India and Sri Lanka, in Southeast Asia eastwards up to Bali. All five of the Bornean species are small (except for M. berdmorei under 30 mm SVL), and have conical body shapes with a rounded snout and relatively long, strong hind limbs. They all share a distinct tendency for a reduction of the length of the first finger. Maxillary and vomerine teeth absent, tympanum usually not visib le. Males pos-
Fig. LXJ Sonagram and osciUograms of Metaphrynella sundana (Bat Caves/Poring, 580 m; 9.00 p.m. +23 °C)
0.6
0.8
1.0
1.2 sec
kHz
4 sec
-
':,..----,
sess a median subgular vocal sac. Eggs are deposited in very small pools. Tadpoles without beaks, eyes lateral, mouth terminal, spiracle medial, ventral, with movable flaps, or with an opening at the end of tbe tube near the anal tube. The shapes of the tail and the lower lip are often indicative of the species. Adults of the Bornean species live in tbe leaf litter layer on the floor of primary and old secondary forests below 700 m altitude. Five species are known from Borneo, two from Mt. Kinabalu.
Tab. 11 Distribution of Microhyla-species on Borneo. Sabah
Microhyla berdmorei (BLYTH, 1856) Microhyla bomeensis P ARKER, 1928 Microhyla maeulifera I NG ER. 1989 Microhyla perparva I NGER & FROG~ER, 1979 Microllyla petrigellQ I NGER & FROG1'm R, 1979
130
• • • • •
Kirrabalu
Brunei
•
•
•
•
Sarawak
• • • •
Kalimantan
•
Microhylidae
Key to the Bornean species of Microhyla 1 No disks and dorsal grooves at bases of toes .... .. ................ ................... ........... .... .... ..... maculifera • Disks and dorsal grooves at bases of toes present ........................................................................... 2 2 3n1 and 51h toes webbed up to disks ............. ..... . 3
• 3 n1 and 51b toes not webbed up to disks ............ ... .
............ ............................................... borneensis 3 I" fmger 1/3 oflength of 3"\ toes fully webbed. ..... ......... ........................... ................... berdmorei • I " finger reduced to a rudiment.. ............ ......... . 4
4 Belly pigmented, but with an irregularly shaped area free ofmelanophores; dark cross bars on hind limbs bordered with light .. .... .......... ... petrigena • Belly without melanophores; dark cross bars on hind limbs without light bordering ..... perparva
Microhyla borneensis
PARKER,
1926
toe webbing 1(1),2(111),3(212),4(3/3), 5(1.5-2); an oval inner metatar al tubercle and a conical outer one; tibio-tarsal articulation reaching a Little beyond tip of snout; kin above and below mooth, with scattered low tubercles with tubercles and low ridges on sides; weak supratymparuc fold; males with subgular vocal sacs; nuptial pads wanting. Dorsum light brown, with dark brown to purplccoloured markings between eyes; large light-edged mark on middle of back; lips spotted with black; broken black stripe on s ide ; hind limb dorsally with crossbars; dark pot on knee; lower urfaces yeUowish, mottled with brown on throat and chest. Head-body portion of tadpoles almost triangular, with very wide nout, flat above, sloping upward from nout to root of tail which is then prominently rai ed; tail tapering gradually from root into a long, narrow filament; eyes lateral, nostrils dorsal, mouth te rminal ; lips not expanded ; tail length 2.5-2.8 times head-body-Iength . Head-body grey, lighter ventrally; fins dusted with melanophore . Total length 22.4 mm (st 32-40). Ecology/Ethology
Microhy la borneensis is an inhabitant of the lowMicrohy /a bomeeflse P A RK~ R 1928: 473 - Type locality: " Kidi District, arawak" (- Bidi District?) Microhy /a bomeensis, P ARKER 1934: 128; M ATSU I 1979: 323; INGER & S11JI!8 ING 1992: 41 -51 ; I N(;~R. STUhB INO & T AN 1996: 362; I"GfR et a!. 2000: 17
Diagnosis A tiny brown frog with a moderately stout body and long limbs; dorsal urfaces of disks of fingers and toes with longitudinal grooves; 3 rd and 5 th toe not webbed up to disks; I " finger les than 113 length of 3Td; tympanum invi ible.
land primary forests dwelling the leaf litter layer on the forest floor and low vegetation « 30 em). It is both diurnal and nocturnal. This specie breeds in small accwnulations of rainwater, sometimes. in wallows u ed by wild boars (m : 2c) . The tadpoles
Fig. 127 Microhyla borneensis
...
Description Males up to 18 mm, females up to 23 mm; TLlSVL 0.62-0.70; head broader than long; snout s lightly pointed, projecting beyond mouth, longer than eye in diameter; canthus ro tralis rounded lores weakly concave; tips of outer 3 fingers sl ightly dilated into disks; fmgcr lengths 3>4>2> I; tips oftoes more expanded than tho e of fingers; toe lengths: 4>3>5>2> I;
131
Microhylidae
are mid-water suspension feeders. Nearly nothing is known about its natural history. Distribution Microhy la borneensis is known from the Malay Peninsula and Borneo. Mt. Kinabalu : Only one specimen has a yet been recorded, a fema le from Poring/Hot prings (555 111).
Mic~ohyla I NGE R
&
petrigena
FROGN ER,
1979
Microhy /a pelrigella I NGE R & FROGNER, 1979 : 318 - Type loca lity: Nanga Tekali t, Kapit District, Sarawak Microhy/apelrigel1o . I NGER, SlUED IN(, & TAN 1996: 362; I NGER ct al. 2000: 17
Diagnosis A very mall, moderately stout, brown trog with long hind limbs; 1<' finger reduced to a rudiment; all toes but the 4'b fully webbed. Description Males up to 16 mm, females up to 18 mm; TLlSVL 0.60-0.70; head obhlsely pointed; tympanum invisible; fingers with slightly expanded tips; 1Sl finger
Fig. L28 Microhy la p etrigellCl
rudimentary; finger lengths 3>4>2(> I); 2 low metacarpal tubercles; tips of toes more expanded than those of fingers, with longihldinal groove on dorsa l side; toe lengtbs 4>3>5>2> 1; toe webbing 1(0), 2(0/0), 3(0/0), 4(0.5/0.5) 5(0); low inner metatarsal tubercle , no outer one; males with subgul ar vocal ac . "Top of head dark grey; a constricted dark grey figure on back beginning at level of shoulder and extending to groin then curving forward in a broad band low on side and reaching just beh ind eye; except for these dark areas, back and side grey white; dorsa l surface of limbs ru set with broad darker bands bordered by light grey to ivory, darkedged stripe; throat, chest and belly dark greyish; belly with wide irregu lar, meandering white area" (INGER & F ROGNt'-R 1979). Head-body portion oftadpolcs oval in shape, flattened above, snout broadly rounded; eyes lateral, visible from be low; nostril dol' aJ ; mouth tenninal; anal tube in ventral fin; tail abruptly tapering subtemlina ll y to a short tip ; tail length 1.5-2 .0 times head-body-Iength . Head-body black above and on sides, whitis h below ; caudal muscle black; fins without pigment except for a ve rtical black band near end of tail. Total length up to 17 mm (st 37). Ecology/Ethology Microhyla petrigena is a lowland species, dwelling the floor litter in primary fore ts where it feeds on tiny invertebrates, mainly ants. Microhy la pefrigena breeds in water-filled rock pothole (diameter and depth < 30 em) along small fore t streams (m: J g). The tadpoles are mid-water s uspcn ion feeders filtering tiny organism . Nearly nothing i known about its Ii fe hi tory. Di tribution This specie is endemic to Borneo. M t. Kinabalu: Only known from the Poring area.
132
Ranidae
Ranidae Ranid frogs are almo t co mopolitan in distribution and ab ent onl y from southern outh America, the West Indie , most of the Aus tral ian continent, and many Oceanic i land . The Bornean representatives of this family exhibit all anatomical features typical for ranid frogs . They have a finni temal pectoral gird le, a sacrum with cylindrical diapophyses, and the omo ternum and sternum are 0 sified. Bony postzonal sternal elements are present, and so are eight holochordal , procoe lous presacral vertebrae and palatines. A parahyoid is absent, and the cricoid ring i complete. Maxillae and premaxillae are dcntate. The fingers are generally free of webbings , whi le we bbings b tween the toes are well-developed (except in the genus Toy /orona D U[lOI S, 1987). The pupil is horizontal. Mating takes place in an axillary amplexus . Ranids are extremely variablc with regard to size and habitus. They have conquered a multitude of ecological niches. The majority are ground dweller , but many can climb as well. Many pecies are close ly associated with water and never wander far from it. Other are, in contrast, typical in habitants of the forest floor, e.g. Limnolleclesfinchi and Rona [uc/t/osa. which may be encountered far away from any water body. Various species e.g. Rona sigllata
and lvferistogenys orphnocllemis. are known to leave the water bodies where they have completed their metamorpho is; they spend their early lives in the forest, only to return to aquatic habitats on reaching maturity. pawning usually takes place in the water, although there are exccptions i"n the fonn of species like Limnonectes finchi and L. pa/avallensis which depo it their eggs on land and later tran port the tadpoles to a water body. The larval development involves a tage of free-swimmi ng tadpoles (except in Tay /orana) . All tadpoles have beaks, denticles, and a sinistral spiracle. Most ranid larvae can be found in ponds, but some have adapted to streams (e.g. Merislogenys, Huia) . Bornean Ranidae are classified as belonging to the genera Fejervarya, Hoplobatrachus, Huia, Ingerana, Limnonecles, MeristogellYs, Occidozyga, Raila, and Slauro;s, and comprise altogether 38 specics at present. Representative of this fami ly are munerous up to an altitude of about 1500 m on Mt. Kinabalu, but only a few species arc present above thi elevation (e.g. Limllonectes kuhiii, MeristogellYs J..:illaba/uensis. Slaurojs tuberilingui ). The 22 species repre~ senting 8 genera which occur on Mt. Kinabalu account for some 60% of the ranid species of Borneo.
Key to the Bornean genera of Ranidae (based on
MANTH EY
&
GROSSMANN
1997)
Skin membrane between the two outer metatarsals present (A) ................ ..... .... ..................... .. ...... ... 2 • No suc h membrane ; tips of fingers and toes spatular (B) .... ......... ... .... ....... ..... ...... .... Iflgerafla 2 Tips of fingers with marginal folds on the disks (C) and usually dilated (D) ............ ... ....... .... ..... 6 • Tips of fingers without marginal folds on the disks and not or only weakly dilated .. ..... ...... ......... ... 3 3 Vomerine teeth present (E) .... .... ..... .. .... ............ 4 • Vomerine teeth absent ..... ........ ....... . Occidozyga
133
Ranidae
F 4 Dark ventro-Iateral line between axilla and groin present (F) ................. ........................ FejervaryQ • No dark ventro-lateral line ............................... 5 5 Dorsum with elongate skin folds, tips of fingers pointed (0) ............... ..... ......... .. Hoplobatrachus • Dorsum without elongate skin folds, if present tips of fingers rounded and blunt (H); males with fanged like process (I) on the lower jaw (except L. finchi. L. palavanensis) ............ Limnonectes 6 Digital bones T-shaped distally (J) ............. ...... 7 • Digital bones not dilated distally ... .. ..... .... Rana 7 Habitus very slender; transverse folds beneath the phalangeal tips (K) ............................. '" Staurois • Habitus not so slender; no such transverse fold .
J~
............ ........... ........ .......................................... .. 8 ·8 Tympanum sunken ........... ...................... .... Huia • Tympanum not sunken. .... ..... ....... Meristogenys
Genu Fejervarya
BOLKAY,
1915
A genu of relativ small to moderatly large robust bodied frogs . The kin texture is relatively mooth, but covered with longitudinal oriented skin glands in the form of folds . The tips of toe and finger are feature less, without any enlargements or di cs. Recently, Fejervarya was elevated from a subgenus of Lilllllonectes to genus rank by DUBOIS & OHLER (2000). They summed up some phenetic differences that exist between Fej ervary a and Limnollectes, such as the shape of the digits of adults.(OBLER & DUBOIS 1999), their types of male secondary characters (BOULENGER 1920) or the
K
differences in the mouthparts of their tadpole (FEI et al. 1991) . Additionally, D UBOIS & OHLER (2000) pointed out the existence of a dark ventro-lateral line from armpit to groin , a unique common derived charactcr only to be found in Fejervarya . More significantly even, the analyscs based on mtD A sequencing ofVEN ES (1999), MARMARYOU et al. (2000), BOSSUYT & MILI NKOVITCIl (2000) and EMERSO et a1. (2000) sugge t that Fejervarya is not the sister-group of Limnonectes (see also genus Limnonectes). Unfortunately, no up to date complete diagnosis of the genus Fejervarya was published so far. The genu Fejervarya comprises 14 species (D BOIS & O HLER 2000), two of which are pre ent on Borneo . Mt. Kinabalu is home to one of these species.
Tab. 12 Distribution of Fej ervarya-specie on Borneo.
Fej ervarya callcrivora
(ORAVENHORST,
Fejervarya limllocharis
134
(WIEGMANN,
1829)
1835)
Sabah
Kinabalu
Brunei
• •
•
•
Sarawak
Kalimantan
•
• •
•
Ranidae
FeJervarya limllocharis (GRAvENHORST, 1829) Ratla limtlocharis GRAVE 'HORST, 1829: 42 - Type locality: Java Ratla Iimtlochalis limnocharis, MATSUI et al. 1985 : 153 Raila IimllocharLv, M ALKM S 1994b: 244, 1996c: 285; H OFF~"'N:< 1995b: II Raila (Lilllllollecles) limllocharis, H OFF MAr>'N 1998b: 89
Diagnosis A small to moderately large terrestrial frog with a very granular back. In contrast to F cancr;vora the webbing of toes is deeply excised, leaving at least one phalanx of each toe free. Description Males 32-50 rum, females 48-60 rom; habitus slender to somewhat stout; head longer than wide; snout round or sl ightly pointed; vomerine teeth present; tympanum visible, about 60% of the diameter; supratympanic fold present; l SI finger longer than the 2 nd ; finger tips not dilated; marginal disk folds absent; fingers without webbings; toe tips not dilated' webbing formula 1(1),2(111), 3(1 /1) 4(3/ 3), 5( 1-2); ilmer metatarsal tubercle oval, outer one round and very small; dorsal skin with irregular longitudinal folds. Males during the mating period with a grey, M-shaped gular pattern; subgular vocal sac entire; l '1 finger with whitish to light grey nuptial pads. Dorsal coloration grey to brown with irregular dark spots; ventral side white. Specimens with a white dorsal stripe are found syntopically with those where such is absent. Body shape of tadpoles oval; fringe of fins tapering to a point; upper side of body olive green with black speckles; lower side white; tail tip striped with black or entirely black; head-body-Iength 13-15 rom; total length up to 45 mm; labial teeth rows formu la 2(2)/3. EcologylEthology Fejervarya limnocharis is a nocturnal ground dweller, usually found in the environs of water bodies (ponds, canals, etc.). It is an exp licit commensal which avoid primary forests . Following heavy nightly downpours, the males fonn large calling communities on the banks of stagnant wa-
From top to base Fig. 129-131
Diffi:~nt
/im/loc/taris from
dorsal patterns of FejervOIya
Poring.':, 135
Ranidae
ter bodies in the morning hours. The frogs are often encountered sitting in the waste water ditches of human settlements. This pecies shows a predilection for lower altitudes. It feeds on a variety of insects, millipedes, and occasionally snails. The larvae favour stagnant water bodies (m: 2b). CaJl: A series consists of rapidly emitted, raspy chirps which are repeated at irregu lar intervals. Note duration 8-120 ms ; internote interval 32-440 ms; dominant frequency 1-3 kHz; hannonics at 5 kHz.
Distribution This species is widely distributed from Pakistan, India and Sri Lanka to Japan, to Borneo and Flores. Mt. Kioa ba lu : Sayap, Poring, MATSUI (1985) reported "one adult female between Layang Layang and Kamborangoh, 2000 m " which probably refers to a translocated specimen.
Genus Huia
YANG,
5.40
6.00
5.80
6.20 sec
kHz 10.67 sec 10,------------------------------------------.
5
:1
:t
:K
'.
:[
Fig. LXTI Sonagram and osciJlograms of Fejervary a Iimllocharis (Peninsular Malay~ia, 40 m; ~ OO p.m l
1991
Taxonomic remark: The species originally included in the composite genus A molops were rearranged by Y ANG (1991) and are now allocated to the genera Amolops, Huia. and Meristogenys. The respective revision was based mainly on larval traits such a the number of continuous lower rows of denticles in the tadpo les of Amolops being three, whereas they count at least four, usually more, in Huia and Meris/ogenys. The upper beak is entire and Mshaped in the tadpoles of Huia, and divided in those of Meris/ogenys.
136
5.60
The distribution of the genus Hula covers the entire Sunda region and Yunnan (China). It includes more or less slender, long-legged frogs which are capable of very long leaps and perfectly adapted to a life in mountain stream habitats. The 1st finger is of equal length, or slightly longer, than the 2nd • The tips of the fingers and toes are dilated, the toes are connected by almost fully webbings. The length of the tibia is > 0.7 times the SVL. Like those of Meristogenys. the tadpoles of Huia are adapted to rapidly flowing stream by a large ventral sucker disk (gasteromyzophorous). The genus is represented on Borneo by only one species which oecurs nearly all over the is land (Sabah, Sarawak, and Ka limantan).
Ranidae
Huia cavitympanum Rana cavity mp
B OULENGER,
(BouLENGER,
1893)
1893a: 525 - Type locality:
Kina Balu Rona cavitymplI/Ium, B ARn.ETT 1894: 203; H ANITSCII 1900a: 73; B OULF.'10bR 1920: 193 ; VAN KAMP EN 1923: 2 11 ; SMITH 193 1: 30; INGER 1962/63: 45 Amolops cavitympanum, INGER 1966: 260.1978: 3 13; M ATSUI 1979: 337; M ANTIIEY 1983: 22; I N .F.R & STUEBlNG 1992: 42; M,\LKMUS 1994b: 224 Huia cavitympanum, Y ,\NG 1991 : 29; M ALKMUS 1992a : 118; H m FM ANN 1995b: 9; INGER, S TUEBIl\G & T AN 1996: 363; ING ER & TA N 1996a: 559; L AKlM et aJ .1999: 32; INGER et al. 2000: 16
Diagnosis A moderately large terrestrial frog with a relatively contrasting colour pattern. Its sunken tympanum makes this species unmistakable.
Description Males 42-52 mm, females 68-80 mOl; TLlSVL 0.73-0.76; habitus slightly stout; head longer than wide; vomerine teeth present; nostril mid-way between eyes and tip of snout, or slightly eloser to the latter; canthus TOstralis slightly rounded; tympanum visible, in a depression on the side of head ; distinct, strongly angled supratympanic fo ld between eye and axilla; l SI fmger longer than 2 nd ; finger tips dilated, outer ones twice as wide as others; diameter of tip of 3r • finger equal to that of tympanum ; marginal disk folds present; webbing limited to basal portion of finger or absent; weak supernumerary metacarpal tuberc.le present; tips of toes dilated and rowlded, slightly smaller than finger tips ; toes fully webbed ; small, oval inner metatarsal tubercle present; outer metatarsal tubercle small or absent; dorsolateral folds present; dorsal skin
}lg.132 Huia cavily mpanwll from Sayap, Sg. Kcmantis lIDO m. ~~~~~~~~
c
~
~
a: ~~~~4~~~~~O
137
Ranidae
smooth, lateral with numerous oval glands, smooth on throat and chest; ventral skin slightly granular. Males with yellowish nuptial pads on I st metacarpal, and a pair of subgular vocal sacs in gular pockets. Coloration above chocolate-brown; light dorsolateral stripe from behind the eye; sides and upper sides of limbs yellowish brown with dark brown spots or cross bars; venter white to yellowish orange; iris light golden above, cinnamon-coloured below; lips with dark spotting. Bo.dy oftadpoles oval, flattened; head-body-length up to 25 mm, total length up to 70 rnn1; eyes and nostrils positioned dorsally; distance between nostri ls larger than distance between no tril and eye; distance between nostril and tip of snout almost twice the distance between the nostrils; lips form a ,large oral di sk and lack pap ill ae; labial teeth rows formula 8(3-9)/5(1) to 12(4- 12)/6(1); both beaks entire; ventral sucker disk covering almost the entire be lly; caudal fin limited to posterior two thirds of tail; coloration dorsally and laterally very dark,
Genus Ingerana
D UBOIS,
1986
The genus Ingerana comprises several diurnal and nocturnal, relatively small species which are recognized by their linger and toes disks being spatulate in shape. The toes are not fully webbed to the disks. The larvae of these frogs are unknown. On Borneo, a single species is represented, which is known from Sarawak, Brunei, Sabah, and Mt. Kina balu.
Ingerana baluensis
(B OULE GER,
1896)
CornuJer ba/ueflsis BOlJLb" OFR 1896b: 449 - Type locality:
Mount Kina .Balu, North Borneo COrlluJer ba/ueflsis. V~'1 K..·\ MPEN 1923: 241 ; SMITH 1931 : 30 Micrixa[lIs ba/uellsis . lliOE R 1966: 255, 1978: 313 ; MA..'lTIIEY
1983: 22;
I NGER
&
STUbBING
1992: 42
/flgerafla ba/tlcnsis. M ALKMUS 1994b: 244; TAN
16
138
1996: 363;
INGER
& T AN 1996a: 559;
I NGER,
INGER
TUEBI:-lG & et al. 2000:
marbled lighter; cream-coloured ventrally. Tadpoles at an advanced stage of development show a light line on the tibia. EcologylEthology Huia cavitympanum is a nocturnal terrestrial frog inhabiting the banks of fast-flowing streams in primary forests (250 to 900 m a.s.I.). Here, it is commonly found perched on rocks or hiding beneath them. Adults wander widely through the forests (INGE R & STU EBrNG 1997). The ma les call individually. Tadpoles live in the strongest currents of medium-sized streams, clinging to bedrock or boulders (m: la, b). Call: The call consists of a weak, high-pitched chirp. Distribution Endemic to Borneo with records from Sabah, Sarawak, and Kalimantan. Mt. KinabaJu: Sg. Wariu, Sg. Kemantis, g. Kipungit, Sg. Langanan, Sg. Mamut.
Diagnosis A small terrestrial frog with conspicuous spatulate finger and toe tips. Description SVL males 20-25 mm, females 24-30 mm; TLlSVL 0.48-0.56; bead wider than long; snout rounded; vomerine teeth present; canthus rostral is rounded; diameter of tympanum 40-50% of eye diameter; supratympanic fold present; 1" finger slightly longer than 2nd ; finger tips spatulate, narrower than tympanum' marginal disk folds present; webbings between fingers limited to the bases; no supern umerary metacarpal tubercles; tips of toes spatulate, wider than the dilations of the fingers; webbings between the toes covering about their entire lengths ; inner metatarsal tubercle oval, no outer one; dorsolateral folds absent; skin granular; males with subgular vocal sacs, but no nuptial pads. Dorsal coloration lightreddisb brown with obscure dark markings; Ijght dorsolateral stTipe; limbs pale
Ranidae
brown with dark cross bars; venter cream-coloured; throat, more rarely also the chest, spotted or marbled with brown; lips striped; iris golden brown with fine black reticulation. Ecology/Ethology Jngerana baluensis prefers moderate altitudes of montane forests where it is primarily found along mountain streams. Here, the frogs can be discovered in rock crevices with spring water seepage on gravel banks, such as the 6-8 m wide Sg. Kadamaian. Their eggs are large (2 .2-2.5 rom in diameter) and unpigmented. The frogs are, however, occas ionally also found far away from water. They are excellent climbers on the vertical rock faces and capab le of long leaps. Distribution Endemic to Borneo, the frog has been recorded from Sa bah, Sarawak, and Brunei. Mt. Kinabalu: Poring, Sayap, Sg. Kadamaian above Kiau.
Genus Limnonectes
FITZINGER,
1843
The genus Limnonectes comprise mainly of nocturnal, moderately to very large telTestrial frogs that were previously ass igned to the genu Rana. The 2nd finger is never longer than the 1st . In contrast to the genus Rana. the finger tips are only slightly or not dilated. Limnonectes is further distinguished from Rana by the absence of marginal disk folds on the finger tips (except in the subgenus Elachyglossa ANDERSSON, 19l6) and the presence of well-developed odontonoid processes at the lower jaw of male specimens. They mostly lack advertisement cal ls and often also lack dorsolateral folds which are often distinctly developed in Rana. These frogs do not usually wander far from water, but there are exceptions, such as L. findli and L. paiavanensis. The genus Limnonectes is present on Borneo with l2 species and thus fon11S one of the most diverse
139
Ranidae
limnocharis do not represent the actual sister group of the remaining taxa from the genus Limnol1ectes
genera of Bomean anurans. Mt. Kinabalu i home to four of these species. Taxonomic remark: The monophyly of the gcnus Limnonectes, as previously proposed by D UBOIS (1986 "1987", 1992). was often que tioned. Based on morphological and molecular data, EMER ON et al. (2000) showed that at lea t L. cQncrivora and L.
in their analyses. Therefore it seems to be likely to place these two pecies in the genus Fejervarya, as previously proposed by D UBo) & OH LER (2000) and actually accepted by Il\GER ct a!. (2000) [or L. Iimnoc/zaris (sec also genu Fejervarya section).
Key to the Bornean species of Limnonectes 7 Dorsolateral fo ld absent .................................. 8
Upper eyelid smooth ....................................... 2 • Upper eyelid with spiked tubercles .................. . ......................................................... malesianlls
• Interrupted dorsolateral fo ld present ..... ibanorus
8 Toes fully webbed ............................................. 9
2 Tympanum obscured by skin ........................... 3 • Tympanum clearly visible or only partially obscured ....................... ........................................ 5 3 Toes not fully webbed ..... .......... ...... .... ............ 4
• Toes not fully webbed ................................... 10 9 Canthus rostralis distinct ................... ieporinus • Canthus rostral is indistinct ...................... ingeri 10 SVL less than 55 mm, males with vocal sac ..... .. " .. .. ....... " ... ,................................... kenepaiensis • SVL greater than 55 mm, males without vocal sac ............................................... paramacrodon
• Toes fully webbed .................................... kuhlii 4 Dorsolateral fold ab ent; lemon-yellow below ... ........................ ....................................... laticeps • Interrupted dorsolateral fo ld present; white below ... .. .... ............. ... ......... .............. ...... rhacodus
11 Dorsal ski n smooth, except for reverse V-shaped marking ............ ,.............. ,......... ,.. paial'anensis • Dorsal skin not smooth ...... ...... ....... ...... ... finch;
5 Tympanum fully visible ........... ............ .. .. ..... ... 6 • Upper posterior third of ty mpanum obscured by skin ..... ........... .... ....................... ..... ..... asperatu.I'
6 Lower jaw with two tooth-like bony projections near the symphysis .......................................... 7 • Lower jaw without proj ections .............. .... ... 11
Tab. 13 Distribution of Limnonecles- pecies on Borneo. Sabah
Lilllllollectes asperallis (1I'
Kinabalu
I
' i ..--f- - .- - I
140
Brunei
Sarawak
• • •
• • • • • • • • •
I
• • • • •
I
I
• • •
I
I
I
- r• I
I I
•
Kalimantan
• •
• • • ---•-•
•
Ranidae
It'~~~
E
~fil!I_g]j
a: @
Fig. 134 A male of LimnonecIes fi'!..chi from Poring is carrying tadpoles.
Limnonectes finchi
(INGER
1966)
Raila microdisca ]illchi I l\cER, j 966: 222 - Type l()cality: Kalabakan, Tawau District, Sabah Rwwfinchi. INGFR & SrLEBING 1992: 42; I NGER, STl. EBINC & T.,,"\! J 996: 363: M AL KMUS I 996c: 293: L.~KJ't et al. 1999: 32 LimnOllecles/inchi, I NCLR CI al. 2000: 16
Diagnosis A small ground-dweller with a conspicuously granular back, weakly dilated toe tips, and not fully webbed toes. Description Males 32-39 mm, females 39-45 mm; TLlSVL 0.53-0,61; habitus rather slender; head slightly longer than wide; vomerine teeth present; tympanum visible, its diameter ca. 2/3 of that of eye; 151 and 2nd finger equal in length; finger tips rounded; toe tips relatively weakly dilated into disks; webbing fonnula 1(0), 2(010),3(010),4(2-3 /2-3),5(0);
outer metatarsal tubercle absent; dorsolateral folds interrupted' dorsal skin with numerous, usually longitudinal tubercles; a reverse V-shaped dermal elevation on back behind the shoulder girdle. Dorsal coloration grey to greyish brown with dark spots; venter white to yeUowish; throat speckled; lips striped; light interorbital band. Head-body-Iength of tadpoles up to 8 mm, total length up to 25 mm ; tails of tadpoles with dark eros bars; labial teeth rows fonllula 2(2)/3(1). Ecology/Ethology Limnonectes finchi is a nocturnal ground-dwelling frog of primary and secondary rain forests , The male occupy calling sites beneath leaf litter on the ground . Most of the time tbe frogs live far away from any water bodies wllich they access only for the purpose of releasing tadpoles, The about 50 eggs are 2 mm in diameter, with a dark brown and a light brown hemisphere. They are deposited beneath dead leaves on the forest floor
]41
Ranidae
within a clutch. The males guard the clutch and transport the larvae to water bodies once they have reached stage 24 or 25 (following GaSNER 1960). Up to 30 tadpoles of 8-9 rom in length may be transported in one batch. A male was discovered near Poring carrying seven tadpoles to an ephemeral pond next to the road along the boundary of Kinabalu Park. This pond is fed only by a small temporary trick le of water, and its water was mu rky and practically stagnant, its banks werc muddy and overgrown to the waterline with low bushes and grasses. Neither the males nor the tadpo les show any morphological adaptations for this type of larval trans location. Free-swimming tadpoles were also found in rainwater puddles on the forest floor (m: 2b). Adults feed on small insects and spiders. Th.is species' altitudinal distribution ranges from 200 to 1150 m, but the population densities are higher at low elevations.
Call: The caJJ consists of a short trill. Distribution Endemic to Sabah. Mt. J(jnaba)u: Poring.
Limnonectes kuhlii (T
CHUm,
1838)
Raila larhlli' TSCIlUDI. 1838: 40 - Type locality: Java Raila paradoxa MOCQUARD, 1890: 148 Type locality: Kina Balu (syn. after BOULENGER 1891) Ralla ~1,hlii. BOULENGER 1920: 62; VA"; KAMPE:\' 1923: 178; SMITH 1931 : 30; INGCR 1954a: 250.1956: 403,1962/63 : 46.1966: 196,1978: 313: M ATSUI 1979: 325 ; [~GER 1985: 48; INGER & STU£llr~G 1992: 42; WONG 1994: 32; HOFFMANN 1995b: II ; IN GER & STUEBINO 1997: 140; LAKI\·I et al. 1999: 33 Lilllllollecles k"hfii. MALKMI.-S 1994b: 225. 1996c: 285 ; INGER, STUEBING & TAN 1996: 363; INGER & TAN 1996b: 75; INGER cl al. 2000: 16 Dicrogiosstls Iet/hii, MANTBEY & D ENZER 1982: 13; MAl\'TIIEY 1983: 22; MALKMUS 1985 : 8, 1987: 280, 1988c: 176, 1988d: 7, 1989: 186, 1991a: 30 Ran(l (Limllonecles) kllMii, MALKMUS 1996c: 285
Taxonomic remark: Limnonectes !cull/ii is cons idered a species comp lex. Morphological and molecular evidence by KOSUCH & OHLER (unpublished) indicates that L. !cuhlii from Mt. Kinabal u is not conspecific with ne ither topoty pical (Javanese)
142
specimens nor those from Kalimantan, Sumatra and the Asian mainland. A second ind.ication for the existence of a species complex are the results of a molecular study of EMERSON et al. (2000), referring to L. !cuhlii from Sabah, Brunei, Java and Taiwan. Further analyses are required to demonstrate if those frogs from Mt. Kinabalu represent a new species or refer to the currently ynonymized taxa Limnonectes conspicillatus (GONTHER, 1872, type locality Matang, Sarawak) or L. paradoxa (MocQUARD, 1890, type locality Kina Balu, Sabah). Comparative studies of and with these two taxa still have to be undertaken. Diagnosis A moderately large to large ground-dweller, rarely observed outside water. It has a stout build, is dorsoventrally compressed, and has eyes which are directed upward in a conspicuous manner. Description Males 44-74 mm, females 50-67 mm; babitus stout; limbs short and muscular; head wide, snout slightly rOtUlded; eye directed upward; lower jaw with two tooth- like bony protrusions near the symphysis; InllJes with vomerine teeth; tympanum invisib le; supratympanic fo ld distinct, slightly arched; 1" fi nger longer than 2 nd ; finger tips rounded; no marginal disk fo ld s; 2nd and 3rd finger wilh dermal fringes; subarticular tubercle oval, but not elongated; no supernumerary metacarpa l tubercles; toe tips disk-like, weakly dilated; toes fu ll y webbed; inner metatarsa l tube rcle oval, shorter than I" toe; outer metatarsa l tubercle absent; dorsolateral fo ld s absent; skin above coarse, almost smooth below. Males without vocaJ sacs, Jarger than fema les, their bony protrusions in the lower jaw a lso larger, and the head wider; no nuptial pads, but a diffe rently coloured marking of their respective positions on the I" finger. Coloration variable and differing regionally; dorsum yellowish , reddish, brownish , or greenish, usually with dark spots, and always with a dark interorbital band; venter cream-co loured, often with small grey spots; specimens with and others without a white dorsal stripe are found syntopically.
Ranidae
c: c:
'E"
~
a:
o Fig. 135 Limnonectes kllhlii from the Headquarters, 1550 m.
Fig. 136 Male of Limnollecte" kuhlii from the Headquarters, 1550 m.
Body of tadpoles oval, slightly flattened; brown with irregular dark spots; tail tapering to a point, roughly spotted in its first half, in the second half with four stripes which are not sharply delineated ; head-body-Iength up to 14 mm, total length up to 38 mm; labial teeth rows formula 2(2)/3( 1).
EcologylEthology Limtlonectes kuhlii is ba ically a nocturnal ground dweller, but not exclusive ly so. It may also be found active on day climbing up to 50 cm high in vegetation. Tbe frogs are almost always encountered in or near water, in primary forests as well as in the surrounding of human settle-
ments. They show a predilection for the banks of water bodies with moderate to weak currents, where they often reside on the lee-s ide of boulders or fa ll en logs , but a lso in ditches and canal s. A morphological peculiarity is a hardened s pinou s proces at the end of the terminal phalange of at least the 3,d toe which is used in defense . By lateral movements it wi ll pierce the sk in of an attacker, and this also applies to tbe hand of a collector (camp. M ALKMUS 1996, MALKYlUS, BR OHL & ELTZ 1999) . The eggs are deposited in the water and attached to vegetation, c.g. to overbanging tufts of grass. They hatch after about 10 days. Tadpoles are primarily
Fig. LXUJ Oral disc of Lilllll Oll ectes kllhlii (Sg. Kcmantis, 1000 m).
Fig. 137 Tadpole of Limllon ectes kuhlii from Sg . Kemantis, 1000 m.
A. Malklnu.
143
Ranidae
found in calm sections of streams near the banks, in residual ponds, and seepages (m: Ie, f, h, i; 2a). The altitudinal distribution ranges from sealevel to 1750 m a.s.l., but the species becomes very rare above 1600 m. Ca ll : pecimens at low elevations do not call. "Males living at higher elevations at Mt. Kinabalu emit a soft tweet" (1:-1GER, STUEBI:--IG & TAN 1997). Specimens observed near the Headquarters often emit dog-like barking and growling sounds. Distribution (see the introductory taxonomic comments) India, southern China, Southeast Asia, Greater unda Islands. M t. Ki nabalu : Sg. Kemantis, Sg. Kinateki, Sg. Kadamaian , Kenokok, Marei Parei , Kiau , Tenompok, Bundu Tuhan, Sg. Silau- ilau, Sg. Liwago, Kamborangoh (up to 1750 m), Pinokok, Poring, Sg. Mantukungan , Sg. Kipungit, Sg. Langanan, Sg. Mamut, thc record "Kamborangoh; alt. 7.200 feet" (SMITH 193 I) i doubtful.
Limllonectes leporinus (ANDERSSO , 1923) Raila macrodon var leporina AND~R SSOK, 1923: 121, 124 - Type
loca li ty: Tumbang Maruwei and North-caslcrn l3orneo, Bulungan Raila macrodoll. S~tlTH 193 l: 30; SMEDLH 193 I : 61 R(lIla bly/hi (not Raila bl)"lhi! B OULENGIoR, 1920). It-. GER 1966: 162 (part.) Raila blylhi (not Raila blyihii Bo t..L~:-J ,ER, 1920), I NGfR & STIJEDrNG 1992: 42; HOfFMAt-." 1995b: 10; I NGER, STUFBISG & TAl- 1996: 363 Limllollec/es bly/hi (nol Raila bly l"ii Boul EN(;ER , 1920), M ilU<.MU 1994b: 226 L!II/I/OlleCles leporina I I'GER & TAN 1996a : 560; It-. GER et al. 2000: 16 Raila /eporifla L\KIM ct al. 1999: 33
Taxonomic remark : J:--IGER & T Al'-l (1996a) examined the type of leporinus ANDER 0 , 1923 and found it to be identical to a ll Borncan frogs previously named blylhii BOULENGER, 1920, but not to be conspecific \'lith blythii from tbe Malay Peninsula. These morphological result have been confirmed by high genetic differences found in
144
biocbemical (Kosu H 1994) and molecular studies (EMERSON et ai. 2000). D iagnosis A large, robust terrestrial frog which can easily be confu ed with LimnoneCfes ingeri. L. leporinus exhibits a dark stripe below the canthus rostralis, while L. ingeri does not. L. /eporinus furthemlOre bas a narrower head, longer hind limbs, and a more sharply defined canthus rostralis than L. ingeri. Description Males up to 125 mm, females up to 175 mm ; maturity on reaching 85-90 rom; TLlSVL 0.52-0.62; body massive; head longer than wide , snout pointed; vomerine teeth present; canthus rostralis sharply defincd; tympanul"n visible; supratympanie fold present; lower jaw with two tooth-like, bony protrusions near thc symphysis; 1'1 finger longer than the 2 nd ; linger tips rounded ; marg"inal disk folds absent; toe tips dilated ; toes fully webbed; inner metatar al tubercle oval, no outer one; dorsolateral folds absent; skin smooth. Males without vocal sacs; with a larger head than fcmale and larger mandibular protrusions. Light brown to reddish dorsally, juveniles with dark pattern; limbs with dark crossbars; whitish ventrally; throat finely dotted ; dark interorbital band present; lips banded with black; a dark stripe below the canthus rostralis and from eye to lympanwl1. Body shape of tadpoles oval, about 0.75 times as wide as long, slightly flattened ; eyes and nostri ls
Fig. 138 Limllonec/es leporinus from Sg. Sasapang.
'"
:>
~~~ ~ <;;
::;
...... ~!".-..'lI
cr:
~~:'-'"'"""", I)
Ranidae
positioned dorsa lly, mouth ventrall y and subterminally; upper lips short; anal tube dextral ; tail muscular; Coloration "graybrown with dark markings, which form an almost zigzag row of dark spots on the tail" (INGER & STUEBING 1997). Head-body-Iength before metamorphosis ca. 36 mm, SVL of freshly metamorphosed froglets 9-11 mm.
Kip un git, Sg. Mamut, Sg. Mantukun gan, Sg. Sasapan.
Limnonectes palavanensis (BOULENGER, Rana p(l/avallensis
B OLLENGER
1894)
1894a: 85 - 'I)- pc loca li ty:
Palawan, Philippines Ecology/Ethol ogy Rana palavanensis, B OW" ENGER 1920: 59' VA,"\1 J(,\MPEN 1923: Limnonectes /eporinus is a nocturnal ground182; S~UTII 1931 : 30; lNGER & SrU~RING 1992: 42; I NGeR, dweller inhabiting the banks of streams and rivers. STUEBING & T.~." 1996: 363; H OFFMANN 1994: 223, I 995b: II ; INGER & TAN 1996a: 56.3: L AK.lM el al. 1999: 32 Resting places are normally not in the water. The frogs can be found at night sitting among leaf litter Rana microdisw palav(wellsis, JI'10ER 1954a: 250, 1966: 222, 1978: 313; MATsUI 1979: 333 near their water body, on banking rocks, or on fallen Hylarana microdi5Ca palavallensis, MANTIi.EY 1983: 22 logs. They spend tbe day hidden in moist shelters, Umnonectes palal'anensis. MALK \1 US 1994b: 244; I NGIlR el. al. e.g. beneath rocks on the banks. They prefer pri2000: 16 mary rain forests, but also accept biotopes in secDiagno is ondary forest. This specie is not present in cultiA small terrestrial frog with a characteristic dorsal vated land where it is replaced by L. ingeri. pattern in the form of a reverse V. Tbe males excavate spawning craters with diameters of 30 to more than 50 em in tbe gravel sediDescription JIlent of shallow sections of rain forest rivers in Males 27-30 mm, females 35-40 mm; habitus rather which the fema les depo it their eggs at night. The slender; vomerine teeth present; diameter of the eggs are uniformly yellowish grey (vs. light with a tympanum ea. 2/3 of that of the eye; 1st and 2nd da rk bemisphere in 1. ingeri). Tbe males have not really an advertisement call, but both sexes emit finger of equal lengths; finger tips rounded; marginal disk folds absent; toe tips disk- li ke and equa l-sounding, deep, soft sounds during tbc matslightly dilated; outer metatarsal tuberele absent; ing season whereby, according to EMERSON & I NGER (1992), the sounds emitted by the males must be regarded as responses to female calls. Fig. 139 LimllOllectes paiavanellsis from Sayap. Sg. Wariu, Tadpoles may be found in a variety of habitats ca. 850 m. although there appears to be a predilection for lea flitter sediment whose layered structure provides idea l situations (m: I d, e). Strong currents are clearly avoided. The larvae are generalized suspension feeders with stomach analyses revealing mainly algae (blue algae, but also euglenoids and diatoms), plant matter, and funga l hyphens. Adult specimens prefer larger prey items including crabs and even other frogs. Its altitudinal distribution extends from 65 to 800 m, but the frogs are most numerous below 300 m a.s.1.
Distribution Endemic to Borneo with records from Sabah, Brunei , and Kalimantan. Mt. Ki na balu: Sg.
145
Ranidae
c:
I :x: ~
I)
Fig. 140 Umnon ecles J!alavl!!lensis from Sayap, Sg. Wariu. ca.~ 50 m.
toe webbing 1(0), 2(0/0), 3(0/0), 4(2-3/2-3), 5(0); dorsolateral fold present; skin above smooth except in the area of the reverse V-marking. Dorsal coloration medium brown to reddish brown with continuous. light dorsal stripes; dorsal . reverse V-marking dark; flanks light grey to greyi h brown, with a tinge of orange; lower side of head greyish wh ite; chest, belly, and lower sides of legs usually lemon-yellow; dark spots in posterior dorsolateral region; coloration of the limb correspondin g to that of the flanks , with dark brown crossbars; light and dark interorbital bands; lips striped with black, usually spotted with yellow or white . Tadpoles mall; tail tapering abruptly near the end to a slim tip; coloration light or dark brown with darker dots; tail speckled with black; head-bodylength up to 8 mm ' total length up to 30 mm; labial tceth rows fonnula : 2(2)/3(1).
146
Ecology/Ethology Limnonectes palavanellsis i a nocturnal ground dweller of the primary rain forests . primarily in mountainous country. The frogs are oftcn cncountered in the leafiittcr layer or in low vegetation (up to 20 cm high) far away from the ncarest water body. The male occupy advertisement sites beneath leaf Iittcr on the forest floor. Adults prey upon ants, tcrmites. and other small terrestrial invertebrates. Ovipo ition take place on land beneath leaf litter. The clutches are comparatively small with a maximum of35-50 eggs and the eggs being rather large in comparison to both the body sizc of the females and other ranid frogs. Eggs up to 2.6 mm in diameter, a light and a dark hemisphere whereby the light one is lighter than in the eggs of L. jinchj. Parental care and transport of larvae like in L.finchi. Free-swimming tadpoles was ob erved in small,
Ranidae
shallow pools of temporary streams and other shallow accumulations of water (m: If, i). The altitudinal distribution ranges from sea level to 1300 m.
Ca ll : Series duration 5-6 s) of20 or more slightly pulsed notes; note duration 50-70 ms; internote interval 140-250 ms; frequency modulation in each note rising from 2.4 to 3.3 kHz with hannonics at 910 kHz.
56.20
56.30
Distribution Palawan (philippines) and Borneo. Mt. ](jnabaJu: Kiau, Sayap, Bundu Tu han .
56.40
56.60 sec
56.50
I'
k1~Z
61 .91 sec
~I;ifil((l •
LXIV Sonagram and oscillo-
5
Genus Meristogenys
1991
Fig. grams of Limnonecles paiavonensis (Gunung Mulu, 150 m; 8.00 p.m.)
YANG,
•
•
•
•
•
•
•
~l(({(((j{( •
••
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t
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f
•
••
mtHtf+.~I#H~~H:~~.; .
The distribution of the genus Meristogenys is limited to Borneo. Jt was not before 1991 that YANG split the composite genus Amolops into Amolops, Huia, and Meristogenys, mainly on the basis of larval differences (see also Genus Huia). The genus Meristogenys comprises small (especially small males in many species) to large, moderately slender, and long-limbed frogs. The head is longer than wide with a rounded snout. The eyes are large, and the tympanum is visible. The tibia length corresponds ""rith more than 0.7 times the SVL. The JSl flllger is generally longer than the 2 nd ; the tips of the fingers and toes are dilated, with the diameters of the finger tips being smaller than the diameter of the tympanum. The toes are fully webbed, and marginal disk folds, vomerine teeth,
and dorsolateral folds are all present. The skin is finely granular to smooth. The reproductive biology could be studied in detail in M . cf. orphnocnemis (MALKMUS 1998). Pairs migrate in amplexus to a mountain brook where the eggs are deposited under water on smooth rocks exposed to the current. They arrange themselves spontaneous ly after spawning to form a single-layered egg disk in which the eggs it side by side connected to one another by the jelly mass. After hatching the larvae form a school which remains stationary in an unusual density next to the now empty egg disk before they eventually disperse after 3-5 days. It is still unknown whether the reproductive biology of Meristogenys is different from that of Amolops or Huia. Like in the genus Huia, the tadpole possess a large , circular, ventral sucker as an adaptation to thei r Iife in streams with strong currents. The head is substantially broadened and flattened below.
147
Ranidae
F~g.
143 Tadpole of Mer~t0!5enys cf. k!..n abaluens!!,
They exhibit an M-shaped , keratinized upper beak which is costate distally, The V-shaped lower beak may sometimes also be divided. The muscular tail is 1-1.5 times as long as the body, The tadpoles possess small venom glands on the body and tail whose arrangements are indicative of the species. The larvae of the species Huia cavitympanwn differ from those of the species of MeristogellYs by having a larger number of rows of denticJes on the anterior labium and by the structure of the lower beak. In contrast to most of the other anuran gcnera the tadpoles of the species of Merislogenys are more easily distinguished than the adults whose body shapes and colour patterns arc very similar, As the morphological points of difference for the adults have often bee'n used in the literature in inaccurate or contradictory manners, we shall refrain from adding an identification key to the genus here , Of the 8 species in this genus, 4 occur on Mt. Kinabalu,
From top to base: MerislOgenys cf. orphnocnemis Fig. J41 A pair from Sg. Liwago after egg deposition, Fig. 142 Tadpoles form a school after hatch.ing.
Tab. l4 Distribution of Meristogenys-species on Borneo, Sabah
MerislOl!ellYs umoropalumlls (MATSI;I, (986) MerisroJ!efl)IS jerboa (G ONTHER, 1872) Meristol!ellYs killabaltlellsis (1'1GER, (966) JlJeristof!ellYs macrophthalmus (MATSU I, (986) Meristof(eflYs orpJlllocllemis (MATSUI, (986) Meris/of!eIlYs phaeomerus (II'GER & GRITIS, (983) MeristoRellYs poeciitls (INGER & GRlns, (983) Mer;stof!eIlYs whitehead; (BOULENGER (887)
148
• •
Kinaballl
•
Brunei
•
•
•
•
•
•
Sarnwak
Kalimantan
• • • • • •
•
•
•
Ranidae
Meristogenys amoropalamlls (MATSUI, 1986) Amolops amoropalamus M AT ' VI, 1986: 623 - Type locality: Bank ofSg. Pa Rilllan, 1300111 alt., Gunuog Tapai Sai. Krayan Country. Sarawak border of East Kalimantan IImolops sp. A, M ATSUI 1979: 340 Am%ps omoropaloll1l1s, MALKMUS 1989: 190, 1994b: 227, 1996c: 283 Merisrogenys (lmoropal(jlllu.~, YA.">G 1991 : 33; M M.KMUS I 992a: 11 7. 1994c: 89, 1998: 22; I NGER. STUEllING & TM 1996: 363: L.~KIM et al. 1999: 33; I NGER et al. 2000: 16
Diagnosis A moderately large terrestrial frog with a conspicuous iris which is divided in brown and golden sections; distinguished from M. kinabaluensis by the outer metatarsal tubercles, from M. mphnocnemis by the not fully webbed toes, and by the smaller tympanum from M. whiteheadi (3/5 or more of eye diameter). Description Males 30-40 mm , female s> 60 mm ; TL/SVL 0.72; habitus moderately slender; head longer than wide;
snout slightly pointcd; vomerine teeth small; nostri ls closer to tip of snout than to eyes; canthus rostralis distinct; tympanum about 80% of eye dianleter; supratympanic fold present; I " finger little longer than 2 nd ; dilation of finger tips about 1/3 oftympanum diameter; marginal disk fo lds present; su barticular tubercle oval, with weak, supernumerary tubercles on outer t\vo fingers; toe tips dilated; toe webbing 1(0), 2(0/0), 3(0/0), 4(2/2), 5(0); inner and outer metatarsal tubercles present, the inner one being long and narTOW; dorsolateral folds present; dorsal and lateral skin slightly granular, smooth ventrally; males with small nuptial pads on 1" finger; a pair of vocal sacs in gular pockets. Dorsal coloration I ight brown with dark dotted pattern; ventral side whitish with dark dots, particularly on hind limbs which also spol1 dark crossbars; tibia dusky ventrally; upper lip with li ght spots, lower lip with dark bands; dark band from eye to behind the tympanum; iris golden in upper part, reddish brown in lower. Habitus of tadpoles oval and flattened, with large ventral sucker; eyes situated dorsolateralIy; nostrils
Fig. 144 Meris!ogenys amoropalamlls from Sg. Liwago, 1500 m.
149
Ranidae
much c10scr to eyes than to tip of snout; margin of upper lip smooth, lower lip with papi llae; labia l teeth rows formula 7(4-7)/6(1) at stage 28 with a tendency to have increased to 8(5-8)/10 ( I) by stage 38; upper beak divided; dorsal root of the caudal fin at end of body, ventrally at the end of first third of tail; groups of glands scattered over the body with each of the granu lar open ings visible; more glands on bases of both fins; dorsal co lo ra ti on dark grey, lig ht grey ventrally. Eco\ogyfEtholol,,'Y Meristogenys amoropalamus is a nocturnal ground dweller of the primary forests closely associated to ra pi dly flowing streams (m: 1abc, e). Its altitudinal distribution ranges from 1300 to 1800 m
a.s.1. Distribution Endemic to Borneo with records from Sabah, Sarawak, and Kal imantan . Mt. Kinabalu: Sayap, Sg. Penatanm, Sg. Silau-Silau, Sg. Liwago (1500 m), Sg. Mesi lau (Mesilau Cave, 1800 m).
Meristogenys kinabaluensis (I GER, 1966) l~o~R, 1966: 266 - l y pe locality: Kiall. Mount Kina Balu, Sabab Raila whiteheadi, (not Rana whileheadi BO UI. ENGER, 1887)
Amolops killabaiuensis
I 890a; 146 (part); VAN KAMPEN 1923: 210; SMITH 1931: 30; WIIITEHMD 1893: 121 Amolops lrillaballlensis. I1\GER 1978 : 313, 1985: 65: MAl SU I 1979: 339; MA~mIBY 1983 : 22: M ALKMUS 1987: 282, 1989: 189. 1991a: 28, 1994b: 227, I 996c: 283; INGER & STUEDJ)JG 1992:42;Wo:>G 1994: 32;] GER,SrUF-Bl:-'G& T IIN 1996:3 13 Meri.flOgellYs k;nabaluells;s, YANG 1991 : 35; MALKMU S 1992a: 11 7, 1998 : 22; HOFl' MA'
Diagnosis The largest species of Meristogenys. It differs from all other species of the genus by the absence of outer metatarsal tuberclcs. Descrip tion Males 58-68 mm, females 75-93 mm; TUSVL 0.58-0.65 in males, 0.49-0.5 1 in females ; head
Fig, 145 MeristogellYs kinaba/uellsis from Sayap, Sg. Kcmantis, ca. 1050 m.
i J:
a: @
"------~..;:
150
Ranidae
widc; body and limbs lender; vomerine teeth present; nostrils closer to tip of snout than to eyes; interorbital distance mailer than diameter of upper eyelid; weak supratympanic fold between eye and axilla; Ist finger longer than 2n<1; finger tips dilated about twice widtb of finger; marginal disk fo lds present; no webbings between fIDgers; subarticular tubercle oval , with supernumerary tubercles on outer three fingers ; toe tips dilated; toes ful ly webbed; inner metatarsal tubercle oval and about 1/3 of length of 1" ""e toe; outer metatarsal tubercles absent; dor£
-
-
Fig. LXV Tadpole of MeristogellYs killabaluellsis from Sg. Liwago showing position of groups of glands.
151
Ranidae
Meristogenys orphllocnemis (MATSUI,
1986)
Amolops orpi1nocllemis MA1 SlJl, 1986: 625 - Type locality: Kampong Bundu Tuhan, ncar Kinabalu, 990 malt., Sa bah
Amolops j erboa (nol Rona j erboa G O~TH E R , 1872), M ATSUI 1979: 338 (part) Amolops from Kinabalu "small", Ii\'GER & GRillS 1983 : 6 Amolops orphnocnemis, I~oE H & STUED ~G 1992 : 42; M ALKM US 1994b: 244 , 1996c: 283
Meris/ogenys orplmocnemis. Y Af':G 1991 : 36; W O'1G 1994: 32; Il oFPM,\NN I 995b: 10; L'1GER, STUEBING & T AN 1996: 363 ; I"GER & T A" 1996a: 560 ; M ALKMUS 1999: 15: L,\K1M el al. 1999: 33 ; INGER et al. 2000: 16 Merislogenys j el'boa (not Rana j erboa G ON TlI ER, 1872). M ALKMUS 1992a: 114
Diagnosis Distinguished ITom M. whitehead and M. amoropalamus by the fully webbed toes, from M. kinabaluensis by the presence of an outer metatarsal tubercle. Description Males up to 43 mm , females larger than 60 mm; TLlSVL 0.7 I ; habitus slender; head longer than wide; snout slightly pointed; vomerine teeth present; canthus rostralis distinct; nostrils closer to tip of snout than to eyes; 1" finger slightly longer than 2 nd ; marginal disk fo ld s present; subarticular tubercle oval ; no supernumerary tubercles, but with light spots indicating their P9sitions on outer two fingers; toe tips dilated ; toes fully webbed; inner metatarsal tubercle oval, outer one round ; skin coarse, with distinct dorsolateral folds. Males with nuptial pads on I " finger and a pair of vocal sacs in gular pockets ; tympanum diameter greater (0.62 times eye ctiameter) than in females (0.42 times). Dorsal coloration choco late-brown , lighter laterally; ventral side whitish to cream-coloured; limbs greyish brown above, with darker crossbars. Habitus of tadpoles oval and com pre sed dorsoventrally, with large ventral sucker; labial teeth rows formu la 5(4-5)6(1) at stage 38 to 5(4-5)/9(1) at stage 41; upper and lower beaks ctivided. The oral disk transfonns into a mouth during metamorphosis on ly after the resorption of thc larval tail. This
152
must be viewed as an aid for the larva to withstand the strong current of its habitat. Ecology/Ethology Meristogeny s orphnocnemis is both diurnal and nocturnal with adults inhabiting the banks offastflowing forest streams. The males can often be seen perched in loose groups at about knee-height on the vegetation-covered banks next to cascades. The heavier females are mostly found on the ground. Juveniles may also be encountered far away from water on the forest floor or in low vegetation. The males call individually or in loose groups while perched on slightly raised advertisement sites, particularly on the banks of rapids or smaH waterfalls . M. orphnocl1emis is the only species of Meristogenys whose reproductive biology is known (see also description of the genus above). Before and during spawning the sexua l dimorphism becomes evident with the large females carrying the much smaller males (53-55% of the females ' SVL) in amplexus to the spawning ites . Once they have arrived, the fema le spends some time in carefully selecting a suitable spot by probing the surface with her hind limbs and ventrum in back-and-forth motions. The adaptation to spawning in strong currents (m: l a, b, e, e) must be viewed as providjng the best possible protection for the larvae from potential predators (such as the larvae of dragonflies and beetles) and competitors for food, as these prefer the calmer sections of the streams. The about 800 eggs of a female's clutch are tightly arranged in an egg disk and held in place by jelly. They arc attached to a smooth rock surface exposed to a strong current. The jelly is transparent when produced, but becomes mi Iky within 1-1.5 hours after spawning, after the animal poles of the eggs have turned upwards, and then changes into a bright whitish blue. The tadpoles hatch after about 24 days, but stay in a dense school for another four days, slowly ctispersing thereafter. When disturbed, communal larvae immectiately sou ~ht the contact with their sib lings, while those older than four days reacted with taking to flight. The known altitudina l distribution ranges from 190 to 1500 m, after LAKIM et at. (1999) up to 1800 m.
Ranidae
Call: (+22°C) The call resembles a muffled crack of a whip. It is emitted sporadically at intervals between lOs to several minutes; call duration 40 ms; the first half ofthe cal l weakly pulsed, the second distinctly pul ed ; pulse modulation present; frequency decreasing from 9.5 to 6.5 kHz; harmonics between 9.5 and 11.2 kHz. Distribution Endemic to Borneo with records from Sabah and Kalimantan . Mt. Kinabalu: Sayap, Marei Parei, Sg. Kinateki, Kiau, Sg. Kadamaian Sg. Liwago: Mesilau , Bundu Tuhan, Pinokok, Sg. Mantukungan, Sg. Mamut, Sg. Kipungit, Sg. Langanan, Sg. Tananansad , Sg. Sasapan, Sg. Manggis.
0.30
kHz
0040
0.45 sec
5.0 sec
,-------------------------------------,
10
Fig. LXVI Sonagram and oscillograms of Mel'istogellYs orphnocnemis (Sg. Kipun!,rit, 550 m; 8.00 p.m. +22 0 )
0.35
j
5 -
Meristogenys whiteheadi (BOULENGER, 1887) Nona Whiteheadi BOULE"OE R, 1887b; 96 Type locality: Mount Kina Baloo, North Borneo Rana whiteheadi, (part) M(X()UA RIl 1890a: 146; BARTL ETT 1894; 203; HANlTSCH 1900a: 73 ; B ou l fNGE~ J920: 195; VA~ K AMPU:-: 1923: 210; SMITH 1931 : 17 A molops j erboa (not Nana j erboa Gt:. 'THE R. 1872), (part) I:-;oER 1966: 264, 1978: 313 Amolops whiteheadi. MALlO,IU S I 994b: 227, 1996c: 283 Meristogenys whiteh ead;, YA NG 1991 : 40; 1 ·GER. STUEDING & TAr- 1996: 363; I NGER & TA" I 996a: 561 ; LAKI).1 et al. 1999: 33 : I NGeR et al. 2000; 17 Amolops whiteh eadi, INGER & GRms 1983: 5
cr.
Diagnosis Three species of this genus, i.e. M. amoropalamus. M. kinabaluensis, and M. whileheadi, share the trait of the 4th toe being webbed up to the distal subarticular tubercle. M. whiteheadi differs from M.
amoropalamus by having a smaller tympanum and being much larger, and from M. kinabaluensis in having a larger tympanum. Description Males up to 64 mm , females up to 93 mm; TLlSVL 0.69-0.70; head about as long as wide; snout round to slightly pointed; vomerine teeth present; nostrils closer to tip of snout than to eyes; canthus rostralis distinct; tympanum 40-60% of eye diameter; supratympanic fo ld present; 1st finger as long as or longer than 2nd ; finger tips dilated; marginal di sk folds present; subarticular tubercle very distinct; toe tips dilated ; toes fully webbed except for 4~'; inner metatarsal tubercle oval, outer one very small or even absent; dorsolateral folds present; skin granular, smoother ventrally, particularly towards throat. MaJes with large nuptial pads on 1&1 finger and a pair of vocal sacs in gular pockets.
153
Ranidae
Fig. 147 Merislogen),s whileheadi from Crocker Range NP, S~bah. Dark greyish brown to dark reddish brown dorsally, occasionall y with darker spots, sometimes with small light dots; ventral side whitish to cream-coloured; a dark streak below canthus rostral is and supratympanic fold; hind limbs with more or less indistinct crossbars. Habitus of tadpoles similar to the preceding species; with a large ventral sucker' total length 6577 mm; labial teeth rows formula 5(3-5)/5(1); upper and lower beaks divided. Ecology/Ethology Merislogenys whiteheadi is a nocturnal dweller of
the banks of fast-flowing streams (m: la, b, c, e) between 190 and 1500 m altitude, after LAKlM et al. (1999) lip to 1800 m.lt is the most common species on Sg. Liwago (1500 m). Distribution Endemic to Borneo with record from abah and Sarawak. M t. Kinabalu: Kiau, Kenokok, Lobang, Sg. Liwago, Mesilau, Pinokok, Sg. Kipungit, Sg. LangaJlan.
154
Genus Occidozyga & VAN HASSELT, l 822
KUHL
The genu Occidozyga is distributed from Bangladesh, Myanmar and Thailand through the Sunda Island to the Philippines. It comprises small, dorsally flattened, and quite plwnp frogs with short, muscular hind linlbs, which are very well adapted to their environments with regard to their colour patterns. The tympanum is at least partly concealed by skin, the snout is flattened, and the eyes bulge substantially. The tips of the toes are slightly dilated to form narrow, roundish disks. While maxillar teeth are present, vomerine teeth are absent. The two Bornean representative of this genus live in or on small puddles, seepage springs, or in swampy area . Their tadpoles can be found in accWTIulations of water of all orts. The larvae are quite remarkab le as to their body shapes and behavioural patterns. They have very slender bod-
Ranidae
ies and tails with the latter being furnis hed onJy with narrow fi ns. The oral disk i situated terminally the upper lip is very narrow, the lower hor eshoe-shaped. There are no denticle , but the upper border of the djsk has a knob- li ke structure. The beaks are heavy and fineJy serrated. The spiracle is sinistral. The tadpoles of both Bomean species are carnivorous feeding on small in ect and their larvae. The genus Occidozyga is represented OD Borneo by two species which also occur on Mt. Kinabalu .
Key to the Bornea n species of Occidozyga 1 Ventral side whitish, fa intly dotted or immaculate; webbing formula 4(010) ............................ laevis Ventral side spotted; webbing formula 4(2-3/2-3) ............................................................. baluensis
Tab. IS Di tribution of Occidozyga-spccies on Borneo. Occido<J'gCl ha/uellsis (130ULE:>IGER, 1896) Occido<J'ga [aevis (GON'J'HI:.R, 1859)
Occidozyga balue1lsis
(BouLENGER,
1896)
Oreobatl'acJllls baluensis B OULhNGER, 1896a : 40 I Ty pe locali ty: Mount Kina Balu, orth Borneo Ol'eobalrachus baluensis. HANITSCII 1900a: 74; VAN K AMPFN 1923: 229 Phl),noglossus baluensis. MITH 1931: 15 Ooeidozyga baluellsis. I'IGER I 954a: 250, 1956: 40 I, 1978: 313, 1996: 238; MATSUI 1979: 336; MAI'(TIIEY 1983: 23 Occido:yga baluellsis, LNGER & STUBING 1992: 42; M ALKM\;S I 994b: 244; I:o.'GER, STUEBING & TA'I 19~6 : 36 ; I NGER & T.~'1 1996a: 561; I NGER et a!. 2000: 17
Diagnosis A smaJl terrestrial frog of stout, compressed habitus, short, muscular hind limbs, and not fully webbed toes. A faint, U-shaped ridge, open below, in the middle of the back. Description Males up to 26 mm, femaJe up to 35 mm; TU VL 0.46-0.57; snout rounded; vomerine teeth ab ent; nostrils situated laterally; tympanum partly covered by skin, ca 0.5-0.6 times the djameter; supratympanie fo ld present; I iii finger as long as or shorter than 2nd ; finge r tips not expanded; ; toe tips di lated; webbing on 4th toe does not reach toe tjp; weakly developed subarticular tubercles; inner metatarsal
Sabah
Kinaoa lu
Onmei
Sarawak
Ka limantan
• •
• •
• •
• •
• •
tubercle oval, outer ODe smalJ or even ab ent; dorsolateral folds present; dorsal skin with irregular glandular ri dges, usua lly a dorsolateral one behind eyes and an inverse U-shaped one between shouJders; numerous small, round warts on sides; maJes with a pair of voca l sacs; yellowish nuptiaJ pads cover the dorsal and medial urface of J Sl finger between ba e and terminal pha lange. Dor al coloration between head and glandular ridge rcddish brown between shouJders, remainder of back light brown; ventral sides light brown with a brown or black banded pattern. Specimens with brown as well a those with entirely black backs are known from ayap. Head-body-Iength of tadpoles up to 8 mm, tota l length up to 26 mm; light brown, with a black streak between lower anterior margin of eyes and snout, and a da rk, vertjcal bar below eye; caudal muscle and low upper fin with small black dots (see aJ 0 description of genus). Ecology/Ethology This frog is a diurnaJ and nocturnal grou nd dwel ler, occurring in montane regions as well as in the lowlands in wet situations among leaf litter, roots, and OD gravel ground. Besides on poorly drained patches in the forests, it is commonJy eneowltcred
155
Ranidae
FIg.
156
l~
Occidozyga baluensis from a small tributary to Sg. Kcmantill, Sayap, ca. 1100 m.
Ranidae
on the banks of mall running waters and seepage prings (m: 2a). It altitudinal distribution rangcs from 65 to 1370 m. CaJ1: (+23°C) The note serie consist of hoar e, rasping, distinctly rhythmic and trongly pulsed elements of50-300 ms in duration ; internote interval 150900 ms; dominant frequency 1-3.3 kHz. Distribution Endemic to Borneo with records from Brunei, Sabah, arawak, and Kalimantan . Mt. Kinabalu: Poring , Kiau, ayap: Bundu Tuhan, Pinokok.
Occidozyga laevis (G "
TilER,
....
,...
..
kHz
50 sec
': ['---.- -.- -.- -.,-JIW-~-i:.-.-:';_.--l- 'i
Fig. LXVl I Sonagrarn and oscillogram of Occidozyga baluellsis (Gunung Mulu, 150 m; 8.00 p.m. +23° )
1859)
OxygiosslIs luevis G ONTIIER, 1859 (1858b) : 7 - Type locality: Philippine Orcidozyga luevis. MALKM l!S 1994b: 244 ; Lt\lUM et al. 1999: 33: I NGER et at. 2000: 17
Diagnosis A small terrestrial frog of stout, compressed habitus with fully webbed toes. This species closely resembles juvenile Limnonecfes kulifii, except that in 0. laevis the inter-ocular distance is equal to or smaller than the eyelid. Furthennore, 0. laevis ha only one tooth-like protru ion on the tip of the lower jaw as compared to two in the case of L. h:u hfii.
Fig. 151 Occidozyga laevis from Taman sular Malaysia.
Description Males 20-3l mm, female 25-48 mrn; TLlSVL 0.41-0.51; head flattened ; vomerine tceth ab ent; nostri ls ituated dorsally to dorsolaterally; canthus rostralis strongly rounded or indistinct; tympanum concealed by skin and therefore invisible; supratympanic fold weak, between eye and axill a; length of 1" finger equal to or longer than 2 nd finger tips not dilated ; no marginal disk folds; toe tip dilated, tip of 51h toe only twice a thick as all others; toes fully webbed; inner metatarsal tubercle shorter than half I " toe; outer meta tar al tubercle minute or (usually) absent; no dor olateral [o lds; dorsal skin smooth in anterior part, with
some sma ll white ri es posteriorly; imi lar ri e n dorsal and lateral sides of hind li mb, in males also on throat; skin on flank with round to oval warts. Dorsal coloration grey or brown with a darker pattern. A morph with a white vertebral stripe exists. Ventral ide cream-coloured, immaculate or with some small dots; ventral side of hind limbs usually with small dots. Tadpoles small , very slender; with a long tail and very low caudal fin; body and margins of the fins with dark pots; head-body-Iength up to 10 mm, total length up to 25 mm.
157
Ranidae
Ecology/Ethology Occidozyga {aevis is a nocturnal species. It lives solitarily in slow-flowi ng water bodies (m: 1t), rice paddies, and small puddles (m: 2b), and are rarely encountered outside of water. TIley feed on insects and fre shwater crustaceans. Altitudinal distribution: 0-500 m. Call: The call consists of an irregularly spaced series of single harmonic notes, lasting 0.03-0.08 s, ended by a lower series of 8-9 short pulses per second.
t..
:::;
:;;
o Fig. 152 Ventral pattern of Occidozyga {aellis from Taman egara, Peninsular Malaysia.
Genus Rana
LINNA E S,
1758
The almost cosmopolitan genus Rana ranks among the most diverse within the order anura. The frogs of this genu are heterogeneous morphologically as well as ecologically. They are in fact so heterogeneou that it is impossible to formulate a generalised description in the same manner as for the other genera. The tips of the fingers and toes are usually distinctly dilated, marginal disk folds are present. Dorsolateral folds are often present and distinct. On Borneo, tltis genus is represented mainly by noctumal terrestrial frogs (although Rana hosii is
Distribution From Thailand to the Philippines and Flores, although it is at present not quite certain if it is the same species throughout. Mt. Kinabalu: Poring.
often found climbing). Most species live constantly, or at least after having reached maturity, in the vicinity of water. The habitats are quite diverse with some species living on the bank of moderately large to large, others in or on narrow streams, others again in swamps or in flooded rice paddics. One species, R. {uctuosa, is in contrast a. strict dweller of forest floors which can also be encountered far away from water. Like the adults, the tadpoles of the individual species develop in a variety of types of water bodies and are adapted accord ingl y. The genu Rona is represented on Borneo by 9 species, 7 of which also occur on Mt. Kinabalu.
Tab. 16 Distribution of Raila-species on Borneo. Sabah
Raila baramica B OETTGER, 1901 Raila chalc01lOta (SCHl.EGEL, 1837) Raila erythraea (SCHL EGEL, 1837) Rana giandu/osa B OULE)/GER, 1882 Rona h OJji B OlJLENGER, 1891 R(ma {Uctuol'a ( P ETERS, 1871) Rana nicobariens;s (STOLlCZKA, 1870) Rana picturata B OLILENGER, 1920 Rona sigllatll (GONTHER, 1872)
158
•
• • • • • • • •
Kinabalu
• •
•
• • • •
Bmnei
• • •
• •
• •
Sarawak
• • • • • • • • •
Kalimantan
• • • • • • • •
Ranidae
Key to the Bornean species of Rana 1 Dorsal coloration conspicuously black and yellow or black and orange ......... signataipicturata • Dorsal coloration different from the afore ....... 2 2 I " finger shorter than 2nd ................................... 3
• I" finger longer than 2nd .................... .. ...... .. ...... 4 3 Outer metatarsal tubercle absent .......... ..... ho.sii • Outer metatarsal tubercle present, round .......... .. ................ ....................... ... ......... ....... clla/conola
6 Supernumerary metacarpal tubercles on every metacarpal .................................. ........................ 7 • Supernumerary metacarpal tubercles only on metacarpals 2-4 .... .. .. .. .. .. .. ............ nicobariellsis 7 Males smaller than 50 mm, females usually smaller than 65 mm ; tympanum black .... .......... .. .. .... ....... ...... ... .......... ............................. baramica • Males larger than 50 mm, females usually larger than 65 mm; tympanum brown ....... glandulosa
4 Web not reaching the disk of the I" toe ............ 5 • Web reaching the disk of the I" toe ..... erytllraea 5 Supernumerary metacarpal tubercles present .... 6 • Supernumerary metacarpal tubercles absent.. .... . ................. ...... ............ .... ... .......... .......... /lIc("osa
Rana chalconota
(SClILEGEL,
1837)
Hy /a clla/cOllota SCIIL"l,I,L. 1837: 23 (part) Type locality: Java RUllO clla/coll ora. MITH 193 1: 17; 1'I(;r'R 1956: 408. 1978: 3 12; INGER & STUEBI'IG 1992: 42; M ALK~1US 1994b: 244, I 996c: 284; ING~ R , STUEB I~G & T~'1 1996: 363; LAX-1M ct al . 1999: 33; Inger et al. 2000: 17 Rona ('lta /co nora rall jeeps. INc,FR 1966: 177 l~v/aralla (Raila) elta/eollota. M ANnl~Y & DFN7H 1982: 19 Hy /aralla clUJ/coila 10. MA'lTIlEY 1983: 22
Diagnosis Rana cha/conota differ from R. nicobariensis in the ratio of finger lengths and foot webbing: In R. cha/conota the I" finger shorter than the 2nd and the I'" three toe and the 5th are broadly webbed to the di sks, whereas in R. Ilicobariensis the 1st finger is longer than the 2nd; and the I" three toes and the 5'h are not broadly webbed to the disks.
Description Male 32-44 mrn, females 46-60 nun ; T LlSVL
0.51-0.60; habitus slender; head longer than wide; vomerine teeth present; tympanum di tinct, 2/3 to 3/4 of the diameter; upratympanic fold absent; I" finger shorter than 2 nd , 1st finger tip weakly, all others substantially dilated ; marginal di k folds present; toe tips dilated, narrower than on the outer
c: c:
i
J:
n:
o Fig. I S3 Male of Rana clla/collota from Poring, 550 m.
fin gers; webbing formula 1(0),2(0/0), 3(0/0), 4( 12/ 1-2), 5(0); inner metatarsal tubercle oval, outer one round; dorsolateral fo lds present; dorsal skin finel y granu lar; ventral skin smooth on chest and throat, granular on posterior portion of belly. Male with yellowi h nuptial pads on I ' finger and a pair of subgular vocal sacs; tympani larger in males than in females. Dorsal coloration yellowish green, normally with brown dots; c ream-coloured ventrally; legs with
159
Ranidae
dark bars, sometimes without; upper lip cream-yellow; lower sides of legs usually reddish. Head-body- Iength of tad poles up to 18 mm, total length up to 50 mm; body oval, slightly flattened, width about 2/3 of the length; mouth positioned ventrally, subterminally; labial teeth rows formula 4(2-4)/3 to 5(2-3)/3( I) ; dorsal fin slightly deeper than ventral fin. This tadpole is one of the very few distinctly coloured ones on the entire island of Bomeo. Body light yellow to straw-co loured, with b lack streaks or dots bClow, in front of, and behind the eye; lower side with many, small, white glands, arranged in two longitudinal sections on either side of the belly. Ecology/Ethology
Raila chalconota is a noctumal inhabitant of the bank rcgions of rivulets cutting through primary and secondary forests, in swampy areas on the edge of cleared areas. The frogs are usually perched beneath or on dead wood or among banking vegetation up to 2 metres above the ground, but usually lower. The species is absent in deforested and other sun-exposed areas and thus differs distinctly from R. nicobariensis in its choice of habitat. Calling males and tadpoles were found between April and September. The brown-pigmented eggs are deposited in groups of 1000-3200 directly into the water. The larvae can be observed swimming slowly above the substnlte, predominantly leaf litter, in calm. water and forest ponds (m: 19, i). They are absent from water bodies with currents. Stomach analyses revealed algae and protists at equal shares, but also p lant particles and fungal hyphens . Adults feed on a variety of insects and spiders. lts vertical distribution ranges from 65 to 915 m, with a predilection for elevations below 300 m.
Call : The call consists of a staccato series of clicking notes. Males form large calling communities at the side pools of mediwn-sized streams and the edges of ponds.
Distribution Malay Peninsu la, Sumatra, Java, and Borneo. Mt. K inabalu: Poring, Sg. Kipungit, Sg. Langanan , Sayap, Kiau.
160
Raila erythraea (SCHLEGEL, 1837) Hy/a elylhraea S CHLI:.GEL, 1837: 27 - Type loca lity: Java Rana eryihraea, 11\GER 1978: 325; HOFFy!ANN 1995b: 10
Diagnosis A medium-sized to large, terrestrial fi"og with the
typical appearance of a "water frog" and wide, light, dorsolateral stripes.
Description MaJes 32-45 mOl , females 48-75 mm; TLlSVL 0.50-0.57; head longer than wide; snout pointed; vomerine teeth present; tympanum visible, ca. 0.75 times the eyc diameter; supratympanic fold weak; ]" finger as long as or longer than 2nd fttlger tips dilated, the largest one about 0.5 times the tympanum diameter; marginal disk folds present; outer finger with dermal fringe; toe tips dilated, but not as widely as on fingers; webbing formu la 1(0), 2(0/0), 3(0/0) , 4(011-2), 5(0); smaller inner metatarsal tubercle oval, outer one round ; dorsolateral folds distinct; ventral and dorsal skin smooth . Males with larger tympani (0.089-0.118 times the SVL in males, 0 .073-0.089 in females), and a coarser skin; weakly developed humera l glands present and nuptial pads on first fingers. Light green dorsally, light brown laterally ; with cream-white or yellow dorsolateral stripes; a dark stripe from canthus rostralis along dorsolateral folds
-
-
-
-
Fig. 154 Rana eryth raea from Poring, 550 m.
Ranidae
up to anus; ventral side and upper lip white; legs ol ive green with black crossbars or rows of dots. Head-body-Iength of tadpoles up to 13 mm, total length up to 58 mm; body about twice as long as wide; eyes situated dorsally, oral disk subterminally, ventrally; lower lip and sides ofupper lip with papillae; lab ial teeth rows form ula 1/ 1 to 1(])/2(1); anal tube dextral; tail convex, pointed, and about twice as long as body; coloration green or brown dorsally, with dark speckles; a dark SU'eak from side of head througb eye up to tai l; caudal fin spotted; later developmental stages with light lateral and dorsal stripes.
Fig. !.5~ Raila elythraea fro~ ~orth_Sumatra.
Ecology/Ethology A shy, nocturnal, terrestrial commensal which also climbs in low vegetation. It actually avoids primary forests and owes its large distribution range to its ability to adapt to situations in and around human settlements. It is usually encountered iu the vicinity of water, very often in paddy fields. The males are often perched on grass or reeds, while the females are more often seen on the banks. The frogs reproduce throughout the year. The tadpo les are found only in stagnant water 'bodies (m: 2b). The diet of R. erythraea includes sma ll terrestrial invertebrates like small mil1ipede , crickets, and ants. This species prefers low elevations. CaJl: Although often found sitting close to cach other, the males do not form calling communities. The call is similar to that of R. nicobariensis, but the call duration is longer, the note length is shorter, and the number of notes per call and the note repetition rate is greater in R. elythraea.
Distribution From eastern India to Indonesia (Sumatra, Java, Sulawesi), Borneo and the Philippine [slands. Mt. Kinabalu: Poriug.
Rana hosii B OULENGER, 1891 Raila hosii B OUUXGCR. 1891 c: 290 - l)rpe locality: Mt. Dulil. Sarawak Rona /rosei. J 'IGER 1966: 187, 1962/63 : 45, 1978: 312; bOGER & STUPHING 1992: 42; MALK M US 1994b: 226. 1996c: 284; H OFFMANN 1995b: 10 Hy laralla hosei, M A ... THEY 1983 : 22; M.,\L K~tU S 1987: 281 , 1988c: 176 Raila (Hylarana) hosei. M.,\LKMUS 1992a: 113 Rona has;;. I )lGER, S TUEIlIl\'G & T AN 1996: 363; .HOfFMASN 1998b: 89; L AKJM el a1. 1999: 33; i NGER cl a1. 2000: 17
Diagnosis A moderately large to large inhabitant of the ground and low vegetation levels with a slender habitus and brightly green coloured back.
Description Males 50-68 mm, females 85-100 mrn; TLlSVL 0.57-0.66; vomerine teeth present; tympanum distinct, 0.75 times eye diameter in males. 0.4 times in fema les; supratympanic fold absent; I" flDger as long as or shorter than 2 nd ; finger tips substantially dilated, outer ones for more than twice the width of finger; tip of 3rd finger equal to or wider than tympanum diameter offemales; marginal disk
L6L
Ranidae
"
.!!' ;0
z
o F.!.g.
~6 J'~a~ ~
10na h~si!. ~o~ ~._Ki"'pUEgi! 1, Pori~g~~O .!:.fl._____________ _
folds present; fingers with narrow dermal fringes; toe tips dilated, but not as much as finger tips; toes fully webbed; inner metatarsal tubercle oval, no outer one; dorsolateral folds weak.. Males with a pair of subgular vocal sacs, separated from one another by a thin septum; nuptial pads present; tympani larger than in females (tympantunl SVL 0.069-0.106 in males, 0.048-0.059 in females).
Usually dorsally bright green, interspersed with darker spots in juveniles, finely pebbled on the back; dark brown laterally, whitish to grey ventrally; dorsal sides of legs brown with dark crossbars. Body of tadpoles oval , slightly flattened ; headbody-length up to 11 mm, total length up to 38 mm; oral field ventral, subterminal; upper jaw white with
Fig. Lxvm Oral disc of R. hash from West Malaysia.
R. Malkmus ~-
162
J
Ranidae
narrow, blackish brown anterior margin; lower jaw uniformly white; labial teeth rows formula 4(2-4)/ 4(1) to 6(3-6)/4(1); dorsal co loration dark grey, unpigmented ventra lly; a ventro lateral skin fold over the entire body. Together with the trongly flattened ventral side this ventrolateral skin fold has the effect of a sucker when pressed to the smooth surface of a rock. EcologylEthology Rana hosii prefers the banks of clear, fast-flowing streams and rivers in primary and little-disturbed rain forests at low to medium altitudes. During the day, the males hide among rocks on the banks while the females spend the days 1-2 m up in the vegetation . Already SMITH (\930) remarked : "The habits of this species are those of a tree frog" , a hint for which are also the extremely widened finger tips. The species possesses a relatively potent skin poison (vernacular name " Poisonous rock frog") which is said to be strong enough to kill a cat. White to cream-co loured eggs (ca. 2.2 mm in diameter) without a halfmoon are deposited in lumps of ca. 500-2000 directly into the water (m: I b, c). The altitudinal di stribution stretches from near sea level to 1500 m . Call: The males call day and night in loose groups from raised sites and are often found in the immediate environs of water, in vegetation or climbing on rock cliffs.
The call consists of either one or two notes, witb the call of one note las,ting 70-80 ms, the one of two 160-170 ms. The frequency range varies and may even vary with the same ind ividu.al. Dominant frequency 3.3-4.5 kHz, decreasing towards the end of the caLI (+22°C). Distribution Malay Peninsula, Sumatra, Borneo, and Java. Mt. Kinabalu: Sayap, Sg. Kipwlgit, Sg. Mamut, Sg. Langanan (750 m), Sg. Manggis. I NGER, STUEBlNG & TAN (1996) listed the species for Sg. Luidan and Sg. Silau-SiLau (1300- 1500 m), records tbat were contradicted by INGER & STUEBlNG (1997) who gave its vertical range as " near sea level to about 750 m".
Rana luctuosa
(PETERS,
1871)
.
Linmodyles IUCIIIO.\'II.\' P ETERS 1871 : 579 - Type locality: Sarawak Raila decora ra M OCQUARD 1890a: 145, PI. J0: Fig. I - I b - Type
locality: Kina BaJu (syn. after
B OULl!NGER
189 1b)
Raila IlIcllIosa. IiANlTSCH 1900a: 73; VAN K AMPEN 1923: 196;
1931 : 16; INGER 19543: 250, 1966: 206, 1978: 312, 1985: 56; MATSUI 1979: 333; INGER & STUJlll lNG 1992: 42; M ALKMUS J994b: 244; INGER, S11.rEAING & T.~N 1996: 363: I NGER & TAN 1996a: 563 H y laratla IIICflIasa, MANTHEY 1983: 22 SM ITH
Diagnosis A medium-sized to large ground dweller with a moderately stout habitus . lts conspicuous, reddish brown dorsal coloration with the light dorsolateral snipes is an unmistakable trait.
Fig. ]58 Eggs of Raila hosii from West Malaysia.
Description SVL 53-60 mm; TUSVL 0.51-0.58; habitus moderately robust; head longer than wide; snout slightly pointed; vomerine teeth present; nostrils closer to tip of snout than to eyes; diameter of tympanum about equal to that of eye; tympanic fold absent; I" finger longer than 2 nd finger tips only slightly di lated ; marginal disk folds present; no supernumerary metacarpal tube rcles; webbing formula 1(2), 5(2), other toes with skin fringes to disc; inner metatarsal tubercle oval, weakly developed; a smaller, round outer metatarsal tuberc le is recognjzable only in about every second specimen; skin smooth, without dor olateral folds.
163
Ranidae
Males without vocal sacs and without nuptial pads, but with humeral gland of ca. 4 mm in length. Dorsal coloration reddi h brown with sharply defined , wide dorsolateral stripes running through the eyes up to tip of snout, separating brown dorsal coloration from black on sides of head and body; flanks and lower side below black area dark grey, lighter towards mid-venter; limbs grey with con-
spicuous light bars; eyes dark, top third lighter with a white band on upper side. I-Iead-body-Iength of tadpoles up to 26 mm, total length up to 65 mm; spiracle closer to eye than to base of tail; labial teeth rows formula 4(2-4)/4(1) to 6(2-6)14( I); coloration of larvae greyish brown with irregular dark spots on body and tail. EcologyfEthology Rana luctuosa is a noctumal, solitary inhabitant of the leaf litter layer which congegrates at stagnant water bodies on ly during the mating scason. Tbe reduced webbing is typical for sem i-terrestrial frogs. The tadpoles live primari Iy in muddy ponds including rh inocerus wallows (m: 2b, c). Altitudinal di stribution 150-1400 m.
-
-
-
Fig. 159 (left) Ventral pattern of Rana /uclUosa from
Bundu Tuhan .
164
Ranidae
C all: The ma les form calling communities at rainwater pools with each male calling from a small hole it has excavated adjacent to the pool. The call resembles the miaow of a cat.
Distribution Malay Peninsula, Sarawak, West Sabah. Mt. Kinabalu : Kiau, Bundu Tuhan.
Raila nicobariellsis
(STOLlCZKA,
1870)
Hyiuratla nicobariensis S TOLICl.KA. 1870: 150 - Type locality: Nicobar Islands Rana nicobariellsis. B OUI.IO"GER 1885b: 389; SMITH 1931 : 9; [:-;GER 1966: 223, 1978: 313; MA L.KMUS 1987: 28 1, 1994b: 244; H OFFM ANN 1995b: II , 1998b: 89; [ "'GER, STLEBIl\(j & T 1996: 363 Rana nicobariensis Ilicoboriensis, INGtR 1956: 406 Hylarana nicobariensis, M ANTHEY 1983: 22
A"
Diagnosis A medium- ized terrestrial frog of slender babitus. The webbing is only weakly developed in R. nicobariensis, distinguishing it from R. chalconota. The ISl finger is longer than the 2 nd in R. nicobariensis and shorter in R . .chalconota.
Descri ption Males up to 47 mm, females up to 53 mm; TL/SVL 0.50-0.60; head longer than wide; snout pointed; vomerine teeth present; nostrils much closer to tip of snout than to eyes; eye diameter slightly greater than distancc between nostril and eye; tympanum visible, about 75-80% of the diameter; supratympanic fold
Fig. 161 RanG nicobariensis fro m Sabah.
F~g.
162 Ra!!.a nicob!!riensis from
Sab~l.
absent; I S1 finger as long as or s lightly longer than 2 nd ; fin ger tips dilated; diameter of finger disks smaller than half tympanum diameter; marginal disk folds present; toe tips more expanded than finger tips; webbing fonnula 1(1), 2(1/0), 3( 1-2/0), 4(2-3/2-3), 5(0); imler metatarsal tubercle small and oval , outer one round; narrow dorsolateral folds present; dorsal and lateral skin slightly granular, smooth ventrally. Males with slightly longer beads and larger tympani than females, and with pairs of subgular vocal sacs, yellowish nuptial pads, and large humeral glands. Colour pale brown with black spots dorsally, darker brown laterally and particularly so on sides of head; upper lip white; lower side dirty white witb a faint grey pattern; hind linlbs with dark crossbars . Body shape of tadpoles oval; light brown with dark brown pattern; tail Witll dark reticu lated pattem; headbody-length up to 15 mm, totallengtb up to 47 mm; lower lip with conspicuous, long, finger-like projections; labial teeth rows fonnula 1.12(l) to 1/3(1),
EcologylEthology The frogs are widely distributed, terrestrial commensals which are limited on Mt. Kinabalu to the margins ofthe national park, but are more commonly ncountered in the surroundillg villages. They are often fOWld sitting in drainages and other artificial water bodies and flooded meadows. They are both diurnal and nocturnal. Males are very common locally, but do not appear to form calling c.ommunities. The reproductive season lasts throughout the
165
Ranidae
year. Their diet comprises insects of appropriate sizes. The larvae develop in the shallow water of ditches and temporary pools (m: 2b). Altitudinal distribution 0-925 m a.s.!. Call: (+21 0c) The characteristic, cackling calls consist of rapid sequences of6-1O loud, sharply pulsed notes of 50-60 ms in duration with a note repetition rate of? per second, an intemote interval of70-80 ms, and a frequency betwecn 1 and 4 kHz; hannonics are not evident. Distribution From the Nicobars through the Malay Peninsula to Bali, Borneo, and Palawan. Mt. Kinabalu: Poring, Kiau.
1.50
1.60
1.70
1.80
1.90 sec
-':lr---- -------=;1
~-
1.-(
Fig. LXIX Sonagram and oscillograms of Rana nicobariensis (poring Hot Springs, 500 m; lO.OO p.m. +22°C)
Raila sigllata
(GUNTHER,
1872)
Polypedoles signotlls GONTHER, 1872: 600 - Typ e loca lity: Matang, Sarawak Rana obsoleta MOCQlJARD I 890a: 147 - Type locality: Kina Balu (syn. after BOULENGER 1920) Raila Signata, VAN KAMPEN 1923: 226 (part); I.J\OER & STUEBIl'G 1992: 42 ; MAlKMus I 994b: 244; WONG 1994: 32; HOFFMA:-'~ 1995b: 12; WGER, STUEBING & TAN 1996: 363; INGER & T...." 1996a: 564; MANTHEY & GROSS~IANN 1997: 116 Ranasignata signata, (part) INGER 1956: 407, 1966: 233, 1978: 3 J3 Hylarana signata, (part) MANTH~Y 1983: 22; MALIC~tU 1988c: 176. 1989: 187, 1996c: 284 Rana (Hylarana) sigllata, M ALKMUS 1992a: 113
Raila picturata
BOULENGER,
1920
Rana pichlrata BOULENGER, 1920: 179 - Type locality: Bidi Caves, Sarawak, Barabas, S.-E. Borneo, and Kina Balu Rana signata signata (not Raila signata (GONTH£R, 1872» , (part) I NGER J 966: 223 Raila (/iy/ara/la) signata (not Rana signata (GONTHER, 1872» , (part) VAN KAMPEN 1923 : 226 Raila signata (not Rana signaUl (GONrnER, 1872) , MANTHEY & GROSSMA.~ 1997: fig. 76
166
uiu
Ta xonomic r emarks: Rana signata and Rana pieturata were for a long time cons idered conspecific, until INGER & TAN (1996a: 564) separated them on the basis of their colour patterns. In R. signata, the orange to red stripe runs without interruption from the eyelid over the back, whereas if such stripe is present in R. pleturata, it is always interrupted. The back itself is marked with many (R. pieturata) or a few to no (R. signata) spots in a colour matching the dorsolateral stripes. Another indication of tbe presence of two rather than one species is the inconsistency in the descriptions of the vocalizations of "R. signata" by GRANDlSON (1972), INGER & STUEBING(1992), SANCHEZ-HERRAlZ et al. (1995), and MANTHEY & GROSSMANN (1997). Since, however, no morphological or ecological differences have been confirmed up to now other than those of the colour pattem, both fonns are treated here jointly due to their great similarity. Diagnosis A small to medium-sized ground dweller with a moderately slender build. Due to the highly contrasting,
Ranidae
yellow to orange red and black colour pattern, R. signata and R. piciLlrata amlot be confused with any other Bornean frog .
Description Males 33-47 mm, females 48-69 mm; TLlSVL 0.48-0.57; habitus moderately stout; head slightly pointed; canthus rostralis distinct; vomerine teeth present; nostrils closer to tip of snout than to eyes; supratyrnpanic fold often absent; I fmger longer than 2»d; finger and toe tips slightly expanded; marginal disk folds present; supernumerary metacarpal tubercles present; toe tips dilated into disklike structures; toes webbed for about half their lengths; oval inner and large rOlmd outer metatarsal tubercles present; dorsolateral folds often absent; dorsal and lateral skin granular, smooth ventrally. Males with a pair of subguJar vocal sacs, nuptial pads, and humeral glands. Sf
Dorsal and lateral coloration black, with yellow or orange red spots; a stripe of matching colour from the tip of the snout up above the eyes; dorsolateral stripes of the same' colour continuous or interrupted (see Taxonomic remarks above); ventral side of males dark, lighter in females; limbs yellow or orange red above, with blackish brown crossbars. TadpoJes relatively slender, oval in shape, slightly compressed dorsoventrally; head-body-length up to 13 mm, total length up to 41 mm ; labial teeth rows formula 3(2-3)/3{ J) to 4(2-4)/3(1); first lower row of denticles interrupted only to a minor extent; blackish brown dorsally, tail pale grey, unpigmented ventrally, covered with smail, whitish glands. These glands form a wide ring around the oral field, a separated round spot behind it, a narrow ventrolateral series between insertion of legs and below spiracle, and a dorsolateral row between eye and base of tail. Rows of glands continued on
to
167
Ranidae
both fin over the latter's full length, bUl absent on caudal muscle. Arrangement and number of these glands differ slightly in the two fom1s .
Ecology/Ethology Rana signata and R. picturata are nocturnal inhabitants of calm sidearms or respective sections of small to moderately 1.20 1.40 1.60 1.80 2.00 sec large (up to about 10m in width) running water bodies. Their occurrence is limited to primary and old secondary forests. The juveniles migrate away from the sites of their larval development and spend their early life on the forest floor, only to return to a calm waters 011 reaching maturity. kHz 5.0 sec They are usually found on the 10 , - - - - - - - - - - - - - - - - - - - - - - - - - , ground, dead wood or rocks, more rarely climbing in low 5 vegetation (up to about 40 cm). The males call individually at first, but appear to trigger others to do so as well, so that large calling communities establish themselves at regular intervals Call: (+24°C) It is at present extremely difficult to (INGER & STUEBL'JG 1997). SANCH EZ-lliRRAIZ et al. (1995) reported on distances of several metres determine which call belongs to which form as both between calling males with the calls not being have been treated as one and the same species in the emitted in choruses. past. According to INGER & STUEBJl\O (1997), the call Oviposition between April and August, preferably of R. picturata sounds like a metallic burr (duration in places without or only a weak current, e.g. in 620 ms, consisting of 4-6 (8) pulsed notes of 50-80 m5 rock basins or cut-off sidearms of streams or riveach) with the intensity of the notes continually deers . The larvae live in shallow, but also in deep, creasing; frequency 1. 1-204 kHz (SANCHEZ-HERRAIZ sections of their water bodies where the bottom is et al. (995). There is, however, a deviant descripmuddy or consists of gravel, often in accumulation of the call by GRANDISON (J 972), who charactions ofleaflitter (I d, f). Stomach analyses revealed terized it as similar to human laughter. There is a for a major part blue algae, besides about equal chance that the latter might refer to R. signata. parts of plant particles. diatoms, and flmgaJ hyphens. Distribution The altitudinal range spans 0-750 m, whereby INGER Malay Peninsula, Sumatra, Borneo, Philippines & STUEBING (1997) state R. picturata to be distrib(with three subspecies in INGER, 1954). Mt. Kinauted throughout Borneo, while R. Signata would balu: Sg. Kipungit, Sg. Mamut, Sg. Tananansad, prefer the lowland . Sg. Sasapan.
168
Ranidae
Genu Staurois
COPE,
1865
The distribution ofthc gcnus Slaurois is mainly re trictcd to Borneo with only S. nalator also occurring on i lands of the Philippinc . It compriscs small to medium-sized, very slender frogs with long hind limbs. Vomerine teeth generally absent, but often present in S. nalalOr at lea t on one ide. Finger tips and toes expandcd into oval stnlcture , marginal disk folds and ventra l cross fo lds present. Toes fu lly webbed; 1>1 fi nger always shorter than 2 nd ; dorsolateral folds wanting, tympanum visib lc, vcntral skin coarsely granular. All species share a distinct prcdilection for rapids and waterfa lls where they can usua lly be seen perched on Fig. 164 Foot flagging disp lay in the genus Staurois; here: S. tuberiboulders on the banks or those pro/inguis from Sg. Liwago. truding from the water. Be ides of the common acoustic advertisement, the males have developed a particular optical display litter covering the bottoms of ba ins or small rivuin which the colourfu l webbings between the toes lets in or forking off from streams. are flashed by strctching a hind limb and spreadThree pecies occur on Borneo, al1 of which are ing the toes either uni lateral1 y or altematingly. As also foun d on Mt. Kinabalu. thi gives an appearance not dissimilar to naval flag signals this behaviour is appropriately termed "foot flagging display" (HARDING 1982, DAVISON 1984). Key to the Bornean species of Staurois Only the larvae of S. nata/or (INGER & TAN 1990) (after INGER 1966) and S. tuberilinguis (MALKMUS et a!. 2000) are known. Their external morphology and choice of Outer fi ngers webbed ............ ........ JatopaJmatus habitat is largc ly congntent, with the body being • Fingers not webbed ........................................... 2 compressed, the tai I long with reduccd tins, the eyes small and situated dorsally, and the skin hardly 2 Outer metatarsal tubercle present; no li ngual papigmented. Differences are on ly evident in the body pilla........... ......... .... ................................. natator lengths at the various stages of deve lopment, and • No outer metatarsa l tubercle ; lingual papilla in the numbers of denticle rows on the lower lip. present ..... ..... ......... ........................ tuberilinguis The tadpoles lead a secretive life among the leaf Tab. 17 Distribution f Statlroi - pc ie on Borneo.
Stallrois lat()palmatlls (BOULENGER, 1887) SUlllrois lIiltator (GoNnmR, 1859) Stallrois tuberi/iflgllis B OuLbNGER, 1918
I I I
I
Sabah
• • •
I
I I
Brunei
TCinabaJu
• • •
I
I I
• •
•
Sarawak
• • •
I Kalimantan I I
• •
169
Ranidae
Staurois latopalmatus (BOULENGER, 1887)
gers are webbed and the snout of S. latopalmatus is very short and blunt, while it is very long and pointed in the other two.
BOULENGER. 1887b: 97 - TYpe locality: Mount Kina Ba100, North Borneo Ixa/lIs lalopa/maills. MocQuA.RD J 890a: 152; WHITEHEAD 1893: 121 SI1IoIIIClll/is /alopa/mala . VAN KAMPEN J923 : 238; SMITII J931 : 17; l l'GE R 1962/63 : 46 Slaurots /alopa/malus, INGER. 1966: 252, 1978: 313; M ANTHEY 1983: 23; MALKM liS 1988c: 176, 1992a: 115. I 994b: 226. I 996c: 285, INGER & ST EllING 1992: 42; l NG£R, S TUEBL'>;G & T AN 1996: 363; INGER & T AN I 996a: 564; LAKI'f et al. 1999: 32; I l'GER et al. 2000: J 7
Description Males 40-50 mm, females 60-70 mm ; TLlSVL 0.58-0.71; habitus slender, but rather stout in comparison with the other two Bomean species of this genus; head longer than wide; vomerine teeth absent; nostrils closer to tip of snout than to eyes; canthus rostralis di tinct; tympanum visible, 1/3 to 2/5 of eye diameter; supratympanic fold absent; 1SI finger shorter than 2 nd ; finger tips expanded into di k-like structures, diameters of all finger tips are greater than diameter of tympanum; marginal disk folds present; fingers not fully webbed; toe tips expanded into disk-like structures, smaller than finger tips; toes fully webbed; inner metatarsal tubercle oval, no outer one; dorsolateral folds absent; dorsal skin granular, smooth ventrally. Males with a pair of vocal sacs and whitish nuptial pads on first metacarpal. Dorsal coloration black with yellowish or white dot , ventral side white to cream-coloured; toe webbings of males whitish to pale grey, dark grey in females. The tadpole of Slaurois latopaimaLus will soon be described by L DAs.
!xaills lalopa/malUs
Diagnosis A moderately slender terrestrial frog with substantially expanded finger tips. The tips of all fingers are wider in diameter than the tympanum, while in all other Bomean species of this genus only the tips of the outer two flllgers are wider. In cootrast to the other species of Staurois the outer two flnFig. J65 Stall/'ois /alopa/mailis from Sg. Kipungit IT. 640 m, Poring.
Ecology/E thology Staurois latopalmatus is a diurnal and nocturnal ground dweller which is found along running waters, often on boulders in or on the banks of streams, but occasionally also in low vegetation. If disturbed, it will leap from rock to rock, or it may even leap into swift water and skip across the surface to another rock. AI night, juveniles are often encountered perched together in groups, whereas adults are mostly found singly. Foot flagging displays can only be observed between 6.30 and 8.00 a.m. duriog which time the males often congregate in group . Their calls can be heard throughout the night and the early morning bours (HARDING 1982, DAVI ON 1984). The vertical distribution of the species ranges from 150 to 1000 m a.s.1. Call: A short, rising trill.
170
Ranidae
Distribution Endemic to Borneo with records from Sabah, Sarawak, Kalimantan , and Brunei . Mt. J(jnabalu: Sayap, Sg. Panataran, Kiau, Sg. Liwago, Sg. Mantllkungan, Sg. Mamllt, Sg. Kipungit, Sg. Langanan, Sg. Sasapan.
Stallrois natator (GONTHER, 1859) /xaills flo/alar GONHIER, 1859 (1858): 75 - Type locality:
Philippines Ixaills gll/((J(us GONTI-lER 1859 (1858): 76, PI. 4: fig. D - Type
locality: Borneo (syn . after BOULENGER 1882) hollis natator, HA \J IISC fI 1900a: 74 Stallro;S gllt/Oll/s. BOUI. E:-
Diagnosis A very slender ground dweller with expanded finger tips and a pointed snout. The external morphology corresponds with that of S. tuberilinguis, differing only in the greater SVL of adults and with regard to tbe colour pattern. Olive green dorsally, with large black spots; dorsal skin homogeneollsly granular (in contrast to S. tuberilinguis which is brown dorsally with irregular green spots and has a heterogeneous ly granular skin). Ventral and lateral sides of S. natator lemon-yellow to yelJowish green, S. tuberilinguis bluish white ventral ly and green with small white dots laterally. Description Males 29-37 mm, females 44-55 mm; TLlSVL 0.55 -0.65; habitus very slender; head longer than wide' vomerine teeth almost always present, but not always on both sides; nostrils clo er to tip of snout than to eyes; canthus rostralis di stinct· tympanum visible, its diameter about 113 to 112 eye
Fig. 166 Stallrois nata/or from Sg. Kipungit 1[, 650 m, Poring.
171
Ranidae
diameter; supratympanic fold absent; 1" finger shorter than 2nd ; finger tips expanded into disk-like structures, outer two wider than tympanum diameter; marginal disk folds present; 2nd and 3nl fingers with dermal fringes ; toe tips expanded into disk-like structures, but smaller than finger tips; toes fully webbed; inner metatarsal' tubercle oval, outer one round and small ; dorsolateral folds present; skin on dorsum and on posterior portion of belly granular, chest and throat smooth . Males with a pair of vocal acs and yellowish nuptial pads on l oj finger; their webbings behveen toes light greenish blue. Dorsal coloration olive green, with large black spots; ventral and lateral sides lemon-yellow to yellowish green; a narrow black sUipe along canthus rostralis from eye to tip of snout. Body oftadpoJes oval in shape and s lightly flattened ; length of tail more than twice the headbody-length, e.g. at stage 41 head-body-Iength lO.5 mm, total length 34 .5 mm; eycs small (di ameter 0.08 times head-body-length), set far back, oriented dorsally, and thickly covered with skin at early stages of development, formation of a transparent "window" from stage 36; nostrils small and positioned subtenninally; tail with very low fins, ending in a broad tip; oral disk situated ventrally, subterminally; labial teeth
Fig. 167 A pair of Stallrois nata/or from Danul1l Valley, Sabah, in amplexus.
rows formula 2(2)/8(1) to 2(2)/ 10(1); spiracle s inistral , with a shOlt, terminal tube; anal tube long, situated medially ; almost transparent, with a reddish belly resu lting from the visibility of blood vessels. Ecology/EthoJogy A mainly diurnal to crepuscular inhabitant of the rocks on the banks of streams and small rivers in primary forests . The frog is also often found in low banking vegetation , but rarely encountered farther than 100 m from the nearest water. Especially yOlrng specimens congregate at night to fonn "dormitories" on larger leaves etc . The tadpoles live predominantly in the thick layers of sediment consisting of leaf litter and mud in sections of streams where there are no currents (m : [d, e). The known altitudinal range extends from I 10 to 650 m a.s.l. CaU : The call consists of a sharp chirp. Di stribu tion Borneo and the Philippine Islands. Mt. Kina balu: Sayap, Sg. Mamut, Sg. Kipungit, Sg. Sasapan. The record "Sg. Silau-Silau (J 540- 1580 m)" by WONG (J 994) is probab ly based on misidentified S. tuberilinguis.
Stallrois tllberilinguis
BOULENGER,
1918
B OULENGE R, 1918: 374 - l)'pe locality: Mt. Kina BaJu, North Borneo, altitude 4200 fect and Mt. Baw Song, arawa.k, LOOO feet S /alll'Ois IIIberiling uis, VAN K AMPE).' 1923: 237: I»IGBR I 962i63 : 48. 1966: 250, 1978: 313 ; M ANTHEY & D l\zER 1982: 12; MA NTIIIIY 1983: 23; M ALK.\IUS 1985 : 8, 1987: 281, 1988d: 8, 1988c: 176. 1989: 187, 1991a: 28, 1992a: 114, I 994b: 226, 1996b: 22, I 996c: 285, 1996e: 11 ; I)
Staurois /Ilberiling llis
D iagnos is A very slender ground dweller with expanded finger tips. Dorswn brown with irregular green spots, heterogeneously granular; ventra l sides bluish
172
Ranidae
white; lateral sides green with small white dots. Distinguished from S. nata tor (see above) on the basis of the colour pattern, the absence of an outer metatarsal tubercle, and the presence of a lingual papiUa (see key) .
Description Males 23-37 mm , females 29-43 mm; TLlSVL 0.49-0.57: habitus very slender; head longer than wide; snout pointed; vomerine teeth absent; nostrils closer to tip of snout than to eye ; canthus rostralis distinct; tympanum visible, about 113 to 2/5 the eye diameter; supra tympanic fold present; 1sl finger much shorter than 2 nd ; finger tips expanded into disk-like structures, those of 3rd and 4th fingers wider than tympanum diameter; marginal disk folds present; fingers withollt dermal fringes ; toe tips expanded into disk-like structures, but not as wide as tips of outer fingers; toes fully webbed; inner metatarsal tubercle oval, outer one round and small in about a third of all individuals, absent in the remainder; dorsolateral fo lds wanting; dorsal skin heterogeneously granular, smooth ventrally. Colour dorsally brm'll1 , with elongate, green spots; green laterally, with small, white glands; bluish white ventrally, usually with dark dots on the throat. Males with a pair of vocal sacs and whitish nuptial pads on the 1st metacarpal; webbings between the toes white. Tadpoles morphologically very similar to those of S. natator; body oviform, distinctly compressed dorsoventrally; tail more than twice as long as headbody portion, e.g. at stage 25, head-body-Iength 3.7 mm, tail length 8.1 mm; caudaJ fins low, tip blunt; spiracle sinistral, short, straight, directed backward; anal tube median; eyes very small, dorsal; distance between anterior margin of eye and
-
tip of snout more than three times as large as interocular distance; oral field ventral, subterminal ; marginal oral papillae small, in one row; denticIes 2(2)16(1); proximal row of teeth on upper lip widely intemlpted medially, inner row of
-
Tadpole of Staurois IlIberiliTlguis. -----------
R . Malkmu~
173
Ranidae
predictably at another site. Foot flagging is shown by the males often right after a series of calls. As the white coloration of the voeal acs is well visible during the calling, they represent another optical signal. Suitable advertising sites are rocky banks and solitary boulders on the banks. The males call during the day between 6.00 a.m. and 6.30 p.m. Single calls are often answered by antiphonie calls, but never to such an extent that ehoruses might form . The tadpoles have a yet only been described from a basin on Sg. Liwago whieh was filled to the rim with leaf litter (m: I d, e). There' they lived photopbobically in an environment with almost no current. The altitudinal di stribution covers 200-1800 m a.s.1.
c c
i 0:
4)
Fig. 169 StaLlrois Iliberilinguis from Sg. Wariu. 950 m,
Sayap.
lower lip with a small gap; lengths of denticle rows decreases continually distally; edges of both jaws narrow, with black pigmentation, and homogeneously serrated. Larvae light brown dorsally, fading laterally so that they appear unpigmented lateroventrally. Tai l with two centTal and two marginal brownish stTipes. The transparent skin of the venter permits to see the intestinal coils and the intensely red gills. Ecology/Ethology This frog is a predominantly diurnal and crepuscular inhabitant of the banks of fast-flowing mountain streams in primary forests , particularly of waterfalls and rapids. Here, the animals are mostly seen perched on rocks during the day while they spend the night, well-camouflaged by their green coloration, singly on the upper side of leaves of banking bushes 50-100 cm above the ground. S. tuberilinguis is very shy during the day, escaping in almost every instance by plunging into the nearest rapid, on ly to re-appear unexpectedly and un-
174
Call: (+ 19°C) Note series of 8-12 s in duration, with 40-60 notes of 80- 100 LnS each, resembl ing the trill of an insect. The frequency rises from an initial 3.8 kHz to 4 . 1 kHz within a note series, with the note repetition rate increasing from 3 to 8 notes per second and the intemote intervals decreasing fr0111 400 to 50 ms. D istri bution Endemic to Borneo with records from Sabah, Sarawak and Brunei. Mt. Kina ba lu : Sayap, Kenokok, Sg. Kinateki , Marei Parei, Sg. Kadamaian above Kiau, Lumu Lumu, Lobang, Sg. SilauSilau, Sg. Liwago, Mesilau, Sg. Mantl.lkungan, Sg. Mamut, g. Kipungit, Sg. Langanan.
-
-
Fig. LXXllJ Sonagram and oscillogram of Stallrois fuberilingl./is (Sg. Liwago, 1500 m; 8.00 p.m. + 19°C)
kH z
9.29 sec
1 0 .-----------------------------------------~
5
Rhacophoridae
Rbacophoridae Rhacophorid frogs are distributed in tropical frica and Madagascar, but have their main centre in the Asian tropic and sUbtropics. Here, they range from lndja to Japan, inclurung the Greater unda Islands and the Philippines. Rhacophorid have a firmi temal pectoral girdle, sacrum with cylindrical diapophy es, omostemum and stemmn ossified, eight procoelous presacral vertebrae present. Maxillae and premaxillae dentate. Fingers and toes di lated into large rusks, short cartilaginous intercalary clement between penultimate and tenninal phalanges present; toes webbed over at least half thei r lengths. Some species of Rhacophof1ls with exten ive webbings on hands and feet enabling them to perform paragliding flights. Pupil horizontal (except in Nyctixalus). Mating takes place in ax,j llary amplexus.
The species of al l Bornean genera, save for Nyctixalus and Philalllus, produce roam nests in which the eggs deve lop to tadpolcs. The larvae are aquatic, with tenninal mouths with beaks and rows oflabial teeth ; tho e of Nyctixalus and Theloderma are non-feeding. Several lineages of Phi/auflls are known for their non-aquatic larvae - their eggs developing directly into frog lets. All Bomean specie arc tree frog , dwelling primary and secondary forests, some species appear to prefer habitat modified by human activity. Rhacophorids are very common on Mt. Kinabalu at all altitudinal levels up to 3000 m above sea level. The family i represented on Borneo by 5 genera with 35 species, on Mt. Kinabalu by 4 genera and 22 spec ies.
Key to the Bornean genera of Rhacophoridae
(after
MANTHEY
&
GROSSMANN
1997)
8
Pupil roundish or horizontal (A) .... .... .. .......... .. 2 • Pupil vertical or diamond-shaped (B) ...... .. .. ...... . .... ......... ............. ..... .. ..... ..... ....... ......... Nyctixalus 2 Skin membrane between two outer metacarpals present (C), terminal phalanges of fingers and toes V-shaped (D) .... .... .... .. .... ........................ .... .. ..... 3 • No skin membrane between two outer metacarpals; terminal phalanges of fingers and toes not V-shaped, but partly slightly notched (E) .. .. ...... . ............ .... .......... ... .. ........ ... .. .......... ...... Philautus
E
3 One or more finger disks with a ring-shaped fold (F) on lower side ............ ...... ........ Rhacophorus • No ring-shaped fold on lower sides of finger disks ....................... .... .......................................... ... .. . 4 4 Upper side, particularly limbs, with numerous smaller and larger warts (G) ...... .. ... Theloderma • Skin of upper side smooth .. ...... ...... Polypedates
F
175
Rhacophoridae
Descr iption Males up to 33 mm, females up to 38 mm; TLlSVL 0.50-0.61; body and limbs slender;. body flattened
dorsolaterally; bead longer than wide; snout obtusely pointed; canthus rostralis distinct; lores nearly vertical feebly concave; tympanum distinct, two thirds of eye diameter; tips of fingers expanded into round or oval disks, no webbing; metacarpal tuberc les present; disks of toes smaller than those of fingers ; toe webbing J (0.5), 2(0.5/0.5), 3(l/l), 4(212 ,5), 5(1.5-2); an oval inner metatarsal tubercle; dorsal skin with spinose tubercles; throat smooth, chest rugose, abdomen coarsely granular; no stlpratympanic fold ; males without vocal sacs; a yellowish nuptial pad on dorsal surface of I" finger. Dor al sides of body and limbs dark chestnut-brown to reddish bro\vll, lighter laterally; lower side whitish with a reddish tinge, beUy sometimes blackish; dark brown portions peppered with ivory-white dots and spots which are reduced in numbers in specimens from high altitudes. Elliptical shoulder spots, a horizontal row of spots along canthus rostral is and above eye, spots on knee and heel joints, and short bars across hind limbs and lower arms are always present, ho~vever, Upper third of iris greyish yellow with a blackish network of veins. 100....er two thirds dark brown. Tadpoles: Head-body portion of tadpoles ova l, a lmost as wide as long, slightly flattened ; snoLit broadly rounded; eyes dorsal; moutb ventral; beaks with black edges, serrated; labial teeth rows formula
Fig. 170 Nyclixalus pictus from Sg, Wariu, ca. 1000 m, Sayap.
Fig. J 71 Ventral pattern of Nyctixa/us pictus from Sg. Silau-Silau, 1550 m.
Genus Nyctixalus
BOULENGER,
1882 .
The genus Nyctixalus is represented on Bomeo by a single species, Nyctixalus picttlS (PETERS, J 871), which is known from Sabah, Sarawak, Bnmei Darussalam and Kalimantan.
Nyctixalus pictllS (PETERS , 1871) [rafus pic/lis PETERS. 1871 : 580- T'ype localily: arawak. Bor-
neo Phi/au illS piC/lis. SMl1'-' 193 1: 19; IN(,ER 1956: 422 Phi/aullls pic/us pictus, INGER 1966: 349 Hazeliapicla , iKGER 1978: 3 13: M A'ITHEY 1983: 23 Nyc/iratus pic/us, M;\LK~I US 1991 : 3 ~. 1992a: 123; t'J(;~R & STUEB,I\G 1992: 41-51: 1I0FFMANN 1995b: 13 ; I NGER, STUEBING & T A)o/ 1996: 363; M ALKMI. S 1996c: 286; M ANTI 'EY & GROSSMA.'1N 1997: 124: L AK.L\f et al. 1999; b /GER et al. 2000
Diagnosis A slender, chestnut-brown frog « 35 rum) with white spots and dots on the back and sides; pupil vertical-oval to diamond-shaped ; fingers without webbings; vomerine teeth absent.
c
I I
a:
o
176
-
--------------------
Rhacophoridae
i
co
~
::i
:i
@
@
Fig. 172 Tadpole (st 26) ofNyclixalus pic/tls from Pulau Pin~ng2 ~llinsular ¥alaysia.
Fig. 173 Tadpole (st 43) of Nyctixalus piCIUS from Singapore.
5(2-5 or 3-5)/3; sp iracle sinistral; tail sl ightly convex, tapering to broadly rounded tip, L.3-1.5 times Lhe head ~ body~ length . Tadpole entirely black to purplish brown, fins and ventral si de of body lighter; total length up to 53 mm; freshly metamorphosed frog lets 17 mm.
frequency 1.8-2.2 kHz; internote interval 80-90 ms) at an increasing volume. This call resembles a muffled tweeting bird call. Hannonics at 4 kHz.
Ecology/Ethology Nyclixalus piCLUS inhabits the montane rain forests between 500-1750 m a.s.l. on Mt. Kinabalu . It is found off from streams in the lower stratum (shrub and bush level) up to 3 m above the ground, but also in the leaf litter layer. The species is nocturnal, leaving its shelters to forage with the onset of dusk. This spec ies is predominantly a sit-and-wait predator. Its diet comprises spiders, hairless caterpillars, and tbe larvae of beetles. Spawn was found in the form of clutches containing 7-15 eggs hanging on the wal ls of water-filled tree boles 1-1.6 m above the ground, and in large empty fruit casks 3-10 cm above the water level (m: 2d, g, h). Call: (+20.5°C) The note series starts off with 2-4 di linctiy pulsed notes (duration 34-45 ms; frequency 1.9 kHz; internote interval decreasing from 150 to 90 ms) which are followed b y 4 weakly pulsed notes (duration 10-13 m ;
Distribution Malay Peninsula, Sumatra, Borneo, and Palawan. Mt. KinabaJu: Poring (500 m), Kiau, LUDlll LtuTIu, Sayap (Sg. Wariu) upper Liwago Valley (up to ]750 m), Headquarters (1540 m).
-
Fig. LXXTV Sonagram and oscillograms of Nyclixailis piCIlIS (Sg. Liwago, 1500111; 8.00 p.m. +20.5°C)
-
...
.ill.
38.40
38.60
38.80
,i , .1
1
J
1
,
39.20 sec
39.00
,
.1 I
kHz
42.84 sec
,----------------------------------,
': f
f , ; ~
f
j
, f , ,
.
~
-
. "
;:
.. II
177
Rhacophoridac
Genus Philautus G1STEL, 1848 The species of this genus are distributed from India and Sri Lanka to China, the Greater Sunda Islands, and the Philippines. The genus Philautus comprises small to mediumsized, usually stout tree frog species, many of which are characterised by a confusing morphological variability. The resulting prob lems are even enhanced by the fact that many species are able to alter their colorations and patterns. The knowledge of the specific advertisement calls is therefore of fundamental importance for the field herpetologist. An attempt to establish an identification key to the species on the basis of morphological traits is for the same reason rendered entirely dissatisfactory. Under these circumstances wc are much rather ineI ined to fo II ow the proposal by DRThlG (1987) to allocate the Bornean fonlls to four species groups whicb are defined as follows: - aurifasciatus group: small or medium-sized species; vomerine teeth absent; nuptial pad small and smooth; ova unpigmented; calls generally pulsed, poorly tuned , with no frequency modulation. Content: P. amoenus. longicrus, l1yobergi. petersi. rejiJgii. saueri. umbra.
- hosii group: large species; vomerine teeth present; nuptial pad absent; ova with dark pigmentation ; caIls 110t pulsed, well tuned, with frequency modulation ; stream-associated species. Content: P hosii, ingeri. - teetus group: small species; vomerine teeth present or absent; nuptial pad large and spinulose; ova without pigmentation; call not yet described; lowland species. Content: P. rectus. vermiculatus group: moderately large to large species; vomerine teeth present or absent; nuptial pad absent; ova half-pigmented; call pulsed, poorly tuned, sometimes with frequency modulation. Content: P. acutus, aurantium. bunitus, disgregus, kerangae.
It must be stated though tbat this categorisation has meanwhile hlrned out to be in need offUitber revision. The species of the genus PhilautuJl are predominantly found in montane forests, between 700 and 3050 m on Mt. Kinabalu . The boundaries of the individual species ' altitudinal ranges are usually sbarply defined at high montane levels and do normally not include wider tTansitory zones. Most species inhabit the lower stratum (10-300 cm) which is where the clutches are also deposited in cushions of moss, rock crevices, leafaxils, and pitchers
Tab. 18 Distribution of Phi/auills-species on Borneo. Philawus acutus D RING, 1987 Phi/autus amoelllts SMITH, 1931 Phiiautu.\' auralltiutll I I\"GE R, 1989 Phi/autus bunitus I NGER et a!. , 1995 Phi/aufus disgreguJI I NGE R, 1989 Phi/aurus hosii (BO lJLENGER, 1895) Phi/autus inf[eri DRING, 1987 Phi/auflls kerangae DRING, 1987 Phi/auflls iong icrus (BOUL ENGE R, 1894) Phi/auflls mjobergi S/.oUTH, 1925 Phi/autlls petersi (BOULENGE R, 1900) Philautlls re{lIf{ii INGER & STUEBING, 1996 Phifautus sauer; MA.I..KMUS & RJEDE, 1996 Philaulus teetus DRING" 1987 Phi/allius umbra DRING 1987
178
Sabab
lGnabalu
•
• • •
• • • • • • • • • •
Brunei
•
• • • •
Sarawak
•
• • •
•
• •
•
•
•
•
Kalitnantan
•
Rbacophoridae
of Nepenthes. Since it is known of P. kerangae, mjobergi, petersi, and SaUel'1 that their larval deve lopment up to metamorphosis is completed within the jelly sbell, it is to be presumed that this mode of development is equally applicable in all the other Bomean species of Philautus. Altogether 15 specie are known from Borneo, 8 of which from Mt. Kinabalu. Remarks: It is at present unclear if P. hosii occurs on Mt. Kinabalu. DRLNG(1987) re-identified a specimen described as " Rhacophorus hosiilBundu Tuhan" as being P. ingeri. Notwithstanding, P. hosii appears in the check Iist by INGER, STUEBlNG & TAN (1996) whereas P. ingeri does not.
Besides light brown specimens with almost 110 pattern save for a dark interorbital bar, there are pale brown ones with broad, dark dorsal stripes forming a sand-glass like marking. Others irregularly marbled with wide crossbars on light brown or bright scarlet. Some species with uniformly reddish brown or moss-green dorsal sides franled with black dorsolaterally and white flanks with fine blackish brown reticulation; lower side usually uniformly whitish grey to rich dark brown. All specimens, however, sbow an oblique white bar extending from posterior half of eye to upper lip; From top to base: Phi/aufus amoenus, between Pondok Ubah and Pondok Lowii, 2 100-2300 m.
Fig. 174 Pair
Phi/autus amoenus
SMITH,
Fi.g. 175 MaJe
1931
Philaufus amoenliS SMITH, 1931 : 18 - Type locality: Kamboran-
gah Phi/aulus amoenus.l~oER 1966: 340; M ANTHEY 1983: 23; DRING 1987: 37; MAuo.rus 1988c: 176, 1989: 193, 1992a: 124; I NGER & Sn.;EBING 1992 : 41-51 ; MAL.IC~lus I 994b: 230, 1995a: 73 , I996b: 20-26, I996c: 286: INGER, STUEBINO & T AN 1996: 363; I NOER & TAN I 996a: 565; MALIC"IUS & RI EDE 1996a: 32; INOER
et al. 2000
Diagnosis A high montane species of the aurifasciatus group with a hoarse, scrap ing voice; lingual papilla present; nuptial pad oval and prominent; legs relatively sbort; coloration and pattern extremely variab le. Description Males up to 24.2 nun, females up to 34.7 nun; TL/SVL 0.47-0.60 in males, 0.45-0.57 in females ; habitus stocky; snout broadly rounded; canthus rostralis distinct, a quarter of eye diameter; pupil horizontal; fmgers with disks, with or without rudimentary webbings; disks of toes a little smaller than those of outer fingers ; toe webbing 1(1), 2{11.5/0.5),3(1.5/ 1), 4(2/2), 5(1); tibio-tarsal articulation reaches up to tip of snout or a little beyond; metatarsals with many small tubercles; skin above and on flanks smooth to granular, weakly granular ventrally; supratympanic ridge distinct; adult males with a subgular vocal sac.
179
Rhacophoridae
Fig. 176 A-.E3 ir of Philat!!lIS GmOellllS from Pondok Lowii , 2300 m in amplexus.
iris yellowish brown to greenish with golden gleam and nelwork of tine black veins. Ecology/Ethology Philautus arnoenus is a typical inhabitant of the lower stratum of the mossy forest at elevations from 1800 to 2400 m, occasionally up to 2500 m a.s.1. Its calls can be heard already before the onset of dusk, increasing in intensity to climax between 6.00 and 8.00 p.m. The males cal l from leav.es and twigs on bushes and shrubs, rarely higher than 150 cm above the ground, occasionally al 0 from roots and tree stumps overgrown with mosses nearer to the
L80
ground. During prolonged periods of drought the frog is often found in cracks in the so il and rock crevices on the banks of streams, as we ll as in cavities among rock slopes. A clutch was discovered in Barnbusa gibbsiae forest at 2300 ill altitude . It comprised 15 eggs of 8-10 mm in diameter. They were hanging on the face of a rock fissure. Specimens kept in a lerrarium presented themselves as exceedingly voracious, trying to devour virtually everything that moved and did not exceed their own body sizes, in particular butterflies and hairless caterpillars, piders, flie s, roaches, but also
Rhacophoridae
wood li ce , ants and earthworms. Suita ble day shelters arc such as epiphytic cushions of moss, holes in branches and roots, rock crevices, empty pitchers of Nepenthes, and fruit husks. Call: (+12 .5°C) The hoarse, scraping note series bears some resemblance to the call of P bunitus and consists of notes arranged in groups of two or three (note duration II 0 ms; intemote interval 150 ms) which are emitted at intervals of 2-3 s. Each group of notes is preceded by a short grace note and is distinctly pulsed. The carrier frequency lies at l .8 kHz with harmonics at 5 kHz. Distribution Philautus amoenus is as yet known only from Mt. Kinabaln where it appears to be restricted to the high montane zone between Powerstation and Layang Layang and to the MesHau region .
Pltilautus aurantium
INGER,
II .iII " '!II
kHz
53.35 sec
1 1 .11. I
.~
': 1 ..----, ... - ... I 1989
Mendolong, Sipilang District, Sabah Philall/lis aurijasciatus, (not Philautlls Qllrijascia/lis S CULEGEL, M AL.KMUS
53.30
56 .88 sec
Phi/au IllS Qurantillm IN GE R, 1989: 239 - Type locality:
1837)
53.25
53.20
1985: 8
Phi/au/us sp., MA LKMUS 1989: 194, 1991 : 30 Phi/all/US "A", MALKMUS 1992a: 125 Phi/aU/liS "B", M ALKMl:S I 992a: 125 Phi/au/us "AIB", M ALKM t:S I 994b: 23 1 Phi/alllils auran/iul1l, Ii OFFM AN N 1995b: 14; I N() ~. R &
T AN
1996b: 66; MALKM US 1996b: 20-26. I996c: 286; M ALK \I US & RI EDE 1996b: 2 1 Phi/aU/liS al/ran/ium mOIl/allum. M ALKM US I 996c: 286
Subspecies: Phiiaulus auranlium gunungensis, M ALKM US & RIEDE 1996b: 22 - Type locality: "Pagaceenmischwald oberha lb des Silaubacbes, 1450 m, am Siidhang des Mt. Kinabalu"
Diagnosis A small species of the vermiculatus group with sharp, short, single calls. Dorsal side light brown with small black spots; in contrast to P amoenus with little variation in pattern; characteristic traits are the unmarked, pale orange anterior and posterior faces of the thigh ; both vomerinc teeth and lingual papilla absent. Description Males up to 27.7 mm, females up to 28 mm; TUSVL 0.50-0.58; body stocky; head wider than long; snout broadly rounded ' canthus rostral is distinct, slightly curved; lores oblique, weakly concave; tympanum obscure; pupil horizontal ; tips of fingers expanded into large disks, with very smal l rudiments of webbings at the bases; thenar and palmar tubercles distinct; di ks oftoes smaller than those offinger ;
181
Rbacophoridae
toe webbing 1(1), 2(1 /0), 3(LJO), 4(1-0.5/1-0.5), 5(0-0.5); a low inner metatarsal tubercle, no outer one; dorsal and lateral skin smooth; tiny tubercles on legs and upper eyelid; chest and belly coarsely throat softly granular; supratympanic fold weak, but nevertheless distinct; males with a subgular vocal sac. Ground coloration light yellowish brown to reddish brown, flanks and sides oHead lighter; lower side uniformly whitish, partly tinged with rosy; occasionally small dark dots on throat and belly; inguinal region and inner sides of upper thighs intensely yellow; pitch black dots of various sizes on back, sides, and thighs, arranged in rows on back; iri reddish brown to yellowish brown with a golden shine and a network of very fine dark veins. The montane populations on Mt. Kinabalu were described as a separate subspecies , Phi/au/us aurantium gunungensis MALKMUS & RIEDE, 1996. They exhibit a tendency towards longer tibia, a relatively wider bead, and a longer inter-narial distance. Their advertisement calls differ distinctly from
those of the nominate form despite the similarities in structure. Their eggs are half-pigmented. Ecology/Ethology On Mount Kinabalu, this species inhabits the montane forests between 700 and L500 m a.s.l. The males form calling communities off from running waters. They call perched on the leaves of shrubs and bushes 50-200 cm above the grolmd. Calling activity begins with the onset of dusk, increases rapidly - particularly after rain showers - to a high intensity and continues throughout the night up to the dawn of the following morning. The calling communities are static to a remarkable extent: five colonies were observed in exactly the same places on the slopes of the Silau-Liwago valley between 1984 and 1998. This species is occasionally found during the day resting with the limbs drawn up against the body on leaves; such specimens are almost white with the dark dotted pattern suppressed (bird dropping mimesis).
Fig. 177 Phi/autus aurantillm gllnllngellsis from Sg. Liwago.
L82
Rhacophoridae
Fig. LXXVI Sonagram and oscillograms of Phi/aulus aurantium gunungensis (Sg. Silau-Silau; 1450 m; 9.00 p.m. +21 0c)
~ ~ ~
2.00
2.20
2.40
2.60
2.80 sec
________
::i ~~~o
Fig. 178 A calling male of Phi/aulus f!!:l'!!.n!!u'!!. a,;!!'a!!li':!,!!l from Sg. L~ganan.
Call: (+21 0c) The note series consists of 14-30 sharply metallic notes (duration 120-160 ms) at a frequency of2.9 kHz and pulse modulation, and an internote interval of 560-700 ms. Males engaged in duets increases the volume and beat of the note series in the second half of their calls often resulting in a final crescendo.
Fig. LXXV11 Distribution of Phi/aI/Ius {Il1mntium and Phi/alltus Imnitus in the area of the Headquarters. 1600-1700m 1500 - 1600m
o •
P. aurantium P. bunitus
183
Rhacophoridae
In P. a. gunungensis the call duration and internotc
intervals are distinctly longer, the carrier frequency lower. In contrast to the nominate fonn where the call begins with a short pul e (5-6 ms), that of gunungensis sets in abruptly and exhibits a weak pul e modulation. Distribution Th i species has so far been recorded on ly from tbe province ofSabab. Mt. KinabaJu: P a. aurandum : Sg. Langanan (700-950 m). P a. gunungensis: Sg. Silau-Si lau/Liwago (1400-1500 m). The taxonomic status and all ocation of specimens from Sayap (1000-11 00 Ill) is unclear at prcsent.
Phi/auills bUllitus & TAN, 1995
I NGER, STUEBING
Phi/autus bunillis LNGER. Sn!ERIl\G & TA~ . 1995: 127 - Typ e locality: Mount Lumaku, Sipitang District, Sabah Rhacophorus sp. "2", M .-'lLKMUS 1991a: 29 Rhacophorus "A", MALKMU S 1992a: 121 Ph/fat/Ills bunitliS. INGER, STUbBING & TA.'l 1996: 363: L:-JGE.R & T AN 1996b: 66; MALKMIJS 1996c: 286; MALKMl:S & RI F.OE 1996b: 24; M~l\l' H EY & GROSSMA~N 1997: 127: LAKIM et al. 1999; l "GER et al. 2000
Fig, 179 Male of Philalltus bunillls, upper Sg. Liwago.
184
Diagnosis A large species of the vermicltlatus group with a hoarse, scraping call ; vomerine teeth present; lingual papilla absent; dorsum green, often with dark markings; venter orange. Description Males up to 4 L mm, females up to 46 mm; TL/SVL 0.52-0.54; habitus stocky; head wider than long; snout broadly rounded, not projecting in profile; canthus rostralis rounded, distinct; lores oblique, slightly concave; tympanum visible, slightly obscure, half diameter of eye; pupi l hOlizonta l-ell iptical ; supratympanic fold strong, rounded; fingers with truncate or slightly rounded disks, rudimentary webbings at bases; I distinct, large subarticu lar tubercle on each finger; inner metacarpal tubercle large; toes with disks smaller than those of outer fingers; inner metatarsal tubercle oval, no outer one; toe webbing 1(1), 2(0.5/0.5), 3(0.5/0.5), 4( 1/ I), 5(0.5); dorsum, limbs, and flanks with widely scattered, small tubercles; venter coarse ly granular; males with a median subgu lar vocal sac and no nuptial pad. Influenced by exogenous factors and depending on the leve l of excitement the coloration of the upper
Rhacophoridae
c c
'"
~
I
n: @
Fig. 180 Ventral pattern of Phi/autus bunitllS from Sg. K~manti s, ca. 1050 m, Sayap.
Fig. 181 Calling male of Philautus bunilUs from Sayap,
ca. 1100 m.
side may vary from pale pastel green to deep dark which often begin almost with hesitation, but then fo llow at ever decreasing panses. They exhibit a green, sometimes uniform, but usually with blackisb to lead -grey, wide stripes and spots; flanks slightly [ising frequency modulation and harmolighter; throat clouded with grey or uniformly yellowish; chest and belly orange, darkening posteriorly; lower sides of limbs Fig. LXXVln Sonagram and osc il\ograms ofPlzilautlJS bun;(us (Sg. Liwago, 1450 m; 8.00 p.m. + 19°C) and webbings also orange; iri s pale go lden, with a rough network of black veins .
EcologylEthology [n contrast to the other spccies of Philau(us inhabiting Mt. Kinabalu , Philautus bunitus is exclusive to the middle stratum and rarely encountered below 3 m above the ground . It inhabits the oak-chestnut forests between 1000 and 1800 (1900)m a.s .1. Its diet comprises spiders, cate}'pillars of butterflics, grasshoppers, and moths. The hardly mistakable voice ofthis species can be beard throughout tbe nigbt, occasionally even during the day, particularly at the onset of a rainsbower. Jt does not fOIlD isolated choruses like e.g. P. aurantiutn, but is relatively evenly di spersed in tbe fore ts. Duets and voca l trios are ollen beard. Call: (+ 19°C) The note series consists of 15-25 hoarse, scraping notes (carrier frequency 1.8 kHz note duration 190-200 ms)
2.00
2.20
2.40
2.60
2.BOsec
kHz
5.0 sec
- ..
,,\ 5
.
,F,I'''
"l'. t,.
,.
I
.' .... . _ ' .. r ., ~~
~0U~
185
Rbacophoridae
nics at 5-6 kHz. The note is distinctly pulsed at its early stages, but i:ndistinctly so towards the end; pulse duration 6 ms.
venter cream-coloured, with fioe black speckling; iris golden in upper third, lower two thirds dark brown.
Distribution P bunitus is only known from western Sabah. Mt. KJnabalu: On Mt. Kinabalu it occurs in the entire Liwago- ilau region between 1400 and l800m (up to 1900 m in places, see also map on page 183). It is furthermore known from the Sayap region (1000 m; Sg. Kemantis, Sg. Wariu), and from the eastern Sg. Mesilau (J 900 m).
Ecology/Ethology Philautus ingeri is known from the upper montane forests. Ma les call at night, from leaves and brancbes 1-5 m above the grmmd.
Pltilautus ingeri D RING, 1987 Phi/ooLUS ingeri DRJ)1G, 1987: 21 - Type locality: Camp three. 1300 m, GUl1ung Mulu, Fourth Division, Sarawak Rhacophorus collelli. (\10t. Po/ypeda/es col/cUi BOULENGER 1890) INGER I 954a: 250 Rhacophorw; hosei. (nol Philallills hosii BOULENOE R, 1895) lNGER 1966: 304 (parI), 1978: 313 (part): MANTHEY 1983 : 24 (part) PhiialitUS if/geri.lNoER & Sn.:EBING 1992 : 41-51 ; I NGtlR & TAN 1996b: 67; MALK,\lUS & Rrr:OE I 996b: 25
Diagnosis A large member of the hosii group with a soft call; dorsum brown; vomerine teeth p resent, lingual papi lla absent; canthus rostralis and iris with disruptive patterns; upper eyelid and heels with prominent tubercles.
Call: A series of about 5 soft "quoink" notes. A call analysis was not yet performed.
Distribution This species is known only from Gunung Mulu in Sarawak and Mt. Kinabalu (see Remarks at the end of the genus account). Mt. Kina balu : Bundu Tuhan (1375 m).
Pltilalltlls longicrlls
(BOULENGER,
1894)
Lw/us /ollgicrlls BO U~"NGER, 1894a : 88 - Type l ocali ty: Palawan Phi/all/US aurifasciatus, (not Phi/auflls alirifasciattlS SCHLEGEL, 1837) INGER 1966: 341 (part); M"'il HF-Y 1983 : 24 (part) Phi/autu,\' tongieru.\', 1NGER & STt' EB ING 1992: 41·51; h;GER, S TL'EBING & TAN 1996: 363: MALIC'I US & RrEoE 1996a: 35: I NGPR et al. 2000
Diagnosis A small brown species of the aurifasciatus group
Description
with a rattling call; vomerine teeth and lingual papilJa absent; simi lar to P. peters;, but females witbout a tubercle on the tip of the snout.
Males up to 36 rom, females up to 50 mm; TLlSVL 0.57-0.58 (males), 0.63-0.64 (females); head slightly longer than broad; snout depressed, pointed, and projecting in profile; canthus rostralis angular and nearly straight; lores almost vertical; tympanluD distinct; pupil horizontal-elliptical; fingers with wide disks, webbing rudimentary; disks of toes slightly smaller than those of outer fingers; inner metatarsal tubercle oval, no outer one; toe webbing 1(1),2(01 0) 3(010) 4(2/2), 5(0); dorsum shagreened; tubercles on dorsum, limbs, upper eyelid, and heels; throat wrinkled; belly and undersides of thighs areolate; males with a subguJar vocal sac, no nuptial pad. Dorsum brown, with a dark inter-orbital triangle; dark bars on limbs; a broad black canthal stripe;
Males up to 23 rnm, females up to 29 mm; TL/SVL 0.59-0.64; head wider than long; snout obtusely pointed; canthus rostralis distinct, lores concave; tympanum obscure, one third eye diameter; pupil horizontal; tips of fingers with wide round disks; disks on toes smaller than those on outer fingers; an elongated inner, but no outer metatarsal tubercle; toe webbing intermediate in extension between P. petersi and P. mjobergi; skin shagreened above; tu bercles and ridges i:rregularly distributed on back, head, and limbs; throat weakly, chest and belly coarsely granular; supra tympanic fold present; males with a subguJar vocal sac; nuptial pad oval and prominent.
186
Description
Rhacophoridae
Coloration and pattern extremely variable in specimens from Sarawak and Palawan; there are specimens with a barred dorsal pattern as well as con color ones, but none with a white streak below the eye or a dark network on a white ground. There is no description of specimens from Mt. Kinabalu.
Ecology/Ethology Almost nothing is known about the life history of this species other than it was found in primary forest between 900 and 3000 m altitude. It is active at night and arboreal, by day it rests under litter on the forest floor. Call: A harsh rattle, similar to the call of P mjobergi.
Distribution Philautus longicrus is known from the Philippines (1200-1550 m), Sarawak (900-1200 m), and Sabah (up to 3000 m) . Mt. Kinabalu: Layang Layang, Sg. Tibabar. INGER & STUEBING (1992) listed this species for an elevation of 3000 m (pakka). Bioacoustic studies, however, promote the idea that it shares the habitat on Mt. Kinabalu with P. amoenus at 1800-2400 m a.s.1.
Plrilautus mjobergi
SMITH,
1925
Philaulus mj6bergi SMITH, 1925a: II - Type locality: Mt. Murud. 7000 feet, Sarawak Philaul!is petersi, (not Phi/autlls peters; BOlJl.ENGER, 1900) SMIl'H 1931 : 20 (part) Phi/aufus aurifasciatus, (not Phi/aulLls aurifasda",s SCHLEGEL, 1837) INGER 1966: 341 (part), 1978: 313 (part); MA1'!TIiEY 1983: 24 (part), MALKMI;S 1985 : 8,1987 : 284, 1988c: 176 PM/aulliS mjobergi, SMITH 1931 : 21 (part); DRING 1987: 38; MA LK.MUS 1989: 193, 1991 : 30, 1992a: 124; I NGER & STUEBING 1992: 41-51 ; M ALKMVS 1994b: 230; INGER, STUElJING & TAN 1996: 363; LAKl~1 et al. 1999; INGER ct al. 2000 Philaullls mjoebergi. HOFF~V,NN 1995b: 14; M,\LKMUS 1996c: 286; MALK.\fU S & RJ EDE 1996a: 29
Diagnosis A highly variable species ofthe aurifasciatus group with a sharp, irregular, rattling can; vomerine teeth absent; lingual papilla present, sometimes absent; females without tubercle on tip of snout.
Fig. 182 Phi/al/tIiS' mjobergi from Sg. SiJau-SiJau, ca. 1550 m.
Description Males up to 24 Im~ females up to 32 mm; TLlSVL 0.52-0.62; head wider than long; snout rounded to elliptical in outline; canthus rostralis distinct, lores oblique and concave; tympanum obscure; pupil horizontal ; finger tips with wide oval disks; distinct rudiments of webbings; disks on toes smaller than those of outer fingers; inner metatarsal tubercle oval, no outer one; toe webbing l{J),2(1 / 1), 3(111), 4(2-2 .5/2-2.5), 5(0 .5-1); dorsum shagreened, sometimes with a subtle hlbercle on upper eyelid or on heel; supratympanic ridge distinct; throat granular, belly and wldersides of thighs areolate; males with a subgular vocal sac; nuptial pad oval, prominent. Dorsal ground coloration pale brown with a tendency towards yellowish or reddish; inter-orbital bar and dorsal stripes usually present, narrowing at mid-dorsum to form an sand-glass like marking; conco lor specime ns are occasionally found, or those with a light, broad medial stripe; thighs with 3-4 dark bars; flanks lighter, sometimes spotted in lower third; lower sides of body ye llowi sh to soft flesh-coloured, sometimes obscurely spotted with
187
Rbacophoridae
grey on chest; iris golden with a dark horizontal bar.
Fig.
1~3 ..!hila!ltu~lI1{obergi
!rom Sg..: S11au-Silall.
Fig. LXXIX Sonagram and oscillograms of Phi/auftts mjoberg i (Sg. Silall-Silau, 1450m; 6.30 p.m. +20°C)
1.50
1.60
.u.. rr
kHz
': [ 188
1.70
1.80
1.90 sec
J.
r
... •
5 .0 sec
EcologylEthology On Mt. Kinabalu , Phi/autus l1!jobergi is a typical faunal clement of the oak-chestnut forests at altitudes between 900 and 1700 m, occasionally up to 1800 I'n a.s .1. The males call mainly at night with maximum intensities between 6.30 and 8.00 p.m., and again between 4.30 and 5.30 a.m, but can occasionally also be heard during the day. They advertise from leaves and twigs of shmbs and bushes, but preferably from tborny rattan thickets, 20-250 em above the ground . On Mt. Mumd Mj6berg found them depositing their eggs in the pitchers of the Pitcher Plants (Nep enthes), no other water being available for the purpose (SMITH, 1925). A female captured on Sg. SilauSilau in August deposited shortly afterwards 7 large, unpigmented eggs (diameter of the jelly shell 6 mm) on a dead leaf in a terranum. Call: (+20°C) The advertisement call consists of a series of 4-8 metallic, sharp, pulsed notes (carrier frcquency 2.7-3.5 kHz; note duration 80-105 ms; inte1110tc interval 70-80 ms) which is followed by a pause of 0.8-1 s and a hOlt, marked final grace note. A note series inclusive of tbe grace note lasts 1.8-2.5 s and is repeated at irregular, often long intervals. Distribution Phi/auius mjobergi is probably the species of Phi/autus with the
Rhacophoridae
widest range on Borneo. Mt. Kinabalu : Common in the entire Headquarters region (1400-1800 m) with additional records from Mesilau Cave, Marci Parei, and Sayap.
Philautus peters;
(BOULE OER
1900)
Ixalrts pelersi BOVLENGER, 1900: 185 - Type locality: Mts. Penrissen , Dulit , and Kina Balu, Borneo and Great Naruna Phi/allills aurifasciatlls. (not Phi/allius ollrifasciatlls CHLEGEL, 1837) iNGER 1966: 341 (part), 1978: 313 (part); MANTIl EY 1983 : 23 (part) Ph.i/autlls pelersi. Si't1TH 1931 : 20 (part); MALKMUS 1991: 30, 1992a: 124; INGER & SrUE BJ NG 1992: 41-51 ; M ALJL\\US 1994b: 231 , 1996c: 286; INGER, SrUEBP.'G & TAN 1996: 363; INGE R & TAN I 996a: 566; MALKMUS & RJ E8b 1996a : 35; H OFFMANN I 998c: 15: LAKlM et a!. 1999: T~GER et al. 2000
Taxonomic remarks: The original type locality is a composite based on specimens from Mt. Dulit, Mt. Penrissen, and Mt. Kinabalu (pakka). However,
the Kinabalu syntype is "an unidentifiable juvenile" (DRING 1987). Diagnosis A stout species of the aurifasciatus group with a very characteristi~, single-pulse-train call; vomerine teeth absent; lingual papilla present or absent; dorsal surface tu ercular; fema le with a tubercle on tip of snout. Description Males up to 27 mm, fema les up to 37 rom; TLlSVL 0.5-0.66. DRING (1987) noted marked differences in the morphological ratios of specimens from various geographica l regions. Similar di ffe rences are also found on a smaller scale in specimens from various elevations from Mt. Kinabalu (MALKMUS & RIEDE 1996a). Head broader than long; snout ell iptical in outline, rounded in proftle; canthus rostralis rounded, lores
Fig. ] 84 Female of Phi/au/us pelersi from Sayap.
189 .
Rhacophoridae
oblique and concave; tympanum obscure; pupil horizontal ; tips of digits with wide oval di ks; distinct rudiments of webbings; disks on toes someFig. 186 A pair of Philautus petersi in amplexus from
Sayap.
what smaller than those of outer fingers; inner metatarsal tubercle oval, no outer one; toe webbing l{l), 2(0.5-1 /0.5-1), 3(0.5-110.5-1), 4(2/2), 5(0.5-1); tibio-tar al articulation reaching up to eye; dorsum distinctly tubercular, particularly on upper eyelid, shoulders, upper side of legs, and be low jaws; heel with a small, spine-like tubercle; supratympanic fold distinct; throat granular or wrinkled, belly and undersides of thighs areolate; males with a subgular vocal sac; nuptial pads prominent. Dor urn brown to grey, pattern high ly variable; flanks lighter; groin and lower flanks flecked with reddi sh brown; underside whitish to grey; iris brown to golden, with a central horizontal dark bar. Ecology/Ethology On Mt. Kinabalu, P. petersi occurs in submontane and montane forests between 800 and 1700 m a.s.l. The males call from shrub thickets, 30-300 em above the ground, often adjacent to rivulets. The
190
Rhacophoridae
day is spent hidden beneath dead leaves and bark. The female 's keratinized tubercle on the tip of the snout is possibly used for the excavation of nests, perhaps in epiphytic mosses. Amplexi were observed in the month of August. A clutch of this species was found on a dead leaf in the leaf litter layer at Sayap. It contained up to 25 unpigmented eggs. Call: (+ 19°C) The call is a 1.2- 1.5 s long train of 16-56 rough pulses (pulse duration 20-25 ms; pulse rate from 27 (in specimens from Sayap) to 46 per second; dominant frequency 1.6-2.2 kHz; weak harmonics between 5 and 7 kHz; no frequency modulation. The calls are emitted at intervals of several minutes.
Fig. LXXX Sonagram and oscillograms of Phi/aU/liS petersl (Sg. Liwago, 1450 m; 7.30 p.m. + 19°C)
6.80
-
-- .. ---
Fig. 187 Eggs of Phi/allflls petersl from Sayap, ca. 1000 m.
7.40
7.20
1--------------.·I.:.I . ..
Distribution Malay Peninsula; widely distributed on Borneo. Mt. Kinabalu: Langanan Valley (800-900 m), Sayap (900-1100 m), Marei Parei (1500 m), Headquarters region (1450-1700 m). The record by SMITH (I 931 a) "Kamborangah, 2200 m", very likely refers to P amoenus. That the " unidentifiable juvenile" (DRING 1987) from Pakka Cave at 3000 m altitude in the series of syntypes represents P petersi, can be dismissed with certainty.
---
7.00
7.60 sec
~I------~------I
I.IUII••I.IIII•••
Pltilautus saueri & .R.mOE, 1996
MALKMUS
Phi/autus sauer; M.ALKMUS & Rr DE, 1996a: 29 - Type locality: Pakka (3050 m), Mt. Kinabalu Philauills cr. mjoebergi, (not Phi/allius mjobergi MITIl, 1925) MALKML 1995a: 70 Philaut/Js salleri, MALKMUS 1996c: 286
Diagnosis A small, stocky, high-montane species of the aurifasciatus group with a soft, hoarse, rattl ing call. Vomerine teeth absent; dorsal side of body brown with a whitish inter-orbital bar. Description Male 21.4 mm; TLlSVL 0.54; bead slightly broader than long; snout rounded in both outline and profile; canthus rostralis rounded, lares slightly concave; tympanum obscure; pupil horizontal; tips of digits with large oval disks; webbing rudiments
19 1
Rhacophoridae
distinct; di k on toes smaller than those on outer fingers; inner metatarsal tubercle distinct, no outer one; toe webbing ](1), 2(1 /0.S), 3(1.5/1), 4(212), 5(1); dorsum smooth; heel with a small tubercle; supratympanic fold prominent tightly bent, from eye to axilla; 2 rows of ridges from eyelid to centre of back; flanks, throat, and belly granular; males with a subgu lar vocaJ sac. Upper side of body, tibiae, and forearms blackish brown, peppered with small whitish dots; remaining portion of limbs and sides of body white to whitish grey, reticulated with black; a whitish orbital bar separates redctish brown of snout from blacki h brown of dorsum; a broad white area between eye and upper lip, bordering blacki h brown of tympanum area; supratympanic fold whitish; belly white, with a umber-coloured, reticulated pattern ; throat yellowish brown; lower sides of feet and hands, and upper sides of first two fingers whitish yellow; central portion of iris brownish Fig. 188 Phi/aU/us saueri from Pakka Cave region, 3000
192
grey; with widcr upper and narrower lower margins in yellowish green.
EcologylEthology Calls of P. saueri can be heard at altitudes between 2200 and 3050 m a.s.l., very rarely though below 2400 m. In contrast to all other species of Phi/autus occurring on Mt. Kinabalu, the males of this species call also during daylight hours, even in bright sunshine. Their diumality must certainly be regarded as an adaptation to the fact that the temperatures at these altitudes not rarely drop to just 5°C at night. This point of view is supported by the observation that calls could still be recorded at 10.00 p.m. in the less elevated mossy forest (below 2600 m). The males call out oflow, very dense bush vegetation. Four pitchers of Nepenthes villosa were discovered in the Leplospermuln-DaClydillln forest near Pakka, in August 1994, which each contained 9-17 gelati-
111 .
Rhacophoridae
nous capsules of 11-15 mm in diameter of p. saueri. 20-30% of these capsules did neither contain eggs nor embryos, but were m 'cly filled with liqu id. The other she lls held larvae at various stages of development. All these pitcher were dead and leaking, although their well-shaded location and contact to the ground secured the necessary moisture for the development of the larvae. It remains unclear, though, whether the contents of the individual pitchers were the product of one or several pairs of these frogs . Call: (+ 15°C)Anote series consists of 2-4(5) pulsed notes (note duration 30-40 ms; pulse rate .150 per second) at intervals of70-100 ms at a carrie r frequency of 1.9-2.6 kHz. paced by a pause of often several seconds, a drawnout "gweeh" sound (duration 90-120 ms) may follow, which may also be emitted alone . The structure of the note series exhibits certain simi larities w ith that of P. mjobergi.
-
-
-
---
I - -- - - -..... It--.... ' ,e .' . ~- - - - - 1
34.60
4I4
34.80
35.20
35 .00
•
•
I
':l
kHz
Distribution Philautus saueri is as yet known only from the montane forest between Layang Layang and Pakka Cave. Fig. J 89 Tadpole of PhilautlJS salleri.
-
-
Fig. LXXXI Sonagram and oscillograms of Philallllls sauer; (pakka, 3000 m; 11.00 a.m. + 1SoC)
35.40 sec
"I
39.72 sec
;. !
lH
• I
Genus Polypedates
TSCHUDJ,
11 I 1838
The genus Po/ypedates is distributed from China and Japan, through tropical Southeast Asia, up to the Philippines, Borneo, and Java. Although CHANNING (1289) demonstrated in a phylogenetic study that Po/yp edates is a genus separate from Rhacophorus. its validity has remained a subject of debate. In contrast to Rhacophorus, a portion of the scalp is fused to the skull in Po/ypedates (except in P. colletti). and the ringfold 00 the lower side of the digital disks typical for Rhacophorus is missing in Polypedates. While Rhacophorus possess webbings between the fingers, such are absent in Polyp edates. The Borneao pecies are relatively large tree frogs with compressed bodies and long limbs. Vomerine teeth present, tympanum visible, pupil horizontaL Fingers and toes with enlarged pads at their tips, toes webbed over half to almost full length. Males with subgular vocal sac and nuptial pads.
193
Rhacophoridae
Tab. 19 Distribution of Polypedates-species on Borneo. Sabab
Polypedates colletti ( B oULENGER, 1890) PO()lpedates leucomystax (GRAvENHORS1, 1829) Poiypedates macrotis (BOUUl.'lGER, 1891) PolliPedates otilovllu.5 (BOUlEI"GER 1893)
The eggs lack pigmentation and are depo ited in a foam nest which is attached to the leaves of bushes in a manner that hangs freely above a water body. Tadpo les with almo t sphero idal bodies, eyes on sides of head, oral disk ventral, strong black beaks and labial teeth rows present; tail with high fins .
• • • •
Kinabahl
• • •
Brunei
• • • •
Sarawalc
Kalimantan
• •
• • •
• •
The Bornean species live in the primary and secondary forests of the lowlands. P leucomystax and P otiiophus, appear also in cleared forests, agricuHural fields , villages, and towns. Four species are known from Borneo, three of wh ich also from Mt. Kinabalu :
Key to the Bornean species of Polypedates Skin of head not fused to skull ; conspicuous Xshaped mark on dorsum.. .... .... .... ..... ....... col/etti • Skin of head fused to skull in adult specimens. 2
3 A broad black temporal stripe ............. macrotis • Temporal stripe absent or weak; often 4 narrow dorsal stripes ............ ...................... leucontystax
2 A serrated bony crest projecting above the tympanum .............. ......... .......................... . otilophus • No such crest..................................................... 3
Polypedates leucomystax ( GRAVE HORST, 1829) Hy la leucomysto..'C GRAVENIIORST, 1829: 36 - l)'pe locality: Java Rizacoplzonls macula/us. (not Polypedates maculatus GRAY, 1834) BOl. L~G[R I 887b: 95; MOC'QUARD 1890: 149 Rizacupizorus leucomystax var. quadrilineallls, (sYI! . after SmINCG~R 1907: 157) M OCQUARD 1890: 150 Rhacophorlls ieucomystax sexvirgata. (syn. after STEJN~GfR 1907: 157) VAN KA.\.tP(N 1923: 249 Rizacopizorus/eucomystox, VAN KAMPEN 1923: 246; SMlrH 1931 : 9; l);OER 1954a: 250; Mi\LK.\.tUS 1996e: 286 Rizacophol'l/s lellcomys/ox lellcomystox, I NGI'R 1966: 307 Polypedates lellcomystax, INGhR 1978: 313; MANlln:v & D~ZFR 1982: 15; MANl'ILEY 1983: 24; MAJ.KM US 1985 : II, 1987: 283. 1988d: 7; H OFFMAN' 1995b: 14; INGER. SWfBI'G & T AN 1996: 363; HOFFMANN I 998b: 89 Polypedates lellcom),slax lellcomystax. MATS!:I 1979: 341 ; LAKIM et al. 1999; J'IGER ct aJ. 2000
Taxonomic remarks: NARINS et a1. (1998) pointed out the morphological, genetical, bioacoustical, and
194
ethologica l differences between the striped and stripeless forms wh ich they observed sympatrieally on the Malay Peninsula and consi dered it justified to grant both species ranks. Diagnosis A small to moderately Large, slender frog with long, slender limbs; specimens from Mt. Kinaba lu usually with 4 narrow dorsal stripes; no bony crest above tympanum. Description Male up to 50 mm, females up to 75 !TIm; TLlSVL 0.47-0.60; head longer than broad; nout obtusely pointed; canthus rostraJis harp, lores not concave; tympanum distinct, 75% or more of eye diameter; finger tips with large round disks, webbing rudimentary; toe tips not as wide as those on outer fingers; toe webbing 1(0),2(010) , 3(0/0),4(2/2), 5(0); an oval inner metatarsal tubercle, outer one small or absent;
Rhacophoridae
~
~::;;
:i @
r-~--------~--------------~---' c c
~
~
.,,.,
a:
I
@
Fig. 19.!J..
P!!.lYEec!..a'!s i!tl!!:o!!.!y!.t£l:.'< ~om Poring, 550 m.
:i
~rB~~~~~~~~~~~~~@ dorsal skin smooth, venter coarsely granular; supratympanic fold present, curved. Light beige, yellowish, or brown above, with dark tlecks or four narrow stripes running down length of back; limbs with dark crossba rs; undersides whitish to whitish grey; upper lip white; nuptial pads yellowish; iris light golden with a fine black network. Tadpoles: Body spheroidal , flattened above, rounded below; eyes dorsolateral ; oral disk ventral, subtemunal; labial teeth rows formula 4(2-4)/3; beaks black, strong, finely serrated; spiracle sinistral; tai l tapering abruptly in the last quarter to a slim, drawn-out tip; total length up to 50 rnm. Dorsum brown to yellowish, with dark stripes at stages 40-41; whitish ventrally; tail grey with dark dot ; fins transparent, dark spotted (YORKE 1983). Ecology/Ethology Po/ypedates /eucomystax inhabits the marginal zones of primary and secondary forests, but also cultivated areas, road sides, and village (where it may even be encountered inside houses). It is a nocturnal frog which may, however, also forage during rajn shower in the day. Its diet comprises insects, spi ders, and even small lizards. The males congregate at stagnant and slowly flowing water bodies and form choruses. It is not rare that a fe-
From top to base: Polypedates leucomystax. Fig. 191 Typical foam nest rrom Peninsular Malaysia. Fig. 192 Tadpole fro m Peninsular Malaysia. F~g. 193 Ta
Bi?tan, lnd~ n~ia..:..
male assumes tbe active advertising by ascending to a branch, drumming against it for minutes and occasionally emitting a croak. It does usually not take long for some males to arrive. Once a pair is united in an amplexus, it chooses a site for building a nest in the branchwork of a bush 10-70 em above a water surface or along the bank of pools on the ground (earth, rocks). The female then expel s from its cloaca a liquid produced by foam glands which it whips to a primary foam with its hind limbs. The subsequent thrust contains the eggs (l50-900) whjch are mixed with the male 's sperm and the foam. The female does most of the nest bwlding and is supported by tbe male only occasionally (KABIS H et al. 1994).
195
Rhacophoridae
Five 10 six days fter ()vipo~i l ion Ihe tadpoles emerge and drop from the foam nest into the water beneath (111 : 2b) . Call : NARt\J ~ ct al. (1998) identified three separate call types luI' each of the two forms (stript:d and stripeless. respectivel y). In the striped fOnll on ly this one has so far been found on Mt. Kinaba lu there an: tWll pulsed call typcs with dominant frequencies at 223R II ", (call du ration 174 illS , 13 pulses). and <)34 11 z (call duration 5Ams. 4-5 pul'e .), respectively. and stac ato call s (dominant frequency <)<)7 Hz. call dUnlliul1 527 illS) . Analyses of the vocalisation); produced by Kinaba lll ~pecimens arc not available (rv1." r~I )l I 9112,1 , M A I ,U I ct al. 19XA, S At\CHF7.- I Ii ,r(i( AII et al. 1905). Fig. 194 PO/lll('da/, ..' /}/(/ cmtis from D..:ramakot. Sa bah.
Distribution This species has a vast distTiblltion, ranging [i'om eastern India and southern China to Java . Borneo. and the Phi lippine Is land ', rrom sea leve l lip to 1000111, on MI. Kinabalu up to 1500 m a.s,1. ;\11 . Kinabalu: Sayap. Kiau, Tenompok. 13ulldll Tuhan, Lllbanc. Kuntiasang. Poring. Whelher the "\.:gg mass" reported frol11 Kambon1l1goh by MATS{.I ( 1979) really belongs to P. lell('()myslUx is qUl.:sliollable.
Po/ypedlltes 1Il1icrotis (BO LJ LENGER, 189 1) fIl jl( I U II \ 13()l l l" ( , I JC I ~~) l ' 2 X2 - Type Inc_ll il,Y : Halall1 Di , lri \'l, <;"ra",uk 11 1/(/( "pi")!'/I" "'I ICU IJJ ' ' /1/\ !iI/ A" ( ,), n . ,Ilk .. IN(,H 196(,: 301' ) 1\"11< 1956: -IDC) IIlwi'""I",,·/i., 111(/<'/'011\'. I Md' ~ 19(\0: J 13 I.J() / \',u 'd a t('.\ II H I (TOI;S , I '(doH, & Sn) B I Nt. 1119 2 - ...t 1-5 I : 1I 1l 1l' ~ ' \\' 19<)5 1>: 1-1 : IN' '!' '', S n '-111M, &. r" 1996: 363:
U/U/( " ,p! lr U "U \'
I. \K 1\' CI
"I. Il)lJ9:
I N(,"~ ~ I 0 1 lOOI)
Dia ~n os i s
A medium-sized to large tree frog with a tTiangular head and a Cilar:lCleristic. broad , blacki . h brown tcmporal stripe . Desc.riptioll 'v1ales lip to 57 111m, Icmalcs lip to 90 111111; TUSVL 0.49-0.60: vcry ~il11ilar in its morphological details to P. I('II('()I/IYSIU.I : fcmalcs have relalivt:ly wider
Il)o
head. and finger tips, but narrower tympani than males; skin smooth. some specimens with tiny bumps Oil back ; olien a low. whitish dermal ridge of skin on outt:r edge or forearm. Upper side of hcad. hack, and limbs tan to brown: flanks sometimes wilh black dots; limbs \vith d;Lrk crossbars ; a broad dark stripe from eye to arm; undersides whitish . \vith or withoul dark dots: throat mottled with brown ; iris light golden . Tadpoles : I lead-body portion oval; ~nollt broad ly rounded; body nattelled above; eyes latera l. visibl- from below: oral disk ventral. subh.:rminal; labial teeth rows lonmda 5(2-5/3( I) or 0(2-6)/3: beaks weakly serrated; tail e0I1ve" 1.2- 1. 7 times F i ~,
195
['O/Ijll't/lil e., l/I{/c rOlis
from Poring.
Rhacophlwidae
head-body-length ; tOlallength 55-60 mill . Head and body dark green, dorsally and latera ll y with light spots, below si lvery grey; eaudallllusc lc with a dark reticulated pattern ; dorsal fi n green, ventral fin white with sma ll black dots (I" oFI 1985) . Eeo logy/E tholog)' appears in low land primary and sccondalY forests. It is nocturnal, preylllg upon insects (particularly roaches and l1cmiptera) and sp iders. Males fo rm large ca lling commun iti es on the ground or in low vegetation (up to 3 111) ncar rainwater- fi llt:d poo ls. Tht: females at1ach large foam nests to vegetat ion (sh rubs, sma ll trees, herbs, grasses) overhanging ditches and rainwater-fi llt:d pools (Ill : 2b, c) at the edges of forests . POII 'f lee/ales II/(/ C'rolis
Call : A long, drawn-out chuckle. the first note di stinctly pronounced . The advertisement cal ls consist or main and short notes, with the fonner lasting 0. 1-0 .79 s and consisting of 1-3 notes (duration 0.09-0.12 t11S , 5-6 pulses per note). The short notes (durati on 0.005-0.0 I s) arc emitted before or after the main notes. The frequency ra nge is \vide, covering 0.4-2.6 kHz (+22°(,) (M A l SU I 19&2b).
I)l'scription Ma les up to !l0 mm, fcmales up to Y7 Illln ; TI./SVL 0.47-0.55; head longer than broad ; snout rounded in prolile, projec;ting; tympanum di stinc;t, oval, 3/4 of eye diameter; tips of fingers expandcd inl o large di sks; rudimental)' webbing betwcen 1" and 2,,·1 lingel's; disks ortoes much smaller than those MOliler fingers: an ova l inner mctatarsal tubercle, no ollter one; toe webbing 1(1), 2( 1/05),3(0.5 /0.5) , 4(2 /2), 5(0.5): skin above smooth . flank s and undersides coa rse ly gra nul ar; a pointed projcc;tiol1 on hecl and o lt en another, smaller one on elbow; a row or 4-6 \vhi le tuberc les be low vent. Yellowish brown above, with num~rous thin, dark stripes running down back from the head; inguinal region \V'hite wit h black lincs and dots; thigh. with 7- 1 I n31TOW black bar's 011 the anterior and poslelior faces: unders ides diny white ; Iluptia l pads greyish or yellowish; iri. yel lowi h grey. Tadpoles: Head-body seclion spherical. tOI orhcad natlened; very simi lar to the larvae of P macro/is: labial teeth rows formu la 4(2-4)i or 5(2-5)i 3; totallcngth 63 1111ll , maximuill up to RO Illlll . Yellowish green above, be low si lvery white: dorsal fill with golden dots. Ecology lE I h ology
Oistribu tio n This frog is knO\vn from the :v1 alay Peninsula, Sumatra, Borneo, Palawan , and tht:: Sulu Archipelago (200-1 50 111 a.s. L). \,11. Kinablllu: Sayar, Poring: Sg. Kipungil.
dwell s the edges or primary fore sts as well as habitats modified by J11;JJ)
Pntypec/aJ£-'S (JJitoplrlls
Fig. 19(, Poll'p<,t/are.1 "lii"II/III.I· fi'UJlI POI·ing.
Po()'pet/ates oti/oplllls (BOULENCiEI{, 1I1/l/(IJJIJlOl'IIs o)(i/lJp/IIIS n OI " l1ol\gol1.
r--. .
1 No:l H.
1893) 11l93: 527 - Ty pe loc>,tit y:
H()rn ~o
Rlracol'iro/'ll.' ",ilopll/l.l. l'vL\ I. " 11 " I ')K'! : 192 PO/YJ!l'dUlc '" olilol'lrll" INt " ·. & S n 1-l1l " ; lIN}: -l t- 5 1: I NI,! R, S ' II 'F ilI No; & T AN 19'.16: :lhJ : 1. ., , \\1 Cl "I. (1 9991 : IN, d'R c l al. (2 000)
Diagnosis A large tret:: ('rug with a triangular hcad and a pro-
truding, sharp point at the angle of the j
197
Rhacophondm:
(seco ndary forests. tree plant ati ons.loggl.:d arcas. even vi llagc '). It i. a lowland species. occ urring fro m sea level up to 500 m. rarel y higher (up to I(Jon m a.s.I.). Its die t cons i ~ts ofinseets (especially arboreal c ri cke ts) and spi ders. aptured spee i m e n ~ emanate a charac t eri~tic. s tro ng odo ur. Males call at night from low vegetati on ( <. 2 m) nca r ' mall pools (111 : l b. c) where the females, similar to P macro/is. alta h th e ir foam nests to low herbs about 30 cm above the water slIIfaee. Call : A series of slurred. rasping not es fol lowed hy som e loud "c hu cks" (IN(oI"\( & . I Ul' Il ING I (97) . A ca ll anal ysis is not ava il able. Distribution
Borneo. ' umatra . M t. K inabalu : Only known from Po ring : Sg. Kipungit.
Fig. 197 /'oit1Jec/lIfes /lli/lip/III,I rrom West Kallfllanlall .
Genus Rlwcophortls K lJ lIl. & VAI\ H AS~ELT, 1822 Species of RIIlI ('()/I//lJ/"lI,I' arc known from the region from Ind ia. Sri Lanka and China 10 Japan, the Phil ippi ne" and the Greater . unda Islands , This genu s incl udes smal l to moderately large. Ibrcst-dwe lling tree Ij'ogs wit h robust bodies and long Iim bs. Most specil.:s po~scss vomcrine teeth, but thi s trait is not constant in al l spe<.: ies. TherL' is no oss i li ca tion of the sca lp whi ch distinguishes it from /'o h~ ){'d{/(('\ . Tympanum visible. eye ' large with a hori m nt al PIIPil : tips of fingers and toes ex panded into largc. round or ova l disb. those of toes not as wide as those of outer lingers; webb in g~ bet ween finger ' and toes prc~ent ; undersides of fingers with circ um-margina l groove~. ome specie ' (I re ab le to pcrfonn paraglid ing Ilights hy spreading their excessivd webbed hands and Icet \ hich thus de-
19X
celcra te, their desce nt (angle of descent less than due to the enlarged surface area . Mal e. with a slIbgu lar. median voca l sac and nuptial pads. RIll/cop/101'I/1 con~ truct foa m ne. te; which arc attached to yegc1ation above nowi ng {I? IIIIKlllimsfri.\. (1'W/()f! III1c1aflt.I·. KlIIllli) or stagnant water bodies (all other specIe .). sometime. nex t to water. Tadpole. are free-s\ im ming and a ti ve ly feeding ; mOllth w ntral. with jaw beaks and lahi al teeth rows; eyes dorso lateral; spiracle slIlistral ; anal tube median or lateral. There are t\vo bas ical ly different morphot ypes of I?hacophol'/ls larvae on Borneo : type I is il. soc iat ed with stream microhabitat s (m: lb. c, d. c. g) and has cup-like oral disks. while ty pe II dwc ll s stagnant watcr pools or water-liled hole ' in phytothelmes (111 : 2b-g) . Bomean spcc ies occur in all trata of prima ry and old secondary forests . A total of 14 spec ie. are known from Borneo, 10 from Mt. Kinahalu . ~5° )
R hacophori dac
Preliminary key to the Bornean species of Rhacophorus (part ly after
I N(iER
&
STUEB I:-IG
1997)
Webbing of foo t green; a dermal projecti on above t he an us ..... ........... ................................ dlllifetlsis • Webb lng of foot at least parti ally bl ae k ........... 2
9 Numerous conical projections on back head, anns. and leg .... ... .... ...... .... ....... . evereffi lIIacroscelis
2 Body green ... ..... ........ ...... ...... ....... .... .................. 3 • Body not green .. ......... ....... ......... ... .... .......... ....... 6
10 White spot(s) below eye ............................... II
3 Outer fingers webbed for onl y 113; SV L < 25 mm
II Upper eyelid with dermal spike .............. galllli
....... .. ............ .. ... .. ....... ........ ...... .... .. ............. kajau
• No such dennal spike ........ .. ..... .. .... .. .... ... ...... 12
• Outer fingers webbed for more than 1/2: SVL > 30 mm ..................... ................. ..... ............... ....... 4
12 Flanks, gro in. and inner sides of thi ghs with blue
4 Outer fingers fu lly webbed ..... ............ ....... ....... . 5
• Without conical projedions .......................... 10 • Without spot(s) ............................................... 13
pots ... ... .. ... .... .... ......... ...... ...... (yufI(Jp"l/(·taIllS • No blue spot s; sides yell ow wit h black or brown spots ................................ .. (lIIglllirostris (part .)
• Outer fingers not full y we bbed .......... ........... .... ... .. .. .. .... ....... .. .................... allg lllirosfris (females)
13 Lower ann and tarsus with wavy fringes of skin
5 A wide sk in fl ap along arm and tarsus; flank s ye llow or greenish ... .. ........ .... .......... lIigropalmaflls
• Without wavy frin ges of skin ..... .... ....... ........ 14
• Flap of skin along aml and tarsus; flank s black with blue and yellow spots ............... r eill\vardlii 6 Webbings of foot orange .... ............... ... ........ .. .... 7 • Webbings of foot not orange .... ................. ... ..... 8
7 Orange network on belly ; forearm with flap of skin along outer edge ....... ... ... ............... pardalis • No such network on belly; no skin fl aps .. rujipes
... ..... ... .. ... ....... ..... ..... ................ appelldiclllaills
14 Inner side of 2",1 fin ger webbed up to di sc .. ...... . .. ... ...... .. .. .... ... .. ........ ... ..... .... ... ..... ... .. /wrrissolli • Webbing smaller ............... ........................ ..... 15 15 . nout pointed, canthus rostralis sharp ... ... .. .... ... ........................ ... ... ............. al/gllliroslri~ (part .)
• Snout blunt, without sharp ridge ....... .Ii/sciaills
8 A pointed projection on heel ......... .. ... balliet/sis • No such projection .... .. ......... ...... .... .... ..... .... ...... 9
Tab. 20 Di slribution of R1wcol'liortls-s pecie s 0 11 Borneo. Kinabalu
Saha h
1927 Rh(/ c{Jphllrll.~ appelliliflllatlls (GlNrt ll'R, 18S!» Rlracop/wrlH ba//leIlJi.~ \'I{,I II, 1954 !!..!!.!!.coplu)r/ls cyltlwplllldatmi M ,,~ III I'Y & STi IOI. 1998 ~!!JI!.lIJ rtlS du/i'ellsis BOllll'''{,1 R, 1892 RhacopllOr/lS elleref(i lIIucro.\'nv.\' B OIJI ,,,{it R. I X96 I RftllC(ljIh urtlsj(lscia/t{l' BOlIU'''{,1R. I !l95 RI/{/copltorus g UllI/i (I Mit'R, 1966) RJUlWjJJl/lruJ harrissIJlli 1 ~lil ' R & II \II I . 1960 Rllflcoplwrtls k" 'Olt DIUNU, 1<)83 Rhacophorus lIigropa/mafus Bot I I/\'(,I:R, 1895 Rhamplwrus purt/alis (Jll/\ 1111 R. 1859 Rhllcojllwr!ls reimmrdl i i (Sell l rid I , 1837 ) Rhllcu"lwrlls ru{if} C'S INlit'R, 1<)66 Rlwcopllllrlls IlI/glI/iros,ris
Alii .
• • • • • • • • • • • • •
I
. .
• • • • • • • • • •
I3nttlci
Sarnwak
•
• • • • • • • • • • • • •
~
1
,
• • • • • •
Kalimantan
.•
•- -• •-• -
~
- I--- -
199
Rhacophoriuac
Fig.
LXXX]]
Distribution 0 Rhacophorus-species ~ Mt. Kinab u
N
1
-5km Symbols' type locality Symbols' localities •
< 900 m 900 - 1850 m 1850 - 2750 m 2750 - 3650 m
200
from literature
Symbols: own or unpubliShed observation
>
3650 m 0 R. R. • R. o R.
*
angulirostris appendicuJatus baluensis cyanopunctatus
+
R. e. macroscelis • R. duJitensis o R. gauni o R. harrissoni
•
R. nigropalmatus
* R. pardalis
Rhacophoridac
RllOcopllOrtlS Ull!(lIlirosfris
J\II L,
1927
N//(/r"o/Jhorll., (/Cliliroslri, YI()« }(IAKIJ. 181)Oa : 151
T y pe
loe" lil y: Kina Balu R/II/L'op/IO/'II,' (/Cillim,I'lris HOI II ",I·K I X')-Ia: KI : II A~IT""
Il)()(la: 7): I '" K ·\\II'I'N 192 .'1 : 260: ~IIIII 1931 : 17: I NI,IR 1l)(,6: 2H1. 1'J7R ' JIJ; \1 ·"TIII) 1993 : 13 NIIIU'Oph(}I'II,\ IIIIKII/imslri.l' 1\"1. 1927: -1 5 ('Ub'li lute name for NI/(/cophol'lls l1Cllli'V.I'lri.' ). 1931 : 150: I N(,I H 8:. T" I'NO: -I : \-l AL~MlI" 1991a: 30, 19l):!a: 122; 1",," & Snl·BI"" 19<)2. -II -51 : M AI~"IS 1'J<)3c: 15, I 99-1h . 229; WIN, 199-1 ' 29j 7; Ilml \lANN I')<)5b. 15. \ 1,,1 ~ \1\ s 1995d: 2-15 . 1996b: 2026. 1'J'J(,c: 2~6; 1,,,, R. S I LII U"(, & T" 1996: 363: M AN"" Y & GK""\I \NN 1997 : 131: M Al ~"l.' 1999: 1J- 19: I. ,IKI'I ct "I. 1999; 1";1K ct al. _000 Phi/lIIl1l1,' hillll1c'lI/allls. (not NlwC'rII'/IIII'11' /"II/t/clI/t/11I1 P I· rt-R' . 1~671
M
\I K~lI
s 19H9: 19-1
1I1111('(}p/WI'III 51' . I . \1 \I ,,-IIL I, 1991 [I ' 28 NI/(/(,(}/,/WI'III " B", ;,,1\I ~ \lt l' 199:!a: I:!:!
Di agnosis
A small rhacophorid s pccit.:~ wilh rcla(ivd y short limbs and a shott, pointed snoul : on ly (he ouler IwO lin gc r~ are full y wehhed: upper port ion or iri . bluc .
Fi:,:. 198 Female of R/WI '(}/ J//lJI'/I ,\
I/fI,; beaks black for less 50'%, serrated: tail elongate, tapering to an obtuse ly poinled tip, Iwicl: as long as body: tota l length up to 75 mm ; head-body brown, hel l and posterior half of tail li ghter. F.cology/Et hology Thi s species occurs in primary as well as in di . turbed IOresls. It is restricted to montane zones with records from between 900 and 1800 m a.s.1. T he Illal es call at nighl. formin g groups around potential spawning sites . The larvae ca n be found ill sm all poo ls form ed by accumu lating rainwater on the forest floor (111 : 2b). oncn in the marginal /.one. o f forl:sts . They arc bottom suspension fecdl:rs
Di stribution Rhaco/,/ioJ'lls hall/ellsis is only known from a /(;w
localities in western Sa bah and north-ca stern Sarawak. Mt. Kinaha1u : Bundu Tuhan (1375 m), Marei Parei, Ilcadquartcrs (1500111), k silau (sec also map on page 200 and 204).
RhllcOphorus (}Illll op II II c/a/u.\' M AI\'TIIEY
M \N 1111 \ & S 11101. I'I'IX: 17 Tnw loca lity: " lid lit' hcr Rand de, Khan Snk "I.onal!'ar" (ea 200 111 li . ). l'rO\ .11/ Sural I hall •• ()"'ri~' I'hanol11 . Tha.land"
IIJltlt'opil",-u , { .'{{/ltlfllll/c/a/" ,
I~hilllllfll~ hllllll( iI/a/lls. ~ not RIll/cop/w rit.,. /n mu(' ululIIs P i l i K\.
I Xc, 7 ) I , •• • 19 54 [\
\1 "
Fig. LXXXVII Somgram and o~ci llograills of NlllImpllfJhllllll!/Isi.\ (Gg. Mulu, 1300 111; 9.30 p.m . 120°C)
1'/1"
2 50 . 1'166 ' 3-1-1 tpan) . 1'I7X . J 13:
1983 : 23
1'1 " .\
RIIlIc..'Uplr(lf'IIS him(lcu/lIw, . (not Rlw('op/WriH /Hma t.'lIll1tus PI Ii
;111: The ca ll is a . ingl e deep rattlc consisti ng of 12- 15 pulses at a constant rate of about 20 pu lses per second . T he first 2 pul ses are Vl:ry 'oft; duration o f the rattl e 750-800 m ': dominant ti'eq uency 1-2.7 k I);" ; intl:nsity modulat ion pre en!.
& Sl FIOF, 1998
.,.1 8( 7)
I N,.I· ••
SI.
"''' N
&
T \N
1996. 3M
Dia gnosis A brown. mall tre~ frog with a slender hod y, long limbs ,uld hU'ge eyes: one or more whi le spots below eye. si milar to R. ga l/Hi. The on ly s pec ie s o r R/i{/('(}plrol'/l ' in the ' unda region with hlue spots on flanks. on po. terior and anterior filccs of thighs, on underside of tibia, and on axillae; no demlal projections; the tibio-tarsal joint reac h\.O~ beyond tip or snout. Ocscri pI io n Males up to 35 mill . females up to 43 111111 : T LlSV L up to 0.63; head as wide as long; snout pointed. 'Ioping backwa rd to mouth in profile: canthus rostral is sharp. curved; lores not concave: tympa-
10.40
10.60
10.80
11 .00
1120
11.40 sec
Fig.
20~
p /lI1 e /lIl// ., .
!! o lotype (fema le) of 1?/lfIcopho l'lls
----.
('I'lI/lO -
.....
kHz
sec
':1'------.
t, t':UlIH,h
207
Rhacophoridae
-------------------------------------
Iiolotypc (female) of IUwC/lI,hon"
CI '(/1/0IJIIl/cllIll/\'
num half eyc di;)mcter: vomeri nc teeth absent ; li nger webbi ng I( I). ::?( 1/0). 3( 1.5/0). 4(0): toe webhing. 1(0). 2(0/0). 3(0 10), 4( 1/0,5 ). 5(0):;\tl ov;)1 inner mctatnrsal tu berck, no outcr one; ski n or dorsum , throat. lin d chcst ~Jll()ot h . be ll y gra nu lar: sup ra tympanic I'o ld curved, bet wee n eyc and axilla; a 11(\1'1'(\\\ dc rma l fringe along ouler edge of both 4'" fingcr and the 5'11 toe: white tubercles o n vc nt ; nuptial pads absl:nt. Li gh t brown abovc , wit h a narrow dark inter-orbital bar; dark markin),:s (orten as crossba r. ) on dorsum ; si des of body and head helovv lim: ca nthus rostrali s - cyc - .'upr:1t ympanic fo ld dark hrown : blu spots o n blackish brown g round (see diagnos is ahove) ; undcrsides whi ti "h. onen dusted with brown ; bcll y y ellowi ~ h : iris go ld en brO\\n . The
20X
from
olllh Thailand .
do rsa l patt ern is highly infl uenced in its intcnsity by thc time or the da y and the rrevai ling mood. The white spots bc low thc cye and blue spot s vary very muc h w ith thc individual. Ta dpoles: Thc lanae have not been identifi ed with certainly ( I MI LI{ & S11n:HI'J(; 1')97). but presumably belong to the morpllllt ype I. Eco lo g~' /E th o l ogy
Rlw('()" hol'lls t'\ '{/1/0PII I/ C III I 11,\' occu r in lowland p.-illlary fo resls. up to an altitude afoOO Ill. but arc ac tual ly rare above 300 m. Males ca ll at ni ght, 1'1'0 111 vegetation (,0-2.5 0 Clll ab vc the ground ncar small streams. Larvae presumably liv ing in forest streams with !I1oderate to ~trong currc nt s (m : I a. b, l:).
Rhacophoridac
-----------------------------------------------------------------------Fig. LXXXVIII Sonagra lll ami osc dlll[(rallllli"Rftllcopftu(TUI/O/JIII/ CIa/II.\" (Gu llun g Lawil. 250 Ill : lJ .OO p.m.)
rus
i-
I
I-
_1._1
14 0 :.ec
II
I
~
Ca ll : (+2 1°(,) The ca ll is co mposed o f 2 short, pulsed ra ttles. sqKlraled by an interva l of about I second ; th t: fir~ t signa l (duration 0 illS) consists orJ-4 pulses. the . econd (duration 110 ms) of2-3 pu lses: dominant freqllency 2.3-4 .1 kl-l/.. Oistribu tion Thi s species is known from the Malay Pe nin ~u l a, and from northern Sumatra and Romeo. :\'It. Kinabalu: alon l! Sg. Mamut . Sg. Kipungit (500-600111) . Fig. 2 11 II rair
The record fro m " Lumu LUl1lll (1600 m)" by I\til' R (1966, 1971\) is probably hased on a misidentilieation .
01'
Rlw('{)p/tOnt"
dlill/('II"is III
Rlruc'oplrol'lls dlllite"s;s B OUU-N(jLR, I 92 1I/l(Icol'/wrt/ \ clllill(,II.H.I III II I I Mol R. I X1J2 . S07
M I.
Dutil.
T~' pc loca lit~' :
B orn~o
Rhaco/J/IOI'II' "1I1i1<'II.I'I' , \V""", ,
t<)1)4 . 29-.\7: "(.1Re I ti l. lOOO
Di a~ n() s i s
A small tree frog with a sharply pointed snout , a slender hody, and slim Iimbs; finger ' almost fully webbed; a whitt: dermal ridge along ouler edge' of arnl anu tar. us, respective ly: small con ical or round skin nap on heel; transve rse, Iwo- Iobed skin fold above anus ; no suprat ym panic fold . Description -1ales up to 36 min, remah.:!> up to 50 111m; TL/SVL 0,47-0,49; head longer than wide: ca nthu s rosl ralis . harp: lores verti cal, not concave; tym panulll 2/_ eye diameter; vomer ine teeth presen t; finger
ampicxus from Dcrama\..ol.
abah .
209
Rh
Ecology/Ethology Nearly nothing is known aboul Ihe life history of this specie. , other than it lives in lowland primary forest" and that males form large grOllpS around forest pools. Spawning si te ' arc on the forest floor (m: 2h, e). The tadpoles arc large boltom suspension feeders Call : The cal l is a short , weak trill , simi lar to the song of a cricket. A call analysis is nol described . Di stri buti on This frog is known from lImatra and 130111eo. Mt. Kina balu : There i' only a single record from Sg. Kipungit, see also map on page 200 and 203.
Rillicophorlls everetti macrosceLis BOlJl.F.NGI, R, 1896 Rlwc(Jl' hOl'/I\ /11Il CI'O,R'eli, 1:1", II "01'1(, I X9ba : 40) T~' I)~ IOf'l lit}: ·10llnl KlIla 13~lll. N. Borneo Rlw('()plwI'l/s IIIllcm.l('e/t.,. KAMPI' 1923: 252: MIIII 1931 : J I 1'lIillllllll' 'f)l r ulll/lI' S\IIT11 . 1~31 : 20 T) I)~ luca li l): K~nokok. MOllil1 Killa Balli. I'\on h Borneo (syn . ~rter I ~(;~" t')(,(,) N"(/c"pl",I'w, sp iculaill' . 1",11( 1954a: ~51 fVwcophonlS ,"'(,I'"lIi I/w cm.\·cdi ,. 1",,1I( 1966: 2'!t>. 1975: 3 13: M AN 1111 \ 19R_: _3: M Al ~\I'" I'IX : X. 19~6 : 3.1987: 2!l2, I'!KK.: : 176. 19l<') 10)2. 1991 a. 28. 19<)2a: I IX, 1994b: 22~, t '1'}~c : X7 : lI oFH1M" 1995 b: 15: M "L ~Ml' I 996c : 2X(, /IIw ('()phoI'lM (,\ '('1"('/1 i. h ',1K & S 11'1111", 1<)92 : ~ I -5 I : IN( d· R,
''I'
/(;'fU 'ol l l lOn/l tll/li/ellsls
frum DanuJl1 Valle
with its foam nest. wehbin> 1(1),2(1/0), }(0 /0), 4(0) ; toes fully webbed up to di . ks: an ova l inner mctataral tubercle, no outer one: ~kin above smooth, granular belo\ : for cutalll:ous appendages see diagnosis above. Bright jade-green above with a brownish tinge down middle of the back, and tiny white spots: a ye llowish canthal stripe ; upper eyelids and snout brown : undersides whitish green: webbinn pa le green, some spec.imen. with reddish brown spots on webbing between ollter two toes: nuptial pads yellowish . The hones of the limbs are green and visible through the tisslIe from below. Tadpoles: The larvae belonging to the morphotype II: labial teeth rows formula 6( 2-o) /3( 1l: beaks blacks in their marginal thirds, finely serrated; tail a little less than twice body length: total length up to 50 111m: head-body and anterior halfoftai l greyish brown ; under 'ides whitish: a wavy, vertical , black lim: on mid-line of tail.
2 10
'TlI rJlNl,
69:
& T"
L .\ ~I\I
1~96 : J6~ :
d al. t<)<)I): 1"01
INt.! R & IAN It
191)(,a: 567, I996b:
cl al. 2(100
Taxonomic rcma rks: Based upon morpho logical di ITerence. , J'JGFR (1966) considered the Bornean specimens a 'cparate subspecies, R. e. l/1a(Tosce/is. thus restricting the nominate subspccie ' 10 the Philippine island of Palawan. The present subspecies has longer tubercle, a wider inter-orhitallllC:lrking, and a slightly mailer tympanum . In contrast to R. (:'. /'l'(:'r(:' lfi . it possesses tubercles on the dorsa l surfaces of the head and hody. Diagnosis A sma ll to mediulll si:led, stocky, monta ne tree frog
with slender, long leg ' and a short blunt , nout: fingl.:r · \\lith n1diments of webbings confined to bases: many pointed tubercle ' on head and neck ; cone-
R hacophoridae
-----------------------------------------------------------
A b ove: F il:. 2 13 Rhu('()phofl/,\ (' 1'1' 1'('/1; IIIUI 'm Sg" KCllIallli , Sayap, ()OO m.
,\'('C'/;,\'
rrOIll
shaped dt:1l11al projections along outer edge of foreand leg. respectively; fellla!t-s with larger, spiculate dermal appendagt:s,
;H1n
Description Maks lip to 32 mm, females up to 55 1111ll ; TLlSVL 0,44-0 ,6l:\ ; head wider than long; canthus rostra l is sharp. curved; lores com;ave; tympanum hal r eye diameter; vomerine teeth present: webbing of toes I (I), 2( 1/0.5). 3( 1.5/0.5), 4( 1-1 .511 -1 .5),5(0); n small inner mctalnr ''II tubcrcle. no outer one; 'kin of the back shagreened with wh ite-pointed tubercles on forehead and eyelid~ . foreamls . and on tarsi; nanks and vent ra l sides coarsely granu lar: some white tubercles below vent; nuptial pad. yellowish . The coloration is significantly innuencec\ by exc it t:ll1ent and exogenous factors and as a conse-
I{ig hl fr om tOI) 10 baSl': Rflll('opfw/'1I.\' 1'. 1II1I1'I'OSce/i,\' Fil:. 2 14 fcmalc frolll Sg. I.iwa go. 1450 Ill. Fi g, 2 15 JlIvl!nill! from Sg. Silau- Silall. 1500 m.
211
Rhacllphoridae
1Ilg.- pla..:e. Their diet illcilldl:~ insects (m(lth~. nie~. gra""liopp..:r,,) and their lanae. a'i \\'1.'11 as "P' deI's. Parts or a prey itcm thaI illa l' prolrude laterally I'rom Ihe nl
Fig. Z I (, h :ll1ak of NIl/it 1I/liI",.I/' fnllll
Sl! Ll\v:ll!(I. 14511
, "', '1(,((/
1IIt/( {'(J\n'"''
111 .
4u(;nc(; i, 11Ighl y variahk:. Relatl\'ely cOII,tanl traits arc an inter-orhital hal'. ;1 largc \:v-s haped to Irapc/oid llIark illg on the OCCiput. and dark erosshars on the hind limh~ . Lppcr s ide or body spoiled in a \ ariety Ill' ~hade'i of bro\\ n. lighter laterally : grOin regioll and inner :-ides or thigl" bright )'e llo\\ wilh mocha-hfll\\ Il 'ipllb; 10\\ er ~ide~ or bod, icmon:ellllw. ll(;ca:-wila ll y peppered \\ ith browil dllh: in"des or mouth and tllll gill: gn.:(;n. E.'(posed to d3ylight ~()IlIC orthe rrog~ change eolllLlrto a whitIsh gree n \\ ith a metallic I1l"0n/e sheell . h-i~ whitish yd low wilh a lille brown m:t\\ork . Femalc>. green abmi.' . \\ nh a largi.'. W-shaped . hlaek dorsal lIIarking ; Ilank~ li ght Wi.'ell 1\ Ith black 'Pol'i and no ye llow inguillal n:gillll: 10\l'cr silk~ ydIO\\.. I he 'iUrl:lce ~1I'lIctlire ant! the coloration award perfect ea muu flage III the rrog'i. e"pec la II y when pen:hed on a ~urilice \\ hich i~ 0\ ergnm n hy Iidlens (lichen mimesi,) Ta dll o l c~ pn.:,umably ~tream-d\\elling ant! t,l' lllorplwl)'pc I. EC4I ICl g ~ /E lh(ll ogy
Thi s I'rog is:ln inhahitant ormonlane primary rores1s at elcvatlOns between 9()() and I X-lll Ill . It i ~ nocturnal. The lIlale 'i call ill colonie'i eslablished in Ihe Vicilllt)' or small ~treams perched Oil hranches and leaves or hu s he~ ant! 'ihrubs 150 to 6()O e m ahove the g round . rhelr l:all al:ti\ ity ~tart~ otiell herore thc Ollset or du~k (rrom 5 ..,0 p.m.) ir it ralll:- . Specimell~ k..:pl in a terrariulll alway~ returned t'WIlI Iheir nightly e:\eUr'ilI)II~ to their I'\:~ I -
C all : '1he noll' ,erie!' COIISI"t~ ot' a ra~plllg sound \\'hlch IS repeat cd 1-1 0 tlllle'i at l:ontlnuall) inl:l'casillg \01i11l1": TillS ,eries is ol:casionally 1()lIowed h) a ~lia rp "Iii di di dl dit" . /\ call allall'si, doe:- not e:\ISl. Distrihution Thi, suh,peeics IS only known from a rew loea litie~ in western Sabah. north-i.'a,lcl'II Sarawak. and Urunci . It. h:in a h a lu : K..:nllkok . Marei Parei . Kiall . Bundu -ruhan. Sayap: Sg. Kemanti'i ( loon 111). COlllillon III Ih..: region hetwcell Sg. Silau-Si lau and Li\\ago illelu(lillg the tribulari..:s (I-lO()-16:'iO m. Sg. Tihah,lr up III IX()O m) . .'g. k"ilau (up to IX-l() Ill). 'iee abo maps Oil page 200 and 20-l.
RllflcopllOrtis /Jtllllti PllIllIlII/I\ g, lllllll
I{"cr.
I-."r"
Ilh ,,, "ld,rll/' l
h (1I It. 1")(1(,
r)
1)('
11H:~di ty : (\kUl:!ioll,;.!
1),,1'" I. S; \I;)"
I M .l H ,,'I... ~1I 'fH I ",
~ ' I II IIN t . ..\'
"4(1
( I Nti l R, 1<)6())
I "
IN'" ~ I'IX'i XI : I" ,,,
K
& 1 \" 11)1)(J
IY\)2 --1-1<'1 \\1 )'" 1(}'J.1
194(,
_t(,~ .
(\1 \ '
" '111'- \l)l)(,l.:
(I
~()-, 7 : I Nl.l U.
2N7, I
' ~, I \ I 1:1
.11 1'1'1'1
Diag nos is ,\ sma ll. bfll\\n. slender lowland specie, wilh n \'el')' short s nout ; conical pf().Iecllon~ on heel and nn upper e)'..:lids. respecti\ ely; outl'r fin ger alllHl'it I'ully \\'ebbed: a \\'hlte ~pot helOl\ Inrgc I.·yes; t ibio-t:lrsnl jOllll IIl)t re;lChing heyo nd eye: nuptial pads nb~ent. D l' ~criptio ll Male~ up III 32. fl;l11ak ~ up to :IX 111111; TLi . VI. O.51-0.5X: head cqunl in lenglh and \\ idth; ~nout hmn!!l y nllilldet!: canthu~ r,)strali s curved. lore~ ~lo pll1 g: tympanull1 Id eyc dialllcler: \. 1111erine teelh ahsent : finl:!er \\ehbing I ( 1-1 .5). 2( 1-I.5iO). ,(01 0.5).4(0-0.5): toe \\'ebbing 1(0). 2(O·()). l(()/()).
Rhacopllt)ri
-------------------------
4( Ii i). 5(0): an oval inm.:r Illetatar~al tuhercle. no ollter one: skin smooth ahove; a pointed. conica l tubercle 011 uppe r eyelid and another one on heel : 3-.t small projection s between heel and base of" outer toe: curved. weakly e"pressed su prat ympani c fl)ld ; throat smooth . chcst and bell y coarsely granular. Upper sides of"the hody brown \\ ith largc dark ~pot~ and transverse bands: crosshars on arm~ and legs; groin and rear or thighs yellow 10 reddi sh orange : ve nt ral sides whitL:; iris goltkn browll . lighter on margins . Tadpoles: The larvae belong to thL: Illorphotype I. I.abial teeth rows forilluia .tn -ell!]( I): heab \1·haped . heavy. black . coar"ely ~erralcd; tail convex, tapering graduall y to a pointL'd tip: lota l length 25 111m : head-bod y portion ye llowi s h hnnvn dorsally and interally, unpi grnL:ll led vcntrally: cau-
Fi!!. 218 Vcntral pancrn of NiI/wIII'/llm/., gellllll lrolll Sg.
:z:
o
K ipul1 gi l. 600 111 .
21~
RllacuplH)ndae
dal l11u~ck and dUI al lin \\'1111 largc dark ~p(l I S: vc nl ral lin pl g lllcl1tk~~ . Illthcation, an: Ihallhe ladpole, urtlll~ :-'PCCIC:-' haw dc\dllpcd a hlllHldal Il:cdlllg ~Iratcg.y a, Ihere arc non-rn :ding a, \\ell as e\otrnphic lan ae ( I ~( " I( I \)92a). 1'\' f) l u~~' / Et 11Il I u~~' gill/IIi i~ n:stric i 'd t(1 prll11ary 1'01'In lo\\' lan d and hill ) re ~lO lI :-' t 150-')50 nl a ,~ . I.) , AI lIi ~hl, III :dcs an: rO lllld lin \cge lalilln (.Hl-liOD Clll ahove Ihe groundl ()\'erhanging rirl'k s Ill' rocky ~lreal11:-' , Inrl11lng loo"e callillg grnllp~ . v\'e I'(\ \l nd ~ p ccimen, (II' thl" , p ccie~ in Ihe IIlId st or a call1llg COl11l11l1ll1ty or Pltillll/iI/S IIl1n llllllllll II I a h: II11l wo-ra tlan llI i\ed r()rC~1. ,01111' .\ ()() II I away ('rom the ncare,t "tream ill the I an ga nan va ll cy, The ,, 111 a II rllaIllIH;" t i" attached to kave:-, (II' \,egela t ion (1\ cr han gi ng i'lHe~t :-.t rea Ill :-. , The larvac livc amollg gra\e l a nd r
Rlll/c(lelllJ/'i/l
c:-.I~
Ca ll : ... inglc high-pitched chirp. f\ ca ll analys i, I, not yel descllhcd . Ili strihutiun gill/II I 1:-. on ly kno\\'n i'rol11 arc\\' loca liti c" III Sarawak and Sahah. !\'It. Kill a ha lu : Sg, Kinatekl, Porill g reg ioll : Sg , Mantukungan. Sg. Kipullgil I :lI1d II (500-(1-10111) , Sg. Langa nan (751l-<J50 111). "cc al"o Illap' (III page _00 and 2(>-\ .
NIt (I('olJ//(I/'I/,\
I'i:,:, 2ll)
1
,,/,IIII/,I/.\
//(/"'/"(111/
11 0)11 Sabah
Desc rip liun l\1ales Lip 10 56 Illlll , rc lll alc~ up to 7() 111111 : l'I ,; SV I () . 5~-() . .sX: head IOllge r Iltan \\ ide: can lhll ~ ro~tr:llis ~ lt a r[l, 11Ire~ almll~1 \crtlcal. \\'eak ly eo t)cave: tYI1lpanum llt"tinct , larger Ih a l1 half eye diamder: \\III1CI'lIlC leetlt prc~enl : all lne:-. \\c hb\.:d lI[l to di~k~ : derma l flap along Ollie I' edgc 1)1' I " loe: a small. ()\allnIlCl' Illclalar... alll1herrlc , Ihl olltcr one: dor,al ...kin smoolh and finl'l) j1,'bhlcd: ~ upralyt)lpat)ic [,lid ~ I ralghl, I'rnm '!c In IYl1lpal1ulll :Ind ~hou ld cr: IIndCI ~ i de' gra l1ul :lr. Uack and upper ~i d c ()I' hcad rcddi~1t hrowl1 : dark rm~:-.har~ 0 11 limbs: :-.Itlc or head sometimcs wilh white :-. p o t ~: \ cillrUIll white. \\ ith dark "pol:-.: I1Uptlal pa d ~ )elltm i, lt : If" lighl 111'\1\\n in Lipper half. dark relitlish bW\\'1l 111 lowcr.
R /1(J (,Op/lOrti S /w rris,wJII i II\'( oI ,t{ & H .\II t. 1959 /(lUII "1,/lflllI\'
tllcali., : " I 1)1\' hIOI1
hcUlt\ ,"1I11
III
1' 4. 1 k
& II \ II I . 1\))4..)- 270 I ~ pl' "I Ih,' B .. ralll Rl\a, I Pllnh
p," ,lh ,
~,II . I\\.I"' ''
IOw( U/'''Oll/\ 11.01/.\\0111 , I 'H, I ~ & ~I II IIU ' l . 1·'·'2 ... 1 11 . 1 ,-: " I III\(, & I ' oJ 1')')1> 161 1" ,,111 \' 1 ,tI 201111
(,lit.
J)i a~ lI osis
moderately large, brown trcc rrog wilh a trian gul:lr Itcild and :111 ob lu ~t.:ly, poinled, ~ha r[l -eclged "nOLlI: thrcc oulcr fingcr~ rully wehbcd: webbil1g betwecn [ingel'~ ilnd loe' brown or black : no dermal projcc (i o ll ~ ()Jl l imb~,
1\
2 14
~
., ;;, " o
(!l
aD
RhaeophnridaL: ----
lingers and to.:;; full y wchhed ; w.:hbin gs black . YL: II()\\- towa rd s tip" of tocs: broad s kin f1ap ~ on forearm , tars LIS , and heel.
.)
'1.,
{'to
• .,. ..".1
.......... , .. ..,.,-. ~ ...,; .•"
r
. . ..~:."",_...c:.
":":" ,
~
.\ ..
\,;o'"'!'.....
,~.",:•
..
,,'
':"' ~" .:"
... .:. •• ,;,J .... !
....;
" ...:::$:
--::.--...:-:t
A MdlklllU5
"il.!. LXXXIX Oral t1 ist: o f lI/w(II/,//(/ntS horn"·(llii.
Tadpoll'S: The larvac hclong to the l11orphol)'pC II . Labial tceth ro\I'~ formula -I-5( }- -I ur 5) . 3: beak lincl y serra ted : tail :-.Iightly Cl111ve,\. tapering ncar end to n
IiI es III pri mar) and \l Id secondary t()re :,; t ~ covering hill y terrain" at low eIcvatiolls. Vcry lillie i" "nown about it s lire hi story. Spee imcns II ere I!.ltlnd in trcc holcs, alllon g ep iphytcs, or o n the Iea\'c" of legeIation , 30- 1}()() cm allllVe the ground . Foam nc"t!': arc attached tll thc har" ahov.: watnlill ed liTe hole", tree trunks. or water tank" limllcd by anastomosing hu tlrcs~es. 1--1111 abll\c th.: gn ,untl I?IIlICIJ/ I /illrt/\ 1J(lIn.\·.wm
(m : 2c,
I~
Ocscription ,talc" lip to ()O 111111. feI11al " up tu Ion n1ln ; Ilj . VL 0.5-1 - 0 . 5~: head as wide as Inllg or s li ghtl y broader; .:allth us rostralis di ~ lin .:1. 10rL:~ sloping, a little .:onelll'C; tympanuIIl halI' eye diameter: inn.:r edge or I " ling.:r and toc , and ollt.:r ed ge ~ or -I '" linger and )'1, toC, lIith f1ap ~ of skin ; an ova l inncr, \1(1 outer mctatarsal tu berclc: S"In aho".: fine ly pL:hblcd , gran ular bGlow, c'\ccpt 0 11 th.: throat ; no s lipra tympallic fold . F.1l1.:ral d gree n abovc , orten ,,"ilh tiny whitc spots; flank s ycllo\1 at inSL:rt io n, of arm and leg; unders idcs \\'hiti , h, with a yc ll o\\' i ~ h ting.: on chest and bell y: iri~ pa le gol d':ll ; nllplial pad s ye ll(lwi~h . Tadpoles: The lanae beloll g tn the Illorphot ype II . [.abial k.:th rows li)J"fllll la 4 - (1( 1--1. 5. or (1)i 2 ( I) or 3-7(2-3, -I . :\ 6, or 7 )i 3( I) or (,(.1-6)/.1 ( I) : bcaks linely ~.: rrated, black in thclr marginal thirds : lail Iellgth nL:ari y twice hody length ; head-body pale grey aholc, while heltl\\'; ta il grey with dark ,pots. El'ol"l!y/ E thol():.:~· Rhll("()/J/wJ"//\ 1I/~ r(l/}(t1I11(/fll\ I" a puurl y kllown low-
land ~ p.:ei '~ occurring li'OIll ~e a 1e1'1:! up to SO() 111 al titude. II d"ells the ca nopy of prllll
)-:) . Fig. 22 1 II/r//( ,,/,/r"1"1/ 1 m g w,Il'/III,'//(\ Inlill Sabah .
L>istrihutiull This Ii'llg is widel y d I ~tn hut ed in all parI~ llf Umlll' O. 1\11. Kinahalu: Knlll\ll onl y 1"0111 the Poring rcgion (Sg. Kipungit). se.: al,o maps on pag.: 2()O and 2 ()~.
R/1lIcopilorlis lIigropa/lIllltlls Botll I' Nit ,/( 1I1'/JOI'lI\ 1,,,' ,,lil~ ' :
(,L' R,
I X95
1It.~J'OIJttlmtl{lI ' B III II t '( ,I Ii,
,. \k.lr
R",.,.··
I XI) "
t 70
I ~ pt'
Diagn"si~
;\ grccn trc.: I'm!;!, \ery large h~ C() 111P;Jr1"1lIl, \\" lth lon g, ~ lelllkr limh~ alld larg..: hand ~ and !'cet: all
H /IlICop/lOrlis pardalis (; [ I I
Ultf/. 'tl/ '/UWU, I'dldt/h" 11 ) 1)1
"I,) ~?
II I)II
'111/ 1(,
I X59
I X,:\t) ,I ('\.:;x , :-n 1'~lh' I .1" Judi !..:, ( ,l it & "', IBI''ljI, IlJ\)_ -11- ." 1. \\ It'\l. l~'(J'::;" I ':; hl'/l'l'l.ti 2000
( 1I ',\ I I II Il.
H O lI ll'n. illld
"
1.\"
niagll(l~i~
i\ ~ ll1alllo lll.:dIUIll-:; IZl'd. ~ klltlcr tll robust. hrowil
Ir.:.: rrog \\ ith I,mg 1':1;' ; 111.: l>lltel tllr.:.: lin g .:r~ arc I'ully \\ .:hl)(:II ; \\ ' ebbll1g~ on hallds alld Ic et ol':lngercd : an orange n.:t\\(lr" IlI1 thl' Iwll y. D('~cript
inn \1 :lIc, up to 55 Illill . klllak ~ lip to 75 nllll; II . . SVL 0 .... 7-0 .... ') ; head Illn gl'r than wid.:, snOUI rounded ; canthll ~ ros lrall s di stilll'l ; lure ... ~klpill g. \\ca"ly Cllllt:aw : t)'mp
Fi:,:.2ZJ NIIlI('o/ ,lu)/'II ' /h l relali., IWIll
P ' )I'II I).! ,
550
111
Fil!. 222 NiI" I",,/ ,il lIJ"/ 1\ 11I!!.m/ llI III/CI/II' I'ruill Sg I-:qllll1Ill , 1'<>1 III ).!
).!II I, 5:-(J
~lIlJ 11 " ",,, II (1()(lOh) \\.:re the lip,! tll d.:scnh.: th.: g liding I1l g hl ut" thl ~ rro),!.. and h (,II{ II ()66) oh~.:nnl : "Olle li'og wa:o- thrown Into Ille :Iir li'om a h.:ight or li\e l1l.:h.: r~ ab,)\c 1:! l\llllld . It Illllllcdi atcl y nghted 11:-..:11' and lIeld th.: halld~ alld l"cd ill the plane or the: hody wllh the diglb wltIc ly spread": ~cc also 1 ·. ~ II · I{"" t\ & K(}IIII (11)l)()). fill' large 1,)al11 IlC"h ;Irc attached to low vegetatlOIl nrdep""ikd Illl the muddy hall"~ llrSlllall pools alld \qtllo\\·... or rhllHl~ alld wi Id boars Oil the ror '~I 110m (Ill : 2h, e) . I Ill' larvae Dlk ll dl'n; lop 111 l'X trclllc l: Illllr"y. turbid \\:llcr (1 \1'01 I{ & Sl lI lllI (; I ()'n) alld arc hottolll ~ ("'pell ~ i()1l Iccdl'rs.
Disll'ihutiun l'vlala)' Pcn i ll ~ lIla , Sumatra, alld Bornco. Mt. h:inabalu: Tllere: '" oill y OIlL' record rrom 'g. Kipungi! I 150\) Ill).
r
..."£o r
..;)
R hacoph ori dne
melatarsa l 1l.lberele. no ou ier o ne: dorsa l skin shag reen ed. coarsely gra nular he low (smooth on th roa t or I cmalc~J: derlllal flap s along ouler edges or forearm and hnnd and o n UlI'SLlS. round nap 011 heel: ~l lpral Y lllpani c fol d low. curved. Colora l illll ve ry va ri ab le: upper s ides tan to reddi~h hro\\n with dark Illnrkings. o nen irregular while. blue. red . or yellow spo ts o n bnek ( r le gs; lim bs w ith dar k cros~bars; flan ks ye ll ow w ilh blaek !l ec k;:: c hes t and helly pale ye ll uw wi th an orange ne twork; throat whitish: iris li ght gu lden . Tadpoles: The larvae be lo ng to the morphot ype II . I.abia l tee th row~ formu la ()(3-() )/3 or 7(3- 7 )13 : !ni l tnpcring to a lIarrow tip: tail lt:ngth ~I i g htl y less Iha n Iwice body le ngth ; total le ngth li p to "(S 111111 ; i1 ea dhod y a nd tail grey to li g hl bl'llwn . sO lll et imes with seve r;11 da rk "pols; l)nen wilh on ly one spot 011 side llf head. Eco lo~~' 1F. 1 hnln!.:y N//(/copllOrt/s !Jare/alis a ppcar~ 10 be a res ident o rthe ca nopy Fi!(. 11.t Rhtlcl!/Jhol'/f\' parr/oli.\ from Sahah . ~ traIUIll . It can be fou nd in pri Illary and ~ecn llLlar v rore~ t s and i~ rathe r common along logging road~ . It i~ Ca ll: ~ h o rt . raspy chllck le. /\ c;1I1 , Inal y~i~ is not ahk 10 perro rlll paragli d ing fl ighl ':: hy sprc
217
Gymnophiona ( 'aeci lian~ arc pantropit:aL elongate, lilllhk~ ~ ;Ilnpill bians ofa worm- likc appearance . They lat:k sternal and pelvIC !:!irdlc~, and the t:o luille li a is either Illa~sive or absent. The body i~ ~egmentl:d by annu lar gro(\VC~ IVllh the IlIll11ber of annu li corresponding to that oflhe \erlebrae (up 10 J OO). In the more pri lni tl\t: gencra thc lown la ye rs \)1' the "kin ellcasc ossilit:d ·cales. The tail. if pre"elll , is very short and pointed. ('aecilia ll ~ arc adapkd 10 drill -burrowing with thei r he;1\ ill' o"sified ~kulb . For th.' samc reaso ll Ihe cyes an: rudllllt:ntary and obscure. altlwllgh Ihey sti ll posscss a Icns and a retina wilh photon:c..:ptors which can distingllish helween lighl and dark . Between the eye :lntl the 1l0sIIil Ihcn: i~ a protrusible sClhory tcil iade . The tcdh arc cuned, present Oil pre lila xII lat:. maxillopalatincs, \omers, dentaries, anti sp lenials. In C(lntrasl to all mher verlebrate" which p o~sess only a single pair of jaw 11 11"t:ic,;, caeci lian s hm'e anolher, "econd pair. LUllg~ arc prc"en t (in e"cept or :1(1'('/(1("//(/(///(/) I\ith the len lung u"ually hcing rudimentary. Caecllian~ rcprudllcc by int..:mal fertilization for \I hich the males arc equi ppcd with a slIlglc. mcdiall , protrll"ihle p..:nl~ (phallodeulll). I-ema le:-. either lay a cllItch or ..:ggs or gi\c birth to li\.e you llg.s. In Iht: case Ill' egg- laving speciL", the femaic guard~ her clu lch unti l it hatches . The larvae arc am phibi c wit h gi ll slit s (spiraclc) alld labial lobe:--. rhey li\'c in s hall ow water and reed al night. Caccil l an~ arc carnivorous. While nUlllerou~ worb e .\i~t O il ~y~lematic-taxonomic and anatomic quc~li()II~. (lilly v..:ry little is known on the n:tlural hi~tory tIl' the s..:creti\e gY l11nophinniall!'> 1\ hich arc also llllly rarel) kcpt in captlvily. Horneo i~ hOllle to onc 1;1 111 lI y. Ihe Ichthyophiidac.
Ichthyophiidae This lill1lil y i!'> v. .·idespread throughout SOll thl:a!'>1 '\ !'>ia ranging from Indi a and Sri Lanka tn Horneo and the Philippines. It cOlltaln~ Illcdlum-si7ed eaeci lian!'> of up to 'iOO trll11 in Ienglh 1\ ilh poinll:d tails and subterminallllllllt hs. The tentacular openillg" ar..: ~i lll ,lt ed mid-way betwcen thc cye~ and Ihe n ()slri l ~. Their primary annu li arc suhdivlded Into ~t:clllldar, annuli (2- ~ . econc\arie~ pcr primary), both of \\ hlch forlll r he\ rons pointill),' toward thc tail on Ihe vcn tral "Ide. The dermi~ ineludt:~ sma llnssilicd scales. The eggs produced hy Ihc species of Ihis t~lInil y arc vel)' rich in yolk . n1CY an: deposilcd in a stTing in moist "oi l and guarded and defcnded by Ihe Icma lc. Eggs attacked by fungal infl:ctions or v.. hich have perished otherwise ;Ire consLimcd by Iltc fl:lllale . The period of development ti'om oviposition to thc emergence of the Inrvac lak..:s 70-XO days. The larvae lead :.In :.Il11phihlc lilc and are eqUipped with well-developed latera l Ilile organs tncuroma"IS) and Ul11plllbllY organs. While th..: fllrm..:f ~e l"\'e as meehanoreceptors, sellsit iv<.: III wakr eurrcnt" and pres~ures, th..: lalter arc elcctrorec..:ptnrs. The larvae have one or lwo gill slits. and low caudal fins . 1 h..: ICiltacles do not appear until "horliy bcl(lI"I: Ih.: completion of the larval ue\clopmcnt , that is aller 1 2-1 ~ Illonth~ .
Adults live in primary and old secondary rorests ill e:t11h hoi..::;, under Icar litter. or beneat h log.. Their li)od coillprises ; 1I1hl"llpOl L~, earth wnrl1l~, and small vertebrates. rhe life histories nfall ~pec i e~ arc nearly unknown . This lillilily i" repr..:sent..:d on 13orneo by 2 genera (( 'alllla("lwcili{/ and /dllllyoplli\) and 6 spce ies, nn 1\111. Kinabalu by one genus (lelt/It.1"IJ/lltis) alld ont: SpCCICS . These 11\0 !!encra can hardl y be di ~tinguished on the hasis or their e'(l<.:rna l lIlorphologies - ('l/lIlIa /"(/('("i/III 11
2 1X
Ichlhyophiidae
gi ll sli[
/---------------Fig. XC Larva
-
tail fin
------
or Ich,hyophi.l'. m:uromas[s
anlpllilary organs o
0
0.", • • • _ . ; ' _ _
• ••
.~
Fig. XCI Lalcral·lin.: urgans in fclllhrophis.
Genu. lchtltyopltis
FITZIN(I1-.R.
1826
The species or [hi s ge nu. are almO~ 1 as widespread a~ Ihose of the enlin.: family. They exhibil all the above mentioned traits Iypical 1'01' the family. FOllr specics are known [I'om Hornell. one li'om MI. Kinabalu .
Iclttltyopltis l,illh"lIpl,,,
1110110('111'011.'\ BU. LKLR.
/1/1111.,,,1,,·,,,,,
Incality: Slngka\\ang. If-llI/n(JI)/"'" IIUIII()( 'hrtll(\ ,
Il",~, K.
1859
I X5'i (I X5~) : ! xx
Ty,,~
ilnnlC() I N I.! J< 1~)(, 2· hJ
~6.
1<)(,h: 1))
Diagnos is Purpli sh black wi th lilly. diffu se. yel lo\\ spol~ on Ihc throat : tenlack~ at uppcr lip. much closer to the eye than to the [h)stn I.
Dl' SC ri plion TOlal length up to no mm: hend as wide ns body: snoul roundcd: sma ll cye::. visiblc through the ski n: body with 2~7 annuli. Inil with 4-6 annuli : I X-25 maxillary-premaxillary 1l:c[h. IY-~I \'ol11eropalatal leeth , I Y-23 mand[bular leeth. 2-~ ~p l e nial teeth . Lggs: unknown. LarvlIc: heau depre~scd : calldal lin dislinct: lotnl length lip to I ()2 111111 .
ECIl In:,.:y/E t hllln:.:y A forest ~ liecie ~ ranging fi'tllll sea il.:vel up w 6()() altitud '. burrowing beneath the duff of 111g~ . The larval! live in small. ckar ('orcs! ~lrl.:ams .
Oistrihutinn vV ide~ pr(;ad on BOrnl.:ll. ·1t. Kinahalu : 1;1I11l[1 an.:al Ranau Di~tricl : Su. Kipullgii.
Sabah
KlI1aoalu
II ru 11':'
Idllh.1'Ojlhis lII'1i('()lIu/'lv T \\ I "I(. 19(,S Idll /.!X!!l,hi., hillll~lIluri.1 T,," ,\(, Il)(,) fdllh.. I/phis dlilill'lIVi.1
r \ \I
Idlfhl'Of/fti.1 1II/II/Oe/If()I1.1
I)\(.
I l)(,()
IIIIHI K.
lli'il)
I
III
•
•
I
Sa 1';'" uk
Kahmantdn
• • •
•
•
21 ()
Reptiles are marked b a skin covcr,,'<.1 With scale. and ru ually have a t;reeping mode oflocum li n. The conservative systemalic considers the class Reptilia a group of r prc entatlVes of the phylwn Vertebrata or vertebr'ltes. These ar alllmais breathin with lungs whose bl)dy temper tun:s largely J pend Qn the temperature ~~5 ofthclf immediate environments. All onoday's reptile evol cd rom ancestors which onginall po se soo two pairs 0 limbs, ending in cia ed finger. and toes. ano a tail. 11l skin in orporates but, . mall numb r of gland and i, lally covered ith a protective layer 01 homy scales, The number, arrangement, and individual 'hap s of the latter arc;, ithin l'ertrun limits of natural variability, ~~~~?:' ind) ',hve of ~ci and genera. Reptiles reproduce lind r thep-rincip]~ 0 internal ~~~~~~~~~~~ fertihlatio~ but there are a lew species known which multiply parthclloge- ~ n~tie lIy i.e. Without th necessity of amaong 0 h\:o lhffercnt sexc (e.g. vithin the numly ('kkkonidaeJ, m specll~s gIve Otl1h to fully deve[opod young. but the va:,t majority of rc:phlel) lay Telatively large eggs. with hard and calcified or parch-like sh lis which contain substantial amot'JJlti of yolk. As a rule, thcs • are d 'iteu on land The clukh are usually neIther guarded nor actively incuhated """""...~, by the parent '. The young hatch after some lime and. re immediately 'clf'upporting. rh rc are to e ce,';/,; of 6000 specie! ofrcpttlc!), systematically grouped in 4 orders. Borneo i inhabited by rcpltles ofthrec orders, namely Tl)codytia (ero odiles). quamata (scaled r ptile). no cstudincs (turtle, tortoises. ,nd terrapins). ~~,;... Cr()C()dile· arc e ily Identifi as Illey are the urtiy reptile 'who, e teeth are 'Jlsibl wh n the mouth is closed. Equally easy to recogni:re arc tllrtles and tortOISes ~_", _ who e bodles are prote ted by a shell. The latter may be domed or low. and hard or soil t the touch. All the remaining Ieptilcs of B meo belong to the order quamata am.! compn e the li7~lrds and the . nakes.
Squamata
(scaly reptiles)
In contra t to crocodiles. squamat .. all l1ave -n. transversally arran 'ed anal slit , and no teeth are viSible when the mOlflh is c1l ed, Males po 'css a pair of copulation organ. (pene ) of which but one i.' employed for the actual mating. The skin i' covered by n prote tive layer of horny scales which doe, not grow with the animal and ha t b shed trom time to time. The underlying new scale cover then' slightly lar er than the previous on . The hypodermi. (lower layer of skin) c ntains pigment cells whieh detcrmin the colouration and colour pattern of a reptile depending on the type and arrangement of these cells. Thi. rdcT c mpriscs two suborders, named Sauna (IInrds) and erpentes ~~~~~i (snakes), re pc tively. The latter have evolved from the Iimrds and Ulcrefore --~..,., repr 'cntamuch unger stage ofevolution. Lizard ' arc usually, but not genemlly, characterized by our limbs and movable eyelids. ' nake.• on the other hand. do neither have limbs. nor movable eyelids, nor ear openings. Certain fo orial groups bnng about complex problem as to their proper allocation to one group or another. The members ofthe snake family Typhlopidae (blind nakes) fOT example exhibit kull structure which are very similar to those of lizards. while the lizard family Dibamidae (blind skink ), who 'e members do not po e s limbs (male with rudiments) or tympani, hare many traits with the nakes.
Oizards) Lizards have conquered almost aU types of habitat . high up on trees and have devel ped the ability t controlled gliding flight, other dwell on or even in the Several of them are e cellcnt swimmers, and a f~w even avoid the ocean . Lizard" feed mainly on live . some spec ies al so prey upon molluscs , worm , vertebrates, crustaceans, and carrion, or have pecializ<,'(i on a vegetarian die t. Our knowledge a b lit how lizards live in the wild j . till relatively limited . ervations are u ually superficial and restricted to brief moment before a lizard tak to flight and disappear into the layer of leaf litter (like many kinks for example), r da hes up a tree trunk with tremendous peed and thus withdraw ' frOm the uriou eye of an obscrv 'r. th r li7Nd . rely on the camouflage effects ftheir colours and colour pattern which blend in with their surroundings and may not give themselve ' away by mo ements for hOllIS on end. 1"0 locate them in tbe semishade of a forest is extremely dillicult and very much dependent on incidental di. eove ry. Very few lizards are !rue to a locality for longer periods of time 0 thata purposedriven earch is rendered extremely difficult ifnol impo 'ibl . ome 110 sp ics oflizards are repee eated on Borneo whi h are systemati . Ily grouped in eight families and thirty- e n genera. Five 0 thes families and twenty-five genera have been recorded from ML Kinabalu (sec Fig. XU). Usin identification key (page 223), they can eil: ily be al their proper families. onsiderably more dlfficult. is the correct allocallon of represcntatives of the Sc inc idae . where generic trait'> ar not a lwa ys externally, and Gekkonidae, wher indicative traits can sometimes o nly be evaluated with U1e aid of a magnifying g la:.s. This is even more tru when it tht: iOl.'J1tifieation of the individual ·pceic:.;. The latter require. the use of a binocular microscope which U",li"""~ the counting of se'lles etc., but a layman may still be faced w ith ins um10untable problems. As la r a. the agamid arc concerned, Ihe generi c identificatIon i somewhat easier beca use it can be based on relatively welldiscemi hle ex ternal tmils. When it come. to spec ies level , however, anulher ki nd of compl ication may make itsel f felt as fe males often exh ihlt all appearance which is enllrely dissimilar to that o f the males of Ihe same species. A ' the pigmcnt cells e mbedded in the hypodcnn is can alter the colouration and the di tribut ion ofcolollrs to a sometimes sub lantial extent by cha nges ill cell ize and arrangement , colour and patte rn traits have only a limited suitability for idenlifi ati on purposes.
Sauna
Distinguishing features in lizards
~:..-_---
rm, lral - - - - supranasal
1l1cn lal
-:1+:-'
(1I)Icma~al)
~I-t---
rrontona. a I prcfrllJllal fronlal supraoculars
---->,->.;--"'1--
frontoparielal pandal
-J.---';l-- r - -
pO\llllCnlal subm
inlcrpJrietal nuchal
Fig. XCII Head scales
or a sk ink
... <.,upracllt3rtl:S
tcmporal~
postnasal nasal rostral
Fig. XCIIl Toe infra lab ials
tympanum
tail lenglh (TL)
Fig. XCIV Place of measurements in lizards
222
preanal porcs Fig. XCV Anal region of a gecko
auna
Fig. XCV I Rhombic imbricalc .,~ales
of an agamid Ii/art! . hOI1l(\geneou~ in ~ h a p c. size. allt! orientation.
Fig. XCV II InhOlllogencllu, scales of an ag:lll1ld liLard . intcnllixed with larg~r O I1 CS .
Fig. XCV III Uranular scales 01 a gecko. IlItcrmixed with larger roundish tubcrc1l:s .
,upracil lary edge
Illll'hal crest dor,al nest
canthus roMrali, Fig. C SOllle ex ternal morphological charact cr~ of agamid Ii/.ank Fil!. IC Kee led scales of an agalllid lizard.
Key to the families of Bornean lizards
(aftcr :v1 '\'ltll t Y &
(JRO\S\IAr-.i'<
1997)
Eyes visible ........ .. ..... .... .. ....... .. .. .. .... ........ .... .. ... 2 • J-::yes hidden beneath sca les (A) ...... .. Dibamidae 2 Head scales la rge. symmetrical (13 ) ...... ... .. .... .. . 3 • I lead sca lcs small. usua ll y asym metri ca l (C) .... 5 3 La teral fo ld present (D). limbkss ..... Ang uid ae • L<.Ileral fo ld absent. lim bs present or absen t .... 4 4 Scales shin y. tail length r 3 x SVL ... Sci ncidae • Scales dull. tail length ' 4,5 ' SVI. (E) .... .... .. .. .. ........ .... ...... ... .... ......... .... .... .... .......... ... Lacertidae
o
5 l:3ody anti ta il wi th distinct rows of la rge tubercles (F) ....... ....... ... .......... ....... .. ....... .. . I.an thanot idae • l:3ody and l<.Iil without ruws of tuhercles .... ... ... 6 6 l:3ody compressed latera ll y. gula r sac and nucha li dorsal crests usua ll y present (0) ...... Agamidac • l:3ody clmlpressed dorso\,entr<.lll y. no gula r sac or crests .... ......... .. ....................... ............ ........ .. ...... 7 7 Pup il \'ertically ova l (H): if the pupil is round . ling"r and toes are lil ig.rane (I): adult ~peeimens < 40 Clll ill total leng th ...... ....... ...... Gckkon idae • Pupil ro und , fing.ers and tues stmd y (J ); snout and neek co n~pi c lluu sly lo ng (K): adult ~ p ec illl e n~ ' . 100 em in total lengt h ....... .... ............ \ 'aranid<.ll.'
I~ J
K
223
Agamidae I he lli,1I 1111111(111 (II a~,l1l1ld Il/al-Li, I' ~' Ilntined to llil' Old World . I Ile~ ar~' r~' pre,cnlcd il1 '''III IH.:a,lern hm1pc·. un lllu, 1"llhe .\Ii IcallClllllllll'lli. III Sill IIhem and cC'l1lral 1\:-'la ,'" till 11111'1 Ii a:-. .lapan. Ilw C'lltm: InLill\U'-lralJ<1I1 ,\1 cill PC' la ).!" . alld III \u'aralia . 011 HOIIlCi' al1d Ih~' 'UIT(ll111dlllg regllln, agalllld li/ard, arc l'haraCil:lved by laleral l ~ L(llllpl"l:,:-.ed hodle:-. 1\ illl tOllr 1\ l'll-dc'\ elJlped Ilmhs_each 1IIIh IiI e clall'ed linger, allti toes. ThcII' dull scaic:-. dll 1101 I'orm a ,)"11metn ca l Ill' c'k ;1i I) dtllc'lcllll :lled arrallgel1K'lli 1111 thl' Iwad _ Thl' pllpil, are 0111\:1)" round . -I he majuril), Ilf a).!: lI llid:-. I'll" l',' a IlTtchr.1I nc',1 .1I1d III a gul;lI ,a~'_ In ,01Ill' 'pec Ic'. t\:l1lale, dilli:r lillie rl'llml11ale .... ill olliel':-.Ihl· 'e.\e:-. :Ire ,Jl d""ll1l1l<11 111 Illl'il apl'l'ar:l1lel" Illal II I, til tlicuh to hdil·lt· thl') hcllll1!-!, I" Ihl' "line ' IXTIl':-- _ I'hl"c Il/ald:-- .Ill' ; illlllhle'(l' lu,,\cI~ dlllrnal alld 111 1.' mil le !II' Ie", 'Incll~ on trel', alld ,hl'lIl1,,_ ~1aI1~ :-- pecll':-
\,r
"I'
(/'11111/111111 '1',\ "('/,111111/1/1).
Thi S 1;llllil) '" Il'ple"l'llled Oil Il(lrnc(' h~ 1l 11l,' ~CIlCJ'<J ;tlld 27 spl'cics. ",ilh ,('len gencT:! (III." ol'lIfticft ;Irl' IIH"I\II ~ PIC :lI1lIClldc11lIC) and I-I ~ p ecIJ.:~ oCl'lJrlillg (Ill r>. IL KlIlahalli
Key to tlu.' Bo.-nea n gene ra o f Agamida e
(111 part a ft er:--'l 1\J1111)
&
Siucs or hod\' \\ Ith largc. ,pr.:adab lc ~ kill IIlCIIlhralle, alld ncd. lappel:-. (A) __ __ _____ _____ . __ . __ D/'{/('o • Sides orh\ld~ \\ Ilholl i sprcadahk: , kin m.: m brallc~ . __________ ________________ ____ .__ __ _____________ 2 2 Sldcs !lf ft e:!d CIll'L'red 1\ ilh eXl rclllc ly large s hi t'ld~ he low Ihe 1)'ll1pallUtll ( 13 ) ___ ____ _______ IIYI'sh·a/otl'.\· • Sidcs of lIead \\ IthOLl I large ~h i cl u s _______ ____ ____ __ . 3
3 Tympallutll clearl ) \' Isibk: ( .) __ _____________________ __ . 4 • I'ym pall L1 11 I \111 i'lh le ________ .__ __ ____ __ ______ ____ __ ____ ______ . 8
... 1'v1a les 1\'1111 a horn -like prnlrusioll ( D). f'emalc, \\'il h a sma ll hUlll P (1--. ) on Ihe li p or Ihe snout __ __ ._ .. _.. _......... ............. __ .... .. ... ___ _____ . ___ _. Horpe.lollrll.\'
• illlih sc\e, witholl t horn -like p ro l rlls i o n ~ or humps Oil th.: lip or Ihl' ~11\)U t ___ _______ __ . __________ __ __ ___ ____ __ ____ 5 ::; Latera l ~k in 1i.1kb 011 tht' neck prese nt ( F). homoge nco us rhombiC dorsa l:-- (;), TUSV L --.. 3: eggs 'pi lldk-,ha ped ( II) __ ____ .____ _________ . __ . IJm l/(;/w('cfll • No lale ral ,ki n lil lus on Ihe nec k. T USV L ' 3: mal ( I) ____ __ _____ ______ ___ .__ _____________ ___ ___ __ ___ _______ _ 6 l'gg~
G
( i R()'S\IA~\J 1L)67)
I\gami dac
(, Vent ra ls > dorsals: dorsals usua lly interm ixed with larger. tubercular or rhom bi c scales (J); if scalation homogeneolls, dorsa ls neither rhombic nor kee led ........................... ............ .. GOtl(Jceplruills • Ventra ls :; dorsa ls ......................................... ..... 7 7 Gular . ilC of males wi th late ral poc kets ( K) , dorsa ls
ofvaliolls shapes, sizes. and orientations. rhombic, tips ma inl y ro unded .. ..................... ClJ mplicilll.~ • Gular sac of ma les wi thout latera l pockets, dorsals vary ' Iightl y ill siLe. rhomb ic. tips truncate ... .. ... .. ........... .. ...... .. ... .... ..... .. ........... .. ... Ps(!wioca/oles
8 1\ small protrusion usually presen t (L), TLlS VL > 1.9. hase of the tai l ro tund in males ... AI'''allioti,\' o protru ion. base of th t: tail in males nattened ahove. TLlSVL < 1.9: ifTL/SV L > 1.8. supraciliary edge and nape with spines (M) ...... ........... ..
M
.... ..... .. ... .... ... ........ ............. ..... .. ... ... .. PillJxoplrtJ'.\'
Genu, AplulIliofis
P LTLRS,
IS64
Specil!s of the genu. Aphall io/is arc di stributed 011 th e Malay Penin s ul a, Sum atra, Born eo . and on many sma ll o ffs hore islands. Smal l agam ids (SV!. up to 70 Illm max .). wit h slendl.: r bod ies and lo ng. de lica te limhs; c nlargl!d. hum p-like scales on the tip of th l! snout or a sca ly protrusio n in two o r the three spec ies o r Ap/w ll iOlis. Lin ing orthe mout h blue. Tympanu lll hiddcn , sometimes indi cated by cniargl!d sca k s. Dorsal sea lcs
small and hete rogencou ', ve ntrals alway. keeled and large r than dorsal s. Nuchal and do rsal crest absent or weak ly deve lopcd . T hese tree- and bus h-dwellers inhnbit primary low land f'o rests an d hill y regions In the genera l vicinity of ru nn ing water. They fecd on smal l insects. Al l threl! species of th is gen u ~ occ ur on Borneo. although prese nce of'AII/ulllio /is II CII I i/'(}sl ri.\· i~ 'ub.icet to sOl11e debate. Mount Kinaba lu is home to onl y one spec ies, .-1. 0 1'1111/(/ , whi ch h a~ hec n rccorded up to c l eva ti o ll ~ or900 III a.s. 1.
Kc. to the Bornean spccies of Aphalliolis A distinct prolnrsion on the snout ........... omala • No or onl y wea kl y deve loped protrusion on the snout ........................................................ ....... .. . 2
2 Fl anks with tra nsverse bands. iris ll f male brown .. .... ...... ..... .... ..... .... ... .................... ..... act//im.l/ri.l'
• Fl anks without tran verse bands. iris of males blue or green .......... ............................................ . It/sea
rab. 22 Di,trihullon of 1 II} /IlII11 (1 li.\-s p..: c ic~ on Il 0 1'11 co. .'Ip //(Jllill /i , (l ct/li l'll, /,.i" '! AIl /Wl/ioll ' /m'(,1I P I II ~\.
.1p//{/l/io/is oma/II
M Olll! d 1'\\11.
I H ~9
1 ~ 64
( L IIlIlI III
.Ii I
Ill' .
1)<9 3)
---
...
Sahah
Ki naba tu
•
•
Brunei
i
•
Sa rnw ak
.,
•
I Kahmuntan .,
~
• 225
I\g.amidae
Aphdlliofis Orllafa (LIIYIII DI. .11: [;1)1' ,
I 893 )
J, '"11 . IX'I.! : 25 1 Ty pe l ocll lil ~' : "I' Ihe Sanda~an - n ay I '\. nl>m~\») " .!"Im/rm, ,m"'//. R" , IIJ I <) I : '12. ./(11)"/11111
01"1111/(1 1 '"'11 '"
" II' 11t~ n~iglth\\urhood
lig . -1 7. SMItIl 1<1>1 : 22. pi II fi g 2. M\" 'IIII 19X-," 2(', 1111 NII \R I1 & SII X." " 19X-I 2-1 '11'11111". \1 \l ~'l\ ' I'N-lb: 2-15 ; I \N 19<13 : 1 .I11/W/"I'II /111"11111 II I .1"",,- 1'130 (syn . ancr tvh" rill \ & (ill(" " 1\ ' " 1'197 anti OIS & 11 ",,1\.2(00) .1111'11/11'-<1 IIl1.II//
·',,"
Fig. 23() 1',,1 a I<; of "'pliUllill/I'
o
g
226
Diagnosis Sl11al lcsl specics ufthe g.enu. A/J/wllio/is (SVL less than 60 111m), Ineonspicuuu ' ly brown; both sexes \vith n dislincl prolrllsion .
111'1/(//(1
Description VI. males 54-55. tCmales 52-57 111111 ; TL l11a les I 10-115 . fema les about 102 mm; lip of the snoul with a tlexible. ollen backward-point in!:! protrusion ; slllall gular sae present; low nucha l crest con~ist ing of triangular ~cale s; dorsal crest separated Ii'om th e former. consisling of small. se rrated scales,
I'rom KUlai . Ka limantan .
!\gamidae
paralkkd by IwO rows of tubercu lar sca le ': conical laterals smaller than dorsals. intermixed with larger scale", Uppersicle uniforml y brownish. head ancl hody sometimes with a faint olive tinge: occasionally \ ith a light . indisti nct ve rt ebral stripe . Fe lllak s with dark radi a l pattern around the eye and with two li ght bands between the eyes; rostral protrusion slightly coni ca l a" opposed to tl allened in males: mak s with a small nuchal sai l and distinctly broaden tai l base. Two eg ' $ (7 , 15 111m ) p CI' cl ut ch, Ec() I() ~~l m l h()l()gy An inhabitant of primal)' lo re. ts.
:c 2
m u
_ . -......... 0
Distrihul i() n Fi g. 23 I Fl:lllait: of Ap/w11 iOli.1 Ol'llara from Dcra ma~ o t , Sa bah. Two localities are known from Kalimant an. i.e. upper Mahakkam river (1.1Illli ilL Jt.l IUI ' 1<)05) and Res , Kutai (photoSa bah and Brunei (D A ~. per" , COnll11 . ), M t. Kinagraphic record K\I()\\'LI \), all th e others lie in halu: Kiau .
Genus BrOIll:ho£:e/lI
K AU P,
1'l327
Of the agamid genera occurring on MI. Kinabalu . ha. the widest di slribution . Its range extends from the Nieobar Islands throunh Mya nmar as the western limit on the ma inland . to Vielnam in the east, and sOllt heast through the Ma lay Peninsul a. to I\'ew Guinea and to the Philippim: ·. im:luding numerous sma ller islands in the whole region . Mediul1l- 'iled to large species (SVL 80-130 111m) with long. slender limb s and tail. Dorsals and latera l" homogeneolls in shape, size. and orientation , Skin folds on the sides orthe neck sllppol1ed by thc hyoid apparatus, Nucha l cre ,t usua lly present or at least indicated, dorsa l crest present, indicated , or abse nt. If dorsa l crest present , continuous with nuchal crest. Gular sac absent or onl y weakly developed , Gular sca les and sca les coveJing the gular sac rhomhi c in shape. Tai l 'Ii ghtl y triangular at Bmllchoce/a
it s ha. e. then roundi sh or sli ghtl y lat erall y eompre ' ·cd . In contrast to all other simi lar agalllids. femal es of this genus deposit spindle-shaped eggs (not confirmed for 8 . ce/e/Jl'II.1'i,\', ,\'111 £I I'(IKliill(l , and danieli) .
227
Agamida\.:
13 ornco is hom e 1) Brollc/wec!1I juhllta (D:\s. pen;. CO I11I11 . ) and B. crisla/clla . The laller occurs both \. . ilhin forcsted habitats and in heavil y di slurhed. \.:ven urban. setting. hut seems to prefer upen areas. In the Hcadqllarters region of Mt. Kinabalu. these lizards ca n be found in fore~t s Lip to altitudes uf approx, 1700 III a.s. 1. In the Pllri n!.: re g ion it \\:\ s found outside Ihe forest (M \\11111 \ & DI ',VI ' R 19X ~).
Brollc/wce/a cristatella (K lJ lIl., 1820)
H A III ' R~Ii\\I\I.
,.1 .~(/1II11 (,/,/,\/,/1"11,,
K."". I X20: I ox
T)' p~
thKIII "11 tX'I'ia .
.,1 ;
\1\ "'1111\
ii.:
x,: IJI
/I,."
lit
R(x",
[)I V I K
,I:. Sh)( ,KI II lwl\ 11)91 ' I XO
l'''i''' '\RIl
N
1'11 5 ' 12 1; "\111 111931 : 10. I'IX2 : IK. \ ·1',\1111 Y 1<):0 25:
I'IX4 : 24 ;
h(l( clo a;,H "It'lla, i\1
M." III ' \ k
II,))!" hocd"
6- 10 (rarely 4-5 or 12) uppe r rows of dorsals pointing IIpwards: dorsa ls/ventra ls 1.5 .................. .. , ".......................................................... ('ri.~f(lfell(l
• 2-4 upper rows of dorsa ls pOinting upward~ ; dorsals! \.:nlrals S 1.2 .......... ..................... .iI/haw
Fig. 233
~
.0
:5 I-
Q
22X
/J1'()1I('/lIIce/tI
cr. ('1'1.\,/(//('//"
11 0 1
C,,/rIlt'S ('1';'/01"'111". Vl'"()i<\Kn I X90: 129. pI. VII : fig. '. Ja;
~1\1 " " "
\I 10.\11"
19~7 '
1991a : 33. ItN2:! . 1111. TIN ItN} : 2. Nhl
Key to the Bornean species of Brollchoce/a
luca lil ):
IIluil'"lcU
(;I<""\I\'N
I'IX- ' 9; 01\
c'
285 . 19X8c: 175.
10.\""
11)<)4 h: 245:
1<.)<)7. 16 1
')1 .. M \I ~~II " I 994h: ~14.
fi g" 12. 1" I 'Jli(>,: 2liX
Oia ~ no s i s
Slender. long-Iaikd, long- limbed . and narrowheaded agamid~ with a low nuchal eresl and dorsa l crest indi cated . Dnr~( tl s di amo nd-s haped. hOll1ogenOl! .. imhriea!\.:. smaller tha n the simi larly 'haped ventra Is. Dcscriplio n
. VL mal es '1<7- 1:l 0. fe males 82-% 111m ; TL males ] 00-440. female~ 290-330 111m. A row 01' en larged
frorn Ihe Ht:ad4uarh:r!>.
I\gamidae
sca les o r var iable lengths behind th e s upr aci liar y edge. Small nu cllal c rest or lanceo la le . erect sca lcs; dorsal crest indicated by a low. serrate d ridge ; 6-10 (rarely 4-5 o r 12) rows or dorsab direc te d up- and backwards. late ra ls backand downwa rd s : ve ntral s about 1.5 to 2 tim es as la rge as dorsals: sca les o n the g ular 'ac smaller than ventrals. Uppe r 'ide us uall y green: whitish or light blue spots vis ibl e o n th e si des. o ft e n fus in g to ro rm ver tic a l ba nd s ' tl an ks so met imes reticulated with brown; s ides or the head green to li gh t blue. occasionally with a reddi sh tinge: o rb it, o r only o rbit e d ge . a nd ty mpanum us uall y blac k; ve ntral suril lees complete ly g reeni sh yellow. g ular sac occasionall y ye ll owis h red. ye llow. or whitish. F. ntire colorati on su bjec t to 'o lo ur c ha nges toward s brow n. dark g ree n to blackish. yellowish. or reddi sh. 1-2 s p i ndlc-s haped egg. (9- 10 " 40 .5-42 111111 ) per clut ch. Eco logy/Etho logy Brollchoce/a crisllIlellll inhabits trees a nd bushes on the edge orrorest ·, a nd urba n area ' s uc h as pa rks. Its vel1ical di str ibu ti o n ranges
Fig. 234 Nn lll c/ro ce/a a i.lw/el/a from the I kadquancrs ill a blad,ish co lour phase.
229
Aga mi dae
fro m ~ea levc l up to approx . 1700 m a. s. 1. O n l'vll. Kinaba lu thi ~ ag<1 mid ha:- abo been fo und in pri mary fore:> t ( M.\ I I<- MlI"> 1994b ). Th e ir sl ight ly sprcadablc ribs cnables it to pHI'aehut e fro m a trce to the ground (M,\ 11If\ & (j1!()~~ M "\I\I 1994). Distrihutio n Myanmar t() ew (i uil1ca , and Ph iliprines. Mt. Kina ha lu : Poring, Kia ll . Kenokok, Sayap. Lobang, Blindu Tlihan, Il eadqllarter:- regio n includ ing the Liwa go Trail.
M ANTII LY
Genus Complicitlls in M J\'.JTII LY & GI{oSSM/\ NN, 1997
MOl1otyr ic genu. endemi c 10 Mt. Kinahalu . desc ribed ol1lhe basis ora sil1gk: male speci mcn wilh a habitu s simil ar to specics of thc genus P SI' IIl/O cli /oles. Un iqllc amon o aga mid l izard ~ due 10 Ihc comb inat ion of latcral gular sac podets and almost granular sc utell ati on o f the gul ar sac .
Comp/icitlls lIigrigll/aris (OT;\ &
1(9 1)
01\ L"- 1 1I~ 11l'. 1<)'11 : 11\5· IX7. fi ~ . 4 T)·pc IMa lil)': Kina ha lll P~ rk (a ll. ca. I "00 Ill) Rm" clwce/lIl11g n g ll/lln ,. vt 'l ' " , 1')1)4h: B5 ( '''/(If,',\ (i ll • . '~ll.) IIIg n g ll/llrlt. \1 " ~ ' " I~ 1994b: ~4 5 (,,,/,,, (,.\. 1I1)!JI,f!. lIhm s
COIIII )/i C: III1 \
lI ;grig,ulllri,'i,
\ '1"
1111 ' III
\ 1\' ti ll
' "'' ' (j ~(l''''\ ' ,\NN
11)97: 165. fi g . LX II
Diag nusis . mall aga mids (SVL less than RO mm ) wi th slightl y heterogeneous do rsals regarding shape, size. and orient atio n. tvlales with latera l I.! ular sac poc kets, nucha l crest, and dorsa l crest indi ca ted : limbs and tail comparati vely short.
Desc riptio n SV L 111::1 le 70 1111ll . TL male 155 mm : nuchal crcst composed o f 5 hi oh, lallceo late sC3 I e~ : dorsal crest ind icated by erect, dis tinctly spaced sca les, continued 0 11 [he ::I llt erio r port ion of the t::l il : dorsa ls of varioLt S ~i i' (;s , mainl y ~ l11 0 11th : latera ls lIsliall y d iam o nd - ~h a p ed, point ed or broad ly rOllnded , difTer-
Fig. C I Ma k nt' C/llI/l llicit l/ " lIig rigll/lIri.l . dra\\·ing. alkr th e ho lol ype.
:n o
I-I IK II >A .
Agalllidae
ing in size. arranged in irreg ular rows: vc ntral s slight ly het erogeneous in size. keeled : sca les on the gular sac vl:ry small. prcdominantly granu lar: shoulder fold weak ly developed: tai l substantiall y laterall y compressed laterally behind its base. Ground co lour of males dark greyis h brown abovc: . evera l whi te spots below the eye. 011 shou lder and nuchal rcgion ; two widc. whitish cross band s on back and upper flank s: lower side predominantl y IVhiti~h: gll iar sac bl ack with a white margi n and whitc spot s. F.col ogy/Elh olngy : Unknown . f)i stri hu tin n :\,11. Kinab ~llu :
Only known from the type loca lity.
Genus Draco LI
'KALUS,
1758
Thc di stribution o f this genus rangcs from India cast to China. Ihe Philippines. and the Indo-A u. tndian Archipdago. Very slender. petitl.! bui ld li:tards up to a VI. of 140 mill . Two large. spreadable. often co lourfu l wings (p a ta~ia) support ed by ex tended. movabl e ribs. Additional skin sai ls on Ihe si des of the nl:ck (neck lappets) and dewlaps in va rious shapes and co l our~ . indicati ve of the specics and often also or the scx. Nostrils orientl.!d upwards or laterall y: tympanum visib le or covered with ~calc~ .
Shapc and co loration of dewlaps and inner ~i d c of the neck lap pet s arc invo lvcd in the recognition of conspecifics and arc therefore or grea t va lue for thc taxonomi st. The pal
T:lh.2J Dl slriblilioll of {)/'(/('()-spcc ic., on Homeo.
D raco a/jill i.1 B .\~ 11 111. I X95 /)/,// ('u cu rtlllfll .1 f)/'{/('()
('il
(,l'is/lIIel/lls
" 111t· ~,
I Xh4
(il ' I\111I R,
D raco j iIll IJrialllsfilllh l'ill lll.I' /)/,(/('() hll " I/I U/()IIO,!!,OI/
Kl III.
l/Iic I'o/elll,1
Dmco III l/xilllll.l· BOlli {) /'{Ieo lII e/al/ op ogoll
I X72
' 1.1 R.
I X20
H ill Ill\(a-Il .
I XlJ]
I X93
II jg,.itlpp l!l/di£· III11/II.~
II \1\ III
II.
Oran} ob.\'(,lIr11s ObS(,II1'IIS 13m LlMoI~, I !-i!l7
-
Drac(} qllillt/ ll e/i,sc;al/I.\ Im l,r:ilJll r ba 11i·"I\II,. 1936 /)"(/( '0 I'I,{(//I.I' ,IIIII/u/rlll/II'
SI
1111 (oil .
I !-i44
I X\.))
I
Saba h
Kinabulu
• • • • • • • • •
•
•
•
• • •
llrun~i
•
Sarawak
Kalimanlan
•
•
•
• • • •
•
•
• • • • •
• • 23 1
Agamidac
Key to the Bornean species of Draco Five patagial rib~ ........ ............ ... ... ..... ... ... .... .. .... 2 ix patagial ribs .............. .................................. 6 2 Nostril directed upwards. supraciliary edge without a thollllike scale ............. ... ... .............. ......... 3 • Nostril directed laterally. supraciliary edge with or without a thornlike scale ..................... ......... 5
3 Patagium dorsa lly yellowish to greenish or bronze to golden brown with 5-6 pairs of dark narrow bands. dewlap smoky grey (males) or pale yellow (females). lower portion of dew lap with greatly enlarged sca les .. .. ................ ohscllrllS obsclIl"IIs • Patagium dorsa lly black or blackish, dewlap without enlarged scales ................ ................. .... 4 4 Patagiull1 dorsall y black with orangt:-yellow spots. in males dewlap jet black with a white portion on posterior part of base; in fem ales dewlap black to dark reddish, base white; in both sexes front portion of the inner side of neck lappets black , basa l portion pure white .... .... ............. .......................... . .. .. ......... .. .... me/(Ill opogo/l lIigrillppefl diclllufIIS • Patagium dorsa lly with black marbling sometimes forming five ill-defined bands or similar as in 4. in males dewlap bright yellow with a large black spot on base. in females dewlap reddish to lightly washed with orange. same also on inner side of neck lappet;, in both sexes ...... .... .... ...... .... .. .... .... .
5 Supraci liary edge with a thornlike sca le. body with latera l row of enlarged scales. in both males and females dewlap and inner side of neck lappets pa le reddish to yellowish, ta il without crest ... .. ......... . .. .... .. .... .. .......... .. ........ ..... .. jimhriatlls jimbr ia/lls
• Supraciliary edge without a thornlike sca le, body without enlarged sca les; dewlap in males goldenyellow with brown anterior edge. tail of males with crest ...... .......... ....... .. ....... ...... .... .... ... . crislale/lus 6 Nostril directed upwards. supraciliary edge withoul a thornlike calc .................... ... ............ ....... 7 • Nostri l directed laterally, supraciliary edge with a thornlike scale ............................ ... ... .... .......... 8
7 Patagium dorsa lly with a pattern of alternating orange red and black bands ............ ................. ...... .. ... . .... ... ... ... ..... ...... qllillqlle/ascillflls IOllg ih"rha
• Patag iulll dorsally with a pattern of alternating green and black bands ......................... muxillllH 8 Patagiu m usually reddish dorsally in both sexes with black spots. or dark wilh green siripes (some males): upper side of body predominantly green (males ) or brownish (females) ............. ('O /'llIl/I1 .~ • Patagium black dorsally with lighter spots. upper side of body bronze eolourt:d or light brownish to lighl greyish ................ ........ . I"O/OIlS XlIlllalmllllS
.................. ...... ..... ..... /w{,lIIolopO}t1l1l micro /epis
'! Dntco ()/"(/( 'U 1l1!i""
(~nillis
B,.,,{I L1 ' IT, 1895
1\ '"11 111. IX'I5.1 ~I)
/J nlt 'fl (tUIIIIIII.\, :\MIIII
1931 - 22
r)p~
(pari ..
Im' alil) : Bonl ~"
Ic mal c'\ ,
Diag nosis prter FhRTt I·n IX95a) "Si milar to IJ co r/llii/ll. but wilhoul larg ' spinclike sca le above tl1\: eye. Gular appendage very sma ll ; in Ic mal es almOSI abse nt." Descriplion ('Ifter B\ll lllll lX'>5a) "Mall: Hac k dllil brown. tinged with green ; three di ~ tinct greyi~ h white Iran svcr~e bands on the back:
232
wing IlH.:n lbranes abm·e brighl brick n:d spotled wilh black. with broad black outer marg ins. a pal e bluish c.rcy line down the centre oCthe belly: IInderside or wing lIlembranes dllil brick red tinged wilh blue. a nd spotted with blackish brov,:n , margined with blolches or black alld greyi;,h while. Adull female Back g rey moltb.l with dark brown and tinged with gree n: wing mem brane!' bronze g reen spotled wilh black , wilh a broad black band o n the outer margin ; g ul ar appendage small . ye llowi sh g reen; chest hlue; a line down Ihe cen tre or belly and undl:!" pari o f hind limbs pale blue. s ides of body g reyish white mottled with black. lInder-
ga midae
side of" win!.? me mb ra ne yel low ish gree n, ou te r lI1argin bille." Remarks: The above quoted description of the dewlap o f the female i5 based o n an error (COIllpare Diagnosis): it actua ll y refers to male ' where such d e~c ript io n is mi ssing. The all ocati on of the n. co rl/l/IIlS females to the taxon n.a/IiI/is by SMI rll ( 193 I ) is based o n the mentioned absence o f a dewlap. Of all spec ies of" Draco inhabiting the Sunda region, the absence ofa dewlap in females is unique to D. ((llilli.l". The allocati on of females referred to by , ~ II T II ( I 3 1) to D. a/1il/i.l" i" done here with reservations, as neither the types o f O. a/iiI/is nor the specimens origina ll y avai lable to MII"II could be examined. The va lidit y of D. q/lillil' is currently uncert ain . I::eology/ Ethulogy:
'nknown.
Oi sll"iblilion Uorneo. MI, Kinaballl:
Draco
'"o
~
I
'l
COl'llufIlS
Kiau .
GC
I"II I'R,
1864
125 Type Inra lit )': B,,, "CO I1N(J 12l{: II 'Nil '
Ora("o ("omll/II.' (31 \1111 It . I X6-1 IJra("o '"01"1111/''' .
MIX
4.>' ,~I)
1900c: 2: S '" 11 1 193 1: 22 (part . ('I)): Itl R,.,u 191 5: 72. MI , ""' I'I ~G : 1'1: EIlIN II\ HIl & S J(X,"IN I'IX-I : 24: 1<)9J: 2: M \I ~"I" I ')')-Ih: ~-l5 ; I ( "~II' cl a1. 1'1'1'): I 'J~, fi g. 2/\ , II, ( '.
r,,,
OmC<J\IJlI(}I'/NII'IUrlllI/,,,. III "NI< ,I'!16 . I X1:
M " l lll' \
I'I X1 .
25
Diagnosis A small to mcd iulll-si7ed species (SVL max . X5 111m ) wi th 6 patag ial ribs and a fh orn like sca le on the sup raci liary edge . ostril s oriented laterally: dewlap tria ngu lar, lip orange ye ll ow in mal es. in females dewlap white: inner si de of neck lappets and dewla p s imil ar! co lou red . j
From fOp fo uase;
n/"(/ C()
con /II/lis
Fig. 235 rcmalc (lcn dorsal patt crn. right vl'n lral paft
dor~a l
pa tt ern. righ f \clliral pattern
Fig. 237 \t1alc (ventral pallc rn o f Fig. 231l Dewlap of a male.
pata~illm) .
233
I\gJl11idJC
234
;\gam idae
Description
SV I. males lip to 71<. females lip 10 R5 111m; 6 patagial ribs; nostrils oriented latera ll y; a thornlike sca le on ~upraeiliary edge; tym panum u, ually not covered with scales; dewla p triangular, with small sca les: neck lappets wi th larger scale, : males and female~ with a sma ll nuchal ncst composed or triangu lar sca les: dorsa l crests absent in either sex: dorsa ls varying in si/e. smooth. 'ometimes intennixed with some larger sca les; keeled sca les on bases of patagium: fringe- like sca les along the rear edges of the thi ghs: ventra ls kee led, abo ut si:t.e or dorsals. Dorsal surfllee of males green or greenish brown. occasionall y with indistinct. dark banel. : dorsum of' fema les li ght brownish with ex tensive darker inclusions; both sexes wi th dark srots on head, nar e. and 011 sides or lhe neck: lower 'urfacc of mal es and remales whitish with a turquoise to bluish centre and sl11all ve nlrolatera l dark (male ' ) or ye llow ish brown (females) spots : patag ium reddish above. more greenish towards the body (males). or brown i, h (females) with dark spot · OIxasionally converging to form bands: a wide dark marginal band sometimes with ve ry Iight inclusions: some males dark with indistinct. greenish longitudinal lines: in males lower surface o f r atagium bluish or reddish with a blue marginal band and large black blolches fusing on the margi ns: in females lower surface of patag ium yellowish white proximally. then somet imes reddish, and bluish with dark spots dislally. dark spots occasionall y fus ing to form a marginal band: dewlap in males with a ye ll owish to orange ye llow tip and a greeni sh white base. inner side of neck lappets \V'ith a COITe, ponding coloration: inlier side of neck laprel. alld dewlap in I cma l e~ \ hitt: wi th very pale ye llowish brown speck les. 3-4 egg~ reI' cilltch.
Draco f imhria/lis fimh ria/lI.\' K l
III,
I ~{20
Om,'" /i1ll/WIIIIII ' KI III . I !C O 101 '1~ IH' l"I' alil) : " llId,a Ilc,lriclt·" 1)' 1'" 10(' ,liil)': \1ala y l' el1ln sul a url c nlal, " ( II I 'NI<, 1'1.1 6) OrtU fJ l illllonlll/" . S MIlIl I'n t 10. 1 1. 1-I "NII\~ I ) & " "",ION 19~4 . 24 , M\I ~MI" 1'/X7 2X~. 1'1'12" 110. 1,,,< 1<)1), J. M Al ~ M l " 1994b: 2-1 5: ilrtl co l illl/'I''''III.I lilllhrllllll.l . M \N 11 11 \ 19 .I ' _5
Di ag nos is
;\ moderalely larg . speci<.:s (SY L 90-1 17 mm) wit h 5 ralagial ribs and a thornlike scale 0 11 the surraciliary edge. oSlri ls orien ted lalera ll y: dewlap Iriangular: in ner side of neck lappe t ~ and dewlap orange (yellow ill . pecimens from Poring. MeCit IRI'. pers. eomm .) j
Uesc r ipt ion SV L m:! les 9()- I 12. g ial
rib ~ :
rema l e~ 102- I 17 mill ; 5 palanostrils d irected lalera ll y: a Ihorn like
Fig. 24 1 /)rll l'ojilllhrimlis jilllhrlllJII., from Poring.
Eco lo gy/l+~ th o l ogy
usuall y inhabit luwland fores ts and hill y regions. Probab ly thi s is a forest edge and canopy sr ee ies alld can be fou lld in totally disturbed urban areas. such as the ground s of the Sabah Museum . /)J'(I CO cuml llllS
Uislribuliu n Borneo .. umat ra. Java . and Bunguran Kina lJa lu : Ki au. Poring:.
I s l and~.
:\1t.
235
!\gn ll1 idae
sl:ak on s lip rac ili ~lI) .::d(!.e; t Y ll1 pan ull1 l1~ lI a ll y without sca le.; dew lap tri angul ar with sl11nll sca les: neck lappets wi th largc r sca les; do rsa ls hl:tcrogcneo us. preuomin nll tl y smoo th. wi th on ly n k w kee led ~calcs int l:rl11 ixed; laternl row of en largl:d sl:a les: fringe -like scn le. on pos ter ior edges of t h ig h ~ nnd hnse nl' tail : ve nt ra l ' kee led. Inrgcr than dorsa ls. \)orl'a ll y l:o lorat io n li ke bark o r tropica l tre's. brO\vn. grl:y. 01ive, and pale green ish, interspersed with darker pa t 'hes; patngiulll 1I1(Ji stinctly stripl:d wi th black and gre ish brown ; lowe r ::.urfnce o r pa tagi ul11 li ght grey, sl otte d wi th dnrk brown (111ales). or light orange with bl ack ::.pots (fel11 ales): inm'r side o r neck lappets ora nge (ye ll,)w in Poring. speci mcns) ; ul:w lap reddi sh ora nge (yel low ill
Draco
IIlllXilllllS B Old t' l\l;I'K ,
1893
Il, ,, ", ", d R, t ~ 9 3 : 522 't'YI.e lura lit)': MI Dulll , 2()()() k " l I~ " r aw "~l IJra.-
25
Di ag nosis rhe Inrge. t ~pel: i es or f) ra('() on Borneo ( VI. lip to 139 11l1ll ) wit h (, patag inl ri bs, and no thornlike scale on supraci liary cdge. ostri ls oriented upwa rd : dew'lap Ir iangul ar. lower port ion reddi sh brow n (males) or l(ree nish g:rey (females): inner side of neck l a ppct~ gn:y to dark grey. J)csc ription SVL mnlcs up to 139 , fe male, li p to 137 mm ; 6 pat agial ri bs; nostril s l11'icnlt:d upwa rd : no thornlike scn lc on the ~ upra ci l ial , edge: tympanum pnrti all y or co mpl etely covered with sca les; dew lap triang ula r. with ~m a ll sca \(;s: nec k la ppets with slightly en lnrged scales along the edges: dorsals very ~ m a ll. primaril y homogeneo us with just :J sma ll se ri es of e nl arged sca les alo ng I ase o f p ata~iu m : fringe-like sca les on the rear margins of th ighs and on ba~e o r tai l: \'cnt rals kce led . larger than do r~ a l s .
236
P or in ~ spl:c imcns). ant erior and lower marg in s light er (fcmales) ; similar co loration in mal es. Males with low nu cha l sa il and c:Judal erc.:st o f small. tria ngul ar scales, dew lap co mparati ve ly larger and wi der. uchal crest uf fcmales eo n ~ i s t ing onl y o f slightly en larged sca les. 3-4 eggs ( 10 " 17 mm) per cl utc h. Ecolo ~y / F. t holog)'
IJl'(lco/)illl iJrill /1IS inhabit s predom inant ly low lnnd
fo rests and hill y n:gions. Distribution Malay Peninsula . L3ornco. Sumatra incl ud in g the s urruu nd ing. sma ll er i. la nd::.. a nd J ava . Mt. Kin abalu : Kiau: Poring.
-----------------D rsa ll y preuominantl y green \\ ith a brow ni sh hl ack pattern ; pat ag ium black with numerous light , brownish gree n, di scontinuous lonp,itud inal lines: lower side o f patagi um almost co lourless: inm:r side o fm:ck lappets gre (ma les) or dark grey (fe male, ): dewlap black al its hase. wi th white dots. remai nin g, portio ns redd ish brown (ma les) or greeni sh grey ( fe males) , Males wi th nuchal sa il and a mllt:h larger dewlap, 1-5 t:ggs (II ' I g Illlll ) per cl utch. Eco lo g~"/ F. t holo::,:y Draco 1110.1';11111,\' prefers primalY ra in forests uplO altitudes of approx . 600 m a.s.1.
I)istribution Peninsular if alays ia. Sumat ra , Na tuna Is lands. and Borneo. \11. Kinllb ll lu : Poring.
Fig. 243 I'emale
or /)m ('o
'(i(, I1/(//Il" from Sumat ra .
I11 l' itllllJl)(}gO Il IIIg r i llfll lell -
Aga midae
Draco lIle/ul1opogol1 l1ig riappelldicll/atlls B ARTLLH, 1R95 /)J'
JII"r /tII'I" ' lId" 'lIlulII ' n ·"l lll' II ,
tlN' a
l) ~
T)p~
tnca tily: KlI Clllll!;. Silr:m"k I)i ll~ n() s i s
A slllall to modera tely la rge species ( ' VL up to 93 mm ) with 5 patagial ribs. no thornlike ~ea lc on the sup racil iary edge. Nostri ls d irec led upwa rd : dewlap tri angular. black : inner ~i d e or m:ck lappets blac k and whit " Desc ription VL males up to Xl). fem ales ti p to 93 111111: 5 palagial rihs: nostri ls ori ent ed up ward : no thornli ke sca le on 'up rac iliary edge: tympanum partly or compl etely cove red wilh smooth sk in : dl:w lap long,
F ig, 244 Ma le of Draco /lW/W/fJIJOl!,1J 1I /lig riIlI'IJ('I/(/ir/II III " frolll ' uma tra ,
('I
23 7
Agamidac
triangul ar: Ileck lappets with en larged sca les: dorsa l" heterogeneous. 'mooth or f ~l illtl y kee led: a latera l rm.\! or enl arged sca les: frin ge- like sca les 011 posterior part o f thi ghs and on base o f tail : veillrais keel ed. larger than dorsal. . Dorsa ll y mostly green . with brownish bands or diamond shaped spots and lighter in c lu sion~: patagi um dorsall y black with nUlll erous ye ll ow-ora ng' spots. these so metime;. wi th smal l dark ce ntre: dorsa l colo ration and patt e rn of' pala ·ia shimlllerin g th roug h. visi ble on the 1Illtkrside. Front porti on (II' inner side or ncck lappets black, basal portioll pllre whi te: in males dewlap jet black wi th a white portion on posterior part or base. in fe males dew lap blaek III da rk reddish. base white, 1-2 egg~ per clut ch. Fcology/ I£I hology f)/'{/('() /11 . lIig rillI1fll!lItii('u/a/II.\· inhab it s usuall y ~ee ollliary and prima l)' lowla nd forest" up 10 d evati 1I1 . o r approx. ()OO m a,s.1.
Fig. 245 Mal..: of /)ra('o 1IIl'IIIIJr)IJogolllli/Lllllalra ,
Distrihution ' lImatra an d Borneo. \11. Kinabalu: Po rin g (NI ( OI JlRr. pel's. <.:o ml11 ,).
lalll l
Fi:.:, 246 Mal..: or OmclJ lI1<,icllllll}()glJll JIIgnfllll)<'lIdiclIfrom Saillubollg. ,'arawak.
Draco obsclIrlls ob.\·curus
111111 1'
Boult: O/'{/('(I
GER,
I RR7
Ho, " "''' R. I XX7b. 'IS
ohSCllrUI
T) I)e
t"cali!): " \1 oulll
Killu Haloo" IJI'IIC'O IJ"."III~'., .
Me"
"' R, KH) 1915: K5: IlJ9-1b : 2-1 5:
I X'IO : 12S, pI. VII I: f, ~ . I. I a. I b: 1'131 ' 10,1 I : I .·\ N 199:l: 3: VIM ~Ml"
<JI Ikil
S.1I111
{)rtlt·",ill'JI/"'/I.' . (1101 O)'{/,,, f",."trl\lIV H, ,,
" , ~ 1')10' 12 .
Om",
IO/'JI1I1\1/1
<';\11111
II ", I~ .
193 1' 10. 3 1
"I", /II'"V, II I '''1(, 1'131.: 2 1
25 : I.II''' ''RI> &
)'''.KI'' 2-1
19(0)
WI It-
, \it "'II' \ 19X:1 .
1(nn: (, 'S (noliJm"'J1 ,,1111/,\ I 7SX) :vI .\ \ 1111 ' \ & l)" NIJ It 19K2: 16: Nil lit!> 8: S"' .RI " 2-1 : M\I "" S I'I"-Ih: 245 IJ,.",,, ,t)lall v 't)lrlllv . 1'101 OraC(I 1'111011' LI\N \I I';. 175K) M '''"I1\ 1<)83 :25
OrliCH nhscuI'IIS oh."t'UI"US. MI I,\ II H"i /)m ('" ",11Ifl.1·,
1.,"
I."",.",.
Diagnosis A modera tely large species ( ' VL up to 100 111111) wi th 5 patagia l ribs; no thornli ke sca le on the supracil ial), edge. oSlriis orien ted upward ; deWlap tongue-shaped. smoky grey (male~) or pale yell ow
23X
Agaillidae
chestnut co loured; lowe r silk or palagium grey to whitish , wilh indistinel bands and maroon margins of'ten fragmented into spots in fc mnles; inner side or neck lappet. maroon (males) or pal e pink (fcmales ); dew lap smok y grey (mal e;.) or pale ye llow (females). 1-5 eggs per clut ch. [eulogy/Ethology
Draco o. OhSC III ·II.\· inhabits primary rorests up to elevations 01'900 m a.s. 1. [)ist ributiun
' umatra and Borneo. ,\-11. Kin abal .. : Kiau . Porin g.
Draco quinque/uscia/us
IOIl~ibarba
111.:-<'< 1(" 1936 'I11/1I1"e/ascialt" /""gihllr!>a II I NNI<" t 'J1(,. I'll T) po Barcilil. Sar,,\\'a~, ~llrll~n nrl/('O (will'l" e/a ('1<1111'. '-;\111111'111 10, 11 . M""I K' 19x3 :
iJl'([ ('()
l o(' a li/ ~' :
70: Em "" \~Il & ~I( ~ ,RI" I \JX-l 2-1 , Il)l)-lo ' 2-l~
lii:.:. 247 M3ic of IJram ohswn/l' oh,c/ II'I/s li'om Poring.
t IN
IlJlJJ I ; '1\I ~~ H
,
lI<.li pnngs.
I
(rcma l e~) ;
;\ medium-sized species (SVI.IIP III 10(1111111) with 6 patagial ribs, IlO Ihollllik.: scale on the "upracili:1ry euge. 'ost ril s poi nt ing upward , dewlap cr.:~ ccnt-shaped. mustard-ye llow (l11ale~) or dull ye llow (females). sa me co lours on inn.:r ,.i de or neck
r(l('(}
{1"il/qlll~/(ls('illfll'
IOflg,ih",.hll.
I\.I ,\Nlll1..,
It)~1
15
[)i agnosis
inner side of ne ck lappet s maroon (males) or pale pink (females). Dcsc riptio n
. VI. males up 10 100. tCmales up to 94 mm : 5 palagial rib.; nostril s orienled upward : no thornlike sca le on supraciliary edge; tympanlll11 entirci y or pal1iall y covered with smooth skin; dewlap long. lingulate: sli ghtl y co n ·tricted at its base, lower portion with greatly enlarged sca les: ned l appet ~ I.vith enlarged sca le;,; dorsal s heterogeneous, main ly smooth : a lat eral row o r enlarged sca les: rringelike sca les on posterior part or thighs and on tai l base; ventrab varying in size. keeled. slightl y larger than dorsa Is. Dorsall y mostl y grey, some limes pale turquoi se with numerous hrowni 'h indu sion s: pat ag iull1 ye llow i:"h to greenish. also bron 7e coloured to go lden brown (female;. with more brilliant co lours); 5-6 pairs ordark narrow bands; rear margi n orpatagiuJll
l a ppct ~.
[)cscri pt ion
' VL males up to 106. females lip III lOX mm : 6 patagia l rihs: nLl~tri ls oriented upward : no thornlike sca le on s upraciliary ed~e; tympanUI11 rully covered with sca les: dewlap long, slender, ereseent,.haped, with small scales: neck lappets with enlarged sca les: dorsa ls heterogeneolls. predol11i nantly smooth : a lalera l row or en larged sca les: frill\.!e-like scales on the rear margin s of thigh. and on base of tail ; ve ntral,. keeled, aboul sa me silt: as dorsa ls. Bri ght green (males) or gree ni"h to olive (fema les) above, speckled with rt:ddi 'h tu din y brown all over.
239
Agaillidae
with a pallern of broad brown band ' (males and females); patagi lim ye ll owi 'h orange 10 reddish . with 5 dark brown to black bands, eaeh with a miulinl.: of (jnc white dots; coloration and pallern of fel11a les more contrasting and brillianl: lower surface or patagiul11 yellowish white. with 4 black bands (males and ICl11ales); inn..:r ·ide of neck iapp<.:lS mustard-yellow (111'I Ie. ) or du ll yel low (females) at base. tiP whitc; tkwlap mustard-yellow with greenish stripe. on base (males) ordull yellow (rcmales). Male~ with low nucha l sai l and larger dew lap than females. 1-4 eggs per clutch. Eco I OI-:~I /E th o lo gy
D. if. /o/lgihorha live in lowland
forests and hilly regions .
., ~
o -
Oislri b ul io n
©
Fil!. 2.tX
f)/"{/("II IIIIII1III1,·!II\CIIIIII.\ IO/lgi!>",.!>".
Genus GmlOceplwlus
KAUI' ,
jll\ . fn)111 llall! ApOi . Hrlln.:i .
1825
The distribution of this genus is limited to the Sunda regi('n and the PhilippIne Island". Moderatel y large to large agal11ids ( ' VL 90-170 111111 , TL ..t:\O III III max .) \. . ith a dl~tim: t 3ngularhon .. rid Ie bl.:twl.:cn I.:ye and no~e (Canthus rostralis) and occasionall y stron!.:ly raised angular 'upraciliary border. Roth sexe~ with gular and shoulder folds, gular sac. nuchal crest. and \ isihk tympani . laks and the ll1ajonty of 1l:l11ah.:s have a dorsal crest {variahle in IH.:ight even within a population. independent (,rbudy ;.i/e). m:hal crest lI~ually ~iluatcd on a nuchal sail. Ihe lallcr covered with row" of lar!.tl.: scaks. il11ilar rows ()(""cak" un the base orlhe dorsal erest. differenl from Ihl.: remaining dorsal sca les. Head covered wilh small, gra nular. heterogeneolls scales. Oorsab
240
'uma tra and Borneo. Mt. Kin uhalu : Ki au and Poring.
and l"l..:ral:,; sometimes almo;,t equal in sii'c. usually intermixed with large r ones . Ventrais diamondshaped. imbricate. and larger [han dOL al. ngle-headed li za rds o 1"1 ell illhabil the C~1I10p _ of rainlOn.:sts at low and Ill..:dium elevations. where they arc uiflicult to locate . Favourite resting places at night arc lhe ollter end, or thin branches. Field observations scem to indicate thai they do nqt wan der too far awav li'OIl1 running waters . Romeo i. home to rour ~pecies. Iwo of which have <11,0 be..:n re 'orded fro lll Mt. Klnabalu . Ta xonomi c comm ent s: After examination of a larger number of s pecimen s or COIl(J("('plialll.1 /)()rll('/lsi.\· it becamc clcar thai GOI/() CI!I' /W/IIS t!ell::.eri ;Vh'Jllll Y. 1991 is ajllllior synonym ofC-. /wr/ll!/I\is (M .\ 'J rlll ·\. unpubli s hed . sec also 1\ 1I1l' \ & GI{(lSS \l \1\ 1\ 1997: 19 I).
!\gamidac
----------------------------------------------------------------------------Key to the Bornean species of Gonocephaltls Supraciliary border strongly raised and angular posteriorl y ................. ,.... ........ " .. , doria e d()ria e • Supraciliary border round .... ............................. 2
3 A row of enlarged sca les bclow the eye ending above the tympanum (ty mpanum/eye > 0.7); sides of body wit h or without light roundish spot. ; iri s of males bright blue ............................. /iugasler • o row of en larged sc al es below the eye . tympa num/eye < 0.7; sides of body usually with light oval 'pots; iris of males brown or light blue ,... ,................................ ,....... ,.... .......... bomeluis
2 Dorsa l sca les almost of equal size, not in tennixed wi th la rger scales; dorsal and nucha l crest discontinuous in male', females with nuchal sai l onl y . ,.. ,........... .. ,.. ,... ,', ...... ......... ,.. ,.... .... .......... g rfllJdis • Dorsals intermixed with larger sca les; dorsal and nuchal cre ts co ntinuous in both males and fema les ...... ,.,., ................... ,.. ,............................ 3
Remark: Because of G. mjoberg i there i only one female specimen known; it is not included in the key. ISKN
STLCBII\(;
Ta h.2" DistrioUlllllI 01' (i()II(/( '('I'/IlI/II.I · spee l c~ on Bornco,
I GOl/ oceph a /u.\· borll clIsi.\ (Stllll(oI l,Il::-l8) (i'lI/lIc('plia/II,1 doria £' t/OrilW GIJI/(}CCpIUl/IIS ;:rfl//{lis
P I II KS.
(U Kil I.
I X7 1
Typr
B ,)'IlClI
M lt- IIII '
•
• t
• • .
•
-I
• •
Kalimantan
• •
• •
moderate ly I:HgC CO II II('( 'I1/W/IiS (SVL up to 136 mm) wilh a round supraci li ary ho rder and a
1848)
\11111 I 'J, I : 10. 3 I: 1 \1 t.;. \ Il " & (i""" H '" 1<)<)7' 17'1. fig . 122.
(jrwut'cplw/II'\ bona' (III'I\ ,
19,)-1h. 2-15 :
•
Sa ra\\ a~
Dia g nos is
1.(JI,/llnt,1 hOl'lll'lISIS I"C~) Slllill ol lo 1~-1 X: -1 . 1'1. III
Io""alil) :
Bnmci
-1-
1925
GOllocep/UlIIIS hOl'lIellsis (S CIII FG LL.
KlIlahalu
-f:-r"
I X-l_-_J___
(i(}lWt'l'I'/IlI/us /iogasl er (CII ~721 (jou()ce/l lla/fI.,· IIIjo/Jt' lgi S~IIII I .
Sabah
small tympanum ; sma ll coni ca l III weakly graml lar dorsa ls usuall . intermi xed with larger ones: malcs and females wilh a continuou:- verlcbral crc~ 1 compo 'ed o t' lanccolatc ~ca l es.
12.1 /Ian.,>
IHIO(\/H/Wlltifll
D I IVI k
(it ' I IIIR .
IX72
(")11 .
aOcr
\ 'f " I III '
&
111'12. 12)
(i(/IIULL'I,fllI"" 'J(JI'Ill't' II\/\ , VI "lll l I &. 1) 1 Nil R 1<}~2 : IX ;
M \N 1111 ' 19X3: 2(,: \1
\ 1 ,\II ,
19XX~ :
(,(//llIn·I"""II., 1· ..lIil (hOI'll(,(,II 'i,I , .
175. I <)na: 13()
IVh N llI1 I
&
DI '1 "1< 1'1~ 2:
IX (imJ(Jel'I,It"III' Iwllll. (IItll LOI,/I1T11' "d/ll 01
\ 11 1<11
&
~ IIIK(l".
19X.l: 25: I III "1\1<1l & S J"I, R I ~ 19X-1 : 2-1: \ -1.\1,'11 s I
IlU7 ,
M ·" " I I \
Desc ription SVL ma lcs II R- 13(J. fema\cs <)0- 1:10 111m ; TL
malcs 261 -3 10. fema le:- 2 15-'275 mm ; body muscular. with an almost round tail an d limbs o t'nwderat e lengths; sup rac iliary hordcr round; co nlinuOllS, high crest of clo~e l y arranged . lanccn latc sca les (fastened to onc another in olin!:? males) . decreasing in height lowa rd s th~ terminal third o r the body. and ending a bruptl y on the ha se or the
24 1
A gamidJc
242
Agamidae
tail ; tail with low. serrat ed sca les (old males); femal es wi th a high Iluchal crest o f lane co late sca les. dorsal cres t considerably 1m er, decreasing in height substanti all y in the anterior half of the body, reduced to small triangular sca les in the poste;:rior porti 6n. DOI'sal s and lat era ls sma ll , conical. usually intermixed with so me larger sca les on th e fl anks; o ften with a rO\ o f enl ar!-:ed scales paral1.:1 \0 the dorsa l ere t: upper sides of limb. with keeled sca les: scales on throat and gular sac convex. not keeled ; ve ntral s smooth but more or I.:ss di stinctl y convex on the chest (young spec imens up to about 65 mm SVL wi th "ccled ventra ls); diameter of tympan um small. rati o to eye di al11 eter 0 .37-0.59. Semiaduit males brown, oli ve, or green dorsa lly and laterally, with a dark. vary ing. re ti ulated pattern ; adult males predominantl y brownish, reticulated pattern often indistim;t: iri s brown or a faded li ght blue: females ru st -red dorsa ll y and dor '0latera ll y: with ma r ' or less distinct. ligh t. ova l spots: dirty white ventrolaterally, with dark reticulation, or dorsa l ide greeni 'h with an almost bl ac" reticlliated pattern; iri~ hrown; gul ar sac light. with dark. interrupted tripes; limbs and tail banded light and dark. llatehl ings green to 01ive above with a tla rk re ti culated patt ern and 'hort light brown or rust-brown bar ' 0 11 the back . FOllr eggs per clutch 01"22 111m length: hat c hlin g~ 35-37 + 03-05 111m (SVL ~ TL) .
:;:
'2 CD
u
o
t:
~~:S~ ~~""":I~i "" @
Fcolugy/l<: thology hOl'lll!lIsis inhabits primary rai n forests up to altitude:" of approx . 700 111 a.s. 1. They li ve on tree trunks and vines, vcr onen in the vic inity of running waters . In case they arc disturbed, they may also retreat into hiding places on the grou nd . Eggs arc deposited in intervals of ' lbOUI 3 month s.
C.
Dist ributi on Endemi c to Borneo. M L Kina balu : Po ri ng, Ki au.
I{i ghl fro m tup til base G(J//!J('('pha/lls hO/'J/{' II.,i., Fig. 253 A fe lllale;: from " g. Kipungit t. -00 111. near
Poring is layi ng eggs. Fig, 254 Ha«;hli ng from Sg.
Kipungit I. POring.
Fig. 255 Ju venile;: malt: from Poring.
243
Agamidac
GOllocepIUl/II., gram/is (GRAY, I R45) Ihl"I,I"',II, g l'llllt/"
1111
(,I"'.
IX4 ~,
2J'l 'IYp e l ura lil." : Rangoon
errOl I
( " " /(' (('Idlill"" gralld..I , M " 1111 \ & 1)1 NI l R I "J~ ::: 1X: \'h ~111I , 19X"l 26: 1 ii i ~ II \ 1(\1 & " I. II,HI ' II)X4 : 24. M \ 1 h \ Il" 1 9XX~ : 175. M " "I1 \ & J)I W IR 1991 ' 3: 1 AI\ IlJ93 ' 4: \1 ,\LJo.\II , 19l).Jh 237.245 Dia l-: no s i~
A larg~ angle-headed dragon (SVL up to 160 111m) with a rounu supraciliary bordcr: dorsa ls homogcnous. vcry small , not intermixcd with I(lrger scales: dorsa l crest ahsent (females). or distinct ly sepa rated 1'1'0111 thc nuchal crest (males). Desc ripli on
SVLmalcs 1.\-4-160. fcm(l les f 1:)- 1:l7 I11m: Tf.malcs 345 -427, remales 305-365 mm , l30dy more or less strongly cO lllpre s ~ ed . rohust: malc. sllmmcr than fcmales; lilllbs rdatively short ; supraci liary border round: nuchal crest or males composed or long. imbricale, lancco latc sca l c~ on an arched sa il : dorsal
Fig. 2S6 Ylak of (//JIl1l1'l'pJIIIJIIS graudi,l from Sumatra.
244
Fi g. 257 female of (/(J1/tJt'l 'pJIII/II,\ Fi!!. 258 (f(JI/()( 'el l /IlI/II.' gram /is , from Sg, Kipung il I, Poring.
gral/(/is ~omc
from Poring .
days':wccks ol d
!\gamidae
crest clearly separat ed from the nuchal cn.:st. abo consi sting OrJOllg, il11bricate.lanceolah; scales. decreasi ng in height at both ends: ca udal crest limited to small. tri a n ~ ular ~ca l es: females without dorsal erest. nuchal sa il with irregu larly ar ran ~e d la rger sca les: dor. als and lat eral~ ve ry slllal I. holllo!.!enOIiS. ·Illooth . without intermixed enlarged scales: limbs covered with smal l. smooth or sli ghtly keeled sca les: sca les 0 11 the gu lar 'ac and ve ntrum smoot h. Um:k or ma les in various s h a d e~ oj' green to almost bl ack: Ilank. hlue. with ye llow spot ting: gular sac ye ll ow to red. st riped with blue: remales ex hibit a dark band with light bor lers between eye and shoulder: bac k brownish, gn;enish. anthracite. or al most black: light. chcv ron-. haped bands: co loration oj' flank s similar to that of males ; co lour or gular sac li ght with dark stripcs. Ju ve nilc, simi 1::11' to females. but flank s pal e and reticulall:d dark . 3-6 (most 4-6> eggs ( 10-1 I -< 2 1-26 mill) pCI' clutch: hat chlings 32-3 5 + 4H-65 mm (SVL- TL).
Ecology/Ethology gra //di., i~ native to primary ra in rorest~ lip to elevations of I.tOO m a.s.l. , on MI. Kinabalu up to approx . 700 m a.;,.1. These agamids are onen found on trees or shrubs imm(;diatcly adjacent to 'mall ri vulets. rapidly fl ow ing st reanl'i. or wide rivers. Males usuall y ~eek the vici nit y of t(;l11 ale ·. Females, but also J u ven il e~. like to perch on rocks in, or nex t to a wa terbody, parti cu larly during morn ing hour '. Ir in dange r, they leap intn water and di ve out of ;,ighl. De leca ti on also takes pl ace in water. Females mature at ca . 20 month;, and begin laying clutches of eggs eery 1-3 mont hs. Incubat ion takes 75-90 day ',
Genu lIypsiclI/otes
Diagnos is Relative ly large agamid li ;tard ( ' VL up to 1.t5 mm ) with ex tremely large sca les on the si des o f the head in combination with a hete rogeneous scutellati on o f throat and gul ar sac.
M AN lilL Y
&
D ENZER,
2000
Monotypi cal genus knllwn onl y 1'1'0111 tlm:e males from Ml. Kinabalu. difl(;ring from all other Asian specie by extremely large pl ates on the , ides oj' th e hea d in combinati on with a heterogeneous sC liteliation of throat and gular sac.
(/O//()cC'f /ill/II .\·
Di stribution Mala y Penin sula as we ll as Pul au Pinang an d Tioman . Sumatra and Pul a u ias. Nako, and Si po ra . BorneLl . M L Kinab a lu : Po rin g. Sg. Man gg is.
Fig, ZS9 Head ur the Il':lltyp.: or f h'p,II('afOl('.I Alllllhllflll 'I"'/,\' ( m aic).
HypsiclI/otes killuba/Ilellsis (m· GK-lJ s. 1937)
.o K ina· Balli vlollnlaills ,
orlh
I1l11 Il~o"
(,,,lOll" ~1I1t"l(Illt el/'i'\. WI .. ,,' III 19(>7: 39: I'" 19')3: & I III, I\) I 1996: IXX : INla .. &. L. '~ "I t'll) : 1·1,\
2: 01 ,
(al"ln kill: _35 I "1"'/( '11 lo!<' I' klllahalut'll l il . :\'t " ' 111 \ 8:. ))1 Nil" 2000 11- 211
"
I: 245
Aga midae
Fig. (" II Mail:
or I h 'psiCII /ores kil/ aha/llt'lIsi.l'. draw ing alter the ncotypc.
Ucscriptioll Relati vely la rgc aga mitllizard (SV L up to 145 mm ) wi th moderate ly Ion!!, lim hs and tail ; dor 'als scalc ' cqual in size and orientation; latcra ls vary ing in si/.e. shape. and arrangement ; sides of' head wit h an cx tre mcly large scale be low the tympanum (abou t the siLe or thc oc ul ar cav it y), pro hahl y sma ll er in fe male. ; nu chal and , epara te do rsa l crest. present at lea!:i t In males. the lat1er proceeding on the tai l. Males with a distin ctly deve loped g ul ar sac , ante n or edge with long. la nceo late ca les; !:ie.ti es on gul ar sac very small. rang ing in shape from ova l to rhomb ic and varying in siLe; throat \ ilh sca le s di fferin g ex trao rd inari ly in shape and size . Di stinct fo ld on the shou lder: ta il
246
swo llen heh ind it. ha. e. then . ubstan t iall y CO I11 pressed laterall y. I lead and body gree n. th e latter v ith a banued pa ttern of choco late- brow n and blac k spots; tail pre do mina ntl y brow n with sma ll grec n in c lusIo ns. I<:cology/l<:thology One spec imen was discovered at ni ght slceping on the lear o r a bush 20 CIll abovc the ground and 1.5 111 fro m a smull stream. Uistribution Endemic to Mt. Ki naba lu . the onl y prcc ise loca lity known bei ng Saya p. 900 m a.s. 1.
Agam idae
Genus PllOxopirrys
H lJ RR EC HT,
188 1
The distribution of thi::; genus i~ limited to the islands of Sumatra and Borneo. Smal l (SVL lip to 84 mm , hut usuall y sma ll er), poorl y known species wi thout horn-like protrusion: tym panum not vis ibl e. Gu lar sac absent or
very small: low nuchal crestuslIally pre. enl. Dorsa l ere. t absent or indicatcd. Dorsa ls small. heterogeneo us, and intermixed wi th larger tuberc ular scales. Base of ta il in males substantiall y thickened, di . tin etl y elongate and dorsa ll y flatt ened in shape. Rorneo i. home to four spec ies. two of whi ch occur on Ml. Kinabalu.
Key to the Bornean species of Phoxophrys (in part afte r I N(.LR Supraci liary border with spines ............ spinipes • Suprac ilia ry border without spines .................. 2 2 Continuous dorsal crest of low, closely arranged vertebral sca les .................................. lIigriiabris
• Dorsa l crest indicated by several widely spaced vertebral sca les .... ................................... ........... 3
Tab. 25
\)i ~ lr ib lll io n
1960)
3 Gular scales and lateral sca les on the tail base keeled, head of males "nonnal" in siLe ............. .. .. .......... ................. .............. ................ bomeellsis
• Gular sca les and latera l . ca les on the tail base smooth , head of males conspieuolls ly large ...... . ......... .............. ... ........... .......... ... ... ....... cepl/alum
or Pllfl\()/Jhn '.,·-specics on l3orl1l:o.
P'lfJx(}phIJ" ~ b()nl eellsi~
IM.He 1960
Phmopfrrrs .' pillicep.,
( 1' )'11 KS.
S\U I'll.
I fi(4)
I"JGER,
•
•
•
Brunci
1960
Ph"wph,ys honw('II., ;, I Nl,) R. 1960: 222. II!!, . 1.2/\ Type Iuc
i\ small agamio (SVL less than 79 111m) without a
suprac iliar s pin e: ve rt ebra l crt:st composed of wide ly spaced sca les: base of the tail with keeled scales laterall y. Description SV L males 63-66. females 6 1 mm : TL males 92- 10-1 , fe males 90 mm : occi pital regio n and sioes of head wit h conical 't:a les: nuchal crest . hort. consisting
Sarawak
,
•
-
1925
Plwxophrys horneells;s
Kinabalu
-
PIIIJxophrys cephlllllll/ (Mo( 01 'ARD. I R90) Plwxoplll:vs lI igri/o bris
Sabah
I Kalimantan
•
• •
1 •
I
•
of ro undi sh (ma les), or natleneo (fema les). coni ca l scales: gul ar sca lcs keelcd: dorsa l crest indicated by some rai . ed sca les: mal es with a smal l g ular sac; dorsal s and lateral s heteroge neous. main ly smooth , intermi xed with larger, keeled , and some tubercu lar sca le ': ve ntral s homogcncous on the chest, keeled, more h cterogc n coll~ posteriorl y and almost smooth : base of the tai l \ ith kee led sca les on the sides. Males brownish to almost black dorsa lly, wi th yellow, ob liqu e bands and a ye ll ow radial pattern around the eye: throat ye llow with a brO\ n ~ tr iped pattern: ventra l . ides whitish , marblcd wi th brown. Young fe mal es dark brown dorsa ll y with narrow, grecn, oh l iquc bands: some en larged sca le ' whitish: upper si des of limb_ same co loured: two light stripe. between the eyes, another from the eye to the ang le of the moth: sides If head and neck with
247
J\gamidac
some green markings: throat whiti ·h. marked \vith a Ih.:twurk o f brown stnpe!>. Youngs rust-red dorsa ll y with a pallerll u[' light brown to ligh t uli ve bands. upper ~ id es of limbs sa me eo loun:d: two whiti sh bars between cye~. anothe r one fro m the eye to the ang le o rthe mouth : great pa rt~ llfm:ek and sides of head whiti sh to li ght ~ reel1 i s h . Inlllai cs tail base ~lIb s tantiall y widened with 4 rows of keeled slIbeaudal s. Two eggs per clutch. Ecn logy/ Elh olugy P. homeclIs;s inh abIt s preferably shrubs. J'hese small agamids have been recorded from altitudes between 1300 and 1500 III a.s. 1. on Mt. Kinabalu . Distributi o n
BOl1leo. Sabah: Crocker Range (I. D AS pers. CO llllll . ) Sarawak : re\!ioll of' theTrusan River. ;\11. Kinabalu : Bundu Tuhan . Sg. Liwago. Sayap (Sg. Kcmall tis). ~
::> ~
'":r:
Phoxophrys ceplwlulII
" Fi!!. Z(IH Young Ih\:
1 -':11l~ k
(M ocQuARD, I1)90)
or ['III}l O/JIII:,'s hOrllecl!.,i.' rrom
II L'atlquancr,. ' ·"/,lra/"/11 1'>1 .. '.'1\KIl . I X')O : 1,0.1'1 VII: Ii!!, -I , T) I} e loca l il): " Kina B alu" ('Y I1 . of (jlm 'IT/II" I./O/'
/'<,l1l1m."OIlIll
Fil!. 2(. 1 M:l lc or 1'llfIxoflhrl'., 9()O
H\
hIlI'llC"IlS;\'
from Kia u.
-1:1
"II'.
1'('II(lra~lJlIIlI ('t-',dlll/IIIU, \\' 11111' 111 '0 1 ~93 : I ~n
PI",.",,,h/'\s n,/J/"dlll/l, hI" ~ 19(,(1 : ~21 (\,,11l1 'I't·t·I~' J . ~·1 " "" \ I 'Jx', : 211. rill NII"K!) & SJO(,H " , I<)X.J : 24 : T" 1\)'1.1: ~t,\I ~\1t'
~.J 5.
1'J9-1b:
1
7:
2X').li~. ,; ~" Nltll\
l'lOOe 2:
/11
R Olltl
&
(iK()SS\l\"N
1'115 : 9.\;
S\tlII'
I'nl : 22
PI"'.\()flhl'l ·.\ IIlgn/ahn\ . (nol ()1(}Cl'llll/\ (Ja/",/a/'(/) 11i.~ri/I1/'n.\
I',
'l "\.
I XM I T \" I 'l<)~ , X. \1
\1 ~ \11 ,
Il)l)~h . ~-I5
[)iag ll osis 1\ rela ti ve ly sl11all aga mid (SVL up to g4 mm) with-
out a ~ upra ci l ia r spine: \'..:rtebral crest of widely spaced scales: base of (he tail with ~ ll1 oo th sca les lateral l . D('scri pl io ll
• VL males 7 1-1\4. f'e ll1aks 59-7-+ 111m: TI. males 126-140. femak s I I I mm : terminal portion of supracili ary -:dge, occi pital reg ion. 'ides of head. and
Agamiciac
kl\\'erjaw covered wit h con ica l sca les: g ular ~ealcs va rying in si/.e. n.:arly diamnnd-~hapcd. smooth to granular, sometime. intenlli'(ed with almostlubcrcular scales; nuchal nesl or 5-R . epara led. thick , triangu lar 'ea k s of various si/e; vertehral crest indicaled by a few, widdy spaced and ereelcd 'ea les: do rsa ls heteroge neou s, inlermi x.:d with so me larger. co ni ca l sca les: ventra ls s moolh to kebly wnvcx in ad ult spccimens. sometimes kceled in juveniles and semi -adults, larger than dol'. al. ; ha se or tail wilh smooth st:alcs laterall . Grt:en ish to a/u re dorsall y. with dark g reen. oblique bands va r yi n~ in width: hea d orten pakr than bac k; Ihroal whitish with SO lllC dark green spnb; ventral si dcs li ohl . marbled wit h dark . Femalcs dull brow il (W III IIIII AD IR93) with a e llow throat. Head disproportionately large in males relalive 10 body size: hase of tail substan li al ly wi de ned, dorsa lly slrongly fl attened with a seri es 01' hard . mucronate sca les along the edges. lalerall y wi th smooth sca l e~: two rows or keeled subeaudals. Fig. 262 Mal<: of Phorophl'l 's ('('plwlllill
11'0111
EcologylEllwlogy Pho1'01Ih,T.I' ('('p llllllllll appcars In he a ~Ill\\,-Illov
ing li laI'd. Whcn disturbed this ~ pcc i c~ wi ll remain compklcl y motionless . They ma y exhibit a ca taleptic behaviour over a period o r :--c\cral minut es which includes a hall~()pen mouth and a vau ll ed tongue. Durin ~ thi s stage indi vidual s can be manipulated al will. These lizards inhab it !> hru b~ and trees in rain (01'.:sts (, 500 lip to approx . 2100 III a.s.I.). Re mark : Int ere~ til1 gly. thc two specilllcn . which subsequenl ly bet:a mc the Iype spet:imc ns were caughl by nati ves in ral -Iraps positioll ed on Ihe grou nd . This Illay indi cate a more terrestrinl lile than Illo ·t ot her BOl11call agaillid linll'ds lead .
Oistri hul io n Saba h: ·unsuron. Sg. Mesilau. Ranau. MI. Kin abah. : Kin u. Kcnokok. Tenoll1 (l nk, LUlllll LUIllLl. Bundu Tuhan, Kamborangllh Road. Kamborangllh . Sayap (Sg. KCllla llti s).
Kamborangoh Road. 1750
Ill .
G;
'""
(J)
I (I
2.tlJ
Anguidae
Anguidae The family of the anguim: li za rds is organized in Ihree subfal11ilies whi ch inhahil Illilinl y the tempera Ie and subtropical region, of the northern hemi sphere. hut arc also represented by some spec ics in south America . Onl a fcw ~pec ies arc kno\,vll from oiliheasl Asia. Apart from pnmill ve four-legged species. various transitional stages kad to more advallcl:d kglcss lonn~ with himll y allY rcmains o r a girdl e skeleton . A II spec ies arc considered to be ca rni voro us. MOSI of them arc (lviparou:" but some givl: birth 10 full y deve loped yo ung. As IiiI' as Borneo is concerned. the fa mil y is represented only by the genus Uphi.\'(///I ·//\· (g.lass li/ards) of the subr;lmil y Gcrrhon(Jtinae whose members are characterized by the abse nce o f limbs. They arc distinguished fro m all othcr legless rcptil es by the presence or lateral 1()lds.
Genus Oplli.\'ulIrtiS D ALO IN, 1ROJ The dl~tribution o f thi~ genus is di sjunct with repn.:scntatives in the southeastern USA. parts of northern Africa, sou theastern Europe. West Asia. and Southea st r\ ~ia . Comparativel rig.id body with slrong latcral folds; no lilllh~ . I lead slightly pointed anteriorly and hardl y di stinci frolll the neck. Fyes \\ith movable eyel ids. car-opening present. Tail fragile with the ability to rcgene rate. Large. rhombic sCil le~ in regil lar 100lI.!itudinal and transverse rows, attached to ossi fied plates . Lateral fold s ex ten s ihl e, with s mall . unossi fieu sc; de ~ adding to the animal's rnobilit y. Only one species is known frol11 13oll1eo and has also been recorded from Ml. Kinabalu.
Opllistlllrtls bllftikol eri Lll), 11i 1)1 ' h UDL, 1905
sals in 16-1 g lon gitudinal an I lJX-1 05 transverse rows. central 12 rows of' dorsa ls kee led; ventrals smooth. in 10 lo ngitudinal row s; a ll eaudal s kee led. Coluur in alcohol (after I»: RoolJ 191 5): " ArOl n ahove; a dark latera l hand edged above with a li gh t band: thi s latera l band is continued on the tail. ;\ small blue spot on the interparietal. indi ca ting the loca tion of the parietal-eye . On thc antcrior part of the back irregular tran sverse seric. of blui sh . pots, anteriorly edged with black . Lips and lower parts pal e ye llowi sh; five oblique dark I in cs on the side of the head, runnin g across the lips. the lirst behind the nostril. the second below the eye. Ihe third at the corner oflhe mouth . the fourth below the ca r-opening and the tifth at the nape : thc latter is con tinued on each side or the belly."
Oi ag nosis A moderatel y large li zard of up to 125 mill SVL. charaeteri7ed by almo:-t equall y si7ed front al and interparietal sh ields which form a wide sulure.
Eco luJ.:y/E lhology Thi s anguine possibly lea ds a sec reti ve life benealh leaf litter ur in Ihe so il. Only three spec imen. have becn found so rar, o r which one wa s di scovered 011 a path through primary rai n forest at about 300 III altitude. The :,toma ch of' another spec imen co ntained the remains of a cock roach and fragml:nts o f the leg o f a rat. Such rodent would. howeve r. be much tuo large luI' thi " liLaI'd , 0 it may he safe to assume that the leg was torn 01'1' a dead rat.
Oescri ption SVL 11 9- 125 mill. TL 375 111m; frontal and interparietal 1<:>1'111 a wide suture; 5 s upraoeul ars; dor-
Distribution Thi . species is endemic to l3orneo . Mt. Kin ablllu : without preci~e localit y data .
')I "')1,
O"h"'"Il'II ' 1i1l1l/~"f''I'I I . lll/ll .II 1')(J5 : 192 T~' rc 10\' alit) : ~·t "lIl1 ' I /H llg K(I~hocl1 g , Ccn"al - Il<1l'11co ( Bllk" I lalll' Kuhll lll!. UII ."" SungJI Karll''', \'ie" Ka i lin alitall) Opili l'tll//'I/\ ""{'IIIA ol en . I (,' K 1'I 5 ~d - 47'1: M \N " II \ 19XJ . 27;
!-til
250
NII \K II..I:: 'i JIJ" RI '
1'I~4 :
_4 . M u"."
,
I<)'I4b: 246
Gckkonidae
Gekkonidae Geckos have a cosmopo litan di stribution inhabiting all tem perate and tropical regions. Often they arc the first reptiles to invade newly formed islands (e.g. from volcanic eruptions). They are sma ll to modera tely large (total length up to about 400 mm) . mainl y crepuscular and nocturnal lizards. typica ll y wi th a granular skin stnleture. sua ll y they possess \'ertical-oval pupil s as well as adhes ive lamellae underneath their digit · (subdigita l lamellae). The armngement and shape of the 'e lamellae are impOl1ant fea tures for the identi tication of . evera l genera. Adhesive lamt:llae enable gcekos to climb on seeming ly smooth. vertical. urfaces. Geckos arc known for a highly evolved autotomy and the abi lity to regenerate lost tai ls. Regenerates arc however different in skin. tructure and shape. Geckos inhabiti ng Borneo u 'ually posses (origina l) tai ls which arc longer than the snout-vent- length . In co nt rast to most other gro ups of li;tards. many gekkon id lizards arc ab le to produce clearly audible sounds which onen arc indicative of a spec ies. They feed on a large variety of invertebrates and severa l species exploi t the abundance of insect. att racted at night by electric light. Many gcckos adapted to manmade altera ti ons of' nature and liw in and around human habitations. All . pecie in the range reproduce by laying eggs which Illay be attached to a surf~lce . Produc tion of several fertilized dutches alier a single Illating process is (Amphigonia retardata) is not uncommon among geckos. Borneo is home to I I genera compri sing 29 geckonid species with R genera and I () species occurring on Ml. Kinabaili .
Key to the Bornean genera of Gekkonidae
(in part atier
M.","THFY
& GRU
SMAI\N
1997)
True eye lid s presen t (A). body co mpre ssed latera ll y .. ..... ...... ... .............. ... ... Aeillroscalabotes • Without eyelids, body compressed dorsoventrally
........ ... ...... .. ....... ... .. ........................ ..... ............... 2 2 Pupil vertica l (B) ............................................... 3 • Pupil rou nd ........................................ Cll emuspis 3 Fi ngers dilated (e) ........... ...................... ........... 4 • Fingers slender, sli m. curved (D) ............ ........ ... . ... ..... ... .... ...... ........... ..... ................ Cyrtodactylus 4 Inner fingerltoe we ll -developed ....................... 5 • Inner fingerltoc rudimentary (E) ...... ................. .. ...... .... ........ ......................... lIellliphylloducfyills 5 Inner finger/toe clawed (F) ............................... 6 • Inner finger/toe without claw (G) ..................... 7 6 Body and tai l with lateral sk in fringes (H) ....... ..
E F
G
~.
....................... ......... ............... .......... CoS-,'lII bO(IIS
• Body and tail without lateral skin fringes ......... .. ...... ............... ....... ........................ .. J1elllidac~lIlus 7 Body and tail with lateral skin fringes (H) ......... ......... ..... ....... ... .. ... ..... ....... ....... .. .... ... P tydlOZQOII
• Body and tail without late ral skin fringes ........ 8
25 1
(id.konidae
H Subdig ital lamellae divided (I) ....... .... ......... .. 10 • Subd igital lame llae 110t divided (J) ................. 9 9 Pai red sublllaxillars present (K). median row of slibealidais en larged (L) .... ...................... . Gekko • Subm
Spec ies or ( 'l'l'/nr/lf('tl'/II.I' occuLTing in the SUllda region inhabil predominantly pnnwry rain forc , ts. but ;,;on1<': ha ve a l:;o adapted II sc<:ondary and bu~h rorest~ . They normall y livc at a lt itudes bctw.;cn ~ca- Ic \'el and ap prox . 1.)00 1\1 , Oil tv!!. Kin abalu up to 2500 1\1 a.s. 1. T h.;sc arc stridly nocturna l gee k()s which o ften leave their she lte rs only a Ii.: w hou rs aftc r ~undOl\ II. They arc fo und o n the j'o rc~t floor. Oil rock '. in l: ave~ , or Oil and in tree trun ks. \cry onen in the vicinit. of runnin g water. Man y s p ec i e~ are able to curl up their ta ib laterall y. I hey Iced t lll a vrl rie ty of in vc rtebrat es. 1-''; \11<1Ies pro duce c lutdle~ oi'u:-.ual ly two roundi:-.h hard-~helled egl.!s. in some spec ics the eggs arc
Ge nlls Cyrfot/{Jcfylus GR,\\'. I '27 (icd. o~
of the ge nu ~ ( :IT/()t!a('~r/lI,\' an; di stributed frol1l SOllthca~t A~i ; l to t\e~\ (iuinea hut abo occlIr in n() rthea ~tc rn Australia and nil the So lol11on Islands . I'vlainly l1lodcratcl y large lizards (SVL of ROlllcan spcc ic ~ up In 125 111m): hody cOl11pn;ssco oorso\'enlrall y. \\ ith g round eo ltlur usually be ing somc shack of browl1i sh: d(lI'sa l >. url;lce marked wilh highl y variahle. dark spott ed or handed path; rn~ . I lead acutetriangular in ~hape when vic\\ed li'olll above. and u~l l ally very distinct from the nCl:k. Pupil \'ertiea lova l wit h l:rena te edges. I-ingl:rs and toes clawed . ~Ic ndcr. 110t dilatcd. their arc hed terminal phala nge~ eom presscd latern ll y. Vcn tral s urfan; of basal phala ngc:-. with enlargeo plates or lalllel lae ill rOlvs. Ma lc:-. usuall with preanal alldior i'cmoral porcs. somc species with preana l gr love>..
Tah. 26
Di ~ tribllci nl1
of ('1 I'/iI"{/ (' II ·III"
-Sr<.:<,; i c~
~ pindk ~ h a p e d .
or the c ig ht :-.pec ics kllown to inh abi t Borneo. four ha\'e beel1 rel:ol'<.kd fro m Mt. Kinab alu . C:l'r/od(fc/.l'/us is th e rerore thc gc kkonid ge nu s wit h the greatest s pel: i fic di ver~i t y o n thi s moull tai n.
on i3ofl1en. Sabah
Q'rl"d{[(·~J'llI.\ 11/I11/(,11,\';~ ( l\l '>t()I I.\RIJ. I ~N O)
( ~I'/'lIJ(I(/{ ·/.I·II/' ('(/ ''<'I'III('l}lus t MdR
&
1-( 1"1 ••
()'l'Tot!u('(yll/,\' (,(1/I.\'(J"r;III/.~ R,. 1'1< 7 1) Q'I"OIlflCfylus
;11 '(" .;
111,,111.\. 1l)()O
(:I'I'todl/( ·/.I'III., /ll1I/1I.1·W/II., ~ R OOlI. 1915) Cyr'()d{[('~J'II/.' IIIl/lslI;; 111,,11>\, 1990 ( :I'I'/orfl/('/.1'I/I.I
252
I M,IIL 195'1< 11 11-.\1 ". 19911
pllln.uIt ·II'
( "'I'/IJdll('/\'/t,,' ' ·IJ.,hi/
196t
KlI1:1batu
Bnll1cl
• 1- •- - - • • - • . •• • • - • • • • j'
•
Saruwa~
Kalimanlan
• - --• - --• • •
•
C;ekkollidill.:
Key to the Bornean species of Cyrlo(/actyills (in part ane r Light narrow net lVork pattern on head and light narrow bands on the hack ................................. 2 • Colour pattern di ITercnt from above .... ......... .... 3 2 With enlarged femora l scales, ma lcs and females with 9- 14 preanal pores fornling a narrow angular series, male. with 0-6 femoral r ores; subdi gital lamellae benealh to urth toe 22-28, supralabia ls 1016; SR-65 ventrais (al mid-body between ventrolatera l folds) ................................... C(l1I.m briIl I/S • Without enlarged femoral sca les. ma les with 810 preanal pores fornlin g a wide angular ~eries, without femora l rores; subdi gital lamellae 2 1-23 ; supralab ials X-I I ; 5X-62 ve ntrals ... lI/alu.I"U/lIIS* 3 Su pral abial s X-l). no femoral pores or enlarged
fellloral sca les. male~ with preana l groove containing two pairs of pores; subdigitallamellac 222(,; supralab ials 8-9; 5 1-5R ve ntrals ........ ........... .
IIi KII)A 1990)
Male~ with preanal groove cont aining 3-5 pai rs of pores (7-9 tota l); subdigi tal lamellae 17-22; supralabia ls 10- 1 I ; 43 -55 vl.:ntrals ... fJllbiqdclIS • Ma les without a preanal groovc ....................... 6
S
6 Back with a pattern of dark blotches; male with 7-8 preana l pores forming a narrow angular ~e ries: subdi gi tal lamell ae 23-28; supralabial~ 1012; 40-43 ve ntrals ..................................... ;1I~ eri • l3ack wi lh dark cross bands .............................. 7 7 Ma lcs wit h 7-8 prca nal pores lorming a widc an· gu lar series: subdigita l lamellac 22: supra labia ls 10- 12; 4R-S I ven trals .... .. ....................... lIIat.m i; • Ma les wi th X-12 preana l pores rortn ing a narrow ang ular series ; subdigita l lamcllae 25-30: suprala hiab 10- 14; 5()-5X ven tral s .......... . .ro.\'hii
C(I I 'ernicoills
• More than l) supra labia ls .................................. 4 4 \-lal es with femoral pores (4-9 on eaeh side). enlarged femoral scales and 9- 1 I preanal pores form-
ing a narrow angular sl.:rics; subdigi tal lamcllae 20-23; supralabia ls 10-12; 40-45 vcntrals ......... . .... .. ... ... .. ..... ... ..... .... .... ......... ... ..... ......... blill/ell.I';s
• Mal cs without femoral pores or I.:nlarged sca les .................. ................................................. .. ..... S
C:rrlo(/adyills baillellsis (MO<.()1 A IW, 18(0) (',n'lIIodc/t tl'll/\ /Jull/t 'n,,, \-Il n 1.'1 \I~I). I :--NO ' 12:'. pi VI I· fig . I. la. Ib, it' r~ r{' tnca lil ) : KlIla Balli (,I 'IJIJWr/U(, (I II" 1t("11 ('/1.\ 1.\ , li lll 'I I Nit! I( I Sl)-ld ' 7 ~2: ))1 i{ nc lIl 1')I:i : II. ~~1I111 IlJ .>I III. II. 2 1. .11 : \I " 1111 \ 19X3 c~ ( ' ,,,,'odClUI " '" halllt'II\;\ , I-~HI f\,1t\l{lt & ~1I1(,I{I- ' l'/X4 . 21 . 2-1 . ~ I \I '~1' ·' IlJX1',, : ~'i. 1'l~'1 . I·) ~. 111'11>\ 1')')0. ')1 . t\ 111 "11 " 1'191 a : .13. 1'1'11 to: (,. II~ . I, 1,1. 1'I'I2a 1211. 1')'1-111 21~ , 2 ~5 . 199(H;: 2XX : :\1 '1\111 1\ 8: (11<1', ... f\I'" 1997 2 19. Jig. 1~ 5 (,oJln/cwlI '/tI,\ ha/J/( ' I{.\/\ . r \N I ~)ln "' ('I 'mllot/". 11
hl\
1II0rl]loralll\', tllol ('1J'10t/W/I/II\' mWJllOIo/lt.\
(,1<\I . I ~.11 ) 11 11\ 11 '" II l'IOOa: 70. IC)()().: : I. III RII.,II I') I ~ 1.\ (parl.) , ~ \ 1I 1 11 1'1.11 : 11. t\ 1" 1111\ I')S, 2<1 ( ' l 'l/otltl (, (IIII ,\ IIIt1F1ltortllll\ . (1101
(, 1<\1, 1 ~111 1 1< 1" 1I1"1I&
I.I 'ml1uc/1I1
'l/lI\':-'P J \lIJH 'O
( 'n '/odo(/yllf\ I/UU 'mOl'lllll,\
:-"' " 01<1' IlJ~-1 ' 2-1 Il)X~ _ . .L lig . "'~a
* The occurrenec or C. tionab le .
IIIlIlarwllls
on Borneo is ques-
Remark: A sati sfactory identi fica tion kcy which lakes both Ill alc~ and fcma lcs into considerati on is cu rrenll y Oul of qucstion !.inee fema les of 1>omc spcc ies are st ill unknown .
Diagnosis .1\ moderate ly large ( '\,ulldIlClrlIII wit h a maximum SVL of XC> mm : no pn:a nal groove; ll1ale~ II illl 9I () preanal p() re~ arranged at an acu te angle in a
shall\lw (leprc~ sio ll . and (1-9 ~e p aratcd kmoral p o rc~ on ei ther s ide or the li1nncr; -10--15 rows of \cntral~ at mid-body bel\\Cl.:n ventrolatcra l fold s; 2 1-23 subdigital lamellae bencath li llirth toe . Desc ription
SVL lip to X(, mm ; tail ,l ig htl y longer: nod y s lende r: dorsa l sea lc s linl.:, inll.:nni~ed wilh larger tubercl cs in 2 1-2 -1 irregu lar ro\\'~; \'cntro latera l ii) lds wcakly dClcloped; -1--1 5 rows oj' small. roundish. im brieatl.: vcn lra ls betwl.:en \ cn tro latc ral fold~: l"O,tral broad with a central g l lll)\ e, in cont act II ith
253
(ickkonidae
nllstli I, ; I 0- 1~ ,tlpl !labia Is. <)-1 0 infra lahials: mentaltriangular.lililowl.:d by 2-3 pairs o(',ubmax illars: I{)urth tOl.: with 20-2.\ ~uhdigital plates. largest on basal phalange : tali dor~ally with ~ mall ~calc~. intermixed with tubnck, : its basnl porli on with X ~cales and 2 tuhercle~ per sl.:glllenl. Males wit h ()- I I prea nal pore~ arranged at an aUlle angk in a shalllll\ d ep re~s i on (-1-6 per si dl.:): pI' 'anal porl.:' widcl . ,eparated. III largl.: kmoral scales, 4-<) Lln each Ihlgh: IClllall.:s \\"itlwut pores. bil l with a selies or enl arged scales arranged at an obtllse angle III a Ilat prea nal region : males with 3--1 pairs oflarge. IC lllale ~ with 2-3 pair, of sma ll postanal tub e rcle~ . Iklge to yl.: I I(l\\"i~h bnl\\11 dtm;ally. with irregular. dark brown Sp(lt,. waved or crenated ham!'" or ~ h ort. broad. longitudinal da , h c~ : head onl.:n with a darh. hrown . almost V-shaped lal eral hand fro m the tiP of snout to the nape: limb, and ta ll handed In li ght and da rh. brtl\\n abO\e : \entral surl:lee alillost unililrml y violel grey.
Two hard-she lled. roundish eggs ( 12 '- 15 mill) per clutch : hatchlin~~ .\ 1-32 I 3-1 -35 mm (SVL I T I ) [ H IKIIl\
1<)<)0).
EC(l I ()g~' / F. t hology
usua ll y inhab its montane fo rest, at altitudes ()('()nn 10 I XOO m. up to 2500 III a.s.1. on \-1 1. Kin a balu (photographic records K
Uis tdbutinn (:1 'r/or/o(",-, -tIlS halllells;s
is endcmic to l3 0mcn (Sabah. Brunei) . ;\'11. Kin a balll: Poring. Kiau . . g. Kcmanti s. KenoknJ... Uundu Tuhan . I.lImu LUlllu . ['(liver Station. Kaillborangoh. St!. Me ila u. Headqllarkrs regIOn. and on the Summit Trail up to 25(1) 111 a.s .1. (K'l()\\ I I S pers. comm.),
254
(il:kkonidal:
Cyrtodacty llis
COli Obrillll S
(PI'.n .R. • I X71) ""II.\Ohrillll' Sarnwak. Ilmncn
(;\'IIIIIOdll, ' {I '/IIS
il) :
1'1 11K' (
(X7 1): 56'1
r) I)~ I''''a l-
I ,)~:"d C) . fig . 5. I I. 1991 c X. lig. 1. I<)l)~" 129 .
( -\'r /(}(/(I C 11'1", 1"(111,\0"111111\ , M\I h.(o.II '"
II l~ II "
1990' lJ3 . ~h l 199-1b' 2-15
\ 11 "
hlJllIlw/IIc..'",n 1\1,\1 "f\Il I.... Il)X<J 196 T) I'l' Inca li!) : " W:ildhJch hillier dell Ilnl Sprlng\ (h()() mI" (,yn ancr 11 1"")\ 1')')0: en)
Ui ~l gnus i s
A large s p ecie~ of CJ'I'/{JdUl'll'IIIS wilh a ma:\ imum SVI. of 125 mm : no prea nal groOVl:: maIL::- wilh 9I I preanal pores arranged in an obi usc angle. and 0-6 ICll loral p o re~ per Ihigh: 65-70 row~ of ve ilirab at mid-body betwccn ve nlro lalera l fo lds: 22-2t1 su hcl i),! ilal lamellae bell ea lh fi)urlh tne.
Ul'sui pi ion SV L up to 12<; 1I11ll. T L lip III 16.1 111m ; bod Y sic nder. bill nluscular; body scak" slllali. intermi'l.ed
Iii:.:. 264
('I
ri'"/(/( /1 ·11/ \
(,1II{ .,,,lInlll/\
with oundish. wcakly kl:l:kd tllherdc ~ in I X-20 irrl:gular rows; \.l:nlrolail:ral fllid Wllh lar),!l:r lubl:rcles; small. roundish . imbri cate \' c nlral ~ In 65-70 rows at mid -body hetween \cnlrolatl:ral I() ftb . Roslral large with 3 ecnlral grom e. quadrangular. in contact wilh the no~ lril s; I 0- 16 ~ up rala hial s. 9- 1J infral abials: l11enlallriangular. !(lilowed b. one pair (l11ore rarely 1\\0 pairs) of large s ubllla'l.ill ar~ . Fourlh IllC with 21-2X subdigital lal1lcllac. largcsi on basa l phalange; lall roundish In cross-secllon. tapering 10 a poi nt . wilh sl1l allll at ~calc~ abn\'e alld 4-X row~ or kecfed luberck~ at It S ba~e . Malcs with 9-1 I prea nal pores arranged at an obtuse angle (~-6 per ~ide). and 0-6 i'cll1oral pores on each thigh. and ()- 7 enlarged femora l ~ealc~; no preanal groove; 2-3 palr~ or large pll ~ lall ~tI tll bcrc1 e~; rcmalcs wilh enlargcd preanal an d femoral seaics. but no pore~. and 2 p a ir ~ or poslallallubcrclcs. D(lr~al sur liJce~ rich chOClllall:-bnl\\'n Wllh darkcr ,eetillns and ~ca lt e re d light dots : ~-X. u~ually indivi duall y ~ h apc d . crea ll1 -co loured. \\ hite or ycf lo\\ bands or cro~s lines; dorsa l ~ ur ral:e of head wll h a
{'rom Poring.
255
( ickkonitiae
eilaraclcn, ll c. Ind \ Idual. li gh t network or 11I1 ~~ : labia Is spoiled \.\ Ith li gh ter co lour: limbs and tail wilh naITOI\ lig ht hanlb: ITnlral ~i d e dirt )' \\hik . belgc. or reddish grey: jllvcnde~ co loured and pal lerm;d likc adults. hut ~uhslan li ally more conlrast ·d. Iwo eggs per clu lc h; hatchlings J I J I mm ( ·\ ' L+T I. ).
(vr1o{/actylll.\' illgeri
Dist r ihut iull rcn i n ~u l ar l'vIala y~ia.
Sumatra, and lIorneo. :\11. h:in aha lu : ('orln g: Il ot S prln !.:~, Sg. Kipun g ll I and II.
256
1990
'rllI\
111,1:('111 11. 11>\. I ')()o 1)(, . lig. 7. N "1)11'>11....,.,1P'''rlng I i ol ,·Pllng . Ranal! ()lstriCt. S,lh"II" nrflld,l"rlul' 11"':"/1 ~hlL\I '" 1\)\)1," X. lig Ih, ~- 6. I"'!~a .
('1'110.101
it~ : " ncal
leX
19'1411 211.2-1 'i, I'J'!h,"2XX
(, o/ndillfl ('I
f:c ul ogy/ Et hnl ng)' These get: ko~ live in pnmar~ rain rorest~ of' lhc 1(1\\ land ~. Tht: prctcr to ~ I" y ncar the g round (up tll appro:-- . I nlelre) llil Iree tJ'llnk~ . between plank root~ , or In holes 111 the tI' unk ~ or large. Sillolltil barh:d trecs in Ihc \ IClnil Y or inlillethatd y nex t to running wata~ . They u~Llall y becomc aetivc onl y lI'e ll inlo Ihe night. They reaCI \ l'l')' ~ellsilivcl y to ligh t and will escape IInmedlalely when disturhed by a torchlight. Felllale~ pn1tiuec dUlciles ~eve ra l limes per year.
11 1"-11),\ .
h/\
UIJ.':i
nT"
199-': .'
r/()t!ui fl 111\' /""'/\111<'(1' I Ill11
19'iX ,
MIl,.",
( 't/'Iut/l/(
11/11' /mhl,qi/j
/1\
1\(.1 It
I'IXl) ' 19.'
I) iag n os i ~
f\ slllal le r ~pccics o f (1'l';ud{/C lylll.1 with a m:l XI-
Illum SVL. of RO 1I11n: no prea nal groove: males wilh 7- X preanal p()re~ 31Ta nged al an acute all!!lc in a depres.-i < n; no fL:nlllral porcs: 40- -1 3 row~ of ve ntra ls al mid-body belWL:e n the ve nlr()l alCral folds : 2.,-21' ~ uhdi g il;d lamellae nc ncath fourth tne . I)t'sc r iptio n SVI up 10 xo I11tll. 'II up 10 100 Ill I II ; body s lender. ~ Ii g h l l y cOJllpr~ssed dors(hL:n trall y; sea lc ~ smal l. intermixed II· ith 17 irregul ar row:- 01' larger. cunical luhercle~ : vL:l1lrolaleral rold~ wL:akly indi cated by a !tm. rounded tu bercles; -10-..13 rows of
(iekkonidae
and on trllnk~ of'young tree~ in the \'Icinity orrunning waler and I S often seen on bran 'hes 0\ erhan!!ing a stream , It can also be encountcred on bu~hes. rocky clil'fs. and on lhe lower pnrtiun~ of bigger trees up to heighb or appro'( . 1.5 mctre~ , When di~turhed , indi\ idual ' produce a hi~~ln g ~Llund Ilhile l1ccing. They hnve al~ll been ob~erlcd flipI ing over the entire antcrillr bod) including the nnns in a way that the body is hanging down rrom the Iree trunk at an anglc or approximatel~ 60", They cling to the ~lIrf;H; e only 1\ ilh their hind limbs and Illay maintain lhi~ cataleptic pO~lIion for ~e\ end minute~ ,
Fi!!.266 I lnder'llk llr CrI'lOdl/ Cl r /lI.1 illg <'ri rrom 1'01'-
Oi Slribllliun Endemic to Hurnco . 1\,11. I..:inahalll : Poring. Sg , Langanan , Sayap. Sg , \Variu. Sg, Sasapan.
Ifl g.
small. roundish , imbricate \entral~ at mid-bod) bel" een \,cntrol:1teral rl1 ld ~ . Rostr:11 large. quadrangular. in contact with no~ trib; I0-12 supraI3bial~. ~ - I () inrralabials; mental triangular. followed by 2 pair~ or ~lIbma\illar~. 2.1 -29 subdigital plates heneath rourth loe. largest lin basal phalange. Tail round in cros~-~ection. with ~mall scaks above and rings or tubcrclcs. I 1-12 rows or granular ~cales bt:lllccn rings at tail hase . [\'1ak~ \1 ith 7-9 pn:anal pore~ U-.J per side). arrangcd at an acute angle In a deprc"sion ; no Il:mo1',11 pon:s, no large rellloral sc;lIe~. no preanal gnH)\'c. Dnr~al Slirfi lCC
oehre tn sand-colol lred or light grey ith a slight tinge nf'reddish. \\'ith 5-6 dark brown hand~ ~l1mel il11c~ rragmented to liml1 large paralerkhral blotcllL' ~ : neck ol'lcn 1\ ith a dark Y- or Vshapl'd marking; dark Slrlpl' rrom posterior edge of' eye often e'(tending to the in~erti\ln or the arlll; dor~al surf~l ce nr l illlh~ lighl brownish \Iith dark ~peckle~; d{)r~al ~urli lce or tail 1\ ith more or Ic~s \1 Ide, dark bands: \ entral surf'ace~ II hitish grey, ~l1llletimes greyl~h \ il)!.:l. One egg (),5 ' 12 111m pCI' clutch.
Cyrlo{/(fctyills lIlil/.m ;; ( "1'11,,/' 1('/1 -/'"
1111 /1 '1 '"
11"",,, 1')')0
II IKID,\,
tOO, lit'
1990
II . 12
J'~ Jll'
I"l'alil~ : " mlhl' I'ar~ I k"dqll.II I" , "I ~ mabal" N .II'''nall'ar~
.111(,00 111m \11 ( r r /OlI,w / d'l\
~1 \1'" 111' 1l)l)1l;
7, Ii . . " 2. 1 { )9~.1 127.
IlJ%r 2xl'
1' )')·lh : 2-1 5 (, ullrducf l 'lu ,
K,n.,nalu. '-,;,b"h "
lI1 a { \ tl ll
11 /(1 1\/ 111,
I
\"J IlJl)
1. .; ;
Oia gnosis ,\ large spcl'ie~ 01'( ·rr/{}(IIIC/I '/i'l . S\' L about 10:" Inl n; no prcJnnl gro\lle ; Illalc~ \Iith 7 preanal pure, arran~ed at an oblu,e angle; no Il:lllOral pore~ ; 51 row, of'ven trals at mid -hody hetll'een venlrolatcral rold~; 22 ~uhdigital plates beneath liHlrth toc ,
II
Ec()I()~~' /E l holug~' Thi~
gecko !.:a\'l'~ it~ day-~hl'lter ;dready with Ihe onsl' t or du~k , II appear~ 10 prefer dwelling on kal;"
Description
SVI. lip 10 lOS mill. TL ~2 111m (regellcrated): bod) s!.:ndcr. slightl) cOlilpressed dor~o\entrally ; scale~ smal l. intcrtni'l(ed \l'ilh larger tuberclc~ in IH irregular rows: a weak lv in!licakd ~ki llf'() ld c()n~isting or low, l'I)und tuberclc~ het\leen a'\llIa and groin : 51 1'0\1 s uf' 'Inall. roul1lli sh , imbricate \ entra I, bet \1 cell ventrolateral folds at mid-bod _, I
25 7
( ickkoni
Frmll lOp 10 basl' Cl'rtudllcll'i//s illll/.luii
Fig.
abuve, and r i ng ~ of low lube rc les: 4-7 rows ur g ranular ~calc~ separali ng Ihe rings al base oftai!. M a le ~ wilh 7 prean al pore~ (J-.:I per si de ), arranged at an obtu se ;ln~lc : 110 Ii.:IIl0rai pores, no prea nal
2 (,l~
fro m Sg. Sd;IIl-Sdall , I ·450
I'llrlrail
Ill .
f ig. 269 Vcmra' vicw.
Oistrihutiun Endcmic 10 Borneo. :Vlt. h:inabalu : Ilcadquart crs region ( 1-l5()-160() 111 a . ~ . I.) .
groove.
Dorsa l surface ye llowi sh brllwn wilh hig hl y irn:gulad y arrallged and shaped dark brown spots and band s on back: head with ~mall dark ~pol s; a dark band from Ihe posterior edge of" the eye sometimes ll1eellllg ib 'lllinterpart un the nape; limbs a nd lail wilh dark band s; ve nlral sur!:lces withollt any pal krn, Il g hler thall abuve. Ec oIU I! ~' /E tholo g ~'
The lew speci men'> rOllnd as yet were di scovl::red more than 2.5 metres ahove g round on Iree Irllnks (the type spee illlen al 7-X 111): unly one male wa s e ncoulll c red (I n Ihe ground (M \1 ~MI IS 1996c) .
25X
GeIlU ,
Gehyra
GR ,\ Y ,
1834
The di stributi on of the polylypic ge llus (Jehrm ex lends from "vladagascar and islands of the Ind ia n Ocean, thro ugh A ~ia, I he Indo-A uslral ian A rcli ipelago , 10 Auslrakl a nd is l al1d~ or the Pa -ilic Oc~al1 . PllPlll; lliolls now also exist in Mex ico and the USA duc to introduction _ Smaller gcckos with vertical-oval pupil s, and dorSllVentrall y cOl11prc. ~cd body ami tail. Skin fi llel y gra nular. sc ns iti ve tn Ihe tnlleh. ,lIld tears easi ly. Fingers a nd t()e ~ oceas ionall y webbcd. broad at
(iekkonidae
their bases. with well-developed . divided scansory pads below. Outer four tingers and toes devcioped normally. with thin . erect, clawed termin:ll phalanges; inner fingers and toe ' also developed lIorma lly. bllt without free terminal phalanges: furnished with tiny. onen concealed daw~ . Maks with pre:lnotcmora l pores (i .e. preana l and femoral pores in a continllou ' series) . Species of the genus Gt'lirm live as commensals of mall in large ci ties a~ wdl as in forests . They Iced on a large variety of" invertcbrate~ . Females produe e~ clutches or two hard-~helled eggs each several times per year. 130rneo is home to only one species which has also been recorded from MI. Kinabaili .
Uelty ra IIlliti/ata 11. 'Ulu /w ,,11t,\
(Wl\ i(j \W-J"\J,
lIIull/lIllI \ \Vllob\ I ,\NN.
I X1:' : 21X
I ~35) 'IYIl t' Inca l-
il) : Manila. 1'1""1111111C' (;" hl"rtl llllili/(I/ti . M,,,,), ,,<1, IX'l1l IJX . II \ N''''11 l'lOOa. 711. IV" I II" 1 l)~J -:! .j : 1 II INI I "(1)& <; "".~I ' I<JX.J 24. M " " ' '' ' 1'192:1: 12(]. I \N 1993 . 4: M " ~.", 1<)941>: 245 I'(',-up" \
mllltlllll" , S \ lI l l1
It) , I :
~
I
Di ag nus is t\ ~mall gecko \\ith an SVL of up to 60 mill . 130dy
dorsall y with tim: scale~. lI'itholit tllberele~ ; skin very easi Iy torn; tai I strongl y eOlllpn;ssed dorsoventrall y. also without any coni ca l tuberc les ; inner lillgers all d toes with diminulive cl:m. s; li)urth toe with about 25 ~ubdigital lamellae o f which 7-X arc divided; male~ with 25-44 preanofcmoral pores .
inti'alabia l ~:
large triangular 10 pentagonal mental . fo llowed by 2-3 pairs of" large sllbmaxillars: earopening s mall . vertica l-oval. Limbs short. hind limbs with ~hort frin!!e~ along the poslerior edges: fingers Hnd toes with basal webbin g~: inner fingers and t()e~ with liny claws ; rourth 10C \\lth altogether 25 subdigita ll amellae. 7-'11 of"which divided ; widened . dorso vc ntral ly compressed. segmenled tail without lubercles, bUI with tind y serrat ed edge~ . older specimens with a broad tail ba~e : basa l segl11ents with 10- 11 . aimosl ~ trai!!ht row~ of ~ mnll. Ilal 'ea les: median s ubcalldal~ greatl y enlarged. Male with a long. curved . \"ely obtu~e row of 25--+4 preanoli.:mora l pores; no poslanal lubercles . I·.ntire upper surface li ght grey to light brown. occasiona lly II1l1ch darker. wilh or without small dark spots and light dots; vc ntral surillce ye llowi sh tu dirt y white. Two hard-she lled egg~ lX .5 ~ 10.5111111) per clutch : hatchl ings 45--+ 7 mm in total length . Ecology/E thology The cre pusc ular and nocturnal geckos live as comlllensais orman in villages and large eilies. but also in t rainforests and open country. '1hey often reside aroli li d and in hOIl;,e;,. on poles. trecs. or fa llen logs. They are common in the lowlands, but a lslI fllun d in the Il ea dqLlarter~ rcgion or Mt. Kinabalu . Fema les produce several clutches per yea r in which the eggs onen slick 10 one another.
n esc riJlI iu n
SVL up to (,0 mm , TL up to (,0 mm or s li ghtly less: hod y compressed dor~oventrally: head oval. moderatel .. di~tind from hod y; dorsnl ~lIr1;lce~ or body and lilllb ~ with small granular ~eale~.no tubercles: 35--+-1 row~ or large. imhricate. In'al -shaped ventrals bel ween weak ventrolateral folds: skin very ea~ily damaged : roslra l qlladrangular, mediall y grooved , ill contact with no~tri l s; X-II ~upralnbials; h_l)
259
I)i~trihlltinll
Sri Lan\..:l . I\ndalll
Tal\\an , 1'v1;i1ay
(iCllll~
Icobar
1 ~ land ~ c a ~1 I~I
Pl'nln~ula ~ l)t(th cas t W
Geklw L,\t
RI
f\;
II.
'\ ell' C;uine:l .
176H
I ~ dt'tnbuted in SlH ll h and sout hup to I\: (I rea and Ja pan . rurthcrIII {I 1\.' al' ro~~ Ihc Indo-/\uslrall
I he gcnu:- (;dA"
\.'a~1 i\~ i a . n~)nh
I'h ilippllll.'~ . !\ascarcllcs, \!laLlrilill~. Seyc hdk: ~. Madagascar. MCXI(.'lI. <. Llba and I \;11I':lIi. \'It. h:illllhal .. : Il<.; adtJuarter~ regioll .
lIr prcanofcilloral Ihlrcs prc~I.'n l. p(1, t;1I1al tllhl'rcles p IT~\.' nt. The . IllIda reg ion. includin g 11 (11"111;(1. is hOlllc to Ihc largc~t spl.'eics (II' Ih i~ gcnu~. the tokay gccko. They arc trce-dllclkrs and li\'c in 1(ln:~I~. but it is not IIIKlll111ll0n to lind thelll Ii\'ing a~ ,,:olllmc nsa is o\' man . They an: 1l1,Ii lll y acti l e at dusk and during the nigh l. hilt ~llll1d llll eS their "pc(.'ics- indil: ati lc calls can be hcard dUrtllg the da). T hcy fCl!d on a large va ri ety nI" il1\' c rtcbrate~ although large specics (If' Ihi~ gl' nll ~ ..:a n also ov..:rpolVer small er Vl'rIl'brales. rh eirciulc h e~l'on,i' lt v p ical l ) llrl\\'(\egg~ wh Ich arc "gilled " to Ihe ~lIr IJce . i'cllIaics may lise the ~amc deposition ~I t e sCI,era llilTll's . Malc~ lISU :tll y grow largLr in tolal length than (cllIa!l!s. or the Ihree splxil's occurri ng on Hllrlll'O. only the silla ll c~t. (J<,""fI /I/(I//an ·hlls. Ita, been recorded (i-OIl1 \I t. Kinaba ill .
Saba" (}('''''" '\;"'!," (I "1\ \1
1".
(;/'/../.." 1II 0 1/IIrc llll ,' (I).
(,, 'H (J 1/1i1/11I (i ~\\.
''' ''(1'
8:
1~ IHI(o'.
) X.H,)
IX-I 2
• •
I ri ~ and bac\.. h'Tecn ........ .... ... .... .. ............... slII;,h; lri ~ brownbh. hack 11 0 1 green ..... ... ... ... .... ..... .... 2
2 Back predominantl y brownish with small darker spots, male:> with .12 --10 prcano fcllloral porcs: sma ll species. , VL of adulLs abolll 100 n1l11 .. .... . . . ....... . ..•... , .............. ........ . .. . .. .. .. _.... '"
BrUIICI
~arawa~
K~,lillld lll all
•
•
•
•
+
•
Gekko 1II()Jwrclms (l)I J\I ( Rl t & 811~RON , I X36)
Key to the Bornean species of Gekko
•
I
k.1Il.1h,du
175X)
/'lain/Oe/rlll ' IIIl own/III' Illralil) : (,d~ "
",\",h' >lII C"
111""<1"'1111 ,.
l~HI ' 11\ 1{l1
,'x.
1) 1 r>.11
nil & ll IlH(II ..... I >-:3 h: JJ5
1') IU'
r \llIhl1ll . t"lh'",,,,.,,
'> \1 1111
~ III( 1' 1 N
t'I.11 · 10 ..\1 ; I '), -t o 2... : I "
1')X1 24. L 1\ 1\ 1... :\11 '"
~h " "11\ Il)\n ,
1'l')·lh 2 to;
III Q 1I{/1"('h II,"
• Hack prcdominantl y bluish to grcyi 'h with reddish spots. males wilh 10-2-1 preanal pores. no fCJlloral p()n~s: large spccic~. SVL of adu lt s about 160 nun ,.... ..................... ..... ... ........... .... ... ...... .. .... ... geck()
I)iagno 'is
Or GeHu Oil Rornco, reac hing a maXllllUI11 SVL or abollt 100 111111 . Dorsa l ~ urraec light br(lIIn to liglll grey ground colo ur with IlllIl1Crull~ light an d dar\.. do ls or ~11I ,,1I spOIS. S l11 a lle~t ~ p e(.'Il'~
(I\:k "(lnldae
Ocsr r iption SVL l11ale:-> lip to 102 111111. TL up 10 125 Illill . 1\:males ~l11alkr; body rohu~l. slightly cOl11pressed dor~ovclltrally : head olal. Illod\:nltd y distinct from Ih.:: hody: dorsal "urf;I\.:C \\llh small ~eal.::~ . illlermixed with larger. l'Oundish tlll)L:n; ic~ in I (>-1 7 irregular longiludlnal I"(1\\S: JO-3l< roundish lentral" al mid-body bdween ventrolateral {()Ill-;: ro~lr;d grooved Illedially. in cunla '1 1\ ilh nostril~ : 10-11 slIpralahiab. l)- I 2 infralabiab; menial small. triangular: lor 2 pair~ oflarg\: slIbmaxillars: lingers and loes nHllicralcly broadenl'd. ~ol11e with '\mall ha,.;al webbings: fllUrih toe wilh 16-1 7 ~uhdi g ital lalllcilae, 4-5 hasallallldlm; di\'itkd : lail ,egl11cnlcd by ring,.; ortlll1L:rd\:s: median . undil ided suhcaudab hmaden\:d . Iloth sexes \\ ilh a euned row or \:nlarged pn;anofemoral scales arranged at an ohtllse angle. larger and accompanil'd by 32-..W preanol\:llIoral pores in males: Il;male~ without pn;anofelllnral pores. sOl11etilll\:~ p l,sessing indenlcd ~ealc~ ; h()th se'(e~ wilh 2. males \)Ceasionall y \I ith J poslanal IUberele~ .
Dorsal slIrI;lI.:e crealll-eoloured to greyish brown. ,peeklcd with numerous blac"ish hrown dt\ls; tubercles yellowish or dark : onen (1-7 paravertebral blackish brown ... pot~ bell\eenned and ba"e of tail. alt\:rnating. with verlehral light lI1arkings : a dark. almost \V-~haped Illarking on th' occiput : golden to amher-coloured 1rI~ wilh a nel\\or" of dark n.:d \'\; ins: venlral surfa\:e dirty \I hite . ()(.;ca,ionally slightly Iranslucent : hatchlings dar" hrown with di~tinct yellow dot~ . dar" lllar"lI1g~ Intii"tincl. Two hard-sh<.:llcd l:ggs, I (J- 1:1 mm III diameter per clulch : hal c hlillg~ 25-30 t 26-3<; 111111 (SV I I I I ). Erolog} / EI h ul og) (;" kko /1I1i/Hlre/llls illhahit~ 101\ land~ and l1lountal1lous regions lip to altllulk~ oJ' 15()() m a.s.1. This »peries i:- COll111101l 111 and around hUIll:lIl habitation, . in plantallon" and in primar;. I<Jr\:,t,. II i~ usually found li\ ing ill pair~ . more rarely In larger group~ . I hese g.ecko~ are mainly crepuscillar and Iloclllrnal and prescnt Ihclllseh \:' a~ I'ery shv. They arc tarilOrial and do IH) tokrate other ,pcci\:" or
hous!."!gl'ckns In Ih!."!1. 1!."!ITilnric". The ca ll or mail;" i" 111l! \ ery loud and consisl" or a coni inllolls s..:rics or ~everal ·'lOd ,"-snu nd s. Females produce cllllch<.:~ or..:ggs ~ewr(lilimes per ycar; and an: firmly attached In a suitablc surl:1ec. Thcy otten reed on ell1ply shclb Oil!."!!."! Ihe halchlings
(iCIlLlS l-Ielllida(.'~l'III.\·
have emerged in ordcr not 10 waste a source of calcium . Di stributio n From Thai land ~nllthcasl 10 Irian Ja ya including the Philippincs. '11. Kinabalu : Kiau.
Granular dorsa l scu lellalion inlermixed with larger luherclcs . Fingers and toes dilated . \\"llh s lender. clawed phalange !". an d free , angled terminal phalanges. Subdigilal lamclla!."! divid<.:d llIediall y. Mal e!> with pn.:ana l or fClllOral pores . Usually li\ ing in and around hou ses. Ihese g.cc ko!> ICe I largely on Sill a II ills<.:ch. Several times per year. female l> attach their c1utehe ~ ( r t\Vn egg. tn a suit able surface. BOlli spccies occurring on lI11rlll:o an: al so presenl on IVII. Kinabalu .
01--1 ' . 1X17
or
The dislribullon Ihe g!."! nus //ell/ll/actl ·/ItS eXlends lI'om ~olliheast Fllrorc Ihrollgh Arriea. t\"ia. the Il1doAustralian /\rchipdago ln the north nf i\u!>trnlia (inlroduccd) and IlUIllCrous Islands or Ihe Paci lie Ocean a~ well as n(lrtheast":rIl south Ill..:rica . 1I{,l/Iidllc~I'//{'\ has been IIltroduced In M!."!xien and thc lJSA . Sma ll to moderat ely large get:kos wilh vcrtical-ll\'al pupi ls and dursovcntrall y cOlllpn:sscd bodies .
Tab. 211
[)i ~ lrihlllion
(If 1/"/IIid{/( " ,'/",\'-sp..;cio.:s 011 Horlll:Il. Sabah
fit-midll ctyill.' In·llI/lII.,·
1)1 \II ·RII
&
HIII R
1:-;.\(,
1-1('m i dllctr/ll.,' .l!lIrlltJ li i
[) I \11 KII
&
H IIIIWI\.
IK"\(,
• •
Key to the Bornean species of H emidacty lus rail roundish. with rows of enlarged tubercles ... ............. .......... ......... .......... .. .. ... ...... .. ...... Irelllllll .\
• Tail compressed , edges fine ly denliculated ........ ............. ..... .. .............. ............................. gal'/lotii
Ilemit/lldY/IIS ji-elllflus D UM I'RIL &
13 IIlRO'-J , 1816
L" llIlIK"N. I X\(.: ''''''"'',1 1')'17)" J'I\"
//("11111/(1 ('/1"/'" /n'II<1I,(\· II I \ 11 KII
' I rir l ~ d Iypl' I "ta l i l ~' ( I
//"/11"/<1'/'/11" !r, ' //{Ilw .
S\1I111
2 ·1 , 1' 111'1 111<11 &:--'1(" .1<1 1\
I<.IX')
11)-1 . 1<)'.1.\ 1" 2-1 5. I')l)(K" 2xx
2fJ2
I~
I'nl '
1')X-I'2-1
1(.(,
R l'-
10, ,I ; ~"' ''' III1\ I<.IX ) ~'. M II~\II "
; t '\ N Il)l)' ·.:
M il ~\I'"
19XXe· 17-1. ( 1)')-10 ' ~ .1 .;.
I Kinabalu -
• •
Hnlllel
• •
I
S:traw:tk
Kalimantan
• •
• •
Diagnosis /\ small hOllsegccko or up to 65 mill SVL: dorsal pattern absent or pre:-.eni and highl y variahk: ~round coloration li gh t to tilinel frolH the bod y: rO~lral qlladran g ular. media ll y groovcd. touching nostrils; 10-12 supralahials: 7- 1() infrnlabial .. : mental large. triangular or pl:lllagonal in shape: 2-3 pair~ of submaxillary shie lds: fingers and loes without basal wchhings: I) - I () suhdig ital lamellac b<':l1eath f"olll1h toe. 7-9 of" which V-shaped and medial ly divided: t
Gekkonidae
Fig, 272
// elllidaclI'/1I.I {i'e/wllls
from Poring,
backward-pointing, conica l tubercles: median 1'011' (,[' suhcaulial s broadl:lH:d , [Vlales with a continuous row of 2(, -3~ pn:a nokmoral flores. l a q::l:~ t ill the cenln.:, i)ors,d s urfa ce in a la rge va rid y or s hade s or brown rangi ng from greyish brown tll ye llowish brown . hi gh ly va ria ble as to lighter or darker: either uniform or ::.ped lcd with dark , o n eil with addit iona l dark s p ot~ or stn.:aks: dark lateral s tripe 011 tin.: hl:ad: tail with or without ra ill t hands: ventral surface whitish to yc llO\,vis h white, vcntral sui'l;,ce of tail ,,;o Jll e til1le ~ reddi sh, .Juven iles coloured like adult s. but with l110re contra still g ra ttems , LJsu all y two hal'd - ~ hclkd . a lmost round eggs (9-10 I11Ill in diameter) pe r c lut ch: hatchlings IX-22 - IX-2.1 mm (SVL+TLl. I::culogy/ l:: t Iwl(l g~' 1/(' lIIidl/eIY/lIs li'e lla/II,\' is the most eOl1lmon hO\l"e
gec ko in Southeast Asia , It s altitudinal distribution l'an~es up to approx . 1600 m C\,".I" but it is less
Fil.!. 273 Mating "I' " ell/u/lICn '///.1
{re I/rill/I' 1'1'0111
POring.
CO il II I1(HI at ekvations abow 1000 II I ,1.",1. It liv's on bou lders. bencath rods or rotting logs. on trees. but is I11()St commonly fi1und on hllildin g~ 01' all sorts and si/cs. It illhabit ~ vill agl:s jll~ t as wcl l as big cities and is usually I:ncount..:n:d in thl: vicin ity of" e lectric Ii ghb where it is draw n to at du ~ k , Although ill prill ci pal <.:rl:pllsnliar alld noctllrnal. th..: di s tin ct cackling ca ll s or male" ca n be heard throughout the day and ha ve carned thi S gecko its ve rnacular nalllcs in Illany rcgions. ror cxamplc " tin gtock " and " tchik ehak " , Preccded by s hort mati ngs. k lllaics prod lice ..:ggs 4-6 timcs per year.
Di stl°ibulioll t\'lcx ieo. Madagascar; lVlaurit iu ~. sout h and cast Af","ica. rrom sO llth India and Sri Lal1k a ea"t tn Taiwan. from th..: lVlalay Pen ill sul a ~llllt h": ;ls t tll New ('uinea. i'urthermore on many islands ni'the Pacific Ocean , and thc Philippincs, 1\'11. h':illll halu : Poring and Headquarters region (around bu ildings and th..:ir imillcdiat e vic in ity).
Cid,koni dae
lIeflli{/actylll s g al'llolii D l \ 11 RII
I hmu /u, fdlt.\ ganrfJltJ
I.. .." li l) : " I ' d '
&
13 1111H) , 11'( 3()
1) 1 \1 '~ II
...~ il IIH{U ......
1:O lli " ITaIlIIL
Ilt'lIlId,11 (1/l/~' ~( r rn (lrll,
S\IIIII
11)3 1
.!~~ ~ 1'1 11 '\ I \.\I(P (.~ \,, '( U, .... I " !' I X 1 11 ~'1
I/ . 'm ll/ol
1" 1.
I ,\~
trill,
\"r.
111'1' I, .
1,'3(t
"lhX
"I y pe
\,111 1 \
Il)X3 ;
['"hil L''',I]
I. .:.! ..L
~ I "",,,
10,1 1. \1 ]~.
ti r _p
1'11)..\11
27, [\·'I \ J f",\1 11'"
2 .J ~
/I(HI rill ~lI. (\1\1 f.. \II ',,! P) I}'1"1
I'vb lcs \\'ith rll\\'S ur 7-1<) re mo ral p ure~ ,)n caeh th igh: I C lll alc~ wit h rIm" nf 15-20 c nl arged remoral "caie::. on each th ig h. "ome s lightl y pilled : both sexe~ wit h I postanal tuhcrck o n each ~i d c . Dorsal SUr!ilCCS brow ni sh in va ri llll~ ~ ha de~. with Ilwrc or k ~ s di slilll: 1 dar ke r. o ik n alsl' \.vit h slll:"1 whili sh ~ P () t s: ve nt ral slIrlllces whi tis h III erealll eo lnllred.
1.2:'
Eculng., / Et hf)l o~~'
\) iag nlJs is .~ , nlall gckkonid , pee i \.'~ II IIh ,I SVL ol' lI p to (,S 111 m differi ng frolll 1111: \'ery s imi lar If. jrl"/Wfll.1 hy serrated c d ge~ or its lail (vs . rin g~ or tu bercles ). Ot'sc ripl inn SV I lip to (») 111111 ., I I. lip to X3 mill ; hod y slender.
do rso ve ntrall y cll lll prcssed; dorsa l ~c a lcs ~ Ill a ll , inte rlllixed wil h a ~ ln gle 11I hen:IL: or gr(lUp" or tuh<.: rci e,: v<.: lll rab ~ 1ll : ill. ~ m ol\l h , ru unLi<.:d , in 29-.1-1 rows bdween wcakl y dc\.c!o p<.: d n ;ntrlli ak ral fo il\,,: head ()\'al, ~ ll g htl y disti nd from the body: rll~ tra l qua drangu lar. Illl'di :tl ly g roov<.:d, 111 l:ontad with n o~ tn ls ; 1::' - 1J ~ lIp ra l ab i ah< I)- I I il1 rra lab ia1:; ; me n(a l large. Iria ng ul ar in :-. ha pe ; ~ p air~ 01' la rge ~ lI b l1l : l xi ll a l '~: li ngcr~ allLl Ines with :-- Ill a ll hasa l \whhin gs : 1 I-I -l ~ lI h di g i( a ll a m c ll ae , 0- I I 01' whi d l di vided ; tai l "lender. dorsovel1trall y co mp ressed. wi th :IClIt C. li nel), ~ er rat c ct edges: dorsal surrace f: lintl y ~eg l n e n tcd , \vit h ~ lllall scales; median row u l' ~u h\.'all d a l ~ hroadc lled.
f-!cIIJidaclyllJ.I·,!!.{I/·lI01ii li\'e, in the low land" and 111()Un-
tallloll" regiuns up w al prox . I.tOO III a , ~.1. wherehy il is more Cllmll 1()n al alt itudes ahovc 1000 III a.s.1. r he nocturnal gecku is fo und in a nd ( Ill h n u ,e~ as \ve ll as In the ll)rests . Rc produ cl"ioll : 'In"t pop ul atio ns :uns ist o r parthenoge ne ti c IC males whie h produ ce c lut c hes o r uSlially :. eggs eal: h several (imes per year. Ois lribulinll From India e:lst to Chi na. from ivbl ay Penin sul a ::;(lll th ea ~ t (0 the Sll iomon Islands. fu rthermore () n severa l Pa 'ili e Oc<.:a ll Isla nd s. Philipp incs , inl roduced III Flori da a nd ;\1e\\ 7cala nd . IVIt. Kill a halll : Pori ng. Ki au .
Genus H l'lIIiphy lloc/acty llls HeJll i/}llrll{){ /(/ c~\ · IrI.l' i~
R II-I t-:.LR,
I H60
di st ri hllted froll l S I'i Lan ka
to ,outh China and the Japanese Ryuk yll b la nd ~.
Fig. 27..
1/t'lllidl l(
,dl/ I x amOfil ~~=-------;
a1. 0 inhabiting the rh il ipp in e~ . the In do- /\II ~ tral ia n /\rchipe lago. and sO ll1e i s land ~ orthc Pacilic Oc<.:an. S lll all. slen de r. dorsove nt ra ll y cOlllprcssed geckos \\III h hOllloge I10U'i. gra nu lar sUllellat ion. Ta i I lI Sllall y sll ll1ewh:II :-.ho rt er than SVL. at ll1a:-.illllllll 01' cqua l k ngt h, Pupi l ve rti ca l-ova l. Ollter 1'0 111' lin gers and t oc~ ()ecasio na ll y wc bhed al b ase~ , di lakd, with I'r<':l:, raised, clawed tel"ln inal phalange,,; inlll:r li ngers and tocs rudimc nta ry. diminilli vc. and lac k raised lerm in :ti phalan ges . but so melimcs w ith claws: basa l suhdigital lamcllae un div ided, di sta l nnes med iall y d i\i ded. Maks wi th prea na l and remora I IlllITS. S p eci<.:~ o f / {eIlJi{J/trllll ti(lf"/r/ JJ \' recd o n all sort s o r tin y insects. Fcmalcs produce clu tc hcs or lI ~ u n ll y [wu sillnII. 1'0 lind ish. ha rd-s he II cd egg~ .
(i.:kJ..nnida.:
Rl1rn':1l i~ h OIll~ o nl v to onc ~ I)CCIC~ Ilhidl h", been n:cordcd frolll IVIt. Kinahalu .
a l ~ll
Ec olo ~ ~"/ l:: tlH)IU I!Y
H cmipliy/lodaclyills ~1'IJlIS typllS HI 1' [' fdR, 1860 " "11111'1"/1" .111111",, /1'1'1/\ Il il
~'". I ~ ('() . 1 ~ 7
·I)p.·
I (>~" I
it~· : ··;\galll ·· I\\ ~" 1
SUIl1.1Iral .lnd " (iO\ Ollnllg Pal t Ul g lJi.I \ ~f · ( ,unl1n ~ Pari.lll g. \Vc ... 1 .1 :1\ 01) " "Jll il'l,, /llIlllu ,.111 \ ""/," \. ~ ., III III 11 10. 2 1. 3 I. I \N 1'1'1.\. (•. ~t \I ~\lI' 1')<)-111 2-1 5 1I('mll,lld/lide/( 1\ I,I' /1'1'"' H'/'U\. \1 \\ 11 11 '
Il) , ' .
I \\'0 hard-,helled cggs (5-6 .5 . 6.5-/\ 111111) per el utc h: hatehling~ 1-1- 17 t I-l- I(l IIlIll I ' VL ~ TL) .
2':;;
This . m<1l1noclurnal gec J..o u~ually inhabits moul1tainolls n:giolls abo\ c appro \ . 1000 111 a . ~ . 1. \1 here it is fo und In ti) re~ls. agricultural an; a~ . and In a~ IIcll as around buildings . Su it able day-~helter~
J)i a~ l1 os i s
/\ ~Illall. re ddi ~ h broil n gecJ..o.of up tn -17111111 SV I. wi th a ~Iilll. roundi~h. a llll ()~t p , ilternlc ~s light brown tail in g rcat Cll nl ra~ 1 tll Ihe blllly ; houy anu tai l lI'ithont lubercl ·S. Descript iull SV I up III -l 7 Illill . TL up to -l6 111 Ill : hotly sl im . dorsov.:ntrally cO l11prc~ ~ed : dor~ab ~m~ ili. smooth . I'!lundish or "n!,!ular: 1111 ~n larged tuherc lcs: 110 \CIltrolateral folds ; Icn tral ~ ~l1loo th . im bri cate, alwul -lO ro\\~ ; hcad OIal. ha rdly distinct fro m Ihe hody : roslral allllllsi pcntago nal. m cdia ll ~ !.:nlo\'cd. Illuching n()slril ,, : X- 12 supralahial s: X- 12 infral ab iak mcntal ~l11a l l . clongat .:. triang ular in ~ hapc ; no or one pa irorsillall subma:-.illars: ling~r~ ,lIld Illes wilh or without ba~al \\ebhin !,!s; inner fing.:rs and II c~ rudimcntary. "ith or \\'ithout tin ) cl ~1\\~ : 12-15 ~ lIhdi gi tal lamellae 1)(' nc<1lh 1()IInh 1O~ . di stal 3-(1 V-~hapcd: wi l slender. rolll1d. \\,Ith line sc ak~. no tubc rcles : l1l e di ~ 1Il three J"(l\\~ o r subcalldals hroadened . \t1nIc~ " ,ith an ang ul ar scrie~ oU1- 15 prea nal pore~ (l -X pe r ~ id.:) . ~.: p anll cd rrom II hic h arc roll ~ or?13 k moral pores on eilhcr ~ide: 3 round p O~ lan a l tuhcrcles (1n eith~r Silk: Dorsal surface reddish hro"·I1 . \\'ilh sl ightl y darke r. fa int. V-shap.:d lI1arl-. in g::. bet II een sides of nceJ.. and tail ba s~; llcc;I' iona ll y \\'ith an intcrruptcd. il1dl ~ lin(;1 latcral stripe o rlhe same colour extending from rear margin or Ihe ~ye III Ihe leg in~erlion; somctimes with li ghtCl" dots ar rangcd in longi iudin a l ~c rie s : dorsa l sur t;ICl: ortaillight to gr.:cnish bJ'l)\\.n. I'c l1tral ~ url 'ace whitish at basc . ~pcckled "ilh brown. dal'k di~t a ll y.
Distrihutiun
Indi a and Sri I anJ..a ca~t tn Ihc .Iapan e~c Ryuk yu 1 ~ lalld s . 1I Iaiay Pcnlnsl ila . (in.:atcr and ~(1 m e of the L e~scr Sunda 1 ~land~ . PhilipPln cs. ell (iuillea. .:\\ Caledllllia. :tnd Polync~ia . i\ 1t. h:in a h,llu : Poring. Ki au .
(JCllllS L epidodll ('~l'III .\'
FII II (il IC 1~43
/.C/}f{/()t/llc/I '/IIS i~ a g~lH l ~ o f ~o uthl.:rn and southeaskrn i\si a. al~1l inhabiling the rhilippin e~. the Indo-r\u straliall i\n; hip~la g(l. New I.e;dand. and i!>lands or the I'aei li c Oce:an . Small. -dender. dorsll\entrally com prc ~~ed geckos wilh a linc. hllnlOgencoll~ly granular sc ul cll ation . Pupd v~rtl cal -(lval. F in gcr~ and tocs occasionally
I L'l)It/Ill/(/( I.l '//{\ 1111;111>1'1.'
11)1' ~1I I' ll
Lep;(I{){III('/r/ll .~ rll l/ l1 l1 l' I/ .,·;.\' () I '
&. H lI lI"lto. . It06) & 11 110; 111 ·\ . II};;;;
-
Key to the Bornean species of Lepidodacty ilis Edge or cOll1 pre~ scd wil findy denliculated; I 114 upper labiab ... ... ........... .. ...... ........ ... . lu}!,uhris • Edge of compressed tail nearl y smooth ; l) uppcr labia Is .... ...... .. .. ... .. .. . .... ... ... .... ..... .. .. raflQll e llS; S
Lepidot/llcty lll.\' nt/wuellsi.\' (}I /\ & II IKII),\, 1988 I ,('pu/odl l( 11/11\
d. 2
'1')' (1['
\\cbhed al hnsc!>. !>Irn ng ly dihl1l'd . usually with lIndi\ idcd slI bd ig ilal lamcllae. lalkr V-!> hapcd di ~ tall y and somd llllCS div idcd; inlier fill gc r~ and IlIC~ lack claw s. t'\ilale: ~ with pn.:anal and I<:moral porc~ or pre:anofCn10ral porcs. Thcsc gec kos feed (In ~ l1lall inve:rte:bratcs. I'cmaks II sua ll y at lach pa ir~ llf hard-slll'llcd eggs to a ~L1it able: ~urface: . Bornco is homc to h\ (I ~r.;cies (I f Ll'/Jit/ot/llctl ·/i{,\'. olle of \\ hich hws on It. Kina ba ili .
Sabah
• •
KlI1abalu
-
Brunei
Sarnwak
Kalim'1II131l
•
•
submaxillnr!>: all fin gers and ioes ddated di ~la ll y: basal wcbbings only indicatcd bctween .1,,1 and fo urlh loe:s: 15 !>ubdigllallamell ac benealh ((Hlrlh toe, di~lill ones V-!>haped anti occasionally divitbl: lail mllscular. dors()\l.:nlrally com presscd. Wllhout I or tuhercles: d(\I'sa l surfilcc (II' lail finely ~ra ll\llar: median ~ u l1(; alldals slightl y e:nlargcd . rVlalc:s with an angula r. e() ntinuoll~ sc:ries or J7 prcanofCllloral pores l.:xtelltling up to Ihe: e:nd orthe thi ghs: a large postanal luberc le on either side . Dorsa l surbee a light shatle ofhn1\vl1 . wilh a dark brown dorso lalera l st ripc from the occiput to the lail: small light dots wi thin th is stripe; ve:rlcbral rcgion with an irrl.:glliar. dark pal1ern : he:ad speckbl \\ilh blacki~h brown : limbs with dark bands:
r01/(III (, II ' I' ()\ , & 111 ... 11).-\ . 19~X : (,I (•. li g I aI.. ralil): "I{anau (III, ' ~5· 1 .. 'i 'iO'N). <;,IIMh .
I . CI ,ulotitldlhl'l"IlIItlftl 'f/\I '
r-.. l " ' III \ &
(' K. ....S"""NN
1997 _-12.
Ii" . 17"
Oi ag nos is ~mall ~ee ko of lip 10 .+ ) 111111 SVL; body and broad Illas!>i\ e lad withoul tubcrcles.
i\
Oescript io ll SVL lip 10 45 mm . TL s li~htl y shorler; dorsa ls I> m:1 l1 . no cnlarged lubercle:s: no vcnlrolate:ral folds: venlrals round . nal: he:ad oval. ~Ii~htly di stinct fi'om body: rostr... 1in enntact with nostrih: 9 'upra labials: l}_ I () in rra labi a Is; mental Iria ngul ar: I pair of largc
Fig. 276 L<'I'/(/odaclr/w cf.
qllartci ~ .
/'
frolll thc Ilcad-
Gekkon idae
original tail with a pattern that corresponds with the hac k; Enti re dorsa I su rf~lCe sOllletil1l<':s uniform gr<.:y hrown . locally Iighter with dark dirfuse stripes: ventral surfaces lighter. ~co log~:f [thology
These geckos have heen rOlilid on the \'iall s or buildings, in rain shelters, and on shruhs and hushes. Oistrihutinn Sahah . MI. Kinabalu : Ilcadqu
Fi~.
277 LepidodllL'lr!IIS IWlllI/ell,li.1 fro m a she llcr ncar Ih e Ilcadqu
Genus P~Jiclro':.OOIl K UHL
&
VAN
I [ i\SS~. I.T . I R22
The di:; tribution or the genus P(I 'c/1O::001l is limited to the southeastern Asia. the westt:rn portion or the Indo-Australian i\n:hipelago and th' so uthern is lands of lhe Philippines . Moderatcly large geckos with dorsoventrally strongly compressed bodies and tails and characteristic lat eral skin frin ges along head, body, limhs. and tail. Latter u. ually slightl y shorter than head alld hod y together. rarely slightl y longer. Skin finel y granular. dorsally mostly intermixed with larger tubercles. Pupil vertica l-oval. Fingers and toes dilat ed distally, with ve ry large \Vehhin~s. Outer fOllr ringers nnrl tlles with c1av,led, rai sed terminal phalanges; subdigital lamel lae broad and undividL:d . Males with preanal and/or femoral pores . 1\11 species arc lo rcst-dwe ll crs mainl y living on trees. but occasionall y one of th ' Ill (P r/lll c!Jp17o-
was observed on the wa ll s ofhuildings in the Headquarters re~ioll of Mt. Kinahalu . '1hL:Y arc equipped luI' s hort parachutin g and feed on in sects. Clutehe. co nsist of two hard-shelled eggs alld arc attached to a firm sur face . Three species live on Borneo, nnc, P. ,.!W('IIJi"Oms, is endemic to Mt. Killabalu .
1'11.1')
Key to the Bornean species of Ptychozooll Side ' of head with skin frin ges, digits nearly fully webbed .... ............ ..... .. ....................... .......... ... ... 2 • Sides of head without skin fringes, digits 1/3 to y, webbed ...... ............................ .. ...... r/w cofJ!to,."s 2 Flanks with two rows of tubercles ........ ..... klf"'i • Whole body wi thout tubercles ...... .. ... hor.l'jieldii
Tau. 30 Distr ibution o r I'll 'cli():(J()II -~ peeie s o n nomcn. Sahah
Pfrcho:IJIJII ,.hucllplllJrlls (HOI
11 ' \1(,1
Killabaiu
I I
hO".I'fjeldii ((O R\\ . 1727J !'/YC/lOcOtJ/I kuh/i S ll I~ I' 01 I< il)()2 PII-c/W::OO /l
• R, I X()(})
•
Brunei
•
Sarawak
K~limalll :lIl
• • '267
P~l'dlO~OOIl rltU('OpllOrllS (i301 11l ' l\ltl1<, I Xl,ll,l) ('( '(~O rho c(I/'/U IIII ' H ill II ,\, II~
lH4Jt) ~ '1 1)1>(.' tCll'ali l ~' : " "'.ld,II11:II.IIl!(IICI "IIl,IIl,lIu . '1 1111 k,L mth ilorne,," (1\ I hu liltl, "I,IJ""II\ I I I "I hI II 1'11111,1 71), rl I IIg, I . l'Ii!'k I . I I I it t 'llll 1')1" "'''
1 )/ 11 IIo_'(U1II rlltu 'o/J/'nn/ \, ~ \III II
lin 1 IJ, \ 1\:'-. I t ll \ & 1)1 NI l U 19K, . ~:'i. I IH'lI'HP& ~1 1)(,HI It)X--I 22. lij.! 2'. J \ \ t ll1 'O\ '9 X';; 1 2 Ij g . ~ ~l': ~ 1 \1 \1l ... 1l)~t} : 1'1', I " III'); , ~ I I"\I' l-lll.lh '.1), ~" "II II I ,'\. (H ~< ' ~""\ I \'\ JI)ill ~ ~ ~ fi g 179 11)1)2
11 Ii . . .,. ' '''
(\ 1 '\11 11 \
J)i ag ll o~ i ~
S111alk,t , pccic, or p/\,('!r/) ::UIIII 1IIIh an SVL 1)1' ma'( , 7<, 111111
and IOC~ tonncdcd \\ ith \\'cbhing~ ovcr ncarly h:tll" th eir Ien!!lh:-.: I O- I-L d i,lall), L1 'L1all y V-:-. hapcd "ub(ilgltal lamellae: ,.,~in fri ngcs 011 ba ~ l' 01' lail spikL'-like. then \\
Iksf ript itlll
SVL nr nlaic:" tip tu 75 1111ll. TL Illuch shorler. i,c. up tu abllttl 51) 1111ll ; clltlrl' dorsa l "ttrfitcc with Iinc granulal' ,ca k s and ,,111<1 11 , rotilld, ~111(l\lth tubcrcles; hrond \'clltrul;ltl'ral "k in fll nge hct\lccll a\illa and gruln I\ith rnt )cd edgc,,; \clltrab granular: head oval. di slind from Ihc hcad : ~i dl' " or hcad II'llh cOllical tubcrdc ~; ro,tt'al hn'ad, without mcdian gromc , ~cparalcd fr()ln llo;O;lrik l) "upralahials: I () inl"ralaolals; IlIclllal , mall , pentago nal; 6 pairs or ~ mall ~ ubma\lII :II' ''; limhs \\'llh ragged , paniall) ~c rratcd lateral ,,~ in I'nnges nn cilhcr :-.ide: lillger,
rhl ~ predominantl y Ilocttlrnal trcc-d weller Ita~ to dal e only hccll rccordcd from altitude,., hetw(;cn6()O alld 1(,00 m a.".1. Ull \111 . Kinahalt l. In thc I kad quarler" rcgilln it i" llLl: a ~ i ('llall y also I()ltnd lin thc otltside walls ofbtlildings. The " para l! hding Ilighl", 1"(1I' which the gcn tl:-. IS rCII\.)\\'l1cd, has :-.0 1;11' 1101 hecn ohs(;l"\cd ill Ihi s spccies . I cl1laks attach their egg" III a stlil;lbk surl:t.:c.
Di st r ihutinn
I' ndcmic to \'It, h:in a halu: Ilcadquarters region and Sg. Kadailiai an «)OO m a,;o;, I.) .
Fil!, 278 !'tn-/llrt llll/ rhm 'II/,!w/'II,1 from the llead'lll" r-
Fil!. 279 Ventral patlern 1,1" !'II'c lllle/IiJ/I r/i/// '"/,/i,,r/ls
leI" .
Imlll Ihc'
26X
Ill:aJtjuarter.,.
Scincidae
Scincidae Compri~ing so nll: 12()() sp 'eies. ~kinks or ~ll1uoth liLa rds rc prc~cn t tile most dilerse ramily Ill' lil:ard~ . They arc present in all tempera tl: to tr(l piea l regions around the world . In mo~t ~pceie~ the ~hll1 Y ~ealcs arc IWlllogcnou~ , roullded and imbricate. Their pllpil~ arc round . and they h ;1\ e ll1ll\ablc c~ elid~ . Some ~pel'ie~ arc noted fIx their extreme ly reduecd or abs,'nt lil11b~ . T he Illllller ea~e l)fien goe~ along 1\ IIh a reduetioll uf' the number or lingc)". and tocs \\ hieh othcm i~e number IiI e. Like gedo~ . ~kinb ha\e tilc ahility to autotomi/c and regeneratc their tails. J lI \'cnilc~ or certain ~peeie~ dilTer ex tremel y rrom adult~ as 1~lr as their eo l ora ti on~ and patterns are concerned (e.g. {JIII'ill "p.). tdl. Kilwbalu is home to predol11inantly diurna l skin k". They arc terrestrial, ~el1lia4l1all C m l(lS~(lrial I.\ ith a preference ror ieI'd" or soil ncar thc ~lIrl ilCe. o t her~ arc oftcn li)lIlld amon g tree buttre s ~e~. yct others inh::lbit the eanop~', the bank" orstn:"I11~ or e\'cn st ream". 011 MI. Kinaba lu ;.kink~ llIay be Illlll1d up to a Iliaximum alt itude 01' appro ... . _200 m (.)/)/ic' lIo/l/()/,/illII.I' Aillllha/itclls;,) . They feed on a lar~e variety or IIl\'crtchralc,,; arboreal ~pecie~are likdy to al~o prey upon hirds ' eggs. and largcr spceies wil i lleca~ion al ly slIppkment their dlct I\ith "mailer li zards. lost skillk s lay two or more ;.oi't-,hdkd e gg ~ _ hll\\eler a f\:\\' gil.e birt h to 1'1111 ) de\\:lopcd )oung. On Bnrncn, thi~ f~lI11il y i~ repre~ellted by .W specie;. in I () genera II llh i( genera and 15 specle~ nativc 10 IVII. Killahal u.
Key to the Bornt'an genera of Scillcidae (ill p
B
2 /\ light \encbra l
~ t rip!:
................ ....... ....... .... ... .... ...... ........ ... ... .... '-illil/ill
• Dnrs,,1 p:Jllern diflcrc:n t from that ... .... ... ..... .... . :1 3 I ~ ub st:J nt ially ~nlarg('d preanal ~t:ak (R), dor~als u~lInlly ~ tro ng l y ke~kd.
.. .. .. ...........
hllmped .......... .............. .
..................... .. ... .
TropitlllpJ/()I'I/.~
o
o or ~el cral enlarged prea nal "c:Jk~. dorsab not humped ................... ........... ..... .. ....... ........ .. .. .. .... .t
.t I frontoparietal •
(C) .............. .... ................
~ rro ntllparietal ~
1:'11/11/(/
:~;
([) ) .... ... .. .. ...... .... .. ...... .... .. .. ... . 5
I,
:; Supra nasa ls presen t ( I '. ) .. .. .. ... .. ...... .... ............... 6 • Su pranasa ls abS~l1t .......... .. ... .. Sphy /l ol/l(l/'pJlIIs 6 l30dy no t l:long:Jte. adpre"sed limb" meel or
OIcrfap (I-) .... .... .... .. .. .. .......... ....... .. .. .. ............ .. .. 7 • Body clongatl:, adprcs"ed limb, do not 111eel « oj ........... .... ....... ............ .... .. ................ ,.......... Ri(lplI
7 •
Dor~als
Dllr~ab
!..eelcd ( I I) ......... .. ......... ..................... .. II smoot h .............. .. ............. L(/I/lp ro /epis
f
. ~\.'.
(
! (~'~\
:.~~ ,
.' . -1'\1;\ \
/
I
.'. _" .
8
.J
cro~s bands (more distinct illju\ cniil:s) (.I) ............................... .... .. .. .... .......... D asill • Dorsum without patl crn, or sma ll dots sometimes arranged in longitudinal series. but not forming di!>tind stripes; often with dark dor~o lat era l stripes or va rillu;, widths ... ............. ... ................ Mahllya
9 Dorsum with
A ptel)'got/oll vi tti/til III EDII.IN(i. 1R6S Ip(CT\ .~odnll
.. lj l /crrg()t/fJ/i i\ a lll11nlll ypi cal genu s cndclllic tll Bol'l1\:\).
tVl11dcrah:ly largc. diurnilllr\:..:-dwcl\c r" with ro hu ~ I , dorsovc ntnlll y cl llnprcs~c d body. hceled dorsal s and long lillil1S (adprcsscd limh\ o\crlap) . Lower eyelid ~caly ; tympanulIl vlsihle . rali round , tail tapering conlinuous ly to a point. .Iu vc niks and adults wilh a ~tnp..:d patt e rn 011 head and anterior part or the back .
Fig. 2l!1I Adult , pcci nll; n of I,'/argot/oll
"'c"
~
:J 1.. ...... . . . . .-
270
\ '1//11111111
I 11111 .... <,. 1X(l) ~() I T) pl"" I Ufa l i l ~' : H"nll'\" l li ~1IlI,lnn;"I1l . Stllllh K,tlilnanianj Ip/c '" r..:,odon 11/f0/111II f\,·1\I "r-.u ,,," 1')X 7: 2X:<, : T " I V9J- 1 \ ' t/{l/(1I111
"llandl<:I II""III,
" I'lt ',:r godol/
I 'II/tlill .
rvl
fh" j(l I'lltalcl. \1 ,\'1 111'
\1 .... \111 ...
1')')..Jh 2..Jh
s... I), N/II(
I t),
"l .
13. fig .
1(, .
r.- l ·"l lll '
1t}~3 .
2(, . r.-l "t..MI '" I()X" 11 1. (1111/>1"0/( '/)1\ \'111011/\ h u 'lI\KII ...~
')14 Hil{J N P) ~· L
22. fi g.. 24. 25
Dia gn osis !\ moderatel y large. robll~t tree ~k ink ofaholll \)0 111m max . SVI . di,tingllishahl e from tree-dw ell in g
frUlI' Sal1 lubol1g, Sa ra\\'ak .
Scine idae
sk inks or othcr genera hy it~ eharat:tcristic light stripcd patt crn on its ot hcrwi 'c dark hcad and anterior body. Oesc ripti o n SVL up to 1}6 mm . TL up to lOR mill; lower eyelid covered with sma ll scales; tympanulll visible: eal'-ope ning sma ll. llva l in sha pe; nostril ill one nasal; su prana sa ls sc pa rat cLl; rro nl onasa l widcr than long. fnrmin g a suture with rostra l and frontal ; 2 prcf'rontab, scparated [i'olll each o th cr: frontal a lillie larger tha n fr o nt opa rieta l" and illtcrparietais IOgcther; fronlOparietals larger than interparietals: nucha ls distinct. ocF i ~ . 211 1 Juvt:niil: spec imen (1(' i t/JII'n '.!.!."t/11I 1'1//1/11111/ 1'1'11111 Poring. cas io nal ly ~ tr o n g l y keele d : 5 su pra ocu lar:--. the ant erior 2 in cont ac t with frontal, second larl;:est. [j nh very ~Illal l : 10 fin d a pair occupy in g a tree toget her wi th their ~-I) silpra ci liars; 7-X sup ralab ials: 6 infralab ials: dorsals and laterals wit h .) or :; strong keels ; ahout o lhpring. They Iced O il in~ccb . SC\cra l times pCI' yea r. fclllale~ deposit their eggs .10 ~caks nruund mid-hody : preanab no t en larged : in forks of hranehe~, among mots llf epiphytic males alld i'cillaies with 2 en larged tarsal ~ea l es : I (1-22 smoot h subdi g ital~ heneath fo urth Ille . rlallts. or 11l:lIca th loo"e bark . Since a Juvenile \Va" found ncar the gro und (M \\1 1111 \ unpllbl.) II lIlay Hend, anlCrior portion of the bod y. and back black : he as"umed that cgg~ arc also la id O il Ihe I()rcst a light, yel l(m ish ~ tripe from the tip of thc "noutt o fl oor. the occiput : a yclk\\\l i ~h dorso latcra l st ripe begi nnillg above the eye and continu ing bron/'c-cllioured Oi s jrihutioJl on the body ; a nother yc lluw ish stri pe fi'om the tip l' ndclllic 10 130rnco ..\'11. Ki na halu : Porill),!, alld Sg. of the Sliout he low the eyc to above the insel1ion of Kada lllai an nt Kiau. th c arlll; I imbs and posterior fl anks hn1\\ nish tn rai ntl y oli ve , wit h sma ll hind, "Iwt,'; and ligh t dots: ve ntral surElce light green to whitish g rccn . JlI\ enil cs with the Sa1l1l~ basic patt ern on head alld an[(:(JCIlU . BracllJ'",e/c.\' rior pnrt of the hod y , IS the adults. but more con0 1 I\ t l:l~tL & I3 mRoN, I R39 trasted: thc remainder of the upper hody is hnlll/eco lo ureu and allllo~t spotless: lim b~ and lail ~illli Bra chy/l/e!£>, is known o nl y 1'1'0111 the Philippines lar. hilt thc fo rmer ~ I ig hll y darker a nd the Inli er and Rorneo . 'Ii ghtly lighter than the p o~terior part of' thc body. Skink s w'ilh short. dege nerated l il11 b ~ , or \.\ i thOIil. 2-4 egg" pe r ehll ch . l3011y long a nd round in cro~~ - sec tion ; tail thid , rOllnd , ellds ill a blunt tip. l~ cu l ogy/E th u l ugy . 1/ J/('Iyg()t/(iI/ "; //(1111111 inhabil~ primary and second Litt le is known abou t the~c "ki nks ot her than that ary [i.lrests including CaVlll/r;1I11 grovc~ along sea the y live in the so il IIcar thc slIrl;lce. SOllle species s hores . Thi~. thermophile. diurnal ski nk ~ h o\\s a arc oviparous. others give birth tll li\'C you ng. Of the nine species known nn ly (Inc. Hl'lldll'lllc/(,\' pre i'cn.:ncc fur Ihe mcdiulll and upper levels of trccs 11/,,1\ . is cn dcmlc to ' abah and occur~ on l'vl 1. and arc rarel y eneoLllltercd Oil the ground or e\en 10\\ Oil trec trunh. They arc shy. a nd it is common Ki naba lu .
27 1
Bl'lIc/t.l'III l!/t'.\· ((PUS HI"II> \. j1 1'U(
111 mcl. ,\
tI('{( \
" ilundu I IIh , II1
II I~
""l
r~
P )S"' S 10 , h Ii !!
IP\ ,,1 ,,'"1
I \ 00111 .
19X2
h(11 ' -; : II
ih ,' 1"'"dqU,IIl "" "I Ih, " l lIah,111I " " 1:lh,IIu. \ :11>:111 , \I.d," ",l'· Ural III m(' /(\ el/I II \ I H I II W ) I\. "'; IC H, J.( I I 1\1 111 .'1" 1"" ·11, 2 1'
In' IClt'alit ., :
j, J~
1- 1 lI ~a r
'allollal ":If"- \11 j\ )~
I
2 ~' .
r\,
!,),)_"
lield nol 1:11' 1'1'0111 thi S PUill1 durin g Ihe' day. I his :lrea I\a~ urig illally ,'owred \I ith primal') ralll 1(lrl'~1. I hCII' rood IS Itkcly \(l C(lIl~iSI "I' ill"CCt:-, thcir larl'ae, and \\"Orlll" . 8/'{/('!t.\,lI/c!,'1 1I/)(/ ~ i, a live-hearing :-.p(.',:i(.':-. , TIll' parat) pc cont ,lInc:d I(llir cl1lbr~. (ls or-l 2--l3 111 III SVL 20-21 nlll1 II l)i ~ lrihuti o "
~g.
l)i ag rwsi~
"milioll . Sabah , 'It. h:in a ha lu : Bundll Tuhall .
Th~
ulIl\, IlIllhk~, ~pccics of ~klnk on lJorncn II Ith a c) lll1dncal . blunllall .
Ot:suipliull SVL 111ak" Lli 111111, 1l:l11ak~ I 1<) n1l1l ; TL malc, X-l 111111, klnal~~ X I 111111. I ollcrcyclid 11'lIh a single largl' SGilc; 11\1 C:IHljlCl1l1lg; l1(1stnl bl'tl\l'Cn ~uJ1rana,ai and lir,t :-.upr:ll:1hl:ll ; no postnasab: Irolltnll:!,ais widl'r Ih:1I1 lon~. lill'111lllg ,1hroad sulllrl' \\'llh rostral; prdhl\llal~ :-'l'parall'd lium l'ach othcr: Irol1tal Iung~r thall II ide: pariclab 111 broad l'llntact hehind inlL'rparil'lal : fro11lop:llidal, :-'CP:II':IICd: 11(\ or I pair lIt' nuchal,,: :; sllprallclilar~ . ~ccol1d largl':-.t ; only ~ supracili:II'" ill thc a11lcnnr (llrlll'l' oltlt.: Cy':: (, ~lIpralabials , lirst \ ':1") l:trgc:. Ihird and l(lImh bdoll cYl': :; II1rralabials: I large postnll'nial; dorsal:-. and Icntral" ('I'c:qu:!1 ~i ·I': . ~ Illooth ; 2~-2.J "c , lk ~ anlllild lIud-body : lOX-ILl ICI1ehr:lI" hetl\ l'CII panl'lal" al1d tall base: :'i -Xenlarg.:d preall:ils: 110 Itlllb~ : tail cylindncaL cndillg in a blulll lip . Dursal :-.lIIl;ll'l' \)1' hC :ld and hody dark reddi~h hn\\\ II , darker p(htcriurly: di~tal port iUII or Ih.: lad dark hlu\lIII\I reddi"h hla 'k: lip \)l'lhl' snout lighter; uClllar sc.dcs darkcr; I·.:ntral "urlrlcc light. :-.p()tlc~ .. neal11-l'Ulnllred : l'lIlhl) O' rcddish grcy uor ... :dl) . ~ nUIiI alld 1,(;llIral "Urf:ICC hght grl'y.
E rn lll)!.\' /E 11141 lu g~ ' The m,dc Iwlol) pc I\a~ collected on the road he1\1 e.:n 1':11Ill11[1uk :llld S()~op()don (1300 1500 III a . ~ . l.) un 1\ 11. Kinabalu at al'llllnJ 2.0() a.l11 . "hile Ihc 1i:l1lale paralypl' 11':1," dug (lilt frol1l a cullil·ah.:d
(I~llliS
/)11,' ;"
(l lia ·(II.·(~lI
(i R\I.
!)lI\1U .,, ' lIl1 r lll c' / 11
27~
IX..j~1
I X\')
(1 '1 II II'<. I ~(, 7 )
I X39
(JIC\Y.
or
rltc di:-.lrihutiun thc gl'nus f)l/.\/ll c:\tcllds rrolll India and ~n Lanka.l'a'l tu lilc :-,olllhcrn parh of Vil' tmll11 , and soulh to .I ,ll:\. '1Ill' gcnu:-. i~ rurthcnllorc prc~ellt 011 ~ (lu'hcm Phllippinc is l:II)(1-,. lodcralGI) large tu large , dilll'llal Irl'c-d\\'cller, 1\'llh rohusl bodies, kceled dllr,als , and long limbs (adpres:-.ed iJmb" OIcrlap). .Il1\cnilc" \I'llh a l'ltaractnl""C bandl'd pattern 1\ hich lildcs 1\ ilh inn.:asing a ~ l: . ()nl~ onc (I)(fsill u !t1"(/('('u) urthc Ihre.: species IIlhabiting Horn..:o has hccn rCl'onlcd lilll11 abO\c :'iOO 11\
Key to the Bornean species of D(tsill (in part after IN(,I' \( & RI«)I\ r-. 19XO) Vcn trals (COU lltcu iIl1l1iu- linc: "rOlll men lal lO \,el1l) more than :-6 ..................................................... :2 • VCll lr:ll:-. 56 ur f'e\\c;r .... .... .... .. ....... .. .. .. .. . II/il'l1("('u 2 Adults wit h 5-R uark nngs bClwCCII axi ll a
S(' llI icil/( '/1I
• Adu lts wi th 8-14 dark rings, pre frlln tals in hroad con lact in mid-linc ........ .. ....... ..... .. ........... gris('(/
Sahah l)lI.\lH J:; r1 .\t 'll ( ; " \I
Dasia
• •,.
Kmah;1I1i
BnlllCI
• "
Sa,., l\v ~l~
• ')
Ka lllllllntnll
I • . )
Seincidac
Dasia o/il'ucea (iRA', 1839 IX39 : .1.11 1)11" Ill ca lil.\' : " Pro n"" of Pliiall I' ,"allg. W", . Mahl) " ' ~1 11,1!.{),\() 111rI o/inl{'cllm. H UI I I /'Iocd'l( I ! H7e ; 2:' I : \1 01 1.tll\I{D 1 ~9()a : 1.13: Ill- ROlli, 1'11-; 201 . ~ M"II 19.11 : 31 1i
\Yak,', 1, 1:lIlt! " i
Oia gnos i '
/1. mo(kralcl y large spL:cies reac hing a maximum SVL of' tlp to 11 5 111m . Lasil y di ~ tin gu i s hahle li'om
all olhL:r L30rnean species o f' () a~ia by Iwo ocelli on the occipi tal region in adults and a not-handed tail in .ill\'t: llilc ~. Desc ription
SVL lip to 11 5 mill ; 10\\('" L:yel id with ~ caks; tympanum deepl y slink in a vc ry small eJr-opellillg: noslril in I nasal ; supranasals separated ; ri'onlOna~al I ..,ider than 101lg, tI"tlall y rorming a SUlllrc with the rostral and till.: li'ontal: prcrmntals usuall y separated (rardy tllliching); front al of' cql lal lenglh or a lillk
longer than Ihe i'rontopariclab and Interparietal" toget her: i'ronillpanetab of equal icnl;!11i nr a lillie longer than the interparietal ; parielab ,eparated ; I pair o f' nu c hal~ ; 4 "upral1nil ar,; 7-X ~u pra e iliars : 7 supralabial s, the lifth and ,i\tli helow Ihe eye ; X infralahia ls: dm""ls II ilh 3, 5. or 7. more nr Ic"" dislinct ~ecls: 2X-30 scale" aro ulld mid-body: -+ 1.t() \'ertebrab helll ce ll the parlet als and \en t; prea llals nOI or only sl ightl y cnl arged; li1l1h~ IOllg. the adprcs~cd hind limb reaches ahout the elbow; 17-2 1 smooth "uhdig ita ls beneath ItlUrth toe; wil 1l1usc l1lar. lon g, lapcrtng conlinunl1~ly 10 a line P0111t : X5 mediall y broadened su bea udab. Dorsa l S UrI ~lcc olivc in \arlllllS ,hades. with indi slim;t cross bands or dar~ spot". ~ll l11C or the1ll with light CC11lres: head llllcn srotlcd wilh blae l..: OCeip111 ~\ilh 2 ocelli . more distinci in1llal e~: ~elltra l SlIrf; ICC in va riably light shades 01' green tll whitish grecn. Black-coloured hodies or hatchlings ~\ith lIarrow reddi sh broll'n cross line s; lilli1h black . band ed light at their bases: dor~a l s urf~. ce or head li ght hi'( \\,11 wi th dark - margined se alc ~; tai l 111li I'orml y
Scincidal'
ye IlO\\' I~h
In reddi~1 hdllild Ihe ha se: venlral ~ur Illce lighl grel'lll "h, 9-1 ~ el va l. I X-Ill I11Ill lon g egg, per dUlch : hal c hlin gs \2 - IX 111111 '-;VI I ~() -5 () 111111 1'1 . Ec olo gy/F: llwlu J.! ~'
/)(/ ~III lI/iI 'lIn' lI
IIlhabll , lowlands and nwuntain(Jus region >. lip III appro~, 1200 III a .s ,1. where Il can he lound In prlmar! and secondary 1(II·est5. ill park s, garde ll ". pl alllallon~. and ('(/'1/(/ril/(/ groves. '1h e~c shy -; kl11 b exhi bil a natural curiosi ty :Jnd claim lerrilol'les \\ hic h Ih ey \'\:l1el1l1:11l1y defend ag; l1n ~ 1 cO l1 specilic~ and o lher liLards, When disturbed the) al\Vay~ run lip trcc~, It is a l11lost illlp()~slbk III ohser\'l: Ihe"e ~kinks for longer pl'riods a~ Ihey arc untirin g for age rs , Once a skink h a~ c limbed lip a Iree , it w ill Vi s it j usl abo ul cvcry Ihickerhranc h. 'hecK ii, rork, and epi phytic piants, probab ly ~e arc hin g I'llI' rood T hey descend tn th e gro und only 10 move 10 annlher trec, The~c li/ards Iced mainl y on anhmpo(i-;. l'l' ll1aics pl'llducc e g.g~ th rec 10 lilUl' lim e~ a yea r. nisirihuliull
Soulh of' 15' N, \. Iyanmar 10 ahollt Vietnam. Mala y Pe nins ula . Sumatra and S0111C offshore islands, ~aluna Island " .lava. and Blll'1leo. :\'Jt. Kinah alu : \\ilhoul preci~e loca lit y data .
(icll us L(("'prolepis FI i'/,lt'o. (oI ' R. I ~43 T he \a~1 disl ributi oll ar~a of thi ~ ge nlls is ma inl y Inhab ited by o nl y o nc \pecics. i .c. !,(/IIII'I'Il"'/)i \ ~ 1/I(//'II.I!,dill{(lII , which", nOI pr~senill n Borneo. The ra nge of /. //II//JJ'o/£'/ )Js covers Taiwan. some islands llrthe Philippln e~ . Horneo, and Ja va. >;everal oflhe I.es~ cr Sunda blands. M lero m:si a . c\\ Guin 'a, Australia (Cape York). and the Sololllon and thc Sanla ('ru/, I s land~ . Iclllbcrs orthi~ ge nl1s arc typi call y KnO\\1I ollly frl)11l a fcw s pec imens (eXCe pl L
Fi/:. 2RJ Jll ven ile spec ime n ll('
with sca ics Iike in /)(lsill and ilpll'IT,~od(l ll : earopcning small. but vis ible : Supranasal>. presc nt or ah~cnt ; dorsal~ smooth . or th c t\\'o s pec ie ' endcmic to l3orn co. onc (LlIlIII' mh'/Iis lIit'{(Il'cllhllisi) has becn rccorded ('rolll Mt. Ki1labalu ,
Key to the Bornean species of Lamprolepis
1IIIlII'lIg dilllllll ) .
. nlall to IllOdenitel y largc . ~ lendcr sk inks; lIsually ca nopy dwellers . Body d or~ovc ntra l l y co mpre~sc d : muscul ar limh:-. lolig (adpresscd lil1lb ~ ovcrlap); tail ~lcl1de r alld relall\,cly .. hort . owcr cyelid covered
274
/)1I\'ill 0 /11 '11<'('11
More than 23 sca le rows aro und mid-body .... ... .. .' ... ,' ... '., ... " ..... "." ..... " ... ,." .. ". ,' ,.". lIiellwellllllisi
•
Le~ s
than 23 sca le rows around mid-body ....... . .. v)'lIcri
St.:int.:idae
£ c()l ()g~,/ E th n ICl :'!J
Lall/prolepi.,· lIiellll'ellllllisi (LIIHII DL J ELDE, 1905) I
'·!:o.' ,,"'
\ \1\
Bl u~ " t
/. lIlil /JrO /I'/ lis lIi(' lIl1 '(' II/lIlI\ i
canopy and rarel y desce nds
I 1\)11 11>1 .1 11.\11. I'IOS : 1')_ '1.\ pc LOllg I:I lu. upper \l nh,,""alll. 1'",1
KaIIlTlalll ~II I' 1I 'L:0\II/1W 1IIt.'1J l1'c.',d'lll \ I. '-;\11111
pro bab ly Inhabit s the c l o~e 10 the ground .
f)i s tributio n Ende mi c 10 Borneo. '11. Kin a h"lu : KI
I 'n I 2,, " I
f)a ~ ;(/ IIIt' lIl1t.'lIhw\t, f\·I \NIIII' I,)~l 26
r Ut
NI I \I{P
&
~ J(Ii. ,WI '
191\4 : 24 Itlm/wo/t'/II.\ mculI('IIJlld\;. T " 1993 : (, !uIIII/ro lf 'I JI.\ mel/I!'!'''!''';,\ ;. \\ \ 1 .... \11 '''i 1()()--lh . 2 ·Hl
Genus MlIhllYlI I-'n I
r>i a~ n os i s
;\ modera tely large sp.;cies reaching an SVL of up 10 72 111111 : 2-4 rows o f enl arged ve rt ebra Is: ~4-26 ~ca ks around mid-body; 18-20 subdigirals beneath (()urth lOe. Dt'sc ri ptin II SVI . up to 72 111111 . TL LIp tn ()(J mm ; lower e )'el id ~ with scak~; car-openin g. very s ma ll. with small l (l bul e~; nostril in I nasa l: supranasa ls ~ m a ll. ~e ra ra ted: rron lO na ~al as wide as lo ng. fOllll ing. a short suture \\ith ro~tral . in COlitact with frontal ; prefrontab lIslially ~e p ara ted (rarel ) in enlllaet): front al ~m~d l e r than frontoparictab alld parietals together; ['r(1n torar iel a l~ smaller than interparietal: paridal~ ill contad behll1d interparietal ; I pair ol" lIuchals; 4 ~ uprao c ulars : ') ~ lIpra t.:i liar~ : fifth supralahial hcitm eye. twi ce as high a~ fourth : 2 - 26 s t.:ak~ around mid-bod y: 2-4 rows of enl arged \ ertcbrab: pn.:a nals slig htl y eil larged; l il1lb~ long. a dp rl:s~e d hind lilllb IOllching Ihe clbO\\ : I X-20 ~ 11l ()ol h sllbtli gital ::- benealh fnurlh toc. Cn l()ur ill alcohol (after '~ I IIII It)J I ): " Bad, light hro\\'lIish-gre.;n with di stinct mctalli c glllSS. an d numero us scat te red hl;lck ~ po f ;" and blat.:\.. ed~l n g.s it) Illost o flh e .. caic~: basl' o rl ail ahm c II lilt black tranwerse han(b : top of hl.:ad hrown.;r than back. scale:: ~tf'<)ngl y olitlilll.:d with hla t.:k: bdO\\ lig.ht ~ke l blui ~ 1t or greeni sh." rah.32 Dblllh"ll(lI! of
:IIII III/J'II illdlJ!!·l'I/.\,{/
IN( a ' H,
1~Q6
Ski nks (lfthe genus iIIalll/.1'lI are di ~ tnbllh; d 111 south AlIleri ca. Afri ca. soufhcll1 and ,()uthea~tern Asia. Mmkratcly large to large. diurnal gro und-d wci lers with ('()bu~t , dorsovent ra ll y cO lllprc s~ ed hodi es and Illu sc ular. well-developed limbs and fail s. Lower eyelid co\ ercd \\itlt ~mall scales in all species inhab itin g lloll1eo: tympanulIl deepl y sunk ill a eicarl y \ isiblc car-openi ng . . · lIprana~a l s . prefron tals. and di vickd frontopancta Is prcsen t. Om. als with s.;\,cral I-.e e l ~ . All o f the four species kll(1\\1\ from Bo rneo occur on MI. Kinabaili .
Key to the Bornean species of Mahlly a ,\ 'Iore than 29 sca le rows around mid-hody ..... 2 • Less than 29 scale rows aro und mid-body.. .. 3 2 Light narrow stripe fro m labia Is to forelimb or beyond; I>l11a lll>pe e ie ~. SV I of adults lel>s thall 70 111m .... .......................................... illde" I'ell,m
• I.ight stripe ab~elll : large s pel:ie ~. S VI of a dult ~ more th nll IO() 111111 ........................ IIII1/f(l'a.\ ('iaf(t J Less than 22 sllbdi gi lal s; dorsal s wilh J ked s;
large spec ies. , VL of adlilb l110re Ihan 100 nllll .... ....... ,...... " .... " ... " .. ,', .... ,.... ....................... I'lIdi.~
• :\1ore than 21 slIbdigl tals: dorsa ls \\'Ith 5 or 7 keels; ~ l11all spc..:ics. SVL of adult s less than 70 111111 ... .............................................................. I'I/g il'em
I/Clhlll 'CI - spc'cil'~ 1111 HOl'llen
B ill >"" ,\
AI .
\I '.
;\I{/b l/)'a 1II11/IWm'illfli ( KI III.IX20j
Mll bl!£11 rullis t ! (H II 'l,d k. I XX7 !l/abll l'a ru.!!if('/'{1 (SI(JII(/I,\. IX 70)
Il)XO
-
Sahah
Kinaballl
• • •
• •
•
• •
Bronci
Sam\\ak
+
• .
KalllllanUln
• • • • • I 275
.:l/a/m.J'a illt/t!prt!IIStt BR( )\\ & l ( \l '\. 19HO \/011/1\ 11 IIldl/III "'"
HI :')\\
S.: \ (
\ 1 \.
ll)~O
1(I\"ali ' ~ ; '·,1"" 1 11'''lh,·,,'1 "I """ I.,,,,. ~llIlll(\,,, \/ohl/la m"llIuII Il1il/e/ (l1ol ItlIc(uti IJJII/lff .Inllilfa f\I, u III \Ill} IX''' •.I 1 I I{' .')II I'll " ltd \ luhll ro lilli/lie Ilrlna/rI , (1l01 I tlilllll i lilli/III IlIlIIo/a
,,_It
\ ar\ IIlg dlstanccs a~ a IInc or rnw or sPUb ; \'cntral "til 1;lce grc~ "b-hille 10 ,laiC \I Ilh hghlCI flc\h y <Jrt:a" on thc chin wdl a~ uSliall y i"llrc and hind 11)\111 rcgi()n~" Iialehiin gs 22 .5 to 24.7 mm ')VL in PhilipplIlc p, )pllial i{)n~ .
,i'
I~:-
('~' I H.·
hl,IIIJ" (i H\\. 1 ~ 4 ))
(, II \\.
1X· I ~)
" "" II'nl ill, ,I ~I " 111\ 1'1~1 2(>. 1'''1'1\1(11.1.: :-'1"
"If'
Iq ~ -1 ' I , I ", 1'''Ii 7. \1 11 ~'"' 1'1'1-11> '-11,
Di ag ll n~ i ~ ~Illall ~pn: it: s (If I/ahul'll. k~s Ihan 70 Illill SVL; 27-34 ~cak, around mld-o(ltly : -W-4li \cak row\
,\
ht:I\\Cell jlanelal~ and ha~e llrtaJl ; d())"~ab \\ IIh 5-7 (\)) ~ccl~. 11<-24 \ubdigilal, bcncath 1i.H1rlh l(lC.
Ill.'sc r i III in n -;Vl.up 10 (,7 1ll111: c:lr-ojlcl1lng \cry \Jl1,JlL lI\lIafly willwut I,'huk\: noslril In I na~al ; 2 ~uprana\al~. ~l:parah:d, IIln ~', lIarrll\\,: {"rolliolla"al a" \\'Idc a~ IOllg. Itl)")1lJllg a \ullln: \\'llh r()';tral alld frontal ; pn: fro III a I" clll~~ J(lgclher or \\'idd) \cparalcd; lI'onlal aholll ~anlc kllglh a~ fn))lIopariclab alld Inlerpanctal logcth,'r: jlanclal ~ lI~lIally \cparatcd by a largc 11I1t:rparietal : 4 largt: "llpraoclllar\: 6-7 (6) ~upralab l al" lifth \cry large and belm\' thc cye: 1I~lIall) h infralablal,,; dllrsa b with 5-7. rarcl y () ~ct:1s (halchllngs and 11Iv(;nik" only ..\ ): 27-..14 (' I) \caks around l11id-body: 4()-4~ (45) \wll:brals ot:tll'cen panctal~ ;l11d ha\t: of tail ; (52) \ ' t:IHral~ bct\\'cenl11cnlal and \ cnl: prcanal\ IHlt or only slightly cnlargcd; IlJnh, lel ali\cl y s hort. adprcssed hind IlIllh 1101 rt:a 'hlllg Ihe dhow; IX-24 (_1-_2) ,,"hdlglIal~ beneath l(lllrlh 10e; lail abrllptl) tapcring behind I" ha~t: . Colour in akohol (arter l3RPI" & AI ('A I \ 11):-<0): "I)nr~al !!IOIlIHI colollr lanni ;.il or grccllI.sh nlilc In Ilghl hnll\n . u~ually \\'ilh 2 to 4 longlludinal I'll\\'\ Di" ~l11all dar~er brown tu blal"~ish blnlche~. "OI11C ~pt:Cllllen~ \\'lIholll paltt:rJ1 ; \ ertt:hral arca bCI\\'cen thc il1Jlcrro\\~ol"ca ~ itlJlally forming a \agud y light hlllJsh \'crlchral 'tripe; narrow light dor~olatcral "Inpt:~ \Olllt:lJnIC\ present from pO\lcrior corm:r (If eyc In lall. In ,,(1l11e speclmcns pre~cnt nnl~, antenorly, \Olllt:t il1ll:" bro~t:n inlll \l'all<:rl:d hgh lt:r bllllchl'~. or \ cr) 1;11111 : lateral \url:ll'(;s dark brown, marked hy a (h .. IIIICI. hght 1I
EC lll ogy/E lhlllu g ~
Thi" ~~111~ ha~ hecn rLcordcd froJ1l altitllde:, lip tll 12()() 111 on Iht: I'hllippillt:\ \\ herc ~pCl'IIllClh \\l')'C liHind on Ihc ground undcrneath lea\'es and III a dry ~trt:aJll hcel in ,el"llIHI<1r~ alld pnl11<1ry li))·t:,I ... lli~1 rihulioll Philippinl:\ alld Homcn (Sabah) . 1\'11 . h:inaualu : withoul l)J'cci~e InealilY data.
Rrmark : I hc \pccilllcn classilit:d by MII( (Jl'\I()l (I XI)Oa) as ;II II/II/I/( '(/I'illlllll I\a~ rc-examincd hy Ihc <1lllhllr. Al'cmdillg 10 II" ph(lhdo-;i, (gi\cn 111 bral:kt:l~ in Ihc <Jb(lIe de,cription) Ihe g.ravid Ii:mall' ("0 nlill SVI. I Xl) Jllm TI.) cou ld he allocatnl lo . 1/ /1111< '/11 '(,1111/ \\'llhoul doubt. 'I hl: nUJ11her of keel~ cou ld nnl hI.' \ cn lied duc to t he poor slat.:: or tht: rt:\pcctllc 'pecil1lcll . Thl: ollly dir!i:rl:lll't: wOl1h mcntionlng wa~ the prescnce nf,mal l inhuks \\hich limn all alnlO"t conlllll H H I ~ nng \·I'llltill Ihe ear-opcning. It can Iherdl)re bc a~suilled thai the skinb dctl'rm lncd as . 1/. 1I111/licl/rilllllll h, SI\.1J III (I (>J I) al\o hl:lon g ,I/. Illtlell/·/'I/.\'(f, and Ihal /If. IIl11lriulnlllllll d()c~ not form pan of thl' l3ornt:an herpctolillln<1 ("ec M \I \1 I. I III<. IIl.\ & '\ \MIH IIJli5 ' 159).
MlIhll.l'lI IIlltlf(fascillfll
(Kl lltl..
I X2())
IX70 ' .... 6 ' 1 ~' ll l' !tH.' a li. ,\ : I1P1 I.IJC '" '.I p k ", dl', ;)!" ,, ' '' ; .I,,,,, (~It I< II " 1'1.101 ,l/"hw(/ "lIIlfI/O\( Ic/lO . ~1f)( I'l .\N Il nNOd 1., 3. III i{( I"1I 191 ""I 1/,,llol'III.'IIIII/II/ ~' (" 24 (, I'i'i(lr 2'1(J
.\ ''-1111'1/\
IIf1t/ll/a \( trllII\ 1\: 1 III .
'IJd,,;alcd
Diag n osis
SVL I I L lip 1(\ 400 1l1l11 largest skil1~ 011 I3(H·ncll. usuall y with con,p ICllOll\ whitt: occllt on Ihe l1an~ s. 30-25 st:alc~ arolllld mid-hod y; dorsa l!'. with 3 (5)
,'c im:iuae
Fi!!, 211..
'\ /lIhlll'll lI/ulll/I1\{'/t1l1l
li'om SUlllatra,
hl'cl~ : -I2 --IX vcr\t:brals bet\\'een parietal~ and oJ' tail : 16 -~() 'iubdi gila ls beneat h fl1UI1h toc,
ba~e
Dl'~criptioll
SV I Illale, up t~1 13 7 mill . fcmalc~ up tn I ~I) 111111. I'L l1lore than ~(){) Illm ; ear-openin!.!. rd,lIi ,dy large. round , with se\er;d or no posterIor lobule, : n(1stnl ill po,terIor portion or nasal: I postna'ia l: supranasa l ~ ~cparakd, (H.:c a~i~lIlall y rorming a ,L1turc: f'ronttl ll:l "d Iwi cl' ;!!-> widc as I 1ng: prefrontal" II~II al ly in e,1ntaet behind frollitlll il,al. l1lor' rarel y separated: frolltal a~ long a~ or ~ I ightly shoneI' th::tn f'ronlopa rieta Is and Intcrparie t:l I to!;!l:t lin: frontoparietals largcr than Illlcrpanetal : panctal, !-eparalcd h) a large inlerpanetal: I pall' tlf' nllehal ~: -I large su praoculars: 5-6 ~upraci l iars: 6-7 'illprala blrd~, lilih helo\\ the eyc: (1-7 inf'ralabials: 30-.15 (u~lIa ll y .\~-3-l-) scales around l1lid-hody: d\)rsa l ~ \\ ith .I. \ ery rarel y:; weak keds: -1 2--11' \ el1eh ra l ~ het\\ cell parietal, and ba~c of' tail: prcanal~ not or , IIghtlv cnlal'ged, len!,!th of' liIlIi)~ c()rn:lallllg \\,Ith
age. the adprc~~cd hilld Ill1lh reaching at Iea'it thc elbo\\', fourth toc With I (1-2tl Sl1looth ~uhdigita" Dorsa l s url ;lee hrm\ ni ~ h 10 01 i\\.: hro\\'n or DII\ e gl\:Y, a lnlost IInicoloun:d or \\ Ilh line blach linc~ : Ihld, dark bn\\\ n lateral slrIpe lI'lIally with hlaek bordered, whitish ~pots : male, and Ii.:lIlale~ olkn ~ cll(\\\ i,h, orang " d 'ep red . or a cOIl1lm1atlon of' thesc co lour, latcrally, t Iroat and ehc~t onen colourcd correspondingly: vcntral 'iurl~lec light beige. yc llll\\ I,h , greenish. or li ght gl'l:y: hatchlin g~ alld .ill\cIlilc~ altogether darker: limbs ;Jnd lateral ~tripe allll(\~t dark brown to black ~\Imetlmc~ with light dotting: dorsum goldcn hJ'()\\ II, each ,ca le hordered \\'Ilh hlach, I i\ e-he
in
,nllth-ca~t !\ ~la ,
It I, 1I0t re,trIcted t,l any
~77
Fi:!. 2115
.l1I~1
h(lIn I/,,/nll"
1II11f/llmnt/!t/
from S,)lllh 'I h;\iland .
parliuliar Iype oj' hahil at and ea n he fOllnd lip 10 apprllx . 111(1) III ;1.:-- .1. (llh~ervation by Ihe author Iln Ciunllng Sihayak . SlImatra). II inlwhll:-' a wide vam.: t)' or forest~ . and CIII ; d ~ll be fOllnd in area s with Ill\\ \·egetallon . This sk ink i~ not very COI11Illon ,)11 MI. Kinahalu. prohahly dlle to a 11Iore e(1I1Ipetit iw si tuation causeu by the pn:~e lll: e oL\:1. /"lit/is Both s pe ci l'~ prcli.:r ~ lIn -e :\pused patches on wider paths and clearings. I he animals arc attcntive. love to ha~~ , and arc e.\l'cllcnt climbers. B C~ ldc s;) wide vanety ()l"illsecl~ lhey prey a lso on smaller li/.anb . Males in p;\rtlclIlar arc hi g hl y krritorial anu defend their krritnne~ \ iguroll s ly again~t other ,kink s. I)is tribulion l' wlll thc :\ndalll an~ and iCllbars through Thailand tn ~()lIlhern ( 'hilla . ami from the Malay Pe ll in:--ula to Ne\·\ (illinea . Palau Islands, and the Philippincs. 1\ 1t. h:ill abalu : POring. Kl au. Sg. Kadam::Ji an . HUl1dll 'l uha1l .
2711
lHlIlJllya rut/is l3(\t ll l·.
(01 ' 1{,
I XX7
I/"hlll(l IIld,S \I, ,, II N"I~. I ~X7c
Il>X. 1'1 \I : li):.\ t ~' lll' 11I ('"lily : " ~ I dldllg I~:lr;"':lh. fj,'rncol and -;11111:11,.,, " \/,,1>111111'1/,11,. :\1<>11.'1 "" t )\I)(),1 t l l. 1)1 ({,"'" t ') I" · I(d 1/"/>11111111.1".1> 1" "11\ 8< J)IVIK l'IX 2. 16: ~"NIIII \ I 'lX' ::!h. ,", IH ' "\HIJ& "1111010 N )\)X"' .
.:.!". ri g ..
f\1\1
\tI 1.... 14.)~ 7 ~H5.
1')XXd 9 . I'N2" 110 . T " 1'1'1) . 7, 1\1\",.,,, J9 ') ~h . 2 ~(•. 1')l)(K 290 . I\1\NII I" & (i"""M.II.. , 19'17.272, lig . I')X
I)i ag llusis ;\ Illoderatel y larg\: ~kil1k (lip to 120 111m SVL) varying considerab ly in colorallon: .'\-l-lX vertebrals b\:lwe\:1l pari ctab alld base or lail: dllr~ab \\ilh 3 k\:ds ; 2X-](1 scales around mid-bod y; fo urth toe \\. ith I X-21 ~uhdi~itab . I)t'scriptiun SV I. lip tll 120 111111 , I'L up to 220 1111ll ; \:ar-opening
round , occasionally with s mall anterior lobu les; no~tril in posterior part or Ila~al ; 2 supranas;\b ; frontonasal wilkr than long. formin g a su ture with
Scincidac
ro~tral and occas ionull y with frontal; prerrolltals uSlial1 y separated by a narrow gap or. more rarcly. forming a short sut ure; fruntal little larger than frontoparietals and interparietaltogdh..:r; froilloparietals larger than inkrparidal; pariel
«)
279
~Clllcl(i:l e
Abole dark hroll n . slreah.c:d II ill1 yello\\'ish bnl\\ n rrolll 'llpercillary hnL'
Fij.!. 2N7 1/"/11/1<1111.'-" /""" lrolll P"'lIlg
1I\1\:l1al, u~lIally 1I'llh :' :-.11'1111); kcd,. 1<1lcral:-. abll II Ilh 7 h.l'l'l :-., '. aroulld 1l1id-h(ld); \ I I'l'rlchr~lI, bclllecil p1 and lilrc-llIllh:-.; belly, I clltml s url;lce or hlml Illllh, alld t;11I s
i.,11 bille: or tail rl.'l ld i:-.h
~a IIlH l1l.·'
'\Imo' t unirorml ) da rk l)llIl' brolln lIith lI eah. ly indl ealt.:d. lig hter dorsnlalL.:ral siripl" alld all eqllilily 1;lint I crkbr~tI ;,trrpl.' In the pos lerior lh lrd nr lhl' hlllly(I\L" "II' &(i R' )""'~ 1 1 \j 1')<)7 alld IIllpubl.). I.alcral " url ~lce or head tinged 1\'Ilh reddi"h : chin. lhroat , and che"t \cllO\I ish grl.'e 11 , re111 ,1111(\<:1' ur Ihc leiliral ,url:lcc dirt ) \\'hite. (}lhL.:rII I~l.', dorsal surl;lec or body broil 11 With nUIlll'rOLl:-. hla ek marks and." Il g hl brown stripe.,. dllr"" laleral Olll.'!-> heginnin g . orange- red Oil laleral >'lIrl;lce . \\' 1111 bl ack spOb. ( hln and Ihroal rallltl" orange. remainder or \\.: ntl':ll ,lII'n lCe J elloll i>.h g rCL.:ll.
""1\11 11 '
or
Eco lUl!y/E t h()lu l!~' ..lllI/lIIrt/ "" .!!,i/('/'{/ IS a l·l.'r1 shy. quill.' rarely l.'nCOIIIl -
lered Inhabllant orprllnary i'orl'sb . II appcars 10 be more conllnon in the Illwlands althollgl1 il has beclI rel.'llrded rrolll up ttl 12()() 111 a.' .1. in the lellggl'r r-.lountaln\ or .Ima. II i, u!-.uall y round alnng Ihe bank , or sillaller ,Ireallls. on clearillgs. or on patl1,. I)i~trihution
lanatwn or Il:maks lancr 13 IRIII II I X95): ",\hOll' hlack"h hrOll1l 11'1111 ['laic hrown IOll gl111(l1 n;1I , Inallon!-> , '1hl'llal grl:c n. black spolted .
((110m
2XO
Nit:obars . 1\' lalay Pcnillsula . ilornco. ')ulllillra , rulau :-.JiLl!->. -knl a\\a llsland:-, • .I;l\LI. \It. h:inahalu :
"on II ),:.
Scincldae
Ge nus Riopa CiR w. I X3l) The di stribution of the genu~ Hio/III eXh;nds 11'0111 continental ;\ liia through the 1'v1:l lay Pcninsu l:) and (jrcatcr Su ncia Is lands tn Sula\\'e~i and the Phi lippilles . Sknlkr. ciongate ground - dwellcr~ with ~hllrl 10 rudimcntary Iill1b~, adpressed Iimbs do not ()\ crlap or evenmeel. Supranasals presclll (\ ~ ab~cl1t ill ~1I11ilar- l ooking ~pccics of '~/J!l('II()II/()rjJlJ/{\' ): lower
I
/(io/,u
h( /lII/~l.rIt!('i
(13 \ 1
Rill/ICI bOWl'i,,,·ii
(
I II .
i "~I1 I1 I< ,
I X6..J)
More liI:tn .17 ~calc ro\\~ arollnd mid-body; a large species. SVI Ilf adlllt~ grea ter Ihan I DO 111m ... ... .. .. .. .............. ... ....... .... .... .... ... ... ........... hl/ll//'fl 'lch'l
• Less Ihan .-n ~calc nl\ls around mid-body ; a ~lI1a ll species. SVL of adults ~Illaller than 61) I11Ill . .... .. .................................................. ... ... ..... bo",,,;,,!:i;
1 1'~:o.\lHJ1t1 H-/IIIt 'Ir . Ilih Io. lIld
n.d"
h ~' l l
,lIl l' T 1>1
Rpu
1I't!,'\"U1tlhOhll ll.1!.1
I
H.~n.\" Ulll hU ll /lno..:,l1
NI Ol'lII>U \\ F t1I,!~1I
I
\l tH
PJI ". :--' \11 II
I X6..+)
I ~p(' l u('ali f ~:
(II I \H,., 1 }\90~1 :
144
fl o ug
l )pl' l uca li t~ :
~ ft-l )
III I{ I"II
~1 \\J II I! ,
\/0(11111\ h , )\\.,./l I,<..:.1 I
((;11\ 1111 ' 1(.
l X()· ' \) 1
1IIIIII.'tt'_\ hOIITill1!1I (i l '1111 I?
19 1:'. 2(, 1. 1 \, 1')q1. h IIn I 1. 1
I~J Xl,
iii Nfl \PI)
26: \1
& '" I. II ,H I
"
Rema rk: The validity of thi s gcnu~ I~ suhlect 10 debate with ~ome author~ a~~lgnil1g the re~pe e tl\, e s lx'cie~ to the g'llllS /.I·,!,!,1I"fJ/llrI (sec GI{l I I{ 1')7 7).
Sabah
• I •
I XlI5 )
Key to th e Bornean species of Riopa
RiOI'll bOJl'rillgii
cyclid c()\en~d with 1111': ~cale s or \\Ith an undivided '·window·' . Tympanum vi~ibk , lima ll. i\ terrest rial to fossorial. diurnal and cr.:pu~cu l ar ~h.inks keding on smaller il1\ert.:brates; reproduction Ihrough ()\ ipo~itinn . Of the two specie~ recorded frol11 l3orn.:o. nne (RiojJiI hOllrillgii) occurs on I'vlt. Kinahalu .
\1 ,,\ 11 ..
19X-l -
1994h
~ 4h
2-i
Dia ~ Jlu s i s
Small skinh. up tn :;X Illlll SVI. : llallh.~ ollen L'lllI)uri'ul (black . ) i.:IIIl\\,. and r.:ddi~h) "ith nUllll'r()U~ lighl dots: -+ sllprallcular~; 2X-30 scall:~ aroulld llIidb\ldy: fl)Llrlh tlli.: \\ IIh 10- 1(, ~uhdigilab ; tall thlch. at ha~e. tapl:rin!; graduall~ .
K'llah(du
•
Bnln~1
S"r,,\\'a~
11-:.IIII11.1I1Iall
• •
OrsCf'ipt ion
S\, I up III 5X mill : TL grealer than SVI : 10\\l:r .:yclld "ith ,,111::111 scaks; ':,Ir-llpl:nlllg ~l1lall, with or \\'itlwul 1-2 antenor lobllk~ : III)stnl In a quadrang ular na~al : sU]lralla s al~ largl:, ill ellllta '( behind r\)~traL rar.:ly s.:parat.:d; prd 'rontal ~ ~ll1alL \\ idely :-.eparat 'd; frol1lop::lrielab separated; panctal~ I;')rmIIlg a ~ lIlL1re behind interparielal ; t"'lally I pair of IllICh;ds; -+ large sliprallcular~ ; 7 ~uprac"lar~ . lil~ t and se\·enlh largc~t ; 6-7 slipralabials, 111'1h large~t and hell)\\ .:ye ; 6-7 infralablab: 2(,-.12 (u~uali y 2X-JO) ~nlollih scales ,lI'llUnd mid-bod) (dor~al~ r~II.:I ) \\ ilh J \\\:a]';l), dc\eloped h.cel~) : :;2-62 \erli.:hral~ heI\\ccn parietals alld has.: (lftad,.t prl:anal~ ~ll g htly \:nl arged: adpres~cd IlInb ~ dn no Illllch : I(Hlrlh to.: \\ ith 10-1 6 ~uhdi g ilal~ ; tad Ihi ch. al ha~e , laperlng graduall:-'. nor~al surfacc yello\\'ish 10 hnlll/l:-hl'l1\\ 11 : fin.: black ~p(ll s pr\:sent (In dor~ab . fonnln g a longiludinal s.:ri.:~ : black dorsolakral ~tnpc I'rom the l:Yc III thc anterior ljuarler 01" thl: I:lil , \llkn broh.en hy ri.:ddi~h :llId yellow /onc~ I S Oll1etllne ~ bordcred \\ ith v,'llo\\' above): a white linc alllll).! ~lIpral:lhlal~ nearh reaching the ear-oj1l:lIll1g: lalcral ,,"r1;ICC of lIech. and hod y u~uall~ 1111111 led " 'llh numerous Ilghl and darh. ~Pl)b : tail I)ne n rcddi~h \elilrulat.:rali) : \enlral ~lIrf~ ll'e oran g.: to Pll1h. . Jll\cnilc ... oll\e bnm n ahlnc'. 2-.t .:gg ... per clllh.:h.
2XI
SCll1eidat.:
a panll'lIbl IYpt.: of' lwh ilal. In Cact. il inh ah ih 1<'1\\ lI~jll ~ t a~ \l'ell a~ Cli ll il'atni area~ a nd rort.:st :-. edges. II is ofit.:11 st.:t.:n on bank,., or slrt.:al11~. Dliring the da y. il i~ ratha ~el.:l'e ti\ e 11\ In g be nea th leaf lillt'r. roc"~. I;tlkn l(lg ~. and ~illli lar con: r. \I:1ln aelivil~. pl' a"~ du ri ng early It()l l r ~ o f du sk. hut it Ill ay emerge to forage a lso III ll\ erca:-,t \I l'ather eo ndil ions, 10
F ig.2!!!! Hilll'/l f"III 'rtllgii frolll Kota Klllailalll. Sabail .
Ecnlngy/ Et Iwl(l g ~ morc CllIlIllllll1 ly fOllnd at 10\\ ,,!e\atiol1 s. rard y lip III 1(,-10 111 <1.:-..1. al1d i!i 1101 bllllnd
Th~ ~"i n" i ~
CiCI1L1S S pl1 t'//O/II0I1JIIIl S 1; 11 / 1"(; 1 R .
I ~-t3
ur Ihe !!t.: I1tI ~ ,C,·, d,CI/O/J/OF,,/tIl.I art: dt~trib IIteu III nlll lil1 ~ l1lal l\ ~ia, the Indll-i\II'aralian ,\rcltipt:iago. ~1I1d the Philippil1c~ . Olkn uirti t: ult 10 di ~ tinglli~h, th esL' nre ~l1lall to mode ralel y largt.: . usually ~Ie nu er ),:l'OlIl1d- alld Ireed\\'d kr~ . -l rll ll ~ lkn:lo pt.:d limbs , lI~l1all y ll\ erl apSpet:il'~
Di strihul iun Indi a (i\ ndam an~). r'vlyal1mar. Thailand. l'anlbodia . La o~. Viet 11:1111. ~o ulh t.: rn (,hinn . I iollg Kon g. JVlalay Pc nin ~ ula , Sillgaporl'. L3orl1t.:ll . .1;1\ a . .'lIlawesi and tltt.: I' hiiJppinc~ I Sulu Art:h lpelag')). \II. Kin"b"lu : willmllt pre c i ~e 10 'a lil ), da ta. ping \I·hell adpresst.:d . Lllwn l'Yl' licb covcn:d with small :-,caks . i\o lo bul e,> aro ulld tltt.: car-opening. [)nr sal~ and \e ntra l ~ lI ~lIa ll ) smooth . No slIpranil~al~, 2 lorea ls. fro nt oparielals and inte rparietal di~t inet and 111 broad conl;I(': 1. Repl'llductlo n in S Oil It.: ~ p eeie~ ovovlviparolls. in othe rs 0\ iparolls. Horneo is hOllle tn 1-1 ~ peeie~. of\\'hich (, have beell rcco rded frolll M1. Kinaba ill . Onl" .'>i) //('''()''/f)rl)''I1 ~ /('I/lliclI/1I1, appear:-, to be endemic to 1\11. Kinabalu .
Kinaba tu
Sahah S,,'lt'"()) )l(l']J/III.\
'1''''
(/ljiwli (I\ot I I 'oJ, ,I ", IX'IX)
'I)Im""I""/\ hlll1'~"j()f l l ll) III 1" II I III . 1')0'1 5.,,11""01111"11111'\ 1',l' a " IJI,"'"" I' I" ,,, R 8.: II '''~II R, 19/,' '\1 '/'CIIO/HI JI plitt, IIt1d' I "1( , 1 U i..'<:.. Il n' \III{, 1%<' .\i"(I"fIlI""I'''//,' /",111",., (111)111 III JI I 1l1 . 1')0<;)
~
• I
•
•
:'pJIl'10mwphu\ mll( ult, fllIlI\- IIM"",19(,7
.\i,h('"('lIIlIrphll.,
1II1I/ li'I/IIIIIII(I(II.,
I
I '\(.11(, (III(.
•
IX<).j)
•
I M'IR~ 1'/5X
,\i'/I('lItt1IUJr/,/JU.\ \h . 'U'~nlr ( liOIII " (,IR. 19I1UI ,\/J/h.' II UIllHI1'hll .... ,\/d/llfll'
( II (H 11" Hoi 1<, 19110)
.\'l 'ltelltllllorph ll '· ICIIUiclIl", (~ IrH. fJI IKI>
2X2
I X'/Oj
•
•
1') - 1(
.\j,ltc 'mOllflll,ltU\ mil/lit/t Ill"" ~\11111... I ()~.:; Sp h (· II 01ll1lIp h fl.\ _... flh/lIl l1 ,\'
I
--
-
• •
-
•
+
•
-
- -
• •
-
-
• •
Kulilllalltan
•
•
.\'plte"olllorl"w\ J..i,,"/J"/lInl . . i.\' tB \" III II . I X') ,)
,\ph .... ntIUUlfl'hu\ mlldtg llt/oll t I1(H II
Saraw~k
Hrunci
•
•
• -
•
-
c--
Scincldae
Key to the Bornean species of Sphel1ofllorphus (in part aftcr Limbs short, adpressed limbs do not meet. tail thick at ba e; dark lateral stripe present; 26-30 sca lc rows around mid-body; 4 s uprao cu lars (collnt cnding with the sca le touching both frontoparietal and parietal); 7-12 subdigitals beneath fourth toe .................................... ill/reili • Limbs long, adpressed lim bs (sometimes nearl y) JI1eet or overlap ............. ..... ........ ....................... 2 2 I'ari etals ill contact with supraoculars .............. 3 • I'arietals not in contad with supraoculars; 36-40 scal<: rows; 12- 14 suhdigitals; tail thick at hase . ................................................ ........... ..... . hallieri
.3 Dark dorsolateral or lateral stri pe present ... .... . 4 • Dark dorso lateral or lateral stripe absent ....... 10 4 4 supraoculars ......................... ........... .... .... ..... .. 5 • 5 or more slipraocu lars ..................................... II 5 Less than 2X scale rows ................... ..... .. ........ . 6 • More than 2/1 scale rows ................................... 7 6 4 longit udinal series of dark spots, 2 on the baek and I along cach side; 24 sca le rows; 21-23 ubdi gital s; ta il thick at hase ............. hlllliko!i'ri • Dark spots on hack not formin g long itudin ,1! ~eries, dark nanks whitish-spotted; 26 scale rows; 2 1-24 subdigitals: tail somewhat thick .............. . ... .. ... ................................................... 11!1I/li('/tI/I.~
7 More than 26 suhdi gita ls; 32-3..t scale rows ....... .. .. ..... ....... .................. ........ ................. ... sh"l/im/i • Less than 20 subdigi tab: 32 scale row~ ...... ... ... .. ...... ... ................................................. //lodiglial/ii II Usua ll y 5 supraoculars (rarely 6): black dorso-
lateral stripe with mall 'eattered ye llowish spot. ; lower lateral parts o f neck and throat nelth cr greyish to light hlue nor deep blue: 32-38 cale rows; 15- 17 subdigitals: lail thick at base .......... . ...... .... ... .... ... .. .......... .. ......... . ........ kill/lh/lllle"si.~
• Usually 6 supraoculars .... .......... .......................
I)
9 37-42 ~ca le rows; lower laleral parIS of neck and throat greyish to light blue (females), blue (males): lowe r parts of flanks ye ll ow ish 10 orange (females), dark with blue spots or blue (males); 16-llJ subdigit
B ACOt'<
IlJ(7)
10 4 supraoculars; 22-24 sca le rOW1>: 1/1-23 sLlhdigi-
tal s .. ..... ... ...... ..... .... ..... .......................... .\Iel/Oll/\· • 6 or more su praoculars: lIlore [han 30 scak ro\\'s ............. ................. ........ .... .......... ..................... 11 11 /\ dark SpOI on cach side oflhe ncck : sca les in 36 rows : I S-23 subdigi tal s ................ 1//(/ (, lti;((I/lI/.\ '0 dark spot on neck : scales in more Ihan 37 rows .............. .................................................. 12 12 Ilcad and body black ahove, with ~ l11all ycllow spots: 41 -42 ~cale rows: 16- 1X subdigitab ........ . ..... ...... . . ............ ... ................ . ..................... hI/liS;
• Il ead and body nor black above .................... 13 13 Lips strong ly barred. black b
wider than light ones. males without blue throent. or if prcsc nl narrower than lighl bars. males with blue throat; 42-49 scale rows; 17-23 subdigitals .......... IIIlIlti.\I[III1I1It1II1.\
SpllellOIltOrplllls {fljiw/i (Bol LI ·t\ (oI · R, I g9~)
L.!'.c".'·"m" "lln'd; Houll N' ,I~ . lu c4.lIil ~· :
·t ·1(,
"Sa\ lI. Nurth
t ~'11\ : '1~~. pt I v· lig. -t '1Ylll' t ~~ '-' :l\HIU " ~!! . \:lV.:1I\
13 ()I"nl' ~'"
tt Y I t·I.)
.~illll·II()IIIII/JI"II.\
,,1/,.,·.1,. T \N 1')<)1 x,
\\.\I ~ ' H ' t 9<)~h Nti
Dia g nosis
/\ vcry small. ~ho I1 -limbed. poorl y known species of .),J/H'I/()lI/or,,/lIIs \\ ith an S I llf abollt JJ mm max .. Ilanks with dark stripes; -t supraoc ulars; 26-30 sca les around mid-body; l(lllrtl1 toe ",ilh 7- 12 ~l1b digilal s: [ail thick al b:ISC, [~Ipering g raduall y. I>esc ri plio n
SVL 33 mm, TL JS mm : car-opening ~J1la ll , round , about diameter ol'eye: nn~tril in I nasal: I'rolltnlla~al widl.:r Ih
2R3
Scin(;idae
Sp h ell 0 III orp" liS (rallo/aem liS 11\( .1'1{ & H O\I\ILR, 1965 Sp/j( 'J/()mIJlJJ/1/I~ ( ~\ 'a/1(dt/( 'IIII1' I NI,\ I( ...X: 11 1.... \llIC 1()() ')
'I) ·Pl'
I "ca lil ~:
,\phI.'1I0IlltllpitU' ~Jh.
I
"SUUt"1 :-',·t;lll Ltb :ttl~ , Iltutllill \ ' rlllo/~il ' IJ1II,~ ,
P H li( ".
I:.:c llltl g ~ / E tlwlo gy:
L IlknOIl 11.
J)i ~ trihllliull
S,J!/( 'lIl1l1/flr,J!1I11 a/fr,'di i~ L' lllkmi c 10 Horn eo
(Sabah) . 1\ 11. I,inabalu :
~ X4
~ g.
1l:sihlll.
s, rVI \I I,I,l I I'... ·.' t l)l) ·~h :
2· l b,
/ . 1',, ,,:OW/lJ1rI \ ,I1';('>,,:,c/ (111I1 ,
no ~ nl a r!:!c: d pr(;anab: a dJl rc:~sc: d l illl h ~ not touchini,!: I(llirth tile II ith 7- 1~ Sillooth sub di ~it al s: tail tilid. at basc , laperlng g raduall y. Colour in akohol (an~r HtJl ll1 N{d I{ I X0X) : " Red dish hrown ahove. 1.\ ilh (I;td,cr spnb 1111 the nape : hlad;tsh S P()t ~ limnin g a st ri pe 011 eac h sid l: orhc:ad al1d bull y. pa ~~ in g throug h Ih l: eyc: whitish hel1eath ."
1 \ , I tJl) ,
11, 7
l) " lri(l,
IX67 .
~f\.l i l1l
In OI / l'g U\lJIf}(1 (' III III/hI)
PJlI.
\ 1I1'/ I 't;ctfu/1i
::: ~I (p;'ll"l)
Diagllflsis Due tll Jl~ coloratioll a \'cry allracti\c (th roat or mak~ hri ghl blue). long- limhed Ire.: skink or the lowland s, 1.\ ith all SV I ll rUp to ()O mill : flallk " with Ja rk "t ripes: 6 suprallculars: 37--+ 2 ~(;alc~ arollnd mid -hody: (o llt'th toc with I()-llJ , uhdi gital,; ; lail lapcring ahrupt ly hehind its basc, Dcsuipl inn SVL 111alc ~ and kl11ak's up to 60 111111 : TL ail nut xo 111111 . 111:des rrolll Sumatra lip to lO S Illill (ZMB 5)\0 12): ca r-opcni ng \crli.:al-o\·a l: no~lril ill I large na sal: ro"tral \\'ider than long. li)l'millg a broad , uturc \\ith rrontona sa L latter wider thall IOllg: pn;rWlltal s 111 broad e()ll ta(;t with eae h other. additionally with frontal and lir"t stlpra(;i liar: rroilta l lullger than its di~tallc (, i'roll1 the tip oC thc "m Hlt ' in eon-
'I'll (l eggs «I '
I I n1l11). hal e hlln g:-. SV I 25-2(1 mm I II JXJl)
111111 .
El'ulo!!y/ EI hHlu!!~
Th l' Ilec-dllci ling sk lll k... hlc III lowland prt Illal} ra 111 f lll·c~t~ . /\ klll.lle 1\ as dl scO\ ered anlllng Ihc bUllr e~!> ur a largc Iree illlllll:dial 'Iy adJ~ l ccl1l 10 a 1'1\ el in Kelamhe (,\ ce h. SUI lI al ra l. II uSl'd a Illlic III Ih e kar-cm crl:d !!l"Oulld Fig. 290 I\ laiL'
or .\/,1i('I/IJllllllplil/\ , \'<111111111'1111/\' Imm
11ll11l .:dla lcl ~ 1l.:,XI 10
Sumalra .
laci with fir ~ 1 J ~ upraocu lar:-.: pari ~ lal :-. In lilll..: contael hehind illlcrparielai: no I1uchals: 6 ~u pran c ular~: 12- 15 s upraci l iar~: 7 (rarel y () o r X) , up rala bial:-. . fi)Llnh 10 :-.ix th he lo\\' eye. firth or sixth largest; (I in\'r alabl~d ~: .17--1- 2 (u,ual ly .IX) SIllCl(llh s ' alL:~ am und mill-body: 7X-9-! I ent ra l:-. het\\'ee n mental and \e nt ; 2 en larged pn:ana k linlhs long, adpres:-.ed hind limb reac hing Ihe a,illa ; Illurth I ll~ wilh 16_1<) \\'cakly keeled sulx li gi t::iI !>; lail lape rin g abru ptly h~hind Ib hasc. Dorsal !>url: lee bnlll /l'-c() l ll ur~ti l llll l ile g re~n: hac k II ith 2 Illngi ludinal rn\\', o r dark !>pots: dal'k dllrsolate ra l ~ Inpe o\'va rying \\'idth hel\\'c~n eye and in ~ertlon or iL'g (occl!>ionaliy exll.: ndll1g (In Ihe lall). inlcrruplcd hy ~ llI a licr a nd hlrgcr. ycll(l\\ is h (II' whiti s h s p(ll ~: \ e nll'llialcral !'o llrrace in relllak~ bri ght orange. ycl lowi:-.h. pink . or din _ ve llo" (rellIalL: ~ and nl aiL'~). each II ilh li g hh.:r in c ln :-.i() n ~: lal ~ ral and \L' ntral s url;lec or head and nec k blui sh grl'y t(l di rty IIhile ( rcllla lc ~ ) or coba lt -bluc. SO IllCIil1le~ c he ~ 1 and later;1I SUr!:ICC (I r hoti) 'olOllrcti likewi se (males ): l'Cnlral :-.urrace din )' ye ll o\\ III ycllo\\'i :-. h g rccn 1\ ith a hlni sh , hinl: (kmaiL'\) or paiL' hlue (males ): I'e ntral !> url;lce lI r l i lllb ~ and lail dark or,lIlgC to ),c ll(\\\ . .llI\ enil e:-. with , illlilar pal lern. bul iL'!>, inlen!>e eo lulIr:-.. dor,al sur f;ln.: ort ail II ilh a rcddi,h shinc. l'e l1lral \ 1Ir!: ICC pcal'lcllinured.
Fig. 291 I\ la1.: ,,1.\i,II, '1IfIIlIlJl'I'/II" lubllng. :-'"r,,\\',,1..
Ihe trunk In hide
( 1'III III/a, '11111\
III
Imlll San -
~
'"
o
2 -"
SCIIlCJ(iac
8:. (;IHlSS 1·\1\\1 I')'n) . DI ' \l/I'R (1 1)')6) oh~er\'\;d this ~ p cc i c!-. II:-ually al he ighh bd\\' 'cn 2 and 'i ilion :-ha ded trees o n Salliuboll g Pe nill SIII
Sp It ell o 111 0I1JIt liS Itillaha/lI(!n ....·;s (13 ,\RIII'I'I. 1895)
1. ~',1!.l)s(mlrl killtlh t illH'II " \'
B \R II "K Ilia llaili. N. Borne,,"
f I I,
!, L)S:r
().,l
Type
I ()('illil~' :
'YJ.:USOlllll ~lflah{1II/{'1J'f\' ~\111 1 1 J9Jl : II , 2..t. 11 ')/,/ieJ/olII(}l'p/w, AIII(I/JUIUt!lI ,;,\_ rv1 \"1111" 19X3- 27 (p~ ln . L I 'BI'NII\RIIi{: ~ lHI,W ' 19X.. j. · 22, fig. :2",. ':..J : :V) ..\il\\Il'i Il):-.tS: t).
I
~~'..,
1')')1 X, VI II
I'>Xl)a . I'll>, I')')I~ . 1.1. 1')<)2a: I \1 : I 11)'1-11>. 2,6 2-10. 1'I'lh," 1'10
\~
~\",
.\,,,hf 'IIf1H1IWI,IIlI\ (\'IIII"hu 11111.). l .l 'R ,HOlIJeI \ 'tll'J.t!.1I1IJW
rVI \1
"Mli"
I 99h ...·: 290
(Ihll I.\ . . ..:,osnwu (1Imltt/a) l 'a";('~(lIlml
P,""s.IXh7)1J1 RII,'" 1'11 <;: 1<J1) (pal'l ,)
1')97).
Di a~ lIo s is
IHstrihutioll Sllln:l tra ,lI1d Homen. Mr. KinalJalu : Pmi ng .
/'\ f~lirl y C011111101l. s lll ~ JiI to l11ediulll s ii'cd. IOl1glimbed sJ..inK ol'the l110re elcvakd Killabalul'cgiort \\ilil a SVL orup to.sX 111111 : !lank> wit h OJ' without
2X6
Sci ncidae
dark u or~o l a t cra l ~ trip c~ : lIsua lly :; S llp r:1oc lll a r~: 32-3R sca k s around mid-body: fo urth toe with 1517 subdi gitals: ta il thick at base, taperin g g radually. Orscriptioll SV I. l11ales 52-5X 111 111 , Il:l11alcs 4.5 -S7 A 111m : ea rope nin g ve rt ical -oval ; nos tri l in I la rge nasa l: ros tral wider thall lo ng. ro nn ing a broa d su ture wi th rrllntll nasal. latt n wider th an lo ng: prerro ntab in broad co nt aet w ith eac h other o r separated by a small shie ld : fro II tal longer th an its di ~ t ance li'o m the ti p o rt he snout, to uching fi rst 3 supraoc ulars (third j ust a litt le): parie tal;; in CO il tact I() r a sho rt d ist:lIl ce be hind in te rpa rieta l: no nucha Is: 5 ( rarely 6) s upraoc u l a r~: 7 s up ra l a hi a l ~, fO llrth 10 six lh bc low eye: 7 illrra lab ia ls: 32-3R s mooth sca k s around mid-bod y: to-XX vc nt ra b betwee n mental and ve nt : 2 e nia n}.ed preanal s: limbs 101l g. ad pressed hind limh reac hi ng fi ngers or elbo\\': fo urth lOe with 1.5-1 7 subdigita ls: tai l thi ck at base , tapering g radua ll y. Dorsa l ~ lIrrace lig ht to medi um bro\\'n or dark brown to blac kish: hae k wll h ~e\' eral longitudin al ro\\'~ o r Illed llllll h J'() \\ 11 to ye llowish spots, occasiona ll y with da rk brown small dots: with or without a da rk d or~o l a t era l ~ trip c fro m the eye In Ihe i n ~ erti o n or the kg (m:cas ionall y continu ing o n the tail ), inte rrupted by smaller and large r ye ll ow ish o r w hiti ~ h SP,)h : ve nt rol ate ral ~ url ; l ce redd ish. werl kl y o range. or greeni sh yel low. int ermi xed with ye l l ow i ~ h to whiti sh spots: ven tral s urf ~1ee o r hcad anu body whi tish ycl lt1\\ . throa t occas io nally ~ peck led with da rk brow n: vent ral s url~lce or tai l Iks hco lo ured to brig ht red or grey. latera l ~ur l ace o f tail occas ional ly w it h the ,ame co lo ur and a grey ~ p o t ted patkrn . 1-2eggs(7 .9· 9.Xnlllltn().-1 / U .XII1 Ill )per cllitch, hatc hli ngs ~ O . I I 27.5 Ill ill to :! 1.5 I 2X.3 111m (SVL I T L). F.co lo g ~-t E I hol ogy
Thi s is a terres trial s kink whi c h a pp car~ to prefc r hi ghe r e leva li oll s li p 10 app ro" . 2100 m a .~. 1. II li ves in the ci u,er surround ings o f the Il eadqu arlc r ~ o n g rn ~~y marg ins o f th ..: roa ds alld ca n als() be fo und in park- li ke stru ctu res a nd ga rde ns. Th e po pulation de ns ity in Ihe Il ea
gioll has gradual ly dimin is hed ove r th e pa st 15 yea rs, a pruba ble co ntrib ut in g f~lcll)I' being th e stce p-ba nk ed wa ter d ra in ~ wh ic h para llel th ..: r oa d ~ . III these cana ls we did not onl y find numero us pc rished s n a k e~ hUI a b n many dead S. ki l/ (/ ha lll C'l/ sis. O Uh ide Ih e Ilcadquart..:r:-. reg ion. th e ski nk exh ib its a prefere nce for lear litter (so me t imcs in cons tan ll y ~hade d area~). bu t ,II ,0 occll r ~ in Ih e vicini ty of ~trea m s . Il owne r, it avo ids roeks of banks and d ()e~ not llee thro ugh the writer. 01 \. 11 1... 11>\. Ko'J & 11 11).· \ ... 1 ( 19X9) ro und an ant nest in a 1~lI lcn log in whic h altogelh er 16 eggs \ 12 cad i) were hai r buri ed . f)i slrihutioll IS e ndemi c to Bnrnco (Sabah) . Mt. Kinaha lu : Mare i Parc i. I. lInlli 1.1111111 . Sg. S il all . il all , Sg. Li \\ ago. Kamborangoh. Sg. vlcsi lau, above Pori ng PIOO 111 ).
Si Jh('I/ /Jlllo rp lllls /iil/aha/II('lIsis
Sp" e ll 0111 01'1''' us III ullisq Uti III tllus IMflJ{ . 1958 ,\ /,h,'l/tWU Jll J/III\'
.n . -1-1
1Il1l/1I ', /ltllfllfl llt,\
(Pill t ) I "
d 1(,
IlJ "X
:! (l I .
lig.
T)(lf I.,,'a li l): " K,a" 13000 kcl) . \1"'"11 ,,"II,a Ila lli . ~ II u ,H I "
,\/ , h CIIOHUJlpllll\ IIlllitJ"'IIIOIIWIlI\ , I 'B I ' " \KI> '-\
I (JX4 .
2-1. \I \I~\I" 1(I X5. 11.1l)l) 2 a . I ~ I , l IN I(,(n (I. I "~ .. ,,, t '>9-11>: ~ -1 h: \-t " "" \ & (. K.." . , , , \ I()(n 27(1. fi ~ 2111 I \ '..!O,\tJlWI I' II ' K,.
IlNIl"
\ 'o ""',t!.tI/ llm ,
IXI,7) 1.13.
(n~)1
11 !.!o.\OIlUl II !t II " " " )
101"/ 4',t:,u/r llll
IXX7.- 2-16 11',111 I. \I •• • " '"'' R' '',II 1')1 :- I'll> 11'.111 I. ~ .. "" 19.11 10. 2-1
II",
'>I
II N•• '
K
(pan, .\/J/WWIIIWlp/UI\
d~ l'cln( ',1:.UIU\ , 1\1 \N 1111' t'\... 1)1 ' " H I ()X2. 17;
.\ /J /' t' I If'lIIlJr/,JIII,' \ '( lrtt'glllllfli
IflrH'g4I1JI\ ,
(110 1
I
I ,'.!,fJ\U nt U II/Ullt/H/)
PIIl W..... IX(17) \ 1"1111' Iq~ 1
'~/}Itt.·nOIll()"/'/IIt\ \O/'( U III'
SJ< H.KI' llJX4 ' 23. li g, SI,IIl,,, ,I/I,,"phul ' I' . \1 \N' "'
( 1101 hId ~
\
&.
]7
19 "X) 1 10
t), Nil" 1'/X2
li ~.
\!I\I{IJ
&
1'. 1(,
Diag nosis ;\ l1Ioderale ly larg..: \ . VI up III (,I.) 111 11l ) ~ p..:c i es Or .~i'''('/I() I I/(}rflllll.\ , highl y ada pted In runn ing wale r ~ : l11 alc~ with a hl ue. le males \\. Ilh a ligh t brown is h Ihroat : no da rk dorso lat<.:ra l ~ t rl p c: (,- 7 S ll praocu l ar~ : -l 2--l() scales aro und m id-b,)dy: I(H lrt h lue \\ ith I (,-23 ~ub d igi t a l s: tai l ta p": rl ng rap idl y bc hi nd the basco
2X 7
Scincidac
Scincidae
Orsrription SVL mal es 64 III III , felll"les (19 mill : TL greater than SVL; car-opening vertical-oval : nostri l in one large nasa l: rostral very narrow, formin g a broad suture wlIh hexagonal frontonasa l; prefrontals pentagonal. in broaLi contact: frnnlallonger than its distance frolll the tip of" the snout. touching lirst 3 ~upraoculars: frontoparietals and interparietal of elfuallcngth : parietals in contact with e"eh other behind interparietal or separated by a slllall shield: no nuehals: 6-7 supraoelilars, first largest , last ,rnallest; 12- 1(1 supraci liars: 6-7 (rarely I{) supral abi als, fOlll1h to sixth helow eye. fi nh and sixth large~ t : :; IIlfralabials: 42·N smooth scale. arollnd I11ld-bod : 10-10 I sca les bel ween mental and ve nt : 2 enlargcd prea nal s: adpressed hind limb reaching the ~holiider: four1h loe with 16-23 rllllndi ~ h , smooth subdi gitals: lail t"pering rapid ly behind the basco Dorsal surface dark grcyi~h brown, wilh 2--1 rows o f d"rk. spot s: lalerall y. a pattern of li ghter and darker spots of varying sil.es: lateral ~ urf~l ee of nape and Ihroat of males wilh a blue tinge: ve nlral surInce \\ hili sh withoul spots. Rl' lIlarks: It is \\orth noting that fi g. -13 (right) of Ihe origin,,1 de ~eriplion by IM,I'KI~ ineonsistclll \\ ith Ihe descn pti on relCrring to Ihc hroad contact o f the prcfi·onl als. II is nol dear though whether this li.llilt has becnillade in the sketch or in the le.\ t. or whelber Ih~ ,ulun: between the prcl"ronla l, varies in lenglh . r.co ln:.:~' lE t hology T he ~e
ar.: diurnal. \ cry shy st r.:alll -d\\,cllers Ihatlive pairs on and hetween rocb along th.: pa l1ly torre nlial Sg. Kipung !t I. Their !l ight di slancc i~ a ~ I ag g.:ring I 0-1:; 1l1etre~. and \\hen ne.:ing they hide in crevices llrjump into th.: water. They Iced on minute in~ccts on the 1110ss-overgnm n boulders. 111
Distr ihulion 1/,,"'i,"llffllllJ{//II.\' is cndcmic to I3nrneo. II. Kin a balu: Pori ng: Sg. Kipungit I. Sg. l.al1 ga nan : Ki au, l3undu Tlihan .
SI,hl'/JUlllllr/lilll.\'
Ahu\'e: Fi g. 29.1 t-.tak
or ,\il /I<'ffIJII)"I"{,hm
from Sg. KiplIn gli I. Bctll \\ : Fig. ZI).t F..:mak nt /lllIf'" li'om Sg. KiplIlI gli I
/ll1I1//\ I{IIlI -
111lI11I'
S/ ,Ir, ' //IJ/lIfJl/lh,/\
/111"'1 " 111<1-
Spilellomorplllls .W /btl/III S I Mi\' I{, 19S9 S / I II"// O/l/lJlplll/\ \/lhOI/II .\
Saragaya
h ," K . I 'I ,X r~ IW l "c" l il~ : S:lIlclabll I )"Inrl, "nh 11m·
I' or~ s l R~ , c,,~ .
n~\J
.\ '" jWII (IJII O lphll.\
1\1 \ 1 ... ' "
"
19 ~7
\uho",l\ ,
l ' II I N H \KU
2Xh. T" 1')l)J · 'I ;
&
S ICH,KII\I
1\1\110.'11 ,
1 \) X~'
24 .
I 'I'I-Ih 246
Dia:.: nosis s mall ~pee i es of ,\/lill'lI IIII/O 1',,/11 1.1' reaching a ' VL of lip to 51{ 111111 : body aimosl \\ ilhollt any pattcrn . no dark dorso lateral stripe. lab ial ~ hiellb conspicuously banded bl ack and white : (, (7) SlIpra ocu l ar~: 3!.{--I 2 scalcs around mid-body: 79-9 1 rows of ve ntra Is hetweCn1l1Cniai and \enl : fo urth toe \vith I X-22 ~ubdigilals: tail tapering rapidl y behind ii , ba sc o Description . VL males up to 5 1 1l1m. fe male s lip 10 5X 111m : TL • SVL: ca r-ope nin g \'enical-o\'al : nO~(r11 in I n:1\al : ro Iral much wider th an IlIn l.!. fo rmin g a broad suture \\ ith frnntona~al , Ihe lalt.:r wid.:r than frontal: prcfronta b in broad contac t or sepa ralcd : frontal longer than it s distanec fr0111l h.: lip llf the snout , touching lir ~ 1 J ~u p raoc Lll ars: p a netal ~ In conlacl wilh each other behind interparietal : 6-7 (usually 6) supraoc ulars, lirst larg.:~I : 14- 17 supra ei h ar~ : 7 ~ lIpral abials. fou rth 10 sixth hdo w ,:y':. \Ixth l arge~t: 5-7 (usually 6) infralablals: ~X - -I _ ,11100th sca les around mid-body: 71) - 1) I rll \\ ~ of ve ntral s between mental and \e nl ; 2 en larged prean a l ~ ; adpress.:d li111b ~ o\e rl ap : fl) lIrth loc \\'Ith I x-n ~L1bdigitab: tail lapering ra pidl y behind it~ ba~c .
Dorsa I surface almost lin ilim1l. dark brown to dark grey, \\·ith indi slin el lighler ~po i s : in mak , latem l S llrl ~ I Ce of neck and bod y ortcn wilh an orange linge: labial s barrcd black and \\hile. widlh o l"the individual hal'S vana bk : \':l1lral ~urfae.: cre:lmcoloured . 2-3 eggs. 10-12111111 in knglh (1 '1 (.1 I( II):;X). El'ologymtlllliogy One ~pecl111cn was co ll ce led far 1'1'0111 a \\ater body rrom LInder a I~illen log in primary fllre~t above Porinl.! Il ot Spring~ .
2Xl)
Scin~' idae
"'ii!, 295 ,\jdl, 'lItJlIl,,/,!I!J/f\
\'(i/)(/II/I,\
Ihun Pllrlng,
l)i~lrihutif)1I Sllil l' 1I1111101'1"IIIS .WhUIlIIS i~
cndclll ic tll Rllrncll (<-;ahah) , 1\11. h:ill.lhalu: Porillg .
Spli ellolllorplius IClIlIiculus (Mo( '()I Il .1..!.IJHJlIUI
2.,
IC'IlI/1{
1f/11111 ~ I o(
T~ lw ,,,,' ,, ' i l ~ :
I \Im,
II( \NI),
RIll' "
\IJIIi"flIIIOrpllll ,~ Idllll( (lIulII . \I ~\I"
I X'JOa . 133. pi \ III Jig :!.
"",:, 1\"ltI
!\ gOHIIIW/t'III1/(ItIIlIU,1>1
\I
I Xl)O)
I>l'sr ri pI io n SVL40 111111. TL 60.5111111 : car-npenll1); vcrlic;iI-ova l: nostri l ill I na~al: rronlnnasal wider Ihan long. forl11Ing a broad ~ulllr(' with rostral and a narrow..:r olle with frollt al. the bller IOllgcr thall its dislanee li'om thl: tip oi'thl: snout and touching fir.'> t 2 "urJ'aoelilars: parictals in t:ont<1ct with each other bclllnd inlerparictal : 4 ~lIpraoeular,,: Xsuprat:i liars: 7 ~lIpralabials,
r-..l
1915 IXO: S r-.1I 111 I l)J) \1 )tn,l '].7, I ." (l}lJl .
\NIIII \
I).
I'I'I-Ih 211.
\l'hIHOlJJtJl/,!rH,\ 1('/1((1('1111",
I
IH Ni l \1<11
&
"II II "H ~
IlJX4 ' 2-1
nial.:lw s i ~
1\ ~111;111. poorl y known species oL\i"wl/fIIl/o/'!IfIl/I'
llr 46111111 SV I : l'olour ill life unknown: 4 supraoculars: 2(, ~cales around l11id-body; 6X roll~ or VL'lllral~ hdl\'e~' n III 'Iltal and vent: fourlh loe with 21 -2 3 ~ uhdl g ilal~ : tai Ilhick at base. tapcl'lng gradually.
2l)()
Selm:idac
rourth to "i,lh below c:ye. sevC: lllh largc:s t: (, ill li'alabials; 2() sllloolh seales around mid-body: 57 vel1chrab between parielal" nnei Inil hase: oR row" or vcntra ls hdwccil sublllaxi llars and venl: 2 enlarged preanal!-,; ndpres"ed hinel limh reaching Ihe \Hist: liHlrlh loc wi th 21-23 subdigilals: tail thick at basc:o lapering graduall y. Colour in alcoho l (aner 1>1 ' Ro()1.1 191 5): Rrown ahovc. anleriorly ~ polled dark brown; a dark dor,olalera l band beginning at th c no"tril. s polleel
Genus Tropidophortls I) , 11 ' RII
&
BlIlRO"J,
I H39
rile genu, hopicl()III/OrllS IS native to the Asian mainland . thc Philippines. Romeo. and Sulawesi. Mndcrah::l y I;lrge "kinks wilh robust. s lender bodies illld rull y dc:veloped limhs. Dm"a l surlace covered with eharacleristic. strongly keeled. hUlllped sca les (e., eeption I . h('CCl/rii) . Lower eye lid with "mall sca les: car-opening Inrgc and clcarly \ is ihle: nostril alll'ays in OIlC lIasa l shidd; pn.: fronlab. rronlOpari -
whili 'h: nanks brown, white-spotted . Ventral surface ye llowish . Re m a rk : I\eeording to own l1lea , urel1lenlS Ihe tail Icngth given by III ({(lOI.l ( 191 5) i~ IIICOITec!. I<:co lo gy/E th()h) ~y:
nis t ribuliun .'>illtelllJ/I/()/,IIItIl., ((,lllli(,I1II1.' is endemic In M!. K ina-
haili . 11. Kinah a lu : Bund\l l uhan .
e t
and interparietal prl! ~e nl. "uprana,ab absent. In contrast to all otherskinks inhabiting Bornen Illese ~pecics posses" a single large preallal plak. 1\ II spl!eies included in thl~ genus are more or le,,~ ~ tron gly adapted to running. water" in :Ind Oil \\'hich they live. 'orlle species arc knOl\n to he IIvc- bcarl'r~ . i'i\ l! specic~ have been recorded rrom Borneo. hili nnly T. IweclIrii OCCllr" al lower a liliudes on J'vIt. " illabalu.
Tropidophortls be('carii ( PI '11 RS, I R7 1) 11I1(,llI ' ''' '," /I" u llri,
Key to the Bornean species of Tropidophorlls Dorsa ls slllool h ......... .. .......... ................ hel'l'u,.;; • Dursa Is kce kd ................. .................................. 2 2 [)or~a l s I",il h :2 keels ... .. ........................... iI/iiI 1111,1 • Dorsa ls wil h I keel ........................................... 3
seale rows around mid-bod y ......... I}('/'plcxlls • More Ihan 30 sea le rows ................................... 4
3 :10
4 32 scale rows,S su praoculars ....... .. ....... h/'llokei • 34 sca le rows, 4 su praoculars ........... .. IIII ('/'IIIJII,I'
Unkno\\ n.
p" , "'.
I ~7 1
I y pl' Inca lil) :
'\7 4
"oro\\a~
livpitiolJ/UWII,\ 1J"f T flrfl \l1H l) 1 \ IOJ I ~Nn. 1 11 ) (mlll/llIl,lwl'll' h{ '{ {(1/'ll ~""" I'n I 10.2]. 11 , I" "" , I'iXJ ~ 7 .1' " 1<)'11 : '/ : Mil "'Il , I <)l)·lh 211>.246 . 1'I'Ihl: 2'}l) Vm ·/lc 'lI h( 't '( 'lInl
I
Iii ' " \ IU.
&
~I U( ,I'( I '
B", , , " ,I
(i 'rl/l idlll,ltllm \ IIlfll (lilliI'd,
19X...J ~
1--1
I X'I hi 7, ~
T) (ll'
. I\orn",," hI 11 alk, " " " II I'I:?.> 777 ) IIO/JidtJphu l'lt,\ IIItJetlJlllJ'clJ III RIHHI 1') I"' . ?7h. t\.1 \N IHI ' 1\) ' , 27; 1\1 I I "'Il , 1<)'1-11>: 21h . 2-1h \mItt 'a mOl -1',anil I'BI ' !I \R III\:" ~H )(. IO r'\ I\)X l ~ ...J \(,h"II{IJIIIII 1,111/\ ' r. \1 ·"" \11 , 1'1. 7 2Xh. I'IX'I 1'1 7 It'I',, lil) : "Kill:! 1I01"n.
\1'/t" III1I1111'/ ,hll' , p o 11 . " I " ""' " l'IXXIi '1 . li g ,I f\1 \1 I.. \ It .... 1\)9\ h :! 1 T~' Pl' I nca lil ~' : ·'I\.If11111 lit I. hllllel dl! 1I Ihll ~Pllll b '" PtHII1g. .• 1111 "iid"'lh"l1~ dc, \1.1"111 "111,,\),,111 ('00 III \IN)" I"'" :tlk,
\p/t( ' IIIJ1J/OI ,,11I1\ 11l/"111l1l/ \
" "'" " & Oil 1'1'1-1) ,\pht'IWIIIIJ11J/w,\ f/ f/UtIlI C'U' \1 ,\ 1'" \II -t 11)1) 2 . 1 I, I
Klilabaill
Sahah fropidop"orw. hecl'l1rii
PI t 11( ....
7i"'I'U/,J/'/uU"u , h't ,,;A cl (;,n l. I J'u/llt!ojJlmn/\ !/'fll',dflJ,htJItl.\
1X71
IX-1 ~
I 11)11 11) ' Jil Il' . I'Ill S JIll( f'1I/'tt ,\ I , I) " ' '" l' , ')' . I'll"
•
~
11111/1111'
I n'I' llit'/ ,/'orrt\ 11('1"1 ' /('\ 1/\ \I\I(lli lI 'I. 1921
Brunei
•- - - •
• •
Sanl\\'ah
• •
I
•
-
1
Kalllll~ULlan
~
~
•
• • 291
SeillLidae
rr
n
I>iag llus is /\ large ~pccic~ ut" Tmllit/()fllrllrtl\ n::lehing 95I)X mm SVL . The unl~' sp - -ies or th l ~ genus on Borneo wllh ~Illnolh dnr~a l s. Orser-iJlliulI ,'V L up III l)X 111111 . TL · 107 111111 : car-opcning
vcrtlcal -pval; head ,hields and body scales SlIlooth; fl"llllionasal wider Ihan 1(lllg: pret"rollials In broad e(lillael or ,eparaled ; frontal substantiallv IUllger thall It~ ,li s tanLc rrom th,~ tip ot" the ~1l01lt, toucillng lir~t 2 ~uprauclliars: rrolltopanctab oeea"inllally :-e parated hy a small sca k: parietal~ rorlllll1g a broad ~utl1n: hehind interpaJ'lt:1:11: ..t -5
Fi:.:. 297 Aliaie' pial!: nf fmIJiJ"IJhurt/\
11('('('/11"11 .
~upram:ulars. lin,a largest. last ~lI1a ll l'st; 6-7 supraeilia rs: 7-X slIpra lahia ls, litHl and sixth in contact wi th eye: 45 int"ralabials: 2X-36 "lIlll11th seaks nrtlund I11ld-body: wnlrals sca l e~ Inrgcst. 5..t-56 row:>; betwccn ~lIhll1;!\.lI l ar~ ami anal pink: om: large preanal pia Ie ; adpressed hind Ilillb reaching at I ca~t up to the wrist: fourth tne \\'ilh 1<)-2 1 ~l1lnllth sllhliigita Is: lai I sl ighlly compressed laterall y. Dnr~al ~urfaee (brk hrown. with ~llghtly lighter bands: lalera l surrace or head and body with light dllts pr "pot~ s(lllletimes arranged in bamb: ·hin and throat whiti~h. th..: latter with dark ~trip..:~ and spots: remainder or ventral 'lIr1 ~ICC ycllowi~h . I::co ICi gy/E thfllu ~~'
The diurnal r IW("('(I/'ii i, thl' l110sl common scincid spt:c les along mountain strcams of Ml. KlIlaha lu up to ..:i..:vatlons of approx. 1000 Ill . It is sellli-,ltillatit: ami II\'(';~ 011 the banks or stn;am~. but is often secn pt:rched on large bou l der~ and 10)2." scaltered ill the stream~ . II~ /light dislancc 01'2-3 metrcs i~ rdatively short, but ir disturbed. thc skink disappears illto a creviec in a /l:bh or .iulllp~ into cascades or pools limllcd by the strealll. Wilh undu lating movcmcnb and limbs allachcd to the body it ~wims swinly 10 another plnec inti\(: ~Irealll where it inilial ly pushes only Ihe nostri Is abow the water surl:lce. Rocky eli /ls overgrown \\.ith moss otlCr suitable resting places li)r the I1lght. Juvclliles ha\e also bccl1located sleeping on kayes overhanging \\ater (D \~ 19(5). It appears that jllvcllill's arc e'H:lusIVC 10 ~mall rivulels where they can be found between rocks and leat" lilter, while the adults prefcr la rger streal11~ . I his may be all illdication that lhe laller also prey upon sma ll li"ard~ including their m.vn oftspl'ing. A /Clllale cxamincd hy ROlli l'i'( ;J 'I( (I Xl)4d) conlail1t:d..t ru lly dev -loped embryos of .' () 111m SVL. Oislrihlllion livl)ic/ofl/wrll.1 hccclirii i~ endemic to Romco . Mt. Kinahalll : POI'i ll!;: Sg.. Kipullgil II. Sg. Langanan, "iau : Sg. I\..adamaian : Sg. Luidan . Sayap : . g. Kcmalllis, Sg. Wal'lu .
292
Varan idm:
Varanidae
Till: distributi on of monitor li /.ard ~ I:xtends fro m Africa thro ugh southern Asia and thc Indo-A ustra li an Archipel ago to Austra li a, Melanes ia, and Micrones ia. These arc mode rat e ly large to ve ry large l i 7ard~ with ro hust lim bs, strong. elawell toes. and a long powerful ta il. The elongate sk ull s of monit or li 7ards are heavi ly ossifi ed withstandi ng the pressure e:I.Crled when swall owi ng large luod items. i\ strong dentition enahl es thcm 10 tea r apart large p ieee~ llf prey. Di stributed ovcr a vast rangl: l11 onito r l iza rd~ have adapted to a ia rl:!e vari dy o f hab itats. but are generall y rathl:r a lilli nal clement o f the Im . . lands. Rl: prescnlati ves of th is 13mi ly llIay be fou nd in
Genus Vamlllls The ra nge of Ihe ge nus
MLR RLM ,
~ i m/llll.\
I R2 0
deserts as we ll a!> in rain foresb . and ~ pl:CICS sllch as I l lI W 11I 1 h(,II:!,II /(,lIsi.l' and I: .W / I'II/IJ/' an: ocea'iona lly co mmensa ls o f 111a n in larg.:r t n wn ~ . 1).: pending on the indi vi dual sii'c monitor li/ards prcy upon in ~cets. amph ib ians, li/.ards. snakes. IUI1Ies, li sh. eggs ora lI so rt ~ . and lIlammals orva ryi ng sii'L'. They are also knO\\ n 10 ICed on carri oll . St:xes arc diffic ult to dis till guis h. but ma les usually grow larger. Moni to r lilarlb are ov iparous; cl ut ch size varies with th L' spec ie!> anti IIl dividuall y. T hree species arc known to occur 0 11 Born eo. Iwo o r whic h have also hee n observed on Ml. Kinabalu.
Key to the Borueau species of Varalllls
is idenlica l to that
or Ihl: family Va raniLiae. rVloderate ly large tn .... l:ry large. muscu lar lizards. Head el ongate in s hape, pup il ro und . Tongue .... ery lonl:! and slim with a dee ply lu rked tip ; ret ractab le int o a ski ll ~ h ea th . .Iu venile~ vi videl, co loured and reveal ing contras ting pattern : mark in gs ill adult s k ss obvious. sometimes [;Iding beyond recognition. The lIiurna l monitor lizards of Borneo are eXCl: Ikn t cli mbers ofte n residing high in Ihe trees. Two out or the three ~pecics inhabiting Borneo have also heen reco rded fro m lower e l eva tl on~ on Mt. Kinaba lu .
Nostril distinc tl y closer to the eye than to the tip of the snout ..... ..... .... ...... ...... ... ................. ...... ... . 2 • Nostr il distinctl y clo. er to tht: tip orthe ~ n o ll t than to the eye ..... .. .... ... .. .. ... .. ....... ... sull'ufo ,. su/valor 2
cc k very long. about as long as the di stance from the tip of the snout to the posteri or margin of the car-opening ..... .. ............. .. .............. ...... r udicollis eek not elongated; di stinctl y shorter than the distance from the tip of the snout to th t: posterior margin of the ea r-opening ................... dlllllerilii
Tah.36 I ) ' ~In b uliun of / {/nlllll.\ -S PCC ' c~ (\11 HorlH:u
I 'u rall llS ,llIlII e,.i fii / '0 /'<11111'\
(S tili J(d I .
I R:19)
m dico /l i.1 ((i nl, I X.J 5)
, 'urallll.1 .\ afl'U/III· .va / WiTIJr ( L.\lH ' t\ II. 176X)
Sahah
Kinaballl
• •
•
•
Smile,
•
Sar~w a"
Ka l llnillll:lll
• • • • • I •
Vatanillac
Htr(IIII1S tll/lll erilii (SCIIII '(,J ' I, IX3\))
l) ia~ Jlu ~i~
I'vlodcratel y large mllntlor It Lanl. IOtalli.:ttgllt ahoul 1."\0 CIll . I lead tdatl\el y "hon, "nuut hlunt : no,Iril \lval tll slit -shaped, dusnbli.:, ~illlated Ileal' Ihe eye : neck "hnrkr Ihan head . nape with large. 1'll1lndi,1t ,cale :-: sli !! htl y larger than dlH~al s: lal\:ral" relali",:ly "tt1all : tim"llIlt witlt .1 -. nat't'o\\ . Itg hter Ct'ns" lines . Dl'sc ri pliul1 Sielltier. IItlllltlnr II/anI,.,; tillallettgth:-: bct\\een 120 and 135 cm , 150 CIll in e.\ n:ptional cases: taillunger than SVL: head ~ hnrt and hroad. a blunt , flat snout: neck slh)rter than head (i .e. tite di~tan e e from the tip "rthe snuuttothe posterior margin o r the IYIll fKlnllnl) : nostl'lls cll1';abl e. oval to s lit - ~haped. dl,,tance rrollltite ttp orthl' ,,"outlwi ee totltrce titlle" a" long ;" that fwm Ihe eye: tympanulll large .
Fil!. 2<Jll I'tiull Sp<:l: IIlI<:1l or / (//'(111 liS
.._...."...........
1111111(' 1'1111 .
-
.,.-
rOllndi sh. ~ Ii !! htl y "link; legs and toe" moderat ely lon g. the l"lter \\'Itlt 'trOll gly curved ' Iall'''': 17-.+ 5 :-:ealcs hetll een the angli.:s or the l1lolllh ae nlSS the occiput : illi Cit" I ,eaks large. larger tit;tll titose lItt the occiput and largl'l' than dorsals : lateral s suhstall/i a ll y "'ltallcr tltall dor" tI ... : (,6 - 102 "cales anlund mid-body: ventrals elongate. quadrangular. weakl y kec!ed or smooth : 7·1-') I lentral" betll cell lite gular !i.,ld alld tlte IIlsertiUl1 or the Iltlldl cgs : 2. u"uall y vcry distinct and largc prl'anal pores: tatl c0mpre",ed Iaterall) IIllh \ ery s mnilialeral :-:eales: "ubeaud ;tI" suhstallt ia II y lar~cr : t1\ (l keeled I'ert 'bml "caic I'llWS rOl'llllng a double cre ~ 1. Sides ~,r head alld lIeck IIl1it(lrtl1l y bmWII : 2 darker dnr~()lateral ~ lripl''' un thl' head betwecn the pllstenor Illarglllol'titc e)e alld the Ilape . IlslInll y I'u"tng 10 rorm a noss band : upper side orthl' L1lld blad i"h to dark browll . with :1-5 narnm . lighter crns~ I illes, "Olllel illle~ brokell itlto S Cl'll~S or spnh: upper "Ide or limbs spotted li ght or with It ght rings: tail hrown. indisllnctl y handed lig ht and dark : \Cnler unirorm 1;lded yc1I('IVI"h hroll'lI . oee:ls il'"all y II'IlIt darker spot" or bands.
Varanidae
VartlllllS sa/wltor .milJ{llor (Lt\LRL:r--l'I, 176X) SI.-IIr" l'u/l 'lIlor
L\I RI Nil .
17hK . 5h
r~ IH'
IIll'a lil}: " AIIll'rIl:""
(I ll l'rror)
1£11'/11111' ,,,"I t t/1m
SIIll,1<1 N
"dnl1or \1 " 1111 \ le/X"\ 27, hU -' II \KO &
l'lX·1. 2,1
l i/nll/II'- \lIhlllo/: I ltI ' " '\ I( U &
19'12,. : 1.12. I<)44b:
~II )(. Kj ,
Il)X-I 24.
(\ 1 \ 1 .... \ 11 ....
~41>
Dia g nos is Very large monitor Ii/an] ma~ ~ i\'c budd ; lotal length lip to 300 (Ill . o~trils roundish-o\'aL u~u al ly situated twice as far I)J' 1~lrth-:r rrom thc eye Ihan rromthc tip orth-: ,,'lllUt ; nllehal scale» 1I0t or on l little larger than the dorsal!>; tad IllU;, ular, Interally eOl1lprc;,~ed , oar- like : dor~ul11 \\ ith roulld, ye lltm . orten m:elhte spots ill 5-(, crll~S rows .
or
Iii!!. 299 Jllvcnile 'pcclmcn of
lil/"flllIl.1
dlll/lI'rilii from
Pllrlng .
.l1I\,1:111h:~
with II1ml: conlra;,ling paltl:rn ; hl:ad and llapl: onl:nllr;lngl:-red 10 rl:d: cro~s linl:s oflh-: body eorre;,pondingly coloured or whitish yc llmv: upper sid-: ()rlh-: blld).la icral stripl:~ on the hl:ad and n-:ck ollen black; lowcr side of the neck with black ~t ripe;, or longitudinally t.:Iongaicd spots: lightl:r Illarkin g~ dominating on the Ilanb: tail with alternating light and dark hands: dark bands ;,Iight ly wider than Ih-:ir li g ht counterparts. ~ .L) eggs (JO ' ~5 tn 25 • (IS 111m, growing to J.\ 50 to 2~ '- ()~ Inlll dUring incubation; hatchlings L)(I100 IOO- IJO 111111 (SVU'I L) [11,\1 'I 11111) IL)L)X] . F.cn ln ~ ~·/E I h n ln ~.v
I ( /lW/lI S dllll/ailii is con~ idered st rict ly arboreal Ii\· ing in coastal lill'l::;'ls. but also in the rain li)re»ts 01' Ihe lowland" and in hill y country more inland . Desp ite ib predilection li)r trees it is a good "Willlnll:r. hUlliin g li sh and aqu ,ltic ~kinb in shallow \\all:r bodies. Crabs arl: a prekrred food ilcm; not on ly the mcat i~ takcn but thl: whole body ineludIllg shell ancr n:l1lmal or thl: h:g~. hen juve niles \\ ill take on larger scnrpions. beetle larvae, spid-:rs. and ;,maller hirds (7.11-1,111< &. H()II ~ II ' I ()<)()) .
lJi slrihuti()11 rvlY;lIlmar. I h;liland , Penilbular ·l alay»ia. Sumatra and Batu Is land;,. Pulau Bangk a, Pu lau Belitung, and Borm:o. 11. Kin a balu : 1'L)rillg.
DcsCI"iptinn Vl:ry large, IXlwl:rful 1110n1lor lii'ards ea~lly -:xl:eeding 200 em. large ~peeimen;, up to .I0() em in total length: aged specilll-:Il~ dorso\elltrally 'olllpr-:ssed: tai l longer than SVL head long, snoul slightly poillted : length orthe ned correlated to SV I : nl:ck shot1er thun head (distanec betwcen the tip or the snout and Ihc po~terinr edge or thc tympanllll1) in juveniles. longcr than head In old specimens: nostri ls large, roundish to mal, c1\l~C to the tip or the snollt. twice the distance or 1I10r-: t(l the -:ye Ihan to Ihl: tip urlhe snoul; t)'mp;lI1ulll larg-:, \enical-oval: limb~ Illoderatel y IOllg: ela\\ ~ Ill' linger» and toe;, s light ly ellrved ; ~X -() () ;,calc!-o bctwccn Ih' ;rngle~ or the ml,uth across the oecipul: ~cales on Ihl: uppe r side or the head rclativcly largc , l1at, and smooth, larger than nuchal ~eale~, the latter usunlly "Iightly largcr than the \l\al-~haped, kccled dorsa ls: laterals lIsliall y ~Iightly smaller than dorsals: 13 7-1 X I s(ale~ arllund mid-body: \'elltrais kee led, elongate. rectangular with round postcl'lor limits, largl:r thall dor:--a ls, ill XO-')5 !'Il\\ s hclwl:clI the gular fo ld and th-: IIl!>erlIOIl of thc hind kg» ; u~llally I di:-.tlnct preanal pore un either :-.ide, ocea~ionally 2. very rarel , ~ : t;lil lalcr;dl y compressed. \cry powerfu l, oar-like. \I ith a double \'crtebral kl:el: subcaud;lb large r than laleral ~ca le s . Enti re uppe r side almo"t black in )'Ilu ng »pccimcns. 1;J(iing III grey with inerl:a~lI . g ag-:; jllvcnil es With
Varanlda~
crn~s ro\l's or brigh' ye llol\.' spot» :lnd/or ocelli Oil the dor'ial 'ildes or the body and lall; nape. Itmbs and tue~ with nllll1l; rou~ 'ililal i. yel low spob; head, body alld wi I banded 1\ ilh y~I I()\\I and hlack laterally. Yelluw l11ar!"lnt!-s Elding lI'ilh incr~asing agc becoillilig Illor~ and l11or~ Indi'ilinel; old specil11clls ortcn plain dar!" grey dur~all y and lalerall y. Venlrally Im glll ycllow (jlll.cnlles) III bded yellowi~h (old 'ipcc imcns). 10-3 0 egg'i (34-4h . 64 - 1t)() 11l1l1) ; hah.: hltngs SVL 120-1.\2111 111 I '1'1. I 65- 1')(J 111m .
Eeu lug) I E th o lu g ~' " ll/WIIIS S. Itd\'{//ol' IS ~ne(lunlered prcdo illinalitly lIear or in waleI' wilh no evidelll pn;diicl': lion 1'01' specific types (11' waleI' bodie~ , and ha~ even becll 1(lutHl ~willll11ing in the open ~ea . II IS Ilot ullcommon to ~nCOUlllcr s i/.eahk populations inhabiting smaller i~lands wilhout I.·cshwaler foraging in the Iidal lones (M ·INIlIl y unpuhlished ()bservalioll~ made in 'andakan Bay). Their powerful. oar-like tail enables thelll to swim and dive cven in rather strong eum;l1ts. Young and semiadult s peCil11ell~ are orten seen climbing. willie larger adulls prerer to sta on Ihe gro und . ~l1nitor lizards do not neeessalily take 10 Ilightl\.'hl:n l:l111f.·ollted . Larger spl:eiIllens In partiullar may del:ide to rather face the opponl:nt and arc able tll inllict severe injurie~ with
Fil!.. 300
Adult specimen or
!(II'£IIIII.I'
sall'll/o/'
1'II/t '1II1/r
Ihl:ir taib us ing it like a whip . Il'gilell the l:hanl:e, they will normally Ilec Iloisll y into the undergrowth or if ellcollllkrcd in the imlllediate vicinity or a water body intn the wilkr. Males arc !"nowil fiJr their combat rituals which include head jer!", di~pla y. l:irclillg. clinch , Virestling pha~es alld evcn lll11unting the defeated oppoIlellt. I-. \'l:l), anil11al which can be overpowl:fed is cllnsiderl:d prey. The diet ()I' jllvl:lliks inl:llIde~ , among others. als~) insel:t'i and frogs whi le adli it s ma y Ilot evell shy away from dwarf' deer. Females bury their eggs or dcpo~il Ihe:111 in cavities in trees : clutch s i/.e depe:nds on age: and body sile or the respedive rcmalc. l lll'lll/IIS s . sail'll/III' appear~ to be fairly rare on Ml. Killahaili . One ~pel:imen was obscn'ed some 550111 rrom thl: nearest water hod " hetwccn Klpungil 1\
!\ndaillans and icohars. to sO lllhern China (K wangtung) alld Il ainan: from thc Ma lay Pl:nin sula sout heasl to thc i~lamb of'Florcs and Sulawesi as wcll a~ 011 l:lluntlcss ofTshorc islands through Ollt the cntirl: rangc and on Borneo. Mt. Kin abaJu : Poring and vicinity.
Ii-om Dcramakot, Sa bah.
Stl(~kes
have colonized a m~lltit.ude of ccologicnl niches. Many :;pecies inhabit shrubbery and ~s !10m are terrestriat a .f:,'Te;tt 'lUmber of them ~\re ra1her ineon piCU01IS and have foss rial habits; some liv in .fte hwai:er throl1~Ybout 1heir Jives, a number of others ven inhabit the oceans . .Many naKesJdT eHent climb -rs lind ase ne equally specialized ~11 snail or slugs, earthworm ', frogs, lizard.• insects, eVeo tennite , amplUbiU!l or (eptilc egg. , or bats. Observing these animals in Jlaturo ol\ell proves rather difticult. As a consequence of lheir position bigh up in the food chain. they have to be mueh more infTequenlthan Iheil' natural prey. In addition to this they spend most oftheil' time concealed and leave their hiding places usually only lor purposes 'ul:h as foraging or mating. Most species are crepuscular and l:<Jn therefore only be observed during the early hours of dawn lind dusk. Due to their noiseless type of 10c0l11otion and Ihe inability to voca lize Ihey hardly give away their presence. Most encounters are therefore pure coinl:idences. This is particularly true in th e case or fo ssorial species which arc mostl y found only by lurning over rocks and rorting logs. SOllle species hunt by ambushlllg prey aud oilen remain Illotion le 's for day '. This habit is usually aided by perlcd eamounage. Whilo most snakes arc oviparous, some give birth to hve young. In a few cases, an active incubation of the eggs can be observed , with th' female roi ling around her 'llIt ch and optimizing the incubal iontell1p 'rature by muscular l:onll1ll:tions. As far as Uomeo is concerned, the 135 species reeordcd from there arc grouped in 10 fall1ili 's and (i I genera. Of these, 7 1 ~lIl1 I1i es in 40 genera and 67 SpeCH.!S occur on MI. /(jnabalu (camp. Fig. XIII , p. 36). Using the identification key, individual s pec ill1 e n ~ ca n easil y be as. igned to a f ~lIllll y. AII()cating a certain speetnlen o rrolubrid ~ n ake 10 the eOITeel genu~, however. proves far more di!1ieult as this is the famil y \\ hleh accommodates the greatesl numher or spe..: ies. In Ihis case it IS usually nccessal)' to evaluate :l l:omblnalion o r w lour and lepidolic trai ts. The sexes hardl y differ with regard to their appea rances, hul IlI ll o~'e ncti c Iransformation ' are a common phenomenon meaning that Juve nilc~ bear diOcrcnl co lour patterns than the adults. This has been tnkcn ml o CU ll, ideratioll in the id":l1tificatioll "cy.
Distin~uishing
features in snakes Fig. ('VI Scalc~ of the alia I region . III Itlt\'Idl'<.I
Fi:.:. ('\' :1 snake .
al1:11
~IIPI';i I : , llIal
lIli"ralahial ro~tral
.
pr<:irolltal I"lI)nt:" mcntal groo\'e slIblll:l:>.,llary
IL'IIlJ1llral J1osttelllJ1nr:11
~l1akc~
d.. I\:l1 - -
Wi th IIll
1 ; lIlg~
r- ----...-
(".:.:k<.l)
~~ ...~t':Vrt; ·."t·r't '~ ('~':-z~r;';:;:
upistuj!lyph
tlllr'al
rear-fangs MlaI..L·'
'n:i1,e~
l>rotcro J,!I YI>h \.\ ith fangs mou nt<.:d 011 :t lI11nwvahk 11Iil.xillary hUlle in the I"ront of the mout h
s oll-noJ,!I~' ph
sna"es \~ith fa ll g~ IIWllllICd on a ro tatable maxillary bOIlI!. so that the 1~lng can he fo lded nat aga ll"t ti ll':
~ ,,"r:::O~f~tl~'C;'~1T1~O~I~tl~h~~~~~? Fig. (,\ '11 Methods lIrcoulltillg l1umbcrof 'calc rows.
Fig.
eVil.
Di ffercnt dcnt itions of' snakes.
Scrpcnk~
Key to th e famili es of Bornean snakes (in part aftcr
M ,\ I\ IIII"
&
GRO';S\1A N N
1997 )
B
A Ventrals diflcren! fmlll dorsals (A) .......... .. .... 5 • Ventrals similar or indifTcrent from dorsals .. 2 2 Eyes visihle. not hidden heneath scales ... ... 3 • Eyes hidden beneath scales (11): head nol sct ofT from the hody. distinctl y round<.'d ... Ty phlopidae ] All scales ve ry smal l (e) , in XO or more rows arou nd the body ............. ............. Aeroehordidac • Scales in Ics!> than )0 rows around the body ... 4 4 5-6 Slipralahials; mental groove present (I) ..... . .......... .... ...... .... ................ .... ... Cy lindrophiidac • 4 Supralabials: mental groove abst:n t (i-:) .... .. .... . ................................... ............. .. .. Anomoehilidat:
[
5 Loreal shield groovt:d: anterior teeth of upper jaw distinct ly enlarged and hinged to fold back (F) .. ... ..... ..... ............. ..... ................. .......... C rota lidae • Loreal shield withollt groove; anterior teeth of the upper j,m not distinctly enlarged ..................... 6 6 Supralabials with pits (G) .... .... .. .. .. . Pyth onidae • Supralabials without pit ~ ....... ..... ....... ... ....... ... .. 7 7 Large. rhombic interparietal shield surround ed by 4 ~ hield s of more or lc;,s equal SiLl'S (II) ... ....... .. .. ... ................................... ....... ..... .. Xc nupelticlac • Scales ofthc head not likc that .... .... " ..... ..... ..... K
G
a~piij~~i1=~~~
H Antaior teeth of uppcrjalV distinct ly enlarged and
syringe-like ( I): loreal shield absent (J) ........ ".". " ..... ..... .."....................... ........... ....... .... ElalJid ac • Anterior teeth llfupperjaw not di st inctl y enlarged llr syringe-like ............ .. ........ ....................... ...... 9 9 Prefrontal shields distinctly t:nlargcd. cun!>iderabl y larger than all other shie lds on the head; intemasal~ fused with na sa ls: no distinct frontal ~h i e ld: di stinct parie tals ab;,ent (K): sma ll body si/.e ..... ........... ........... ........... ................... Xenophidi idae • SGJiation of head not like that .... .. .. Co luhr'idac
I
fo-~/ K
29')
Colubndae
Colubridae This 1;lIllily COIISlillilt:" llit: t!fLl UP llr rCPlile" \\'illi llit: g r..:atesllllllllher of' gcncra alld spe..:i es llf pnxlominanll: harllllt:,,~ ~lIakc~ . They arc ra tlia helcrot:enous \\ ilh regard 10 Iheir appearall<.:e'i. t:c()logie~ . nlld n:prndllcl i\'..: modes. SOIilC spccie . poss..:ss vellom glalld~ and grooved bngs. hul Il:w po"e a rea l 111I"t:al to an adult. hea lthy hllman hcing . This i" wh y llie demarcatioll of Iii is {;lInil y or~nakes a ga ill~1 olhcr~ is for a large pari ha "ed on dClltllion . A" a rult:. I:o l uhrid~ ha'oc a g l y phou ~ teeth (\\'ilh 11 0 grooves or ca llais. "tl"lle willbrids" ) o r arc npi~lhoglypll\lus (wilh grooved fall gs ill the po'ilt:rior upper .i ~1\\ . " rear fan gt:d sllakes" ). Their \'ellt ral "<.:al.;,, art: a lway~ 01" a di 0(; re II I size th all the sl:al.; ~ on the upper hod y. I'il organs or other tlll:nnal sells()r~ nil Ihe SIIPIII arc absent. alld neither arc there rudimcllh or limbs 1(1U lld in llic s \..eieipii. I'he divi ~ loll pI" Ihis 1~lmil y illto vari (lus s ubfa milie s. their dadi stil: relaliollship~ . the II llersubf:lll1i lla r dCl11an:atioll . and Ihc sy:-.tematic positioll of' particu lar ge llera sti ll req uir..: furlher re~c areh .
Key to the Bornean genera of the famil y Colubridae Nostrils poi nted upward (A) ............... ........... ... 2 • Nostri ls pointed to the sides (8) ............. .. ......... 6 2 Pupil round .......... .. ..... ......... ......... Opisthotropis • Pupil vertica l-oval (e) ... ..... ..... ........ ..... .......... .. . 3
3 5 supralabial s .. ... ..... .............................. Fon/Ollia • More than 5 supralabials ...................... ............. 4 4 Kee led dorsa ls (D) in more than 40 rows ....... .... ..... .......... .. .................. ..... ... .. ............. HOl1la/opsi!l
• Dorsa ls in less than 40 rows ............. ......... ........ 5 5 Dorsal s smooth ....... ... ... ........ ..... ... ....... £lIhydris • Dorsals keeled (D) ................... ......... .. CeriJcrll.l" 6 Pupil round ...... ........ .. .. .... ..... ..... ... ..... .... .......... 17 • Pupil vertical -oval (e) or a ve rtica l slit (E) ...... 8
• Pupi l hori zontal
(F) .......... .. .......... ... ..... .............. 7
7 Ventrals buckled (G): vertebrals not enlarged ... . ... ...... ... .......... .. ............... ............. ..... DI)'ophiops • Ventrals rounded: vertebra Is enlarged (H) ......... . ... ............................. .. ................ ........... A lruetll/lu
8 Asymmetrical . cutellat ion of chin withoul mental groove ( I) ......... ...................... ..... ....... .... .... .... 9 • Symmetrica l scu tcll ation of chin with menial groove (1) ............................... .. ........................ II 9 Subcaudals divided (K), 15 ( 17) dorsal sca le rows ................. ......................................................... 10
• Subcaudals cntire (L), 13 dorsa l scale rows ...... . ......................................................... Ap/upe/lura
.Ion
Co lubridac
111 One pre-c hinshield anterior to 2 pair of chinshields (M) ... .............................. ...... . /trIer/wIlls • No pre-chinshield anterior 3 pairs of chinshields
N
(N) .......... ...... ........ ........ ... .............. .... ...... Pareus
11 13- 15 dorsal sca lc rows at mid-body ... ....... ....... . ... ... ............................................. ... DrrocalalllllS
• More than 15 dorsal scale rows at mid-body .... 12 12 One row of small scales betwccn prerronwls and I'ron-
tals and supraoeu lar shields (0) ... ... ... S(o/i('~kiu • No row of sma ll sca les between prefrontals and frolltals and supraocular shields ..... .. ............. 13 13
Tail ex tremely long; dorsals strongly keeled (P) .. ... ... . ...... .. .... .... ... .. .... ... ..... ..... ... .... .
Lept/lroplri.~
• Tail ofn onnal length : not keded or weakly keeled dorsals .... ........... .... ... .... .. .... .... ....... ... .. ........ .... . 14 14 Alllerior maxillar teeth enla rged, separated from subseq uelll tecth by a gap (0) ............ . Lycot/tJII • Anterior maxillar teeth not enlarged ....... .. ..... 15 15 Eye modcrately large to sma ll ..... .. Sle}(Olllltll s
• Eye very large ... .. ..... .... ... ........... ..................... 16
Q
~
16 Verteb rals enla rged ( R) ....... .. ........ .... ........ HoiKU
• Vertebra ls not enl arged ...... ...
P.W lIIIIIIOdYllustes
17 Head distinctly set off from the neck (S) ...... 18
• Head not or hardl y sel ofT fro m the neck ... ... . 26 18 Laleral sca les di fTcrent fro m dorsals, ve rtebra ls larger than dorsals, in 3 rows (T) .... X('I/Odl'rlllll.l'
• Body covered with uni form dorsals .... ........... 19 19 Ventrals and subcaudals round on their side~ .... .... .. .... ..... .... ..... ... .... ... .... ... .. ...... .. ........ ... ... ... ... . 20
• Vcntrals and subcaudals squarish on Iheir side. (U) ........ ................... ......... ..... ........ ........................ 24 20 2 or more loreal shie lds (V) .. ......... ........... P~vus • I lorcal
~ hi eld
(W) .............. . .. ... ..................... 2 1
21 More than 170 ventral scales ........ ................. 22
• Less than 170 ventra l scales ... ........................ 36 22 Dorsals in 17 rows, more than 130 suhca udals . .... .. ... .. ....... .. .. .... ........... .... .. .. ...... ..... . Xelle/uphis
V
• More than 17 rows of dorsal cales; if in 17 rows, then less than 120 lIbcalldals .. .. ........ ....... ... . 23 23 Colour or tai l in distinct contrast to colour of body ... . .. . . . . . . .. ... . .... . . . .. ... ..... . .. . . .. .. . .. ... . ...... . GOIIJ'OSOIllII
• Colour of tail not in distinct contrast 10 colour of body ......................................................... /:;/(/1''' ('
301
l' lliu hrJ dac
24 Vertebrals largel than dorsals (X) : 13 15 dorsa l locale rows around mid-body ... ... . /)(,lIdrelllplJ is • Venebral not largn than dor~als ; 17 19 dorsn l sca le rows at mid-body ................................ ... 25
y
25 17 dorsal !>ca lc rows at mid-body .... Clt rysopeJe/l • 19 dorsal seale rows at mid-hody ...... GOIIYlJphis 26 Lore,,1 shield pre~ent (Y) ........................... ..... 30 • Loreal shield absen t (Z) ................................. 27 27 Mon.: than 13 dorsal sca le rows ..................... 28 • I] dorsal sca le rows; prefrontal and internasal scales as onc shield (AA) ... ............. ('a/am aria 28 7 or m()fe slIpra labia ls ........................... ......... 29 • Less than 7 supralabiais ........... Pscudorabdilm 29 Ml)re than 7 slIpralabi als ... ................. . l.iol'dlis • 7 supralabials ...................... ...... Oli~(/doll (pan) 30 Supralahials in con tact with eye (138) ........... 3 1 • Supralabials not in contact with eye (C'e) ........ . ...... .. .. ... ....... ............. ........... .. .. ......
Z
AA
~EQI
I~vdrahlabes
3113 dorsa l scale rows at mid-bod y ...................... . ................................................. .... GOlIgyllJ.w lI1t1
• More than 13 dorsal scale rows at mid-body 32 32 5 or 0 slipralabials ......... .................................. 33 • More than (, slIpralabial ' ........ ..... ................... 34 33 Dorsals in 15 rows at mid-body .......... Elapoidi.1 • Dorsals in 17 rows at mid-body .. Oligoiloll (pan) 34 Less than 135 ven trals ........ ......... O/'I!(/('(IJIlIII/(S • More than 135 vcntrals ...................... .. ......... 35 35 Less than 75 slIbcaudals .......... . Oligo(lolI (pan) • More than 75 slIbcalidals ................. SibYll ophis 36 Anterior dorsal scales smaller than those at midbody and arranged in obliqlll': rows ................... . ..... ............................. ... ............. PSf'IIc/UX(!IIOr/o/l
• Dorsal ca lc, unifonn ..................................... 37 37 Last 2 teeth of upper jaw distinctly enlarged and separated by a distinct interspa..: e from the anterior tt:eth (DD): the nape may be spread to form a hood whe n molestt:d (EE) ....... .... .. .. Mucropistlwdoll • Teeth of the upper jaw withollt or with small inter. pace ; the nupc cannot be spread ....... ..... 3H 38 The current knowledge of the genera A lllpiliesllla. Rll/lh{loph i.~ . and Xell ll('/,/,ophis does not permit to di stinguish tht:m on the basis of external traits alnne. Please refer to the species' diagnoses.
302
BB
~ DD
Co lubri dac
Cie nLlS A /w ellil/a L II\K,
1807
The genus ( /1/(/ ('111//(/ is di stributed th ro ug hout the entin: Orienta l region , ran gin ' fro m southern China to India and in lil e soulh 10 Ih e Indo-/\lI 'lnti ian I\ n;hipelago . It is al so present on so me of the Ph ilippine is land s. i'vI o(li.:ral ely i;.lrge. very sic mkr tree 'nakes: body s lightl y compressed lateral! . , head cl o n ~ate and distinct from the body, wilh a sha rpl y ddined Can lh u<; ros lrali s. tapering 10 a point : eyes large. with horizo ntal pupils whi ch cont rac t in bri ght light : nostril;. sit llaled in Ihe posh:rior part o r a singular nasa l: IS rows or imbricate, un kee led dorsa l scales: vertebral
Tau. 37 Di stribuli tl ll or . 1 11/(,' " I1I1 -s p cc i c~ on
sca les s light ly enlarged: vcnlrab rounded, sOll1dimes with a wcak lateral keel : lail long, sul1ca uda ls arra nged in pair~ : hcmi pe ni s Sh0l1 and nOI j() rked. The spec ie. o f the genus .'1 1/(11'(11//0 an: diurnal and livc c hi e n y in bushes and shrubs through whi ch they can gl ide with Qreat ele!!a nee and <; peed. It is on ly occas ionall y tha t they d e~ce nd III the grou nd in search of prey. Their f~1Vo ur i t e j()od is li/arlls. but they also take frogs, snakes. bird <;, <;Illalllllalll 11lals, and even fi sh. I\ s filr as is known , all s pee i e~ give birth to livc yo ung. These arc "ndse co lubrids" whi ch do nOI posc a threa l to man. F. ight speci es are kn own . two o f whi ch occ ur on 1J0 rneo, onl: 0 11 1"11. Kina ba lu.
I ~()rnc() .
Sahah
All ac/II //(/I;,s('io /(//(/ A lwl'flllla pl'l/Silll1
( r IS( I II'R ,
I'.-a.\ illil
I
K inaba l"
• • i •
1~~51
( H, '1 1. 11:>27 )
I1ntne ,
Sara" ak
K'lil mallla ll
• •
• •
•
usual I bri ght. grcen witholll an y pat lel'l\ ; nllted fo r il s ex trelllel y poinl ed snoli i.
Key to the Bornean species of A /wellil/a I) () r~ al
surface usuall y grce n, as an exce pti on brown or grey, bUI alwa ys uni 1'01111; dorsa l side of head without markings : ana l ~cll t e usuall y divided ....... ...... .. .. .... ........ ...... ..... ...... .... prasil/fl prtlsillo
• Dursal slIrfllcc brown 0 1' grey. with ill-defined dark crossbars: dorsal si de of head stippled or ~ po tl c d with black. i.c. marked; anal sc ute ent ire .......... .. ............... ....... ........... .............. ... ... ....... . f ascio /a/a
A /Ul elulla prasilla prasilla I)n o/ lln,
/ J/H,\ IIW.\
}',,,'\('n h l
P /"(I\ I/I(/
t/" " '11I1/"
Bu ll', 1r.?7· 5·l 5- T) I1'" l un, l i l ~' : Ja\a 19J I : 10. J~
S\II I II
\ '1" 199 1: 336
("·IIS"h'.
SlI l lIlN'o
!)n o / I/II,\ /" '(1,\'11111 \ .
( 13ll ll, . 1X27 )
F lU
'I" \ l'IxJ : n: 1\1
N II \I(II
\I " " "
I ') 'J ~h .
2·lh;
& S'CI4:in, 19)\.... 24
Oia g nos is Lxtre mely slim s p ec i e~ (a lnll>st 2 Illt:ln:s in lcngth , but nill y bet wl:l:11 120 and ISO e m on
Description I lead distincl , with ~ harpl y edged Canthus ro ~ trali s. I la rge preoeular (rare ly l'o llo\\'l:u by a second . ,' mal ler one ); 2 pos l oc ul a l'~ ; usuall y 2, mo re rarely J lorea!": 2 I 2 or 2 13 tc mporals: X- I() (rarely 7 ) supra la bial ', the -1,1, to 6'h (rarely -I 'h and 5'" or 5'" In 7' ) to uching the ..:ye ; 9- 10 (rarely X) inrrala bials ; do rsal sca les in 15 rows. smooth : I Xt} -2-11 ve ntrals; anal sc ute llsually di vi ded: 1--1 1- 1()() di vidc:d su bca uda is . Up pn ~i d c: no rm all y lea 1'- tn ye ll OWi sh gree n. but a lso d in y white, light grey. ye Il O\\ . oran ge. light hro wn . I'aw n-eo lo urcd, o r blUi sh g re..: n. The \·ari at io n in colour appca rs to be. a t lca ~ 1 pa rtl y. pa rIl clilar 10 certa in popul atio ns. It i:- th c:rl: ro rc li ke ly Ihal no t all ort hc menl io ned colour \'ari e tl es occur o n Horneo . A I \\ ay~ w ith bl ac k a nd w hi t i ~h . <;trc:ab whic h form b a n d~ on Ih t.: Inl e rslitl al s kin of the a nt e ri or th ird o f til \: body: \'c nlra l si d\:s pa l.: grecn wi th a yc llll\\ ls h \\ hlle ~ tn pe alo ng the s ide .
30J
Cllluhridal'
1//(/<'/11//" v ~lriti~s fWIII
JUVl'Iliic Il lrllean s peei mcns l'x hihit ~ horl. obliqlll:ly arranged , bright hille ,Ireak . (markings) on the si des or the nl'l'k and the anterior hall' or th e body. Thi~ ,pecil's is ()vo\'ivlparou~. and a litter lIlay COIl~Ist or I tn I ~ Il\e young. The ofbpring m<.:aslIr<.: ahout 40 to 4.1 cm. Eco ln ~~I /E t h () l n:.!~ /11'0 .\' /111/ is I(Hl nd in lowland area~ fro II I lev<.:l to ahout 1.170 m altitude. \\ ilh a predike-
.,tlll/ L'lIiI/O ~ea
.104
Thailand
1'1'11.\'/1/(/
(left)
and
in di ITcn: nl colollr Malaysia (ab,l\c).
W~sl
lion for th<.: region s at low ckvations. It inhabib tropical \\'l't lo\\' land fores ts and tropical weI montane fi.m.:st~. but is often t()lll1d ill secondary tropi cal foresh. in trori<.:al dry, op<.:n forests, alollg forests path s alld tracks. in dense ly vcgetated shrublands. plantations, in bushy areas, on hedges and trees around houses. and in gardens and other \·egl'ta t<.:d ar<.:as as wdl. Thi . ).nake is invariably assoc iat ed with dense vegctation , from tall grasses to shr uh~ and high tree fo liage. It is mainly arboreal, allhough juvcniks may nlsll bc round foraging on the ground .. trictl y diurnal and very active. A . /II'I/sillil fced, primaril y on li 7.a rd~ and fi·ogs. a lso laking small birds. small arborcal mammals and olher snakes. ThiS spec ics is mild in di sposi tion . il opens its mou th when threatened . but it se ldolll altCl11pt. t(l hlle. It is harmless. but its bite has becn n.:ported to produce sl ight local sy mptoms such as intbmm;)li on at the bite ).i te . Hch ing and mild rain .
Colubridae
The snakes have an interesting threat di sp lay in whi ch they exll:lld the ton gue hal fway and kave it eXlended for as long as they fee l di . turhed . The lo ngitudinal groove heron.: the eye on the side o f the snout makes it possib le for thi s species to see straight ahead and toells 011 its prey in ~Il:reoscopic vision . There arc ditTen:nt observations ahout the mating of thi~ spec ies. 11 r-- I/!>o (2000) observed mating in the late anernoon, last ing 1)0 minutes. Bl ll..\ &
Oistribulioll 'I he ~pec ie s rangcs from southern China 10 Illdi a. and south intn the Indo-Australian /\ rchipelago . It al so illhahit s some of the PhilipPllle i s land ~. It is also widely di stributed on Borneo. !VI L Kilwhalll : Kiau . Rundu Tuhall.
Genus Ampltiesma D UMleRt!.. I3IBRo & l) t I Mt~ RIL. I ~54
Thiny-se\'en speci es arc known in total (\) \\ III et al. 19l)1) , 4 occ ur on Borneo, 2 (lr which Oil Ivll. Kinabalu.
The genus AIII{,hieSIIIII is widel y distribut..:d throughout the sou thern, ea~t..:rn and southeastern parts or Asia ranging fi'om PaJ..istan and Illdia eastwards III ea~lenl Chilla (ilH.:lusi"e of Taiwan), north into the ~llu l hernmost port ion ~ 0 f Ru ss ia and Ja pa n. to Sumatra in the ~o ulh alld . I ila we~i in the sou lheast. Sma ll to Illoderalel y large, t..:rre ·tria l. semiaquati e, harmle s~ s nake~ ; head di~tinet li'om the hod y: eyes la rge with round pupil s: interna sa ls widen ed anteriorly, n()~tril s positioned laterally: dorsa l ~caks kce kd in 15-_ 1 rows, wilh or without apical pit s: allal scull; divided , subcaudals arranged in pairs: hemipcni s and Sulcus "perma l ic us undi vid..:d : maxi liar teeth arnlllged in a continuo us serie~, increasing in length posteriorl y, the IW'O posteriormost teet h ma y be abruptly e nlargeo (\11 \1.1\"'1 ' 19(0). The snakes inhabil lowland andior montane s ituali ons up to about 20()() III alti tude. They live in primalY as we ll as in secol1lJary rorests, Iheir rringes, alld bush land s. A Illore or less strict adap tation 10 moisl habitats i~ noticeable by their prefL:rn;d occurrence alollg rive rs, lakes. ponds, w:t terholes, or ~lI1a ll rapid Illountain streams. They reproduce by laying eggs.
Taxonolllic COIIIIII('nl: ._ '1"11 (, (191)1 , 11)1)4) recorded : tlllplti('\lIIl/ sio/allllll from 13mneo and rrom MI. Kinabalu, but thi s ob\ iously con~ t itllte~ a misidenti licatinn oranolher species a lthough ill S as ye t unclear which . . 1. SIO{UIIIII1 is a species l" ~tnhuled considerably tnnh(,r1o the north . S II 1111 1\(, & 1,<.1 I{ ( II)')') did 1I0t li st thi S s peCle~ any l11ore, but abo did not otTer an exrlanatinn.
Tab, 311 Dislrihulion of , IIIIIJ/1I1 ' WIII-'I,,;.; ics UI1
IllIlKI (2000) recorded that maling started in the early mornIng, la~tin g lor 5-6 h()ur~ . (iesta liont akcs abnllt 5 monlhs (LlI ·K(;\IAf'. 195(l).
Key to the Bornean species of A mpiriesllla 19 dorsa l scale rows at Illid-body ."."""",, .... .. . 2 • 17 dorsal sca le rows at mid-bod y"."." ......... .... 3 2 Snout wi th a broad white stripe "." .... fla vijrrJ/ls '0
: IIIII)hic.\ II/{/ /rell(//I/III
..11II/)lI i<'.I'II/
R,
... .... " ........... ............ " ...... , ....... ",.... .\·/'I'UIII(I('(!/IS(!
• 164-166 ve ntrals; 11 2- 11 6 subeaudal s ... ......... ... . ........................ " .................................... /i'ellu IIIIII
Hllrn ~o.
( n OI I I
",ot K,
I X93)
,·llllfliliesJlla .\ lIf(tll'aCeIl Se (( j. I~ 1111
K,
I X72)
KlOabalu
•
IXX7)
(Dl "". I '):!])
p(!le r .l' ii
3 134-156 ventrals; 52-1 12 subca udals ................ ..
S;Jbah
A 1IIf1i1 ieslll II .flal'ili·ulI ,\ (13(1"[1·",.1
white st ripe on nout ........................
•
---
•
-
•
I3runci
•
Kalimflnlan
Sara"ak
.
• • • •
• t
• 305
( ' lllllhrida ~
Amplliesmo .ITm'(lroll s (B( H II l i'(l/ ll d
i.) : 1\1 1.
""",h.II"
/ r0I'/( luIl0lll\ " ,,"" \t.l " ~ X , 1)1 R (J~
.v,I//'"
x.t
11,\\ " \(,' It , IXX 71>
<)(,
I X~7)
1\ 1'~
IOl'a l-
1\",,1> illl ll ',O
Iltn//nm "
f\..I tK 1.' 1 \f(1I
I >\90 .1 I-HI , pi
l)
lig_ 2. a-c:
I '11'1>' ~ (l.\, II \NII " " I!JOOa, 70. Sill " m il I l}() I a til I l) I i ':1. 7:
/]""1/10"\ 1)" ""
14.J -l t ) +
11'( ,r~ l{,
1'12 ,, 2,
I ;HI N I I:\l< 11 t\.
~ 11111
"'''I,kl'
I ')"
10,3 2:
III
11 .\\"
19X4 - 2 ...
:Valn', ( 11l ~ . ~l!d j I I(nl/nUl\ , "" ' 11 11 ' J9"J ' 2X 1IIII' /I1""'1tl /T,/l'I/"' /I,\ , " " ' ,II \(, 1'1'1 1: '-1 7: 1\1 \I , ~ \\ , I')t).j h' 2-l h . 1'/1)6l' ' '1', /1,1\ ,x (, """\I\'N I')'l? "1
," I)
S IIIIII\' "t;.:
I ~, ,, ,,
1'149
I(,~
Di agnosis Small. ~klltkr s pe c i~s , eas il y id e nlili cd by Ihe while slnpe acr(l~S pre f'roilia Is and f'ronlal. 19 dorsnl " calc~ III a row' a l mid -bllll y; 75 e lll llI a '( , (SI I IH I1\( , & 11\ 101 \{ Il)'}!)) . Descriptio n Olle loreal : 1-2 preoclilars: 2-3 po;-,Ioculars: 24 J temporals : 7-9 ,;upralahials. IhL: 4'1< and 5'" or thc 5'1< and (,'" touchillg the cye: 19 dorsals at mid -hod y, weakl y kt:t:kd : I ·H,-I S7 \' cntral~ ; all:ll scutc entirt:: X7-IOJ subea lldal s. di\'idt:d . Ciret:'lli sh dor~all y. With a vertehral row llr ::;pOI S: two con S pi C ll()lI ~ whilt:' ~Pllt:" on either side or ro\\ s
or
c
!Ii '0 7
« g
Fig,306 P0I1 rai i or · 111111"
1<',\' /1 11 1
IllI n /mll " rrom Porin g.
Sg. Klpull gil I.
the hody. arranged parallel or alternating. the lower one in wlltact with the ventral scall:s: upper side or hl:ad grceni~h \\ ilh black pallcrn : prerrnnlab entirely while. frontal wh ite anteriorl y; lahials whit.;, bordered with black ; chin and throal white: ventra l side white . with large . irreg ularly s cattered black blotches. 1110re densd y and more lincl y spoil cd towa rds the tail : lIndersid . of'tail black.l:aeh slIhcaudal scale with a !:trey spot. 'I his ~ p ceie s is ()\·iparous.
Clllubrida.:
[eulogy/E t holog)' Found along small , clcar, Ilowinu walcrs in primary or sL:L:o ndary fon:sls. rhL: 'nakes arc ol"'-:n SL:..: n sw imming wilh the head held hi g h. tilllphics/I/{/ f/111'i/i"(}/Is is bOlh di - and noclumal. II fl;..:ds on rrogs, Ih.:ir .:ggs, and ladpok~ . TI\ O s pccimcn~ wcrc oh~er\'cd al an allilude or ahoul600 III a.s. 1. in a rapidly flowin g, dearslr..:alll. One or these s n ak.:~ was basking on a rock, the olhcr wa~ ~ubmcrgcd clinuing 10 a fi!ll':l1 Ir.:c trun" (M 1'1111'\ & (j1(()~~M ·I 'I 1!J97). M II "'~llS (1996) discovered a ~ pccill1en al around 10.00 :1.111 . on the confluence or thL: 1\\'0 Kipun gi t ~ Iream s. basking somc 2,5 m high on a gingcr plant. Anothcr nne lVa ~ I()lllld by him al arOlilld 1.00 p.111 . on th.: Sg. Kadalllaian , ami yet anolhcr at around 3. 00 p.m. on Ihe Sg. Liudan. Both the laller quic"l y ton" to flight into the stream , cros-;ed over to Ih.: opposite bank and disapp.:ared amongst a thicket or roots. Distrihution ,llIIphic,w w/hll ·i/iwl.\ is a BOl'llcan endcmite . Mt. Kinabalu : Kiau ; 2 100 1CCI (11 \t\ II\( II I ()O()a), Sg. Kadalllaian (1000 Ill) , Sg. Liudan (\300 Ill) , Sg. Kipungit I (M 'I'l1111 \ unpuh l. ), 011 Ihe eonflucncc or the two Kipungit streams (500 Ill) .
A lllphie.\'IIIlI SlII'lII V{lCeIlSl'
\IU 'OH 'IW('U\I\ ( , \ ' l illI( ,
rrofJu/uIJIJIII,\
flU rropidollOllt,\ Hhl(
Sanlwa~
(Gl.
amI l:a\1
"Iaflf\ PI II
1-( ....
'1III ' \{ ,
1:-: 72 I X7 1)
I X72)
'::;;l)6 ( II t ' W nalllt'
'1~ pl' 1""" li l y:
Il\ dl~~
Pili H", IX 7 1 ".7" (nlH I III(lC'lIlalll\ '>.1r:"\ ,.~ ,IIld 1.1, 1 IIIti "., ('11\1\, BIH II Nt 01 I< I X '7 l)
liHI'U/OIlt}ltI,\ maclIlLlft"
1 III
II";,
Ty p.' Inca lif) :
IX(,cI)
TroplilwIfIIU\
'art/ll'O.
IXl)O,, ' IclO;
III R'1O)\J
1<)17 X5
rrtlpit/olltllll\ OIl/Cultllll,\, /j 'Opidmw lt/\
,\,,1/,/\ ,uru .·u('(',,,,, I "c l ) ,
,> '" II'
:\;11//'/'
\I II'
\1'1'41\1 '//1/' //\' " . 1>1
SUKJRI'
Il)() I a 5:-\ II "" . . ( II l'JOOa 70
"illl I It I/{I)
Sarll\cln'll\l,\ ( S IC!),
1<)11 (1).)\)
10. 1 1, ,2 I)~ . III\ NII\~11
XO.
8:
19H4 24 1\1 \"Ij11l1 \ 19X J. 27: ~I1 I II1INl' It'91 I'J'J-Ih _ N ; SII" III", & IN',I ~ 1<)1)1) . 170
.·fmphi(,MIW \(waH'tln' II\'; ,\,
3c1X ;
tv'"
~Ml "
·fmphw,\ IIIi1 ,"lril\\'oA(,II\I\ , i\1 ·\1 h.~11
"'i
11}9~~h
2... 6
l)iagnosis S lender with a long tail: b e~ l idcntified by its colour paltern, C 'pl:L: ially Ihe I,elltml pattern i~ c haracteristic. 17 rows of dorsal scalc~ at mid-body; maximum length 7X cm . l)l'scri ptiu n One km~a l : I ( ...m.:l y 2) pr,,:oL:ular~ ; 3 (rarely 2) PllSIOculars; 2 ~ 3 or 212 kmporal s; X (rardy 7 Dr 9) ~u pralahial s, Ihe 3,,1 10 5'h or thc -l,h to 6,1, ill contact
Fi g. J07 . I II/flilie.'II/ll .'lIl'lIl' 11('(' //'\1 ' fm lll Sf!,. L iwngo. ~''"'''--""-'
.
"'"
~
.. z
<.:~-..-..--
307
~
Cuilibridae
with the eye: 17 nil ~ orlh) r~al "ca ic" at mid-body. all strongl v keeled: 134- 15-1 ventral~ : anal se ule di vided : 52- 11 _ ~ubcaudal s. dl\,ldcd (SI UI\ i-l,fH 1')22 ). Oli\e-broll'n ahme. with a ~er i e ~ ordark Iran s ver~c hal'". two ~ 'nes ur ~/ ellow. red , or orange spnls llr siripe,,; ,)1' blad,i~h abolc. \\.llh urange ~pots alllerinri).: :1 ~ ene ~ Ill' black spots on each side ; head Icrllli cliialed \\'ith hlae k; labiab yell(l\\ or red 1\'Ilh broad black :-uHlle:-; 1(lI\er slIrfilce chl'ckered with black ;1Ilt! yello\\'. ~ lll1lelll11eS with the black predUlnin ;ltln g. .Il1l·e nik " hrp\\,n : head ahol'e dark gre y: Iabial s willie 1.1 ilh hlack ()lIllin e~ . ;\ clutch l'\lntaln~ 4-." egg".
I:: cohl"\'/ F: th o lo"\' ,..,. ~.
This diurnal 10 crepu~eula r "nake has bc:cn 1'':eorded fn m a long clear ~treall1~ in Illontane us wel l a~ 101\ land reg IOns. but is al so rOlilld filr awa y rrom water. Its diet i" hased 011 rrogs . Iheir larvuc and eggs, alld fish . It ;,. biology is a ltogC:lher poorl y )..110\\
n.
Di strihu t io n
is lIal il e 10 iJornco and Peninsular :",Iulay"ia . :\,11. Kinabalu : Kiau. Lobang . Kellokok . 1.1I1llU LUlllll, Blindu Tuhan. 2100 11:1.'1 (II \ " ' l ll 1900a), IIcadqllart e r~. Sg . ;Vksilau. Sg. Lill'agll. I.iwago trail (1500 Ill). . 1/lIIJhi(' \111i/ .I'lIl'lIlI ·(II '(,II.I'( '
I{{' m a rk : Some aUlhors li"t 8lJ1,t:1I IIII1/W/I/(/(,I1/U/ll Ii)!' Ihe lilulta of' I\ornc(l, bUI there IS no evidem:e Ihal IIIIS species would occlir on Ihe island .
(jenus Ro~~a FI'I II N(it R, I X26 The gcnll" lioig{/ is distributed OWl' a vast geographical area r:lIlg lllg from Iran and parts the I<-lllller LSSR in the II'C"I righl Ihrough to China and Tai lI'an in Ihe ea"l , cO\'ering the whole nr . 'olliheast ;.\ sia. IlIdllllcsia. the PhiltppilH:s inlhc ~ou\h, tIl thc northern portion ul' AII,,(rali a and the .'ollll11on blallcis. I.on g III Illodcratel y long. sll:mlcr snakcs with long tatl !-.; bod y cOlllpn.:sscd lal ·rall y. head largc, vcry distinct I'ron, hIll y: c, cs large. Ihe pllpil a I'enkal s lit : enlargcd ~ roOlcd re;lr fall g s pre~elll ; 17- J I dnr~al :-caic rows ;tllllid-body; dorsal~ "l11onth . anal sClilc u~uall y enlll'l:, . ub audals arranged in pairs. The "lakes of Ihc g l:nll~ lilJiga arc nocturnal and arboreal. rhey lI:ed Oil frogs. lizard~, bird:.. alld their cggs. ~ll1all mamilla Is, and even ~nakc;,. . They 1'1.:producl' hy laying e gg ~. Altogether J() ~ pe e ie~ arc kno\\'lI . livc which occur on Borneo , alld l<-lUr Ihese have also been recIlrded f'rol1l I'vII. Klnabalu .
or
Key to th e Bornean species of Boiga 23-25 dorsal scale rows at mid-body .. .. ('j'IWt!Ofl • 19-21 dorsa l sea le rows al mid-budy ............... 2
2 19 dorsal scale rows al mid-hody ........ t/l'llpic: ii • 21 dmsal scale rows al mid-hody .................... 3 3
/JIJigll (/e /ldropll i l ll HniY,I1 dmll/(';:' /1
IIIIIII'{,(I' II .\
(\3<) 11 1'< (31 )11.
-
(\3" 1
I !Q 7)
/Joigll j(l.\ pidL'(I I I) I ' ~ lf 101. Hllm, '" & Bllig ll
.lOX
/Iig,.i('/'p~
IIMoI R. 1)\%)
I(;"r-IIII K. I X(d )
1)1 ~ 1I 1< 1 1 .
IX54)
II
ilh yellow rings on hody and lai l
•
\)or~ u ll1
brown or grey, no rings on the ho Iy .. 4
or head wit hout markings, brown or ulive .... ........ .. .... .. .. ... .. .... ..... .. ..... .... ...... lIi~,.iceps • Upper side of head wi th dark. light-edged markings .............. ... .............................. jllspit!ell
4 Upper side
Sah:lh Blligll tTl/ fIIl ol/ I \l " II . I X27 )
black
.. ............................... ..... t!efldrup!lilll IIIIIIeel en.\·
or
or
J)or~ulll
--
• • • • •
I
Kinabalu
--
• I
I
• • • • •
Bnlnei
• •
Sarawak
KalimaIHilt\
• • • -•- • • - • • .)
.)
( 'oluhriual?
/JoiX{I
Oi,,,,,,
' :1I1Ot/1I1I
C:VIlO(/Oll
(BOil, 1827)
BOil . t1L 7 5-1') " y pc t" ca lit~' : S'H11alla \I ~ ' " .; I <)')2~ : 116, I \)')4b 2-16
B"i ~(/ ('1/10t/OIl , VI
Oiaglltlsis Very large, but slendcr \\ ith a IOllg head \'cry di~ tinct Ii'olll the neck: body laterally strongly eOI11I ressed: easil y identifi ed on thc basis ofhui ld alld co lour pattern (sec be 111\\ ): 23-25 dorsal scale rows at mid-hod y. Adu ll specimens o f 200 to 2-W el11 an: average. but ncarly 2XO CIll in length may be rcached .
2-+X-290 vc ntral s: anal scute ent ire: I 1-+- 165 subca uIn rairs. Dorsal co loraticln usuall y ye llll\v i ~ h hrown \\'llh a variah le da rk (i.l? reddi sh hr(l\\ n. dark brown , or blacki sh brown) ba nded pattern which beeollle~
lb l ~,
Fij!. 3U9
JlI"c nd c HfII,!!J I
n
/I(Il/1JI1
from \\'c\ 1 Mala ysia.
Description Rod y stron g ly compressed laterall y: hea d ve ry elongate: an teri or teeth di stinctl y enlarged : I 10rea l; I large preoc\llar: 2-3 poslocllia rs (SII ')'JI '(, I \{ 1')22 ): 2 I 2, 2 I 3 or 3 1-3 telllporals: X-I () suprala hial. . usuall y the -+'11 to (J,I, or the 3,,1III 5111 (rarely 411 ' _7 'h. 41h -5"') in contacl wilh the eye: 23-25 dorsa l sca le rows al mid-bod y: ve rteb ra Is distinct ly enlargcd;
.l Oll
( 'oltlhridac
morc d ell~dy :u raile-cd posleriorly: lail \\ ith an allerllalln!! pallcnl III' \·ny dark aild yellow co loJlrs I' \n'ptlllilal ~ pc clmen~ may be light browni~h grey \\ ilh browil hand~ or dark g rey \\' Ilh hlack ish grey band~ . I he harld~ arc parlly bordercd darker. I lead ",ilh Ihe ha~lc colour 01' the d()r~tlJll. lighler III Ihe lahial rq:! l(ln : a hlad ~ln IK' bel ween tile eye al1d Ihc angle or the nllluth . .llI \'enile~ "ho\\ Ihe "allle Iype or palieJ'll , htll arc "uhslal1ll
bite. Ir alarmed, II Ilaltclb Ib ned vertically, displayillg thc hrighl YL' II(,,\. -=1l\(lIlr Ili'lhe Ihroat. 'I his (\pi~lhoglyphou~ ~nak-= -=all Inll iel palllful b lt e~, and el1\'c nOlllaIIOII" oi' hlllll:lnS \~ilh mark'd cililical clTccts ha\ e been reported . One specimcn \\as caught at nighlln a "ma ll 1Il:J II 1ma II r;lp "cI III I ilL' arca or tlte park or I'llrillg (1'v1 \1 f,.~11'~ 1992) . Dist.-ibutiull /Jui.1.!.1I ( l'I/udUI/ IS dl"nbllted from the \-lalay Pe-
lIinsula to Ihe l.e~ser Sunda I s land ~ . II al~1l (lCCllr~ unlhe I'lttlippine~ . 11. h:illabalu : Poring (501) m) .
Eculo:,!yfEthulug)
Boiga tlelldrophila
BlIIg{/ ( ~ I 'J}II"lIlIlllhabl\~ re!!ion" nrlO\\ allitude and
I" u~u ; dly found In IIm:k \'egdalion along clearIng" and on the edgc~ of primary and ;,.econdary tropical wei rures I'> , open dry rore"ts. l!'Oplcal illOlllane l(lre,h. III rllhber pl:lIllalions, alld 'Ill"e tu \'dlage~ . Thl;,. ralher ,Iuggl"h ~nake i" pred<)minanlly arboreal and an e.\ cdlenl ellll1her, hilt II lila ), be aCII\l' abo unlhe ground . During Ihe day, thl" n()ctulnal ,peCles retreat" ill Iree hole" and bccolllc" aCliH: ;11 du~k . lIoigll ('''l/ut/oJ} i" onen dlscm ered Oil hi anche" 0\ erhanglllg "mall creeks, or e\en "eell ,,\illll1lin g acrll~~ 1;lirl y wide I·i\,ers. II reed~ l11alnly on bm.b (Illciud ing ~ma ll rU\\ I) and Iheir egg,. hili \\ ill also lake Illamlllal<;. li /an ls, alld ~nakes . Thi" species i" \\ idespread. but nO\\ here LOnsider~d COlllnlon . It I~ Illli re:dl y a ggre~sl\'e , although ralh-=r vaJ'lablc in di"p(l~iIIOn , and may ~trike and
( BOl II 1>'!'\4hlflIlOlllrlll'},U\ l~t)(H.
71
I)
pt.'
(d
I{,
d('I1,"ujJl"I",
({f1lll'('/l'IIS
1X96)
\.11
,,1111, '(' /('11\ H ilt I I \~ ,j 1<,
IO f:l lil ~ : HOllh.·O
U"lgtl ddrdlofJllI'/(/ "~p o f\ 1 \N 1111 ' 19S"\ · 27 NrII~a dtJlltlt"'/~/JJld I \"I IIII ' ...X III \II}I 1('X2: 14:
,I..
~I"
.• , N
I 'IH '
~.J.
\1 \
~ \ 11 ,
I 'HI \ 1I\HIl
t'I'I.Jlr 2-1(,
I>ia!!"os is
Very Illuscular and large: hody laterally ~trllngly cOl1lpr-=ss-=d: readily di'lilll.!lIished i'rOIlI (lt lter species occurring III the same region by its eon~picu OU" ycllo\\ ihlad handcd pattel'll : 21 dorsal ~ca Ie rows atmid-hlldy. II. t/ (/I///CC/('//\ dill i.:r~ fwm Ihe other suhspec les /Juiga del/tlro/llii/a with regard 10 Ihe nUIliber or hands and the c(lIOr
or
I)csc ripliClII I'i ~.
3 10
J JU An al:Jrlllcd
1J(//~1I ITI/tll/()II fi
M,II:lys l:I.
One loreal: I prencular: 2 JlostoLular~: 1-3 anterior and 1-4 posterior 1i:1 111)(lrab; X. ran:ly 7 or ') , ~lIpra labia1s. u"ual ly thc y
l'o lubr idae
Crt.Hl1 POrlllg .
I{il!hl ahoH : Fil!. :112
POrlr~1I
of Ihe sp.:cimcn from
1I1 L' IOI' ,
RighI helow: Fig. C IX J)r:millg "rille hcad ,cli.:s from a speci llll'n !"WIll Ihl' 11<)-Coll,xlion ,
11I r..:. is mosl (onslanl. Upp..:r labia Is ydlow. wi lh fai rl y' hnlad black l1lar)! ins: 1111'0:11 yl'illlw. so me st:aks with bla(k lip ~: b..:ll y black wi lh latera l ro\v ' spot s (VI)\,II , 20()O) . R( ,1 · 1 I(, I II (I ~(J3 ) IT porlt.:d al1ll1l1
or
or
or
:I II
( 'olllhnda.:
Eculugy/F.I h n l og~' BOI,S!,(/ d. (//111('('1(' 111 inhablt~ Ilm land tropical Wl:t Itll ·.:~ts and mlingroV('~ . It i" usua lI y I'uund in vari-
l)LIS iluuded alld f'ur.:st.:d ar.:as, ClllS.:d "wamps and lIlar:;lll:~. along Ihe I ':l,!.:tated ban),." l)1' rivers.
Iii!!.
31~
~
IJ IllI/gII jll\'l lidl'lI fmlll Thailand
nle biolo!!y nflhi~ pal1 1cular ~ ub~pecic~ has Ilol hecll described in parlku lar. II i~ preslimahly ~illlilar to that of Ihe other stlb~pecies . MI N1111 \ & I)H,/I K ( I YlQ) round a "pecilllen in the l'iciniiY of a river IIi1iCh. at Ihe time or Ihe ob"uvalion. II'''" resting on Ihe g.round . It ,"h~cq"ently accepted a {{clJlidacfdlls fn ' /wlill a~ Illlld . OiSI.-ibulion rhls "ubspeeic" is cildellllc to 1I0111eo. II i~ re,);'onahly COllllllnn alld ha~ been reCllrded rmm all parts Ill' the island . 1\11. Kinahalu : Pnring (WO Ill) .
(DL t-tl:RIL
SoiKt/ .it/spidea BIBRO:--J & () t lMI RII. I X5 ~ )
11'1.<:11'1,11'1"'111 /""1>1,/, ·,,,,, \)1 \\1 '" 11 I ()l); I) pl' I" ra lit ): ,1"';t
. 1I 1111{4 'N &.
I)",,, kll .
I X'i·11>
Diagnus is Comparative l ~mall: ~Icnder; head ~hon. distinct, blunt: body laterally strongly compre>.scd: IllOst ea~ily idellti tied 011 the basi~ or it" CO l lltl' pattern (desuibed below) . 21 dorsal ~ca le rows at midbudy. Ma:-. imum length ahout 150 Clll .
Co lubridae
the vici nity of vill ages . II is ruund in dal11p ,111(1 wt:ll-watered biotopcs sut: h as flo oded ror ·sts. mar. hes. ~ \vamps . and along ri\t:r bank s. Boigo jaspit/('(I is arborea l ::t nc1 nocl urnal. Its diet consists mainl y 01' 1iza rds. espec ially geckos. hut it also takes sm::t ll mammal s. snak es. and bi rds and their eggs. It s an ti -pred::tt or ht:haviour inclu d e~ hi ss ing and tlatten ing the Ilcck lal era ll y whilc simu ltaneously expa nd in g it ve rti ca ll y. Whcn sei7cd it will bi le rcadit y. A female has bt:en reported as laying ils eggs in the nests of tree-dwel ling. termite~. 3 Illelrt:s abuvc the ground . Distrihution Boiga jaspid('{/ is disiribllied rrom the Malay Pt:ninsula sOllth and cast 10 Java. Borneo and Sumat ra. "It. Kinahalu : Pori ng (IIC)-Co ll ccl ion) .
rig. J 14 Boigll jll.I/,id"1I frolll Thailand. Oi,)S'" lIi~J"/ C C ·I).'
(;1 \T IIII< .
IX6J : 1, <1
' I \p~
teu'alit) : ·· ~. a'l
1I1dl~ S "
I)csnipliun
Eye very largt:: I lorea l: 1-2 pn.:oc ul ars: 2 postoCli lars: 212 (rarel y 2 · J [S II ' JN I (;1'1< 1922 1) Il:lllporal s: H-9 sLlpr,dabial s. the .I'" to 5'" touching Ihe eye: dorsa ls in 2 1 row~. verlcbral s dist inctl y' en largcd. wider thall lon g: 243-267 ve ntra Is: anal scutt: t: nlire: 140- 166 subcaudals, di vided . Dorsa ll y hrown to reddish brown. wi lh two row:.. of" dark spots or hal'S Oil Iht: sides and a reddi sh grey vertebral stript:. Ihc latter spottt:d with dark hrO\vlI 10 its si dcs: laterall y with a series of wh ill: spots wh ich Illay t:ncroa 'h on the vcrlt:bra ls: head of the ~a ll1e w lollr as the body with a da rk spoiled pal tc rn . the largest spols haw li ght margi ns : a dark slT-ipc \vith a lighloLitlinc on the lIape: wn lral si de bright yt:llow to ye ll ow ish in il s antt:rior portion. Eld ill g into \·" h il e towards Ih c tail. occasiona ll y black anteriorl y
Thi s species is round fro m sea lev!;: I 10 dcva tion s of ablllli :-\00 m and inhahil s primary fore"ts and
I)ia:.:nosis
Largt:. but rai rl y skndn; body lal erall y (;o mpres~cd : most casi ly idenlilicd Oil the basi .. or its co lour patkrn (sec be low). 2 1 dorsa l sca le rows at mid-bod y. Maxil11um leng.th 175 elll: 1lI,i1 cs larger than rem ales. Description
Onc lorea l: I preocu lar: :2 poslocu lars: I 12. 2 I 3. or 3 I J tcmpora ls: R supra labia Is. the .\'d to 5'h in contact wil h Iht: eye: 2 1 rows or do rsal sca les at mid -body: ve rt ebrals di slillcll y t:n largcd: :240-265 vt: ntral s: f\ n~ 1 SC lltt: t: nt ire: 13 7- 1:)4 subcalld:il s. divided. Dorsall y uniform brown in various shades. wilh li ne dark spcck lin!,!.: vt:rlebrally two longit udinal rows of"h laek spols. usua ll y eovt:ri ng the firsl sca le row ncxt to Ihc vertebrals: upper side or head oli ve: su pra labia Is while or orange rcd to rt:d. \. . ith the dark coloration of the head bcginning righ l abovc: tai I above sOlllcwhat dark er thnn tht: body: lower p<J rl s orhcad and subl ab ials whilt:. or ydlnwi sh to tb: p ye llow: vC lll cr rrlllll bchi nd Ihe tl1ro;ll 1 ~l inl
] 1]
( 'u lubndal'
Colubridae
ttl BOil ' ,
fi!!.317 .Juvcnile Buigll lIigl'ic"p., !"rom Thailand .
pin~ish , ventral seale~ wea~ly ~pec~lcd on thcir sides. bOlh coloraliolls inl 'n~iry towards the lail and continlle on Ihe lower ~ide llr Ihe lail ; underside orlai llip dal-k grey currcsponding 10 Ihc colour of Ihe specldes. In juvenile~ . Ihe colour or Ihe head i~ idcntical to that orthe hody (V()(01 I in prcp .). J to 5 eg!:!~ (41-45 x I 7-:~ I nll11 . ,,)6-:'\6 g). h<1tchl ings Illeasun.: 39()--DO Illlll (BL '11 \1\ 20(0).
Ero logylEt lwlo g~ ' inhabits tmpical ralll fore~b in both kl\~lalld and Ilwuntainou~ regions . It is oliell I
L>ist ril)llt ion /Joigll lIigril'(>I).1 is di~trihutcd frolll thc i'vtday I'c-
ninsl1i<1 ~o llth ll'ard~ Itl .lava .. ·,lIl1atra, and Rorneo . Mt. Kinabalu : Poring (I H)-Cui let:t ion).
1827
The genu:, ('a/all/l/ria i" nati,e to the region frolll M, arllllar in the northll'esl and Chin;1 in Ihe north. thro ugh Vietnam. Thailand and Mala y Peninsula. sout h 10 Scram. Sulawcs i. and llie rhilippille~ . Sma ll snakes: head wl:ak ly set oll'li'ul1l the body; budy round in ernss-section: tail ~hol1: eye distinct. Ros(ral rounded: prclhll1tals paired. rused \\'Ith the illternas;ll~ and in contact lI'ilh the su pralabial~ : parietal meets wilh thc I;bl supra labial: enlarged parapanctal present; 4-5 supra labials, the 2'''' and 3,,1 . .I"'. or 3'" alld 4'h touching Ihe eye: nasal single, sLirrounlill1g the no~tril: no IOI·eal. F1II1her pliolidlltic trait~ include 0- 1 preoelilar. I postocular. no anterior Icmporal scale. and 5 suhlahia ls. f)or~al scales in 13 r()w:, throughout body lenglh; SIll(l(\lh : 11(\ apical pit:-: ., again~1 the hand i1,)lding il (S 11 '1III 'J(, ( ' 11\(;11< I')()'») . ( 'cI1ain spcL!ics appL'ar tl\ be I;tirl y COl11lnun in some i'cgi()n~. while ()f()ther~ unlv a fcll' speci men s havc ever been found . Their dieis consist (\1' \\(lnns, in~l'ets and their lanae. possibly ~tlso ~ lnall li/.ards and their eggs . The ~nakes arc U\ Iparoll ~. with the egg~ being deposited in lear lilter. Filiy-ont: specie>. h;1\c heen dcserihed tll helong to this gc nus. 21 of\\hich OCClir on Ilorne(l including the I I spec ies or Mt. K.ina b.du .
.' 15
( 'olllhri da .
Key to the Bornea n species of Calamaria
•
],,1. or 2,,,1 and 3'". supralahials cntering orbit .... 2 and 4'" su pra labia ls cnte rin g orb it .......... .. . 7
13 AIlliost all vcnt rals withou t dark pi gmentati on . .................. __ ................ ............ ..... ........ ........... 14
] ,.1
ab~cn t ............................................... 3 • Prco<.:u lar present .......................................... ... 5
2 Preoc ul ar
:\ Sllp raoc ul ar and p o~ t oe ul a r fll!>cd into a slIlg le shield ..................................... ...... ..... g/,lIl'illi lll ll • Suprao 'ld ar dis tinct fro m posto<.: ul ar .. .... __ . ____ . 4 4 Mcnt al to u<.: hing. an terior <.: hin shi elds . ______ . !owi
• Mental not tO ll<.: hing anter ior chin !>hields ________ . ....................... ........ .... __ . __ .... . __ ........... __
schm idti
5 2 infral abia ls touching an terior <.: hin shields ...... ....... __ ........................ .... ................ ____ . hO/'II('('I/.,is
• 3 infra lahials to u<.: hing anterior chin
~ h ie ld s
.... Ii
6 Narrow longitudi na l stripes Ill id-uorsal ly; tai l short . tapcring abru ptl y at end ......... ha llers/H'i • No Illl)gitudi na l ~ t ripes mid-uorsa ll y: tail lo nger. ta pering in posterior ha l r ...... __ ........... me!lIIwta 7 /I light longitud ina l stri pe on ~ca1c row 2 and 3
boruered below by a uark str ipe on lirst row .... . ..... ....................... ..... ...__ .. __ ... ............... !/lm/w /t:;; • Stripes. ir pn:sent . not as above ....................... 8
8 I'reOl: ul ar absent .. __ ... __ .................. ____ . ______ . ______ .. 9 • Preoe lilar present ... ______ ............... .. . __ .............. 10
9 Most or the dor~a l ~ea ic s dark brown wi th a li gh t netwo rk or ye llOWis h wi th it dark network ........ .. .. .. __ . __ ... .............................. .............. rehel/ fischi
• liaell dorsa l scale above the 2,,1 row dark orown withollt a li ght network .. .. ..... __ . __ ..... s. sc"'e.~e/i 10 Mcntal not tou<.: hing the anterior chinshields ... II
• Mcntal touching the anteri or chinshields ...... 18 II Dorsulll wit h wi de dark transve rse crossbands of four or more ~ca l e~ in widt h ........... __ ..... hic% r • ()or~ lI lll lac king crossbands .. __ .... ... ................ 12 12 Ve nt ra l and dorsa l co loration dark brown wi th a bold. cont inuous. white lateral stripe ... ftltera/i.~ • Vent ra l co lorat ion dille rent. from dorsa l colorat ion .......... ............................. ....... ... ...... ........ ..... .. . 13
3 16
• All vell tra ls hchind the lirst qua rter o fbouy wit h darJ.. pigme ntati on at least on their latera l tips .. ..................................................... .. ..... ...... ...... 16 14 Eye large r thall ui sta n<.:c eye-mouth .. ....... ........ .. ....................................... ... .... .......... /('I/ ('ogtlster
•
I ~ye
slllalle r than di stall<.:e cyc-Illouth ........... 15
I S Prcf'rontal not touc hing .I,d ~ upral abia l .............. . . ...... .................... ........................... .... s. sc"'e.~e/i
• Prefront al
u ~ u al l y
touching
]'J
slipralabial .. .... ..
. ............ ..... .............. ................. ... ... ... ...... h;c()!or d o r~a l sca le rows on tail at 14 or more suocaudals from tip ...................... 1'\ '1'1'( ' 11; • Reduc ti on to 4 dorsa l sca ic rows on tail 13 or l es~ suheaud" b frOIll ti p .................. .. ............ 17
16 Reduction to 4
17 Tail enu illg in a blunl lip ...................
V;,:~ II/([tll
• Ta il taperi ng to a poi nt .. ...................... moc/est(1 I H !-Je ll y with dark eros 'bands, each mo re than one ven tra l s<.:a le wide ...... .. ...... ...... ... flllllhr icoilletl • !-Jelly not cross-banded or wit h ba nds narrower than one ve ntral ...... ..... ...... .. .. ............. ............ 19 19 I:.ye two- thirds o f eye- mouth d i!.tancc ........... .. ..
... ... ........... ... .. ..... ........ .. ....................... hillel/ illS; • Eye eq ual 10 or greater than cye-Illouth dislalll:e .......... ...... .... ......................................... ........... 20 20 Dorsum wit h longitud ina l dark and light stripes ..... ............ .. .......... ....... .......... ......... ..... gri.n vo/di • Ml d - d o r~a l reg ion not striped ........................ 2 1
2 1 Ta il thieJ.., tapering abruptl y ncar the end to a blunt tip ...... .............................. .. ......... ........ ..... jll'lI kke; • Tai l tapering graduall y ... ........ ....................... 22 22 Vent ra ls of males l es ~ than 145 in nlllll b",r; of females usuall y less than 104 ............... sl/fl/ ellsi.\' • Ven trals of males 145 or more in num ber. of fe males usuall y more tha n 164 ....... grllbolllskyi
<. 'n luhridm:
Tah,·W nl ~ lriblilillll of Ca/lI/lIaril/ . s p ccic~
Oil
Ilorne", Kinai1;ilu
Sahah ( 'I1/al/lflril/ !>al/l 'r ,I)'."i IN,al( & ( 'u/al/lurill biclJlor
1)1 ~II ~II , n IH~'"
( 'a/l/II/aria hOrl11" 'III'I,' 11111 ( 'a/al/lal'/a
( ' I 'c l vll l
M\I(~ .
1')(,5
( ' DI III
~II.
• •
11':5-1
~II( , 1 ~6()
BOI I I.M ;1 ' 1(. I W) J
~
Cu/al/luria ;:rablJII'skyi h " III K. I ~~5
i"" 11111l-
I N72 ) (lIlilll/tII'il' ;:ri.nvlI/tii L, ,I Ilw.,a. I ')JX ( 'a/l/lllaril/ grtwil/il/I/I (
Ca/II/llaria IIil/(,lIil/.li 1",,1
I( /\:.
+
• • • • • •
I
• • • • • •
l'vl IK\ . 1965
Ca/lIl1l/1ria lu(eruli,,' M 11(,<.''',1 1(1l. I N9() CU/lIl/1uria 1£,lIcoga,~t£'r Bill ~I~. 11':(,(1
-
Ca/II/i/l/r;1I /01\ '; 11',, '1 1"'.I~. I I':N7 ( 'II/lll/IlIria IlIlIIbricoitlL'l' 13,,11 I" 111111. I, :! 7 ( 'II/all/ar;a /1I111/W/t: 1
J\ r-..IH KS .... )~.
C"/l/II/l/r;" 1II..tIll/(}/l/ J 1'\,
192J
Cil/II/i/(/r;l/ /II'II!.!.";
!)l \ II' RII •
1 ~) -1
I. lIl1l1
CII/IIIIII,ria s. selllel:£'l; \)1 \11 ~II , ll llmo" &. J) II~ II KII. I N5-1 Cillilll/(,rill sdlll/iflti rvl {{t/{IIIII/rill slIllIl' IIS;S
II( \
8: I",,, Il- 1955
T \ \ I OR.
Culul/lar;a \'irglilata 11011
192~
'" 1\,," , I ~ 27
D l, \I(:RII., BI LlRON
' I~p c
localil) :
.
• • • • •
&
D l ·1I1 ER II.,
• • • - • • • • • • • • • • • • • • • • • •
~
-
K,ilmwl1 lan
~
• • • • •
+ I
•
• •
• • • •
ing Ihe anterior chill shidds: 11')- 1(,l) vcntrais; I X2X slI hcaudals ; tail Ihiek, ending in a blunt pninl.
Calalllllrill hic%r
( 'lIlrmlll"lll hItO/(H' D l \IHHI.
I •
1 ~6 2
13ll1k"" & DII'II KII . III' ,1,1 1111. I X9) ( '''/''II/l/r;(/ ,-,' /w I/III'dll i-lllll-.IH . I ~(,()
Ca/al/ltll'ia I/Iot/estll
• •
"a r~\\'~ k
Hn.lllci
I RS4
BIIH(I)' & Ot \ 11 k ll. I X5-ht 7X
(l Oll,.."
hll 0/0': \11 H 'C.I\I\KU I Xl)()a I L'-: IhK l I I II I Xt) ')(' 1'/1 7. 111 '> , 1",1"& 1 1~\ 196): 1·1'/; I') X~ 27, ~I " "7. 111\1 ~\)', 1')'/-111: 2<111
( 'U/""/(/"'O
X.j . :-'lI lllf ll(ll Il)()la : ('~ ;l lI R..""
,1.\",,"'
niaJ.:llllsis Third and f o urth ~ lIpra l ahia l ~ <:lI l erill g prbil : preoe lilar pre~e nl : ll1<:lllalnollouching anleriorehin sh ields: parapariclab surrounded by 5 sealc~ and shields; prefrontal touehin g 3,,1sllpralabi al : no dar" s qllare~ on v<:lIlral s: nasal s ori en laled fo rward: max . lolal kn g lh 45 CIll .
Lh'srripl ion
Colour ofadulls uniform dark hnm II ahtl\'e. or w ilh obscure bl ack hands slIp<: rilllpt1s<: d o n ground colollr: da rk pigillen iali llll ll';ualiy ending abruptl y on seak rows 2 or 3: ~calc~ or I ' and 2,..1 row~ sometimes wilh hlad. an leT-ior COrll..:rs, forming a conlinllous b laek str ip..: in ~()IllC specimens: head dark brown ahove , wi lh or wilhout IWl\ obliquc da rk
Fi g. ( ' X <. 'iI/' III/ e/l '1/I />1('''/''''
"'''r'';... ,.J'
.fI.t!A... .. ~
~'.
Fi ve slipral:1hials. Ihe ,I '" ilnd 4 '1 , enh.:ring orbit 5 the l a rg<:~ I . I " Ihe 'l1lallesl ; 5 ~ u b l a h ials , first 11ouch-
th
.11 7
halld~ "Lros~ Ihe yL lUll lip ; 1elltral s ye llo\\', 1I~1I ally Illlllla Clilale , rard~ ~Iwt l ed 1\lIh hlad.: (ali hl'llWn ab\l\e. \\ Itll d;lI~ hand,,; under"ide pj' ta li ydIUl\ . \\'Ith or I\itholll a !i;II'k llIedi ,II1 il II\.: , .lUI \.:nlk~ lla\e a yellnl\ groun d colour \\ ilh dark CIos~ball(I ",
, hic llb ; paraparic1al ~ IIITOIIIl (k:d hy 5 ~c;de~ alld ~h l eld~ : redllct lon to 1 i.)lIrdor~ ,tl ~cak nl\\ ' ~ at a pO in t 12 or more ~ubcauda l " I'rom the t<.:rminal ~c lll e : llIa\ , total kllgth 4(1 ,){ clll , l)csni plioll
Fi ve ~upralabia l ~ . .3'" and 4'" entering o rh it. (h ' 5' the l arge~l ; 5 ~ublabia l ~ . lir~t 3 touching the anlcrior c hill ~llIe l (b: I. II- I ')D \elllral~: 2()-_'> "lIbcaudak tad long. tapering graduall y to a blum tip. ( 'nlmll dar~ bnm n abm e: caeh lit)l'sal seale IIlth a d :lr~ IICI\\(lr~ ; ... cal1cred dark hn.l\\ II or hla 'k spO I" [i.lrIll ln g da"hes. PI' shorl l inl'~ 2 1'1 ({""I 11J1 7 1 "". :Vh ~ l lI l \ I') ~ l ~po t s an teriorl\;; \entra l... uni l()l'In yelkl\\ PI', lIlore _ 7, orten, II ilh varying amollnts llrbl
Dislrihulioll Thi~ 'pcc ic, IS (btnhilled 011 Ihe i, l alld~ pI' Jal'a and IlOrl1l'O , \11 , "jllabal .. : I\ltllllUt more precise Incalitle\.
or
1
or
or
F:c() l o~~/F:t holo!-!~'
The ~ pe ('ies I ~ preduminantly a rorest dwel ler. 11 inhabits the lowland a~ II e ll as montane regl(lI1~ lip to a lt iludes or abo ut I~O() metres. Further d<:la ils on its eco lngy arc unavai lable . Disl riulit ion
" en" ~
Thi~ "pecie~
CD
is c ild ellli <: 10 Bo rnell ,
\-11. I
a:
maiall . Tcnompnk Kcnnknk ( lOOn Ill),
.31~
Colubridne
Ca/al1l11ri{f J?risIVo/di Lm.' EI{I DGL,
1938
Ca/amarilllumhnl'oi,i£-u ,t},I'ISH'olt/1 I .O\'j RIIX, I • (l)3X: ~~ 1
Illca lit),: LlIlll:ln Rl\~r; lI"rnCll CD·HI
1IIII1dll
nlh~n : r\'101l111
k~l)
( ',,101llr/rio IUl1Ihri('o ir/ca g ri,,\,.oltli. I I I I-l
Colullluriagri"l1 'o/t/i. h.l il l( 27: ~hl "'I'" II)~S : III . 111'l ~a: 13:'. 11)1)-111: 2.17. IMoiK &. T IN 1'1')(,,,
_'_,'I:
Typl~
Kin:1b"llI : 'Jt.llh
\.\s
11.}_0° 570
& (\1 ,\1{'\ 19(15 : t)~ : ~'I \'ll1 n 1')X1 : II)X7 : 2~7_ I'IX'I : 1'17 _ 11)1)1:1 _ 1.1, 2-1(), I')')('L : 2'1 1. Sn I IIINI, 1')<)1 70 : <; f1IIiINt, 8< h ','1< 1'l'llJ : UII
Diagnosis Third and fo urth su pralabi ab c nt ering nr hit : prc()c ul ar prescnt : mental lo uc hing ankrior ch in shields; wlour blad:.ish bl"Own above with a narI-OW whitc line betw ecn sll cccss iv.: seak rows: imIl1aclIl
Ikscript ion I' ivc or si.\ s upralabia ls_ tit.: 3,,1 and .+,1, t.:n tcri ng th.: nrbil , 5'1, lite largt.:st: 5 sublabtab. firs t J touching tht.: antt.:rim c hin shic lds : both pai rs or c lt illshi.:l d ~ Ill Cc till !,!. at m id- linc: 155-19::! \entra ls: I.:I- I X
subeaudal s; tai l thi ck. taperin g rmm ha sc to a ~ha rp point. Co lour dark brown or black above ; dark portion or sca les wit hout network: hl ackish browil stripcs on th t: ccntral two thirds of each sca lc J"O\\ abo vt: the first. ye ll o\Vi~ h stripes on the t:dges Or ildjacelll scali.: rows; scal es ( II' Ih e lirst r(l\\· ye llow, Im macu late in Ihc anterior part 0[' th e body: t.:ac h sea lc usuall y with a dark spot in the posterior h:1II' or th c body: hcad dark brown above: sll pralabial ~ yellow in the Ill\\t.:r two third s: head below imm ac ul ate ye llow: all obl ique hal' runnin" fo rward alld lip I'rom th t.: gtll ar region 01110 Ih e rear part 0[' tilt: pal·ietal :.;: vt.: ntral s immaculate yc llow, "ometimcs with itTt.:glllilr ~ma l l black spot s; slIbca lida b ye llow. usua lly with a ['ai lH lig-zag line mid-ventrall y_ Ecolog y/ Et hnlnl!~' i\ species urthe suhmol1l anc and montane prima ry rnrcsb li vin g he neath kar lil t T anti ill rOiling trt.:e Sl lIlllpS _ 111 ~.:Irlld i: n ee th i~ s n ;l~e wi ll ~t in g with t t~ tai l tip _ ClI/wlll/ria gri\lllIidi i" a common victim ( rt hc road traflit.: ill the I \eildq uart ..-~ ,Irca K inabalu Park .
or
Fig. J 19 ( '''/CiI/III1'1l1 gri.l l m/di fmm the I k ~d4 I1a J'l.:r" .
3 19
Ta \tlllOlllir rOllllll l'llt : Sce la.\ \ Ilnillic C\lIl IIIll:1I1 IllIder // 1'1/1'11"/(1"('\ /woe/milla/is . Oi:l l! lIu ~ i~
I hlrd and I'tlllrlh ~u pra ':Illerilll! orhit : preoCiliaI' p r ',\:n1: m\:ntal nO! lou chlng antcriol ch ill "h icltk h\ld ~ dark bl'll\\ n d(lr~ally and \\:ntrally wilh a con llnu ous lateral whill: ~l l'Ipe O il Ill .: 2,..1 ,llId .\" "c;tie ro\\'s for the en llrl: knl!th ul'lllc h(ld~ alld 1;111 : llIa\ . itHal kngth 2<) l' m. labi:iI~
1';
--
Ocse ri pt i4111
E
(J)
.)
1) ; lr a p a riel al~ ~urr(l unded
.... ~-'"
by S ~cab, and shiel ds; (1l'eocu lar pn:s\:lll : S su pra lah la k Ihe.\ ·1 and ..j ll . e lllcr-
i\1 \1 I-- \It , ( I l)l)(,C) round an ad ult Illa le ' 1)('ell ll~1l
Disl rihul iun T hi s ~pee l l:~ I' endelll ic to BOI'lll:o , 1l. Kin a h a lu : 1'l:l1lllnpok , 13untlu Tu han ( 1.\70 Ill), Kamba rall!,!.llh Road (I :'i:'i()- 17"O 111). I iwago Irai l ( 1500-17"() In), St! . I ihahar, I leadLJuancrs (ISOO- ISSO Ill). I lIllHI LUlllu (1."2-) Ill). Sg, I iudan , \1l'silau (I XOO Ill) .
CII/lll11aria latera/is
I R90
\~ I I. IX')O.I 11(,. PI . fig 4 __I tI 1-..",.,h.IIII. iI,"".'o 1, .\ 1)t \11 11, IX(}Oh 1" 4 I ~pl' I Ol":lliI ~
( '411rIllhU' /tl lc u { roh , ~1. )(
I )",· I ,,~a l i l "
un)t .\RD,
ot
~ltll"l1
( 'OIIllIl(/I/(Ilo/c l'{ '/I \'
Vllu III I 1-..,""h"llI. 110",,'0 ( -( damort., /oll ·r, di, . H, It II :'\ 1d
H.
1Xt'~k
~-t 2.
n \k ll I I'
J20
ex I
(lIlt/IIIII/ 'llI 1,11, -1'111"
Ill)!. thc orbit . 5'1< the largesl ; S o.;ublabiab. lir"t .1 touchillg anll:rior ehi n sh iel ds: 140-151 v\:lIlrab; I ()-2] ~ ub\:aLldal ~; tail lap\:l'i ng graduall , to a point I'rom Ih~ \:Ild 0 I' Ih\: body. Colour abo\'e and belo\\ dark hlackish brown \I Ilh a I nn~l lud i ll a l whill: lin\: frol11 behind the C)C mn Ihe l:lllire kn)!.lh of body in ~ca k rows 2 and J.
:
Disl rihlllio ll I Xt) 'r
X.I, "I II II ItK il 1901a ('2. 1'1 R, ,," I'll' I/lX . ~ '11I11 1931 12 . IN,.1 R .IC 1\1\1<\ I'll» 116, ~h "' " 11\ II)X.' 2 7: ~"I " I\ "" & ~ '"' ''' N I'IXI 2~ .... " """, I <)9 I .1 .1<1. \I \I .. \(1 , l'I'I·lh. ~~(,
Fi!.:.
rh i~ ~pceles
has h':l:11 rc\:orlkd 1'1'0111 .lava a nd On UOnll:O it has become known so lilr only from 1\'11. Kinahaill . \\ililoul mor\: prcei !-.c H orn~ll .
Incalille~ .
Coluhridae
Ca/alllllria /ellcogllster I3 LEJ'KER, I R60 ( 'lIllIm(fn (lI( //I( 'o~("/( 'r B I II ~ 1 1f.
1N60h . 2, 5. 2XX &. 29)
~' ()('
ngno~is .
I",·alil.\ : Amp,1I I :I"allg. SlillIal,a I Xl) ...k . ,4 I : B \M I I 1- I I IX'i )c: X4: Sil l"""" 1'1111 :,- /Ie . III R""" 1')1 7. 1/>-1 , S 11111 1'1.' 1. 10. II. C~ . I ~,,," "- \1 \", 11)/15 . (" . Iig. Ill. ~1 \,1 111 \ 1')X 1. ~X : I' II""\R " 8: N I l)X4 24. S II 'II "'" I'NI ' ., -'I. \hl ' " , 1')<)41> 2-1 7
( '"Iulllarill
I ClI( Og (/ ,\ /t "',
H ill ' I I N( d
Eculo~y/ Ethology
This spt:cies is an illhabitanl or lowland rain forests. hll1hcr detai Is are un!..ll\1\\ II : ~ee gCllerie di -
I{
<;" .,"'
Uhlgnusis Third and fourth ~upralahia l ~ enter orhJl ; 111elltal 1I0t Illlll.:hing anlcrior c.:hin ~hlellb ; paraparietal surrounded b., 5 shie ld s and scales; no dark pigmentatioll on the \ elltrals: l11a\ . \lltal kngth 22.3 1.:111. I he on ly L30rI1ean species of CaIII li/ar/ll with black hall~rings on till': neck and 011 Ihe lail.
Distrihu tioll This spec.:ies occurs on the islands nr L3orI1en and Sumatra. with a doubtful record rrolll.l ~l\a (I N!,) \( & M AR\ I ()6:'i) . It. Kin ahalu : Kiau (91 5 m) . 1'cl1ol11pok .
Ca/alltaria /1I11tbricoidt!(1 H OI I
Preoe Lilar present : 5 sLipralabials. the .\',' and ~ ' " cn tering orbit. Ihc 5'" the largest. 2"" the next largcst: 5 sublabi als. rir~t .1 touc hing the anterior chin sh ields: 126-157 ventrals: 12-2(, subc.:auda ls: tai l tapcring [ 0 a sharp point ncar tip. Ciround c.:n loLir dark hr(l\\ 11. dor:--al sc.:a lcs usually with a light or dark network ; body with or withoul dark sl ripes, the striped paltern eons isting ofa nar1"(1\\ dark stripc along Ihc cdge:-- uflhe \cr!ebral and paraverlebral seak rO\\·:-.. a wider stripc on thl' adjacent hahes of the ~ " and 5' h scale rnw~ . a stripc along thc adjac.:cnl hah es or the 2'" and ]''' seale row~, and a Ihin Jar\... linl' along the upper l'dgl' of' the I I scale ro\\': IOller tl\o-Ihirds or the 1" ~calc row cre:lIlI -c.:oloured : a hlac\... collar llf 2 to.5 ~ caks In II idth, ~cparalcd i'rolll parietals hy I tl) 3 ~c:llc~ :lnd sOllletillle" hordered po:-.lcnorl), by a naITll\\ yellOl\ ring; a dar!.. ring around the hod; at thc hasl' oi' the tall , usually :l sec.:olld hlac.:\... nng Ileal' its lip: hc:,d dark brown Illid-dor~ally with a ligh t streak running throug h the cl'ntre or the prei'rontal. the lateral edge (II' the slIpra(lclilar. and the lateral POr!illll(lf Ihe panetal : a dark IOl1gitudlllal st ripe along the adjaccnt portion" ni' the prerrllntals alld slipralahi:IIs. cOll i IIlUI Ilg hehilld eye alol1g Ihe adjacenl l'dgcs nr the parietal and supr:llabial scale,, ; rem:lInder or Lipper lip alld lIlI(h:rsidc or hcad crealll-l'lll,lllrcd ; ventral s and subertudals l'i'l'am-cIl loured : tail 1\ ilh a dar!.. lI,id- \elltral stripc ill ~() 1I1 C illdi\ idllal" .
H Olt.
I X27
('olall/(l l';(/ IlImhr i nuc!l'tI II. IleII I 111 I . BI Iii
luca lily: Dl'script ion
in
.
1X27 ~~HI
' I ~ P('
Ja\ a
( 'I1 /a lll arw (( '''"I/'''(,~II. \11
I ~(J() . I
1", 6
( 'II /all/ a r m \ ," I'IJIl /ur", n , Si ll I 11IIUI 1~)Ola
(, I,
H. <,JII\ I(lI
IH
Root 19 17'
1'i-l , <; \11111 1'1.1 1111, 11 27 . 32 Clt!a U/a r w IlImhn nlll/cli I w uhn ('ou/(,lI, L. )\-1 KIIU,l
Il)
\X.
-u
Cola mario IlImhric(lidc,/, h(,11{ & \1\ .1<\ 1" (, ", 7". fi ~ :!o. & S IO' " " l'I X·1 e ·l . S,' """ 1'1' ) I 140 . 1-_ 11' Nil M \I .. , It ~ I 'I'I-th. ~· 1 7
,"I)
{ '" luJl/ono \ '('nllll/Jnlll,~
\ '("'I11I/l1nll" ,
f\1 '\,1 11t \ 19 X~ ~ X
Dia gnusis
Third and fourth ~upralahial~ eiliering orhit ; preoc.:L1iar presel1! ; n1ental IOllc.:hing anterior chin shields; paraparietal surrounded by ~ or 5 !->ea les and shields: bel ly ye llow \\ ith hlae!.. ern:-.sbar~ wider than the \\'idth or olle ventral; rna\ . Iota I length ('~ . 2 cm. Dl'scr iption Paraparid
This snake is found in hill y areas up to ailltude:-. oj' ahollt I~'\() III ill hllliand tropic,J I(n'ests and trop,eal
321
CnlubJ'ldac
Fit!. :'1 I
( ', IIlIlIIlIrlil 1111111"'/(11111,'1/
frolll SIIII1,lIra.
and ~uhlropil:al IlHlJlI
Thi ~ S pC C I C~
322
Calamaria modes/a &
I)l IMI· i{II. B IIli{()'\J ( "ltIIJWlltlIIJtld" .,/a I) , MI RII.
I) I'"
I XS4
II l1m"N ,\. 1)1 \111111. I X'io1a : 7~
lural;1.\' : .l a,,,
("II/umana 111 0.14."/0, ' UI ' ll\lUI
\1
D Ul\ tt',KII,
\I 1-,,,1 '
& t\., :\1{ \ I (j(l) ' I JX. fig ..HI : Iqx.+ . 2-L . IIIBI " I, 199 1 3 40 .
'Nc ,I k
& :'-,IIHdO 1'I'I4h. ~-I 7
N
(alamtll'tulmll('/lM\, ~·I \ ' I III '
I'n(t : 2~
f>ia :,: nosis Third and j'ourlh sli pralabial s enlering. orhil: preoe ular prese lll : II1clllal Iwl touching anterior chin ,hlclds: nasals orienlaled laterally : no lighl stripe on sca le rows 2 alld 3: ve nlr,li s with dark pigllleniatillil. al lea sl l a l~ ra ll y: no ob li que dark
Cl1 lubridac
stripe behind the eye ; tail taperi ng to a point: max. tOlal length 45.5 cm .
sca le row without dark pigment ; n a~ab orient ated latera lly; ma x. lotal k:ngth 46 Cill .
Desc ription Paraparidal slilTolinded by 5 or (, shie lds and ' calc~; 5 ~lIpra l ahiab . the 3hl and 4'h enteri ng orhit . the 5'h the largest : 5 $ublabi als. firs t 3 touching thl: an1cri or chi n shil:fd s; 140- 154 ve ntral s: 12- 1<) slIbeaudals; tai l ta perin g to a point ; (ventral a nd sl.Ihcalidal coun ts for HOll1ea n ro rlil a tion ~ only) . L30dy and tail dark brown or black above; sca les with a light network and an occasiona l dark Ion ·itudinal ccntral streak . or sca les so lid brown except for a li ghtlll.: twork at the edges. or scales so lid black with sma ll ye ll ow 'pOb scattered along body: In some spec imens the dark centra l streaks unite to fo rm short lines; sca les o f 1'1 rolV with or wilhout light et:ntres: head blae k abo ve with yellow spot '. or yell ow ish brown wilh blac k spnb; supra labi a Is spolled with dark; underside ofhl:ad ye llow ish with dark spots ; ven tral co lorati on hi ghl y \'ariable ; amount of da rk pi!,!ment on \ eilirais \ aril:~ from confinl:d to the lat eral edges to cmeri ng the entire width o f the \entral~ : the light pigment is confined to a naITO\V medi an ·tripe or to a senes or small semicircul ar SpOb .
Desc ription Paraparil:t al surro un(kd by 5 or (, s hicld ~ and sca les: preoeul::lr present or absent: 5 supralabi::ll s. the 3,,1 and 4,1, entering orbit . the 5'" thl: largest: 5 sublabia Is. first 3 tOllching the ant erior chin shields : 129-1 liO ve ntra Is; 19-44 subcaudals: tail long. ta pering throughou t it s length to a sli ght Iy blunt point. Co lour dark above. li ght below; sca les above I" or 2"J rows withoul network. dark hrown or black : head yellow abovc and below or bl ac k above and yell ow be low. or various Illt enl1edial e cOl1dili ons; ve ntrals and first one or two ,ca lc rows immaculate ye ll ow.
Uisl.-ibllti on This species occu rs on Java . Simalue. and Borneo. :vlt. Kinahalu : Bllndll Ilihan ( 1370 Ill). knOlll lwk ( 1·.\30 Ill) .
[c olo gy/ [ lholo :,( ~:
i, round 111 hilly area~ up to about 1000 III ;dt itude. and Inhabib dlll.:ll y tropical wet fore:-.ts and tropi ca l wei mllntane forests; it pro babl y ai,\) \lCl: ur~ in plantat ion, and t.:1I 1t ivaled areas . It i:-. a terre~trial or ,elllir()~sona i. ~ecrelive ~ nak e. mainl y at.:live at l1ight and onen tlllilld among forest liller or under rllcks. t;tll en lo g~. and dumps nf dccay ing \egel ati on . It !'> diet IS Co III posed lI1a1l11y of worms. ~ I\l g~. ill ~l:c t larval.:. and 11 al:-.o takes ~ l11a II frogs . ('a /u lI/a/';a s. sch/('g(' /;
Disldbul ion The nOl11inate subspecies ran ge~ fro lll the Malay P e nin~lIla In Hortlell alld Sun1alra. 1\11. h:in abalu : withou t pre ' i~c loca lit y dal a.
C"lalllaria sdtlegeli .\·cltlegeli & DI IMER II. , 1R54
DlJMf:RIL , BIHRU ( 'lI/fI/IIII/W \,
Me,!!.""
T.I IH' IU(,lI lily:
&.
f)[ \ 111<11 •
M AR:>'
(1tl'11
( 'a/omario hi('%
D Ilf\lIl< lI . 1 ~5-l). y lt .t<.it 1\ 1{ 1)
l I ) l,f\UI-
('(//0 111111'1 11
1')') 1
1~
&
1'J(; 1 R.
I tJ55
• H IIUl( IN
I X9Ua : I .~ _: " 1\11111 P).l l : J~ l ')l,X -n
( 'a/all/una '.' h'lInwj '"hcda. LO\ I KIIH,) ( 'a/o own(l
Calall1aria Scllll1idti
I Xqa' X I
1l1111l~(l
( 'a/OI llaria hic%r
&.
1)[ \11 «11 • IlI ItR' IN
., ddc',t!,.elt ,\c 'ldc ''..!c''' . I N lo! K .~ \,1\1(\ I t)(») : 1h :! ,.'d«',t!.('!t. h u N!I\101 & ~I()(.I
HINt,
J)ia gnosis Third ::lnti fourth ~ upralahial~ ellt ering orbit ; melltal nol touchin g anterior ehin ,hield,; di stim:tl y bico lo ured ; dark dOI'S::l 1 p igme ntati on en din g :lbrup tly abnve Ihl: I" seak row : \'ell tral ~ and I"
c. -lIlamwtcl ,",'11111/(/11 ~ l \H\ & I'C d K. Il) ~.'- Il)7: lit! . 27
I ~ I U.·
luca lil ,·: Bilildu
Ilihall . ~ t llUlli I-..11l"h.tili. ',H'lh I hlrn~" (el/'"I1I1/II/ ,1'11111/(/11 h,,," ,I;c ~ t \"\ 1'1('" 71. llg. I. ~ t. \NI III' I t)K1 :::!X. I iii Nit \WIJ (\ S'! X. IH"" I')X ,I ~ ..t f\h lt-..M I I' IC)Xl)
1'17 1'J').1h ~lK. 2.17. 1'1'I.ltI III l'I')(1l' 12 7. :-'111111 "'. 194 1- J... l. \:1 \N IIII"I &.: (' KII'SM\ .... ' jlJ97 ~ll I)i a~ n os i s
Four , "pralabial s. Ihe ~", I and .1,,1 cilt eri llg orbit : l11a x. total len gth 2X CI11 .
( 'o ilibrida~
aJ'lHllid () . (JO p.m. It li as busy em"sing a road that cut through a ~ ubmnnt a nl' Fa gacea primar l'l)('(':sl wilh vc ry ~ tr\ll'lUred terra in . ;\nnther ~J1<.:cimcn \Vas fi)tllld in Ihe \ icinity nrthc Ilcad quartcr~ oj' Kiliaha ltl Park, and yet anot her 1111~ was SC~1l cros~ illg a road at 11.00 p.m . (VIII (0.. \ll oS 19,)6e) . Onc ~pel'illlell had an <.:arthwUI'm ill it;, sl(ll11a ch. another i'ed \111 ~~Inh\\'orms in captil it y. /)istribuliull
Thi s s pecie;, IS ~ I n <.:ndemiie or MI. Klnabalu . :\'It. Kin a halu : Bundu Tuhan (1370 Ill) . Ill.:adquarlers ( 1550111). Kamborangnh Rllad ( 15()()-1 (,00 mi .
Fil!' .H2 ((//(/II/(/I'i(/ w'hllli,IIi frol1l Ihe I k a dqllar\ c'r~ .
Desuiptioll
and ~~ak~; pr~n':lIlar a h~': III : .\ ~lIpralahials , Ih~ 2" and 3,,1~n I~rill g orbil . th ~ .\,1, th~ Illn g ~ ::. I , rarely th~ .\,1. entu s Ihl' orbit, 100 (1'v1 \I (o..'H.., 1l)L)-Id); 'i "ublahiak thL: lirsl .3 tOliLhing th~
~urf(lund~d
by (,
~ 1l((: ld~
ell/alllllria slI/lI ellsi.\' T 'WI
OR,
1922
I " 'cHe Il)~ ~ IXl) I') IH.' luca lit y: (agayan Sulu . p h,IIpl'"'" ( 'fdlllllflfttl/'1 '1111/('hlin S \ II I I1 . 1931 : 27 I ~ P l' JU l' a li t): ..... I.ltl . I\lI K,"ahalu ( ' )11 1Ilkr h <'( I( 'x \1\1< , 1')(" I ~ ~I ( u/umaria \Ulll l/ I/,i " I ' td k & 1\.1 \ I~\ I ~)(l'" I .::!J . fig. ..) I : I H I N il \ I{ II & '" It I t K I \. I 9 ~ ..I 24 "i I II, U I 'd, I t) l} I ' , "I I :
((l/e/Hlor;a \IIII/t' '' \ /\
!,"",J,
I lIIo.M" 1')').lh 2·17 ( '" lclllllll '/Cl / Jt
IId/c '/'tOT I \1 ,'\
III '
Il}:\~ ' 2~
Oia g nosis
I hiI'd a nd fourth slipralahials entering orhil : prellcu lar pr<.:~~nl: l11ental touching anterior chin . Im:kb: parapari ctal ~lIlTt)llll(kd by 5 shidds and ~eJle:-; cal'h dorsal "ca ll.: I,;,ilh a li gh t nelwork : a C(1l1lil1uou~ li ght ~ tripc on the se ak~ orlh(.; first row (1111 , : no dark-edged o;addks
E culu :.! ~·· / E thoIH g)
-I his , pCCles kan;" li s hiding place (1nly "t nighl or
.12.\
Dcsc riptinn
Parapa I'll: Ia I surrollnded by 5 ,hi~ld~ and sca les: 5 ~lIpralahials. Ih e ) ,,1 and -Ith entering orbit. Ihe )'1, Ihe larg.est: 5 suh lahia ls. lirst 3 IOllchlllg the antcrior ch in ~hleld,,: 121)- 1M, v<.:ntrals; 16-2 6 "lIbcaudals: tai l tapering graduall v tll a hlunt lip .
Colubridae
Culour brown abovc. each dm;.al scale above the I" row with a fine dark network; ;,,(;altered s(;ales with dark (;enlra l sputs forming dashes or ~horl lines: first 2 or .3 sca le row;. white in Ihe anterior poni()n ; scales or I" row with light centres. forming a continuous white ~tripc right to the end of"the body: head dark brown abo\ e with sGlttered dark spots: a dark stripe along the upper edges or the supra labia Is. remainder or ;.upralabia!:, ydlowish. wi th or without dark SlIlures : head yellow. \\ ith sma ll dark spots un th . lab ia l :-,uture:-,; ventral~ U~lI all y uniform ye llow with dark lateral (;llnlers. som(; specilllcJl~ with ,mall dark spots in a broad series down Ihe (;entre of the belly: under~ide or tail ye llow. with or withollt a dark median line . Uist ribution This specic~ i:-, cndemi(; tll Ilorneo and C3Qav::1Il SlIlu Island . .vlt. Kinahalu : Sayap. Kiall (')1 5 1l1). Kenokok ( 1000 m). 811ndu Tuhan (1370 m). Tcnompok ( 1430 Ill) . l leadquarters.
Ca/alllaria I'irg ll/alll [Jot! ( '(1 10111111 111 \ (rJ,!.u /tJlfI
ca tit~· :
til H OII· .
II n, ul in F H" II . I X27 - 5-Hl
I X27 I ~I)C'
10-
.1 ;,,<1
( 'u/llllUlnOI 'il ).!u/(I(u , I IU III IlI .111 1111
IXt)Oc: 2:'i.. J .
I N4, 1 1( &
[\1 \1<\
1<)6" : IXll . li g 52: LIII " II \KD & S III\,KI t.. Ji)}i·L 2 L S I1 IIUNl, 1')<)1 : .1 -1t : /-h, ~"l" I'I'/·Ih. 217 :-" ." I"" K & \ '1\K\ ( I ')1" I X7) .,h.'lil lit . n," i'l,,,ol" h,'I\\ ,','n fi nd ( ' 1JI1J(!t.·\la. and 1111.::-') IhUtYIIlY \\' lIh or II h):-. 1 " P t.'I.' lIlh..' l h fC h...-rn..·J 10 a:-. ( . \H'gU/UIlI III
( 'a /aU/ to W \ II'g Il /alo ( ' III fU l n1i1
the
5"1. Ihe largcst: 21k1 the ~ec()nd large~t : 5 ~ublabial ~. first 3 touching Ihc anterior chin .;h lelds; I (Jr)-260 ventra ls ; X-30 sub(;
O(;(;llrS on Sum:!lra . Nias. Ihe Riall ;\r(;hipelago. LJornell . .Ia\
11"..' 1:1I111t.: "'111 ("-,,' B O lli ' •• 1 J( (I ~N-k) .
('a/amana hrtIC /n 'lII'o !ifu 11 ,'( ,1 H: . IX t)1h ..lXI it~· : MUIIIlI KlI1abalu , '.>l1h Il" rn ~"h y ll
"r~' lll' Illfal -
.1lkr h ,,," & ~h "\
19(.) : I X!»
Gellus Cllf}'sope/l'tI SCI II
( 'a/lillI/trW /'ruc IIntra, S ~ II I1 '
l'ill
1.( II
I, I X26
~I II I I ( ' HI I
. 1: tvh K\ 8:
It)! ~ 1a (,2: III 1{4.H IlJ I C) 17. I ()7. 11-, .1 K 1')5'1 192: \1 \NI III·, I')SI. 27
\)iagnosis Third and i'ourtlt ~upralabial~ entering orhit : preocular present : mental touching. or nOI lOu(;hing. the anterior chill ~hield s ; parapandal surrounded by 5 til' () ~hieiLk max . tOlal length 37 .4 (;111 . J)csuipf ion J>arapariet31 surrounded by 5 or (, ~hields and scaks: 5 supra l ahia l ~ . Ihe 3'J and 4'" (;ntl'ring the orbit . the
I'ile distribution or the genus ( ·/IIT.m"dC'll exlcllds rrom Indi a and Sri Lanka In tile west . :-;outhern Cililla ill the nurth . to Ihe Philippines. Sul;l\\('si. Timor. ;\ru , and the Moluccan bland~ ill Ihe ea,1 and south . Small lu 1I1odcralely large. elegant: hody ~lcllder. tail long, head sli ghlly depn.:ssed and distinct from body. wilillarge eyes and round pupils; dorsal sca les \\ ilh api(;al pits; velllra),; keeled and not(;hed . the last olle Lbually divided ; anal ~culc and Mlbcaudal,. al;.o di\ ided .
.1 2:'i
Colubmlac
I he ~nakcs or thc :!c nll " Chrl'.\0l'e/('II arc " ridsl; co lubml s" (I.l:. \\ ith an o pl ~log l y ph o u s dcntition) , hilt th ~ir \-cnOIll docs 1I"lIa ll y not ca usc scvcre er. fccls in hUlllan~ . The ve nlral and ~u h ca udal ~ca lc s arc kecled late r.. ally, ihu ~ cnab ling the , nakcs to clilllb with grea t agilit y o n Vl: l1lca l tre~ trunk s. Somc specics ar~ ab le to " Il y", i.e. th.: can ta~c III a d e~cc ndin ~ (para .. )gliding Ili ght \\ ith the rrbs spread widcly "0 that the LllHkrsidc I~ shaped to work like a parachute. Tire "PCCIC" ur thi s gelllis arc exclusively diurnal and arboreal. The " i"l;ed on lizards. frog~. slllall mamilla Is, hlrds, and s nakes. They arc ()\'iparo lls. Altogether 5 species arc currc ntl y recognized. 2 or wh ich occu r on Horneo, and both or the latt er are also pn.:se nt on MI. KII1
Key to Ihe Bornean species of Ch'J',mpelea Be ll y smoky grey between the keel. , ~ lrarpl y contrasting with the yellow or whi te Ilank s. mid .. dorsal region red 01' orange with pai rs of narrow hlack band~ \\ hich each frame a ye llow onc; I X120 I vent ral s .............................. ... ....... .. ..... pelill.\· • Bell y yc llowish green: a Inngitudrnal rllW or dorsa l spot s which eOvcr 4 ~ca l es each may be pre~cnt: sca l e~ otherwise bla ck wi th a li glrt green or ye llowi. h green. point ed central ~ poIS: a banded pattern on ly in jU\eniles; 19X-2.l9 ve ntrals .. ..... .. ... .. .. .... ... ..... ..... .... ..... ... ......... .. . pUrlllI;s; pllrlll/;s;
Sab:.h
Cltry.\·ofJe/l'tI IIII/'llt/;,,; fllIl'll/I;s; II, Ill ' ' " HI "' . Cltry,\'/I{ll'Iea {le/;/I.\' t 1., ,,\1,\1, ' . 17:'iX)
I X~ 7
Chrysopelea paradisi paradi..,; 130 1J :
( '/IIT,\ "lh/('O
"H'atil~' :
p urtU" ' 1 II
in
130 11 :.
B t lll
111
I
1827 BI )lI.
IX 27- 547
'I'~' IJl'
J.""
('hn ',\,,/w!C 'tt pe l/deft \t I Ju r arh " . (\It I
KI I" '" Il)(l X
20 2
• •
I
Ktnahalll
I3rune.
•
•
•
•
Sar;lwak
• •
Kalil1lantan
• •
cae h " tlmver" consisting of 4 deep red or orangecol (lured scales. but ~ lI c h patt ern Illay also be ab~e nt ; upper Side or head blackish gree n wilh ~ev c r:11 light or reddi sh hands: vel1lra l si de ye llowish g reen : vent ra ls typi ca ll y with black ma rgi ns (M"R,I \IS 1%)<) .
A clutch cont ai nin g lIJltll R whi te eggs . .15-40 In Illill (LI'vI & LII I <)Xl») .
J)iagllo~is
/\ ~lendcr , co lou rful trcc s nake: do rsal sca le s wea kly ~ eele d . with a pi ca l pit s: I !)X-:~J') vent ral s. Ihe laslllrwhich di vided : ma'\ . total length 120 Cill. hut most ~pccimen~ arc only het~\ecn nO al1d XO el11 long . Dcscl'ipliull One loreal : I preoc ul ar; 2 p Istoculars; occasionally the Inreal and the prcJ'runlal shields arc fu sed; X- IO slIpralabia ls. the 4'" to (,II" or the 5,10 and 6'10 . ill eontae l With the eye: 17 rows of dorsals at midbody. ~ ('e l ('d, with apica l pits: 1')i<-2J9 ve l1l ra ls, the las t one di Vided : li nal scu te di vi ded; I ()(1-1 4<) ~ ubea lld a l s. div ided. Hl acki~h green above; dorsals black with green spots; u~ u a ll y with a Ilmn:r-l ike pattern on the back.
326
Emlng~' / I<: I hl,logy
/)(I/'(I{Ii.l'i /wradi.l'i i~ found from ~ea level to elevations above 1300 nl. It inlr"h i t~ tropi cal wet forests. especially thei r clea rings and edges. tropie:11 dry forests, wei and dry Illontallc li)rcsts. ma ngrove rorests, opcn bushy areas. plantat ions. and c ult ivatcd arca~. It is also often enco untered in gardens and dose to human habitations. rhi~ species is arboreal, entirc ly diurnal , and ve ry active . It i~ :111 o ut standing c1lmher. li vin).!. ill tree lil li:1gc, buslll;s. and ha sica ll y allY type o rl all alld thi ck Vl;~ etali on. where it pl'\)g res~es or glides with incredibl y f< lst, elegant In ovell1ent s. This species fce d ~ chiell y on l iza rd~ . cspecia ll y agami ds and gekkoni d ~. but al. n o n fro gs. "lI1all lI1al11l11:1 l ~ includill g bats, and birds.
CIIITS(}/Je//:'{/
Colubridae
Fi:::. 323 Chrl"slJpe /ell paradisi /illradi.li from I hailand .
ChIT,I'IIIW/C(I I ), 1}(1I'lItiisi i~ r;tther aggn:ss ivc. stri\"·
ing and biting viciously when hand led , A tl ea~ t nne case of mild enveno mati o n ha:; neen reporh.:d ,
g rey area along th..: mid· line of the ve ntral s heing p;trticularl y di ag nosti c. This specics is ( vi parnus.
Oi slribuliun Thi s subspeeic!'> is nati ve to the region bdween Mya nmar a nd Tha iland in th e north and Ja va. Slimatra. and Uo rneo in the south , :vlt. Kill ahalu : Poring (1IQ·('o llcL:tion) ,
C/llysopelea pelias
(LII' AUI~. 175R)
( '''/11/'<'1' I,eliel ' L" , ,, I -<. 17.<'X : ~ ~ ·I Type Ifll' a l il~: " In llIll"," CII1:1\rJ/,e/ell l'/III ,\( w hlo rlJ. (\·1 \I ~"" 1' I C) X~ 10
Chll', ,"/,,'/"" 1"'/"" , ~hl " ' " ' 1')~ 7 ' '2',1.7. I'N·H,. 2-1 7. 199(,, : 2'11.
SIII ' BIN,.
1')') 1 .1·12
L>ia ~ n os i s
Small , ~ lcnder. 11\ a x , lengt h ;tbout 75 em . but us u· ally less than 65 em;, I X I -:~O l velltra ls. lhe la st om: usu;t ll y di vi ded; mostly easily iden li fied on the basis or its coloration with the sharpl de marca ted
.127
(SII.li'ol (,I R 19221: 17 r(l\\,~ 01" dnr~als al Illid-bod y: I X 1-20 I velltrals. the last (Inc: onen divid 'd : allal sc utc dividcd: X)- 120 su beautlal s. di\ i(h;d . Uppt: r ,ide or head \\ ith 2 or.1 red tra n~ vcr~e bars: dors ulll dcep rl'd to llrall ge red with d ark-c d ~e d . li g hl (i .e . whill:. blUi s h whitc. dirty II hilc. or ye llowi s h hl"llwn) tranwerse llIle ~ which do not rea ch down tll louch Ihe vClllr~tI, and which become more oh,cure po,tainrl y: sides ufblllly bnl\\ ni sh to br(lIvllish grey \1 IIh ~mall. whili,h hlue or ye llowish spnb: chin and thrllat ye llowish white to greyish white: \·enlral si de olive grey to ~mo"t: -g re y ht:twt:t:n Ihe line blac" \ cntral "eels. s harpl y delimited a gaill ~ t thc lel1loll-ye llow , tripe Ll ll the late ra l edge;, of Ihe \'t:I1Ir:tI>. (1'v11 R II 'i'o I<)hX).
r
e
u
I::cul()g~'/ l::t huICl:"'Y
(J)
( '/11 :\".\111'1'/"11 {IC/ills inhabil~ IOI\·land tropical v,el Fig . .ns a. ]26 ( '/II '!'wlw{m II<"itll.\ I"WIll Poring .
Dc~cript inn One lorea l: I pre 1I':\llar: 2 flo~t oellla r~: 9 sUflralabials. the -t tl. and 5' , or the -t il' tn 6 'h • 111 uJlltacl \\ ilh Iht: eye: 10- 1 I sliblabial s: _ I 2 or 2- 3 tempora ls
Fig. C"X II Chn 'Wll w{('1I I'<,/i(/.\ I"rolll Sg. Kipllngll I.
lore~t s and plant atioll>. lip tn hOO III allilnde . Thi s sllakc is arbl real. diurnal. and ac:li\l; amOllg ruli age and bu~hes. but also un the gro und in Ihid vegc!atioll . It reed~ chiell ;.- on li /.anls and rrogs . I \I KI\II S (I <)R 7 ) found
Dist rihution ClIITso/ >c/ell pc/illS is dislributed from M;ilay Pe-
ninsu la tn Borneo. Suma tra , and Java . Although Wide spread It is conS idered rare thr0ughollt ils range. Mt. Kin ahalu : above the Kiplingil waler/;111 (600 Ill) ; Sg. NIa 111 lit . Pori ng (500 m).
12X
Genus Dellt!re/aplris 13m
1.1- (iF-I{,
I X<)O
rhe gel1us D elltlre/a/l lti., inhahib large parl ~ o f Asia and Australia . Moderately large. vel)' s lel1der: tail long: head elo /lga ted. di ~ tin c t from the body: eye~ large. pupil s round : 13- 15 dorsa l scali.: rows at mid-body. ob li que. imbricate: vertebra ls enlarged: \entrab kccled laleral ly. nOlched: a/lal scute diVIded: ~lIbcaudals in pairs, keeled . The diurnal snakes arc arboreal and call be roulld in primary f"orcsls a~ wel l a, 111 cu lti\·ah.:d area~ . They feed primari ly on amphIbians and li/ards . Al l species are o\'iparou:-. AIJllll l 20 specie, arc knO\-\ n. of II hich 4 li\'l' 0 11 130r/1(:0 including 2 on NIl. K inaha lu. Taxol1omil: comment: SII IIll"(; & 1:--< 011< ( i')t)(») pn.:sent under the name of {)(,lIilre/( III/iisjimlloslIs two ph () to l.!.ra ph ~ \\ hich undoubtedl y show Ikl/(lrL'I(I/I/ti, CI 'lIl1oclt/llri, . Irthe photogra phed ~ peCillll'/I
J)"ml l'dllph;\ ra uJ,,/iIU'flW\ nllIlJ" /;' H'IH" .\ (
t K"
III (. t-' \y &.
II
(,I. II. IXJ -I 1111 IX-'~b-IIi .; 5 J: 1'1 XI 'I) Ill' 1 ()"' a li1 ~ : \Ill( g l\ ...·Il. Sub :--l' 4lh..' ntly d~ ""I~ ILJkd d ~ "M:olil\l"iI, Iblhll1"I " by \11, '1 & II, ,,,,,, (I'n~ 2X I)
/1/(/( '11111" , (//U/"/1I1i'1i1t1
Ihtl( ' lul/a ('ow/v /tf/. 'flliI ,
I
II I ' " \ I
I
t
S II II
Key to the Bornean species of Delldre/uphis 13 dorsa l scale rows at mid-body ....................... . ......... ,..................
IClNl,
& '" Ii II ,~ I '" I 9X-l ' 2-1.
Oiagll.,si~
Rcadi ly identiii ed hy th e cOlllbi nat ion or a ~ I riped pallcr1I il nd th l' imbricate arrangeme nt of ~ea l e,. 13 d(lrsal sca le row~ at lIlid -blldy: \'crll:bra l, cnlarged. A rl'eord kngth of I XI) elll is known. but 140 C111 i, a normal adult s i7.e.
C{//U/O/illt'lIfIlS C([IIl/o/ill eu llI ,~
• 15 dorsa l scale rows
<11
mid-body ...................... 2
2 A light latt.:ral streak. hordcred abovl' and belol\, by blaek ~ treaks ..................... pictlls piCfII,\ • No latt:ra l streak .............................. ................... 3 3 More tilall 170 ve nt rals: l')'e very large: the anlerior part of thc body red .............................. !()rlllll.\l/\· • I cs Ihan 170 \,elltrals; eyt: nlo(ll'ratel y lart!c: Ihc
anlerior part of the body yel low ........... strial/ls
\RI)\\ h hoi.
Dellt/fe/aphis callt!o/i"eaflls ('ulu/o/illeatlls (GR ,\)'. 1834)
/)cndreltlphi\ ('t lltdo/tJlC'cIlU\. i tJ\I IOOl 1<)91 : \-12 . h ' ~." s I'l') III 2-1 7
callle III fac t from Borneo. /J (' I'(lIIIICh/II/'is wlluld be eoniirmed as a nother spec Ies for Ihe i~land . Unillrtullatcly no localitv dat a lias disclost.:d for the ph otographs .
1x q
I
Sanah
f\11l.th.llll
BnUl""
"lr;t\Vil~
K .l ll1ll3man
• • •
•
• • •
•
• •
•
• •
• •
Ih'sniption SnOlll hllllll: I pn:oclI lar: 2 po~ton!lal'~: I narrow lorcal : 212 tcmporals: I) (rarcly 10) '"pralahials. the <;'" ilnd (,I". l1lorl' rarl'i) thl' 4'" to () n Illllchlng Ihe l'yl': dur~al ~ largc. ob liqucly arranged in ollly IJ row, : \'cl1ebra ls cnlargl'd: 16.\ - 1()() \'<:/1lra l ~ . ~.:clcd and notched : anal SClIll' di\ Idcd: I) 1- 170 sllbcauda ls, div id.:d . I3l'Owni~h or greyi~h ye ll (lw d()rsall y, wi th hlack siripes cOlllposed of Ihl' hl ack .:dgl's of thc scale~: a ye llow latcral stri pe framed by t\\'o hlac~ ~Iripl's. the 10\\ cr olle runnillg along thl' outcr cdge~ of the \l'l1lral s: upper side of head n 'arly llllll()J'Jll hnl11/l' to browII: occas iollally a hlac~ ,treak from the rl'ar cd~e ofthl' l'ye Ll\,er thc nape llnto Ihe hack wherc il I~td cs out ; lips ycl IO\\ : vClltra l ~ idl' ye llow : lail
Colllbndae
liil!. :\27
/)/ ' /II/r"/II/"II\' ClIlIc/O/III/'lI/IiS ( WIl/o/im 'ollls
Imlll 1'1Iiall Lall!,'bwi. P.:ninsula r t\ la lays l:l .
striped: a Illedian dark line along thc undersi de of the tail. l' lut c hc1> of 5-1\ eggs scn:r,, 1 I imes per year. Ilateh ling:-. lIleaslln.:d ca. J 40 mill in length . E l'ulo g~ IE tll()ln g~'
is ro und rrom "ea level 10 e \..; va tf(lnS or 1110re than I (lOO III with a marked predilecti on I"lx hilly areas , It inhabits lowland Iropi ca l wd ami dry r()re"t ~. tropical llpcn Illontane roresh. clemings and edgcs, bushy areas, plantation s. cu lti va ted an.:as, g:Jrdell s, and i" also filund in the \'icinity or h()lI ~cs, Thi s agile ac til'e ~pecie~ i ~ arhon:al and diurnal. It lives in tree li lliage. in bu she~ , shrub". and other dense \cgetation, It Il:eds chlcll y on li/anb. espl:c lally aga mit\" and gdJ.oniivc. bit ing rl:addy. but is absolutel y harmless. /)"lIdrdlll'''l\ (". n ll/do l i ll l'lI/ IIS
Di stributio n c, ClIlIl/O/ill( 'II /IIS IS 1\ idespn:ad and comillon throughout ils range. II llccurs from the I)c llllr(, /II/," is
330
Malay Peninsula sOllth to Sumatra. Ann 11:0. and .lava, and al~o inhabits the Phi lippilf(;s . .\'11 . Kinabalu : with0111 eX:Jl:1 IOl:ality data .
Dem/relllp"is pidllS pictlls ((j 11- 1 IN, 17X9) ( o /lIh c r p i<'lIIs (i\ll llt\.
17.
t)
, Ilhsetjllenth de" g llafl'U :" l'11f'l eel c d 10 1I01l)co h)' :'>,11 (I'IIlX )741
111(,
T~' I)(" l m:;lI ity :
NOlgi\t;n: ( I X~ 7: 53 0 I: (aler \1'130: 302) and ( I'v ll '"
J",,, hy B, 'II lU I ,/\
l!elle/n ,/,'/ I/' I> I'"
/11 \, 1.11\'1 ~ ll x ,1 I 'nx . ( 1, S l l '1111 '<, I 'It) I : ) -1 2: VI \I ~ \11' 11)\)..(0: 2-1 7 /lelle/re /,,/>1,, \ l·!. /'1< /,, \ \1 \1 ~ " , , 1<)<)2,, ' \.1(,
Oi ag lHls is
Vcry slender: ea~i1y reeog.ni:ted on the basis or its lat eral ~tr i pe , honkn:d above and below with hlad., and the radiant blue interstilial skin which bCCOIlKS well visible when the ~ nake innates its throat and body. Dorsa ls imbricate, in 15 rows at mid-bod y: I cl1ehrals enlarged: Illa x . total Ic.:ngth 125 CIll , but
Colubndae
Fig. .n R
/)('IlI /r(,/III I /II S I 'IU II \' II/('(IIS
from SUlllatra.
uSLIa lly onl) ahout 100 elll long; females substantial ly larger than Illales. Desc ripti o n J\llales with re latively l::Irge r eye" th:lIl fcmal e~. I preOl:ular; 2 pos t() e lll:1r~; I IOllg Imeal : 212, I I I or I , 2 lelllpllra Is; 7- 10 supralabia ls. the 5'h and 6'" (rare ly the 4'" ) in contact with the cye; dor~a l s in 15 rows atll1id-blldy; vertebra Is enlarged : 1(,3-200 ve ntral s, keeled ; anal ~cutc di\. ided ; 11 2- 16() s ube:1I I (IaI~ , tli\' ided and keel ed . Hronl:e-colulired above ; a yellowi sh lateral stripe up to the tail. hordered with hlad. abo\'c and helow; he::ld brown ah()\e. lalerall y with a black , tripe frolll na sal to nape . behind which it disintegrate~ into s pot s \I. ith blue , more rarely green i"h hlllC, ~ pot s in helween ; upper lips ye lillw : ve ntral side whitish ),ellow to grcenish )'eIIO\\ : t::l il uniform bronJ:e-e ulolll'ed . In se lfllelCnce the snakes Inllate 10 present the bright bluc inter~titial sJ..in . Scvera l cllItehe~ per yea r or 3-X eggs (22-JX.) X.5 - 1 I 111111 , O.l)()-2 .72 g); hatc hl ings measure 202-3 0J mm in length and p n,,~ess thc same en l-
our traib as the adulb . Inc ubatlun d::l)'s .
taJ..e~
X5 - l l6
i<: l"olog)'/ Ethulogy
This ~ lIake is f(Hlnd li'om sea !c\d to e!c\'atlons of ll10re thall 1,)50 Ill . II inhabits llpen lowland trupieal ,vet and dlY loresb-, tropical and slIhlropl Ca ll1l11ntane forest". especially clearings and ed ge~. and abo ~en lb-
Fig. .U9 Iklll/n'/lIplll\ 1,/('111.\ I'I< 111\ 1'10111 TIt:lliallll.
3.1 1
Cnluhridae
lands, llpen ~walllp~, . ,iantatinns. cII III\·all.:d areas. the v icinily nl' ricc Jlnddi e~ , garden-;, and the vic ini t) of hOllses. /)(,IIi/I'('/(// lliis fl. fli('l111 oftc n li \.c~ close 10 river-;, ponti-;. and 111 other \\ ah.:rcd environll1enls. This ngilc alld aetive s pee l e~ IS chIefly arborea l. being. i1l0~t al homc in tlee lil lia!,!e, in thld, hll,hes and othe r dcnsely vegcla led areas through \\'hich It g l ide~ ~ II, in ly, althuugh il olk ll hUllh Ib prey Ilil the J;1'llll lld. 'I hi s snake is sll iell:- diurnal and I'llnd (II' basking. It appe;lrs to ked Ilearl y c'(clu~iv l' ly \111 terrcstrial alld ;Irborea l 1I'0!!" but lilards hav,e also bccll reporh.:d, \Vhen th reatelled thesc anima l, display a defencl' he hal In\ll' lI'ltiLIt i, primarily ba~ed on colour. The:-
ge nll ~ /;'Ilip/Il' c(\ mpri se~
1<'33
The gellus /:'/ilfi /II' 1-; I ery widely di stributcd. ra ll gIll!;! thrl\llg h Ilorth and ,>()uthecnt ml Ameri ca and large part s or Eurllpe . It is f'lIrlhenllorc prese nt th roug hout the tro pi ca l and h.:lllpcratc rcgions or ," ~i ; 1. Variou~ aut llllrs l'o ns iocr the Ilcotropica l species as bl'longing til ,eparatc geller;\. I.arge to l110deratel y large speC ie, II ith sle nder, nlu sl' ular bodi cs . I kad Il at. e lungate. slightl y sct n(rti'ollltite hudy : eye~ mOlkratcly large with rollnd pllpil-;: 2. rarl'ly 3. Plbtocui:tr,: loreal pre-;ent: lelltmb ~cclcd; allal 'l;lIlc dll Ilkd or entire: ~u hca ll dab arrangcd In 1\\'0 WII', . I he largel y crcpusc ul::lr to nnctlllllal, occasionally n ervo u ~ rat sllakes oneil di splay an aggres-;ive dcICIIt:c behaVIOur and arc very agilc , The variolls species inhabit a vanety ol'hahilats. ranging lI'om plantatiOlb to primary rain li1 resb. the ve rtica l distriblltitln extends I'ro m scn-kveillp tll ;tllitlltles of above 2000 III a.s, 1. Their diet t:lInsists of small mamm nls, hird~ . rc:ptiics, 'IInphlbians. li ~h , alld evell ill-;ecls. 1\11 spec ies of naf,h, · arc ()\'iparOll~ .
Tah. ·U Dl slnhlllillil
01'
UII/III1'-sPCCIO::S
(Ill
Tax ono mic COlllllle nt : Certaill alllllllrs list /:"/lIfJ{,C as native 10 Ihrnco, but this is evidcntly h;lsed un a CllllrU~I UIl vv ith L/a/I/Il' 11(/m/il/"(//a, /\ ccnrdil1g tn Sllil 1 / ( 11)')(,) FIll/IIIi' (,ITlhl'lll'll docs not ocellI' Oil l ~ llIllCll . In a rcccnt rL'1 iel\- (111111 "111 ,1(1.1 H 20() I) the genus I\as ~plil. Duc to ~cveral I'easons we hLTe ~eL'p the <.:on~erva ii vc ta xoIHlIllI ":.
I'rrl/lI'I (rll
Key to the Bornean species of E/aphe I\nal scutl: divided: 2 su pralahiais entering orbit .. .. " ., .. , .. . " ... .. " ., .. ... . ,' .... .. ,., I{(ellillra g rahoJVskyi ~CLltc ~ in g l e: 3 supralahials entering orbit .. ..... ... ... .... ... ... .. .......... .. .. ...... .... ... ' .... .. ... .. ......... 2
• '\nal
2 Dorsal scales stron gly kee led , latera l scaks weekly keeled , no black erossband on the nape: supralabials 4-6 cntering orbit ... ... .. jlal'fllil/ eoro • Dorsal scales weekl y keeled, lateral sea les smoot h, a black crossband on the nape: slipralabials 3-5 cntcring orbit .... ......... ............... ........ .... .... r(l(/iaia
Bllrm;'o.
h lup" £, j TIl I'(Jlil/ l'lIIa (S. 1111'.,1 ' 1, I X3 7) /:,/,,/1111' /'ell/ill ill ( 13lJlI" IXn)
1;'l lIfJ" l' la el/ill l'll ~rahtll,'.\'/,, 'i I' '''
I) istri hu lio n Tlte di stnbu till n range~ from India ill the west tll Ii o ng Kong and the rhilippill e~ in the east while strctc hing tll S lll a\\'c~i in the sou th . M I. Kin a h a lu : Sg. I.allga ll
Depending Oil winch gel1(.:ric conccpt is applied. the 2-1 to 40 :- pcc i e~ . Hornell is hO\l1e to threc ,pcc le" tVI'll Ill' which occur 011 \ ,11. Kinahaili .
Genus E/aph e Ftr/IMoI ' ({ in \.-\1 '\\011 ({,
enla rge the surl;lce lll' tlte ;lIltel·ior bodv by late rall y Ilattcilillg the b'ldy and show ing th..: bill\: in te r-;t llial ,k in , T h i~ beaulil'ul ~ peci es i ~ wide'pn:ad and nne ol'lile 1I 1ll~ t Cllllllllun -; na~ cs in tropical I\ ~ia , It i-; \-cry dOCIle, but ma y hite readily and v'lciously \\'11I:n halldl ed : it i-; enl1rel), harmk'ss.
III K , Il-iX:;
j
Saba t.
Kinabalu
Brunei
Sura\\'al.
• •
• •
•
•
.
-
- J-
• •
I Katill1antan I
• • •
( oillbridae
E/ttphe fTavo/ill eata (St
III I (il L,
1837)
('()/Nf,e'" f/ol'o/illt.'a/ln ~( 1111 (II I.
I XJ 7. 141 (a~ } lHJI1g \\t ("llIh<'r III<'IWIIII'/I.' 1 I) pc loca lil ): .I,,\;, and SlIll1dlla, by IllIplil'.'lion 1JIII,I"'llal',,III/I,.lIa S.,IIII 1'1>1 III ~t \\11I1\ Il)XJ ' _X, l 'IUNII\!tD (f\.:. ~IH401HN I«JX-l 2~: ~1111t 1't.1 19 lJl : 3-lJ. ~ 1 11 ",11,1<1<1-111' 2-17, S, 1111 / 1l)'I(.: 12." 1:1£,/)11(' m(,/lIIlIIrtl . L( 1\ 1).(11)•• 1 19JX
-n
rl'/I,II<' ,'IT,llI'llI'll, ~t \ 1
"It"
1'1'1-111:
S\l 1I1I
)1)\1
\2, \1 \\11 11 \
I4)X,
2~
c~7
1:'11I/1ht ' «'JTlhrlll tf.\, S Il I HI " , I l)q I
l..tJ
Diagnosis Mod<.:ratdy large to large, but relati\('ly , lender: head (lnly slighl ly set olrrnlilithe body: eyes large: \,cntrals kcekd: Ihe -1 110 to (,'" ~upralabiilb in contact wilh the eye: average totallcngth abollt I SOcm, ma)( . tntal length I RO CIlI. Dl'scription One prenclllar: 2 IXlstllclilars: 212 or 2 I:; tempora ls: X_I) sllpriloculars. the -Iii' to (,,10 touching the .:y.:: 10- 11 sublabiab: II) d()r~al scak rows, s111O(lth or
sligh tl y keeled on th e side." h.:ekd along 111ld-dorSU111: II)X-2-12 \'<.:l1l r;II.,. k<.:ekd: allal ~eu t e entire ; XO-I I () slIbcalidals , Dorsu111 brol\'ni~h (lll\,': In the anlerior Ihird ort hc body: r )sterior third or the body unil\lI'Ill blach \\ ith a bluish gloss: a ye llow to orange-ctllo ured \crtebra l ~tripe with slight ly darher marg in~ begin:; on the nape radll1g posterio rly : antcrinr ~Ides or the body with u~u;lll y paired , while,blach or hla ch ~P(ltS: intl:l'sli lial ~kin \\ ith a reticulat.:d pattern; l1<.:ck with a broad hlach stripe : a broad black ~t rcak !'rom heh ind the .:ye to Ihe angk 01' the mouth : a IUII'I'O\\' blach streah run" through the ~upra l abia l shie lds: the patlcrn n1h while ante1'101'1 1'. darker to\\';:Jrd~ the tail : ullder~ide or thl' tall dark grey to blal'h , ,1tl\enlles p osse~s the same type Ill' colour pattern. but dl"play substantiall y more intcl1~e colours , Several chltcht:" per year l)I'U P tll 12 cggs each t 5162 x _J'2 5.5 111m , about 21 gl. IIll:uhatioll tahes I07- IOl) days,
3.13
( 'olu hri d: lc
E cul og ~'/ E llwlu g~' f l lll'lJlil ll'lIll1 i~
l(lI l11d hCI\\ eell sea level and cle\';l l io ll ~ of al k: a~ 1 1370 Ill . It Inhahits trop ical weI Il) rl's l ~. e~ pcc iall y Iheil Ill O~ l o pC II par" ,,"ch a ~ ck ariug", edge:-., and Irack s id e~, opl' n lroplcal dry fme"ts, s a\ a nna ~, ~l'I' lIh- and gra:-.s lalld:-.. pl alllat 1l 1l1~. culi ivakd an:a. . , nce p a ddi e~ . and suburban areas. It IS ' n Illlilld clu:-.e to hUlllan hahltat ions, hlddcn under pd e~ ul \\'uod. and It frcquclilly e lll e r~ h(l usl;s in ."earch nf prey. I hl:-. ,,\\,111 and ratherdllsivc 1>na \..e IS prl; d ll lllin a lll l~ t c r re~ t naL prclc rri n!,! a thic\.. Vl'J.!.elatlllll cu\e r, alld sell II -arbo real. where il pro\'l:s 10 bl; a ~ ki l rlll cli mbe!'. 1:1(//1//(' f l (/l'fIlill (,lI /U is tliurnaL 111 (l~I I ) :JC II \ l' III the early Illurn illg ll1' lai c all crlHlolI h (\ ur~. T hi " ~ pec i es I\:e d ~ on rodents. cspccia ll y rats a nd IllI Ce, and nthcr ~ Illall l11amllla ls, and (Ill hink including "lIa ll lowl: ju ve ni k~ also la ke li/ ards alld alllphlhian:-.. II IS an aggrcss iH; ~ n akc, which :-.tri\.. e:-. lien:e lv \\ hell a rnll ~cd . It hit e~ readi ly alld vic io us ly, and Illay inflici a n.: I:tl i\'d y pa inful bile.
1:'/1I1111 ,'
J)i ~ l.-ih\lliun
T hiS ~ p el'lc~ ranges rl'lll11 Vi etnam thro ugh T hai la nd a nd th e M;t1 ay Pe ll ill slIl a to th c i ~ land ~ or Ho rn Cll. Sumal r;\. and Java . Mt. Kinahalll : Ki a ll, i3ullll u Tlihan ( 1.'70 111 ).
on
h'/uplie laC'llillra graiJolI'sky i
I XX 5)
(Ft "(,111 J(.
/:'/aplll\
g/ dh01! '~1'I " !:"il IIIIC
I XS:, : 5('. I'l l. IV,
l i~, _ "t
T ypt·
l uca l il~ : B.ltlll LIPli ca\c Ileal 1 \~ n g,Jltlll , K.dllll.llllan. ~l'Ia t , 1I1
(Solltlu...·...... 1 nnrrH't)) . . tlul n ill,th,,·i r"'!t.uflh:.I,\I- Holl ll·ll).
1I\d) 11 " 'II I )
F!tlllllt '
l\.· ... pn
.·-
1','"
/(It'1II111 '0 , lUI NII\ I{ I)
1'/'/1 - \.1-1. 1'/'/-1 : ,).\\ : 1\.1\,
& ~
~ ! lltd{l ,.
,9X.! 2 1:
SI PIIUN(,
"s l'I'I·lb. ~ 17
rtll/ >hl'lrlt ·Jtllri/l'l e l l g nd'"I1 \l. l'l ,
1\'JI III ' & I )I NII~ IQX)
II t,.'tfll,Jh'luf 'HllfrOJ:roh(Hnl.q .
' Ill (11 .15 ( '· 11
1 \'W 11 1I
19~ .~ ~2 X:"( 1111 / 1t)\)(,:
Co lu bridae
Ta xo nomi c cO llllll e nl : In th ' existing literature,
Balu Haru ca v<.: is ufkn give n as the type luca lity onl y (<.: .g. '(" 11 1.1"1 1<)1)(; ) although l ' I ~ClI I I{ (I XX5). in the original d e~c ription , referred io severa l :.pecimens originating from tv..'O Illca lit ies (the other one being Barabei). As no 'ingul ar holotype has b<.:en fi xed as yet, bot h o f these have tll b<.: considered type localiti es. Di ag nosis
Large, s knder: head slightl y set 011' rro m the hody. ;\ charac teri sti c trait is its do rsa l strip<.: whi ch i~ parti cularl y d istinct in thc pllst<.: ri llr th ird or thc budy. Anal ~ c ut e divi ded; the 5" and 6'10 slIpral abials in contact with the eye. This ~ l1b ~ p ec i es reaches total kngths of he tw <.: en 170 and 190 cm. Dcsc ripl ion
One prcocul ar; 2. ran.:ly 3 pos t oc ul a r~ : 0- 1 ~ u boc u l a r~: <) slIpralabial s: the 5'h and 6'1, touching th e eye: 10- 13 subl abi als; 25-27 dllrsal sca le rows at midbody. th e ce lltra l o nes slightly kee kd : 275-2X5 ve ntra b , k<.:<.:lcd; anal sC llt e di\ id<.: d ; 102- 1l-t suheaudab, d ivided. (;rollnd cul uur slat e grey to gr<.:y ish brow n: head co loured like th<.: hody. indi \ idual dorsa l sca le ~ with dark margins whic h togeth cr \\.ith the lighter inl<.:rst iti ,iI sk in rorm a re ti cul ated pattern ; a light vertebra l s tripe bcgi ns in th e anterior porti ll n o r the body and becomes morc contras ting toward s th<.: tail : the rc ti cul ated sides dar ke n towa rd s the ta il and eve ntua lly tran sform into a broad bl ack latcral stripe: a broa d dark p (l~tllc ul ar stripe behind the eye: sup ra lahi als white to ye ll o\\'. occasiona ll y sli glill y orange : belly yc ll ow tll ye ll owish white with a ~ tri pe ru nning along th ~ ve ntral kee ls, thi ~ st ripe he ing broke n in th ~ ant .:rillr part o f the botl y. .Ju ve niles an: li ghter \vith a lIlon: eo nt ra~ tin g, ladder- like patt ern . About l.'i egg~ in a clu tc h. [e ClI() ~y/ E lh n lo ~y
F:/ilJlI! C /(/(, lIi lll'll gmhOlrsk1'i inh ab it s mai nl y reg i () n ~ n fmiddlc tLl hi gh ekvatio n ~ frll ll1-tOO t(1 at least ~ I ()O m, and is lillind chie ll y in hil ly and rugged reg ions co\'<.:r<.:d with tropica l \\ et and dry forests, tropi ca l and s ubt rn pieal lll on tane ftlre~t s . de-
ciduous and pine forests, and in rock y g ras s l a n d~ . It l i ve~ in cl osed or open fore~ts. on clea ri ngs, in rocky and bu shy area~ , scrub lands, pla ntati ons, on th e perimeters o f culti va ted area~, and also in th e vicinit y of villages. It i ~ also olk n fo und in th e vic inity and ins ide th <.: numerous ca~e s o f karstic and lll11 e~ tun e geotypcs , where it can be observed li vi ng in absolut e dark ness lip to ,cvera l kil ometres from the ent ra nce inhahit ing crev it:es of thc wall and roor (e.g. at ( ;unllng Ap i in Sa rawak) . Thi s unus ual habit mak es thi s ~ p ec i e~ one o f the fc \\ true cave-dwe ll ing s n ake~ . Thi s rat her shy but s wi ft s na ke is bas ica ll y te rrcs trial , oec llrring mainl y in moist, thi ck vegeta tion, bllt al so has arboreal habits, prov in g that it is a s kilful c limber. It is both diu rnal. avoi d ing di rect >. unli ght , and noclUrnal. I be die t is large ly cOl nposed o f mamlIlal s, cspe ' ially rode lll ~ such as rat s and sq uirrels, and o f' hirds an d bat s. S p~ c im <': ll s liVing in caves seelll to feed essentiall y upon ba ts ,lIld s parrows and the ir eggs . It i, an aggress ive snake, strik ing at li ttl e pnl\oeation and hiti lll! fie rce ly whell given a chance tn . Dis I ribul io n Thi ~ ~ ub ~ p ec i <.:~
is onl y knllwn rro m a I'c\\ Itlca litic>. in the moun tai n ra n gc~ or SU llIat ra and Borneo. Me. Kin a b a lu : Kanlhor;lIlgoh Road ( I M)O111) . Il ea dqu a I1 er~ . 1.1111111 LUl11u (SI J1 I'IIIM, I')')-t), BlIntill Tulian .
The ge n u~ C;()lIg l'/OSII II/ O is di stnhllt cd fro l11 Tha iland, thro ugh l'vl ala y Peninsula to Java. Sumatra. and Borncll . Sl11all, s lender: head nnt dis tin ct rmlll th c body; ta il long ; eyes l110deratel y large. pup il ro und : a sl11alll orca l ~ llI e l d I S prescnt ; n a~al ~hie ltl ~ dl\'ided; I J rows or dorsa l ,cales wit hou t apical p i t ~ or k ~e1 s; anal scutc di vided: ~u b t:a ud a l s In two rows. I.ittle is known ab,)ut the bi ology or thcse terrestri al snakes. They lC<.:d on i l1\ erte hra t e ~ and l i /.ard ~ . Their nlode of reproduc ti on I S IInkllO\\ n. 1\\ 0 spec ies have been Ik scnbed for t h i~ g~ nL1 s. and both occur on Ilorneo, one nn ~l. Kin;dxilli .
-'.35
( 'o luhndal'
Ta:\onul11il' fU l11ll1 l .lls: LI \ II (,' ( 19(1-1 ) has sp lit ti ll' Oril' nta l genus /. /()I ''''li ~ alll' tOl'lIlll illto t\l'O !!ro ll P~ (Ill the ha~i~ (I I' IlIe n :--IIC and In urplwlogic;d cha racIl: r~ con:-tanl among ~ pl:c i l'~ \Ir thl' res pecl i\ e g roups. lie I'e lc rrl'd the nominal ~ pec l ':s ( 'onllll'lIlI hll/ioc/('I/'{/ Iloll " I X27 and , Ih/llh('.\ /() lIg iClII (( iI /\ PI n ' I{ S, I ~7 1 tn thl' ge nu s ( ; (l II,!!.r/IJ.\()/JU I FII II ' J( a Ie I );43. 1.1'\ 11 () ( I ()64) \l' a ~ Ihe 1:lsl r":\, I..:\\'\.:r or Ih..: ~lIlit h cas l crn I11c lll bcrt- or 1111', PO\lrly ,t lllhed g.rou p. a nd hl:-- arra nge nlc ill I ~ 1() lI m\ I:d h..: r..: .
Key to the Borncan species of GOllgY/OSOlIIll Alltaior part or hod y with 5 mangc slripc:-: 7710.\ subeaudal s .. ......... ............... ... . /O Il.1! II' II I1i1111J1 • Anterior part or hody with ycll()\\' spots or v. it h three' ye llll\\ stri pt:s: 5X - 7') su bcaudals ... ........... . .... ... ............ . ... ....... .... hllfilldeil'llll/ hllli(){fl'il'llll/
S"bah IX ~7
(;ougy/o" I)/II1/ IItI/ifll!t'irulI/ blllillt/cirulII He'll , (, o lI,1 !\ '!f " ilNIO IlIlIg/C 'lIlJeltllJl 1 PI Ilk" I X? I )
GOlIgy/osoma baliotieirum ha/iotieirlllll B( )II ' , I X27 ( 'on u h '/ltll. a ltudl '/l"I l
Btt 1, 1:-<27 ",,)
T~Jlt' I(){"ali'~ :
'''Ioh( ', /!c /liodl lnh .
(\ 111( I)! \1< 1' I XC)O, !
I J7:
(,I Ihlohl"
)1
1:1\ <1
"\ 11I 1! ' lKP IYOI . I '
(i l lll,l,:'}U,\OI I111 " u IlOt /t-II 't1
/t,,"uIIl 11'11'. (\ 1 \N " II \ 11IXJ ' _X: 1'1)014 _4. ~ 1 \ 11 \1'" t ')')4h ~ 4 7
} lopt'/In
I.l o / le'}I /.\ 1" lI tu ch 'lI e/, \,1 \1 .... \ 11 ... ll)l) 2 a
", 111'11
•
I3 rUI1,'1
•
S:lI':I\\'ak
• •
Ka llll1anla ll
•
stl'lpC~, s li pra la hia ls lig hl \\ ilh b l;ld ; margi ns : hell y orangc or y c ll (l\\, I ~ h \\ hitc. w ith or \l'itholl t fi ne da rk SP()t~ : th e o ll tel' edges o r Ihl' \' cl ll ral~ co lored Il k.: th.: d() r ~( 1I1 1. I're::-u l11a bly 0 \ iparo u ~ .
El'lI lu g~' / Et hullI g~
Rr '(I! I I (II -: I ;t) "\ \ 11 111 I'} " I IU '"'2 <;011:,:,1"'''11111 /l( ,liudl"lnllll "\ \ 11 111 19,1 :'2 h rllllJdltll\.
• •
"- il1ilbalu
\I{I)
&
~ 1f)t,KI N
I '~L 1')t)~ l h . ~l.( )
Uial?,lIosis 'I hi s ~ IKC I ":S 1:- c a ~i l y idc nti fi ed on th e ha s i ~ o f it ~ tlm: c i'(H\ ~ llr ~ 11(l 1 ~ on the an tel'lor hody. 13 n l\\' ~ o f dorsa ls: I Imca l prcsen l. I'll ial kn g lh G I. -1 0 CI11 .
Th is s mall s p cc i c~ inhahi b r..:gi oll s \If ItIW and Illoderal ': c k va l i O Il ~ fro l1l s..:a kwl lip to at Ic a~ t 1500 III a nd is Ihu ~ fOllnd in hoth 10\l'Iand lropica l \Vel rnres t ~ and Ir\lp ica l and s ubtro pi c al \V c t l1l onlanc fo rc ~ l s . It is l11 0rC coml1lon in hill " a nd ru gged a rca~ . (;O llgr/WO II Ill It h(/ /iodl'in/III i ~ a k rr":~ lri al and ra lher st:U'cli\t: ro rl:st-dwd kr. li\'ing
Fig. JJ2 (IIlJIgY/" WI/IIU !J,,/illc/,'/mlll /tetIUJ(/"IIW II frn lll LJ lu ""II Da\. Lllllak-Lntimau Wildlill: Sanctuary, SarawaJ.. .
Ikscripliu ll
T\\ o pre(lc ul i\r~ : 2 p O~ I \lc (ll a rs : I lor..:a l: 1 12 or 21 2 te illpora k 7 supra labi a is. th e ,3'.1 and -IC!' touching the ..:yc: X ~u hla bia l s : 13 dnr~al sca lc nJ" :- Ihro(l ~ h oul thl' Icllgth o r thc hod y: d\ ) r~a l ~ smooth: 1151-+ 5 \ t: lltr;ds(SII ' 1-1011{ 1922, KOI', III\1 19-1 1): anal ~c ul c di \ Ided: 55- 79 ~ lIhL'a (ldal ~ (K u p.; 111\1 1941 , VI \1" ~ 1(, 1l)')2:1. I) \S 1l)<)5, 1),\ \ 11) & V ()( il l 1g e)() In 1\\0 I'lIW:-' . l ;ppe r Side dar!.. hm wll , wi th ~O Ill C sma ll. dark-borde r..:d, ye llen\' ~ p o l ~ o n the napc and thc a ntu'iur portion of' the hody \\ hich llIay he rllsed tll lOrlll
a c:: o
"
" iii 0)
".!'l~:iffl:t1
a:: @
Coluhridac
Hit, JJJ
GUlIgl'/().\'(1I1I1I ha//(Iclcinml h(l/iodl'il'llll/ f'1'(11ll
on thL' !loor in among lear lilter, undL'r rocks. :lnd henea th decayi ng vcgd ali on . It i ~ diurna l a lld rcported to reed ()Il invL' rt ebratl's and li/.ank i\ 'cord illg to 511 11111'J(, & 1'J(.II< (I'NI)) it, dil'! consi~ts l1lain ly of' s p i d c['~.
Gen lls GOIl.l'ophi.\' Rot lLi
VrI :R.
I R9 1
The ge llLls (lUI/yo/litis i~ native to Borneo and I'enin~ular IVIa I'lys ia . Vlnderatcly I;u-ge. s lclllkr; I hod y long. latera ll y CO Ill presscd; head e lollg-atl:. di sti nct emil) thc neck; cycs ll1odcJ'atci y large. pupi ls round ; dorsa l sca les kcclcd. with apical pib. in I') row~ at mid-hod y; vcn tra ls with lateral !..ecls and nLltches on their ~ idc ,: tail long. with kceled and notched slIhcau(htb in t_vo roIl S. These a rc rare trce s na kes the low lallds and mon tanc regions II hn~e hio logy is poo rl v knOll n. I he gc nlls (;(/ ll1 'Oflhi, is mon o t~' pic \I ilh the only :o-pec ic, livini! Oil Mt. Kinaba lu.
or
I\.clabit Ilighlamb, ' arawak ,
Distrihut ion ThL' distribution ran ge ex"'nds ti'olll I'cl1ill sula[' t'v'b laysia to I3nl'l1eo. Slimatra. and .J ava. Mt. h:illaha lu : Kiau. Poring: Il ot Springs region (S()O Ill) and bl'!lveell Sg. Klpungll I and Sg. Kipul1 gi l II ((,00111).
GOllyophis marg arita/us ( P"" 'I(S, 1'1< 7 1) (,-. 1111 '0 " "1110 IJIcll.1!.. l/l'ItoillllJ
PI 11 il ....
I }i71 . -:-. 7 ~
'I ~' IH'
III C:l lif ~' :
<';,Ir'''''"k Diagllu~i s
I I) dorsa l scalL' rOlvs; ~ c ele d . IHll<:Iil'd velltrals ; COIl bv the bril::ill ye llow rlllg~ al'll ulld till' body ill it s posll:rior part. lax . tlllal len!,uli 170 Cill .
~ p icll(l li s
Desc ription One l~lI'ea l : I prel1cular; ::> po st('clI l a r~: X-9 ~ Llprala b ia l ~. the 4 '10 to (,11, or Ihe 5'h and (" lo ill cOlll ac t \\'Ith the eye; 1 1 01' 2 - J tcmpora b: d\lrs al ~ III 19 rows, ked cd. \1 il h apical pils: 2.;O-2-1LJ \,(,l1tml s, laterall y
.1.17
( 'olllhndal'
Fil!. JJ-t
(.1/1/1
(/1"11.\
IIlIlIglln/IIIII'
Ii-oil I I ar.mak,
~eel.:d
and notched : anal ~C Il1L: di,i,kd : IOX- IJO in pair~ , latera ll y keeled . [)nr~a l gnllllHI cu lllllr hn .ilht green, eat:h ~cale \\ Itlt hlad Illarg lll ~ : posle rlllr part or the body and tail wllh ~e\ ' eral urange-euloured hI deep red ring~ u j' which the po~terlt)r Illll,1 one~ arc the most bri ght n n e~: head (lII\C gree n wilh a black slnpc betwcen l'ye and nape and hct\lcen Ihe pariet al .;hic lds: 1111 dersidc~ ~, e ll(l\\ . \enlrals lI!t..:n \\'llh hlad margins and an or:lI1!!l'-CO lollred dash, ~lI h c alld a l ~,
Ecolng~' /E t holngy
Thesc snakes arc arhorea l and inhab it flln::s ts n f the 10\\'lands and Illontane rcgions lip to altitudes or2!JOO III a,'d , They ha\ c (he ability tn asce nt even apparent! , pcrft.:t:t ly sillooth trec trunk" In captivIty Ihe only !i)od acccpted was li sh, It s biology i~ poml )' knOll II.
J3X
Uisl rihutinll rhi ~ spccic::. lives In Pcninsular I'vlalay"ia and on BOI'l1Cll , Mt. Kinahalu : Puring: S!! , Kipllngil I.
Fil!. 335
(iOlll 'O!,III,' IIIlIrga/'illllllS
fro m Sg, K ipun g il L
Colubridac
The gellus (J(JII.I '()S()/IIC1 i~ di~lrih"led !i'om Myanmar. Laos and Vielnam. IU lJali . Sulawe~i and Ihe Phil ippine I~Iands . Largc and slender. nlllscular: body ~Iightly compressed lalerally: head skmkr; pupil round; 2.\ 10 27 dorsal scale rows ;llmid-body; dorsals wilh elongale apica l pib ; loreal shield al kast twin: a~ long a~ hi gh: anal ;.c ute and ~ubeaudal~ divided . The crepuscular and noclurnal ~Ilakes lile on the grou nd and on tree". The gt:IlUS comprises t\ln species. l'ne of which lives on Bornen and on MI. Kinabalu .
COIIJ'0 .\ ·Olllll
("o/lIh('r
f!"I'II<,
(uJ'cepIUllulII (i3()It,
''''/'/I11/IIS
<1.111
1\0 11 .
I X27 . q 7
T~
P('
11-127)
IOl'alil~:
I'I'II"/il. !-til '" "'" '" Sto)( "U ~ I 'IX,) : 2.) IY91 J . .U : 1\1\1 h.r-.!t i..,
(;0'" UHJJ1/a O\I 'r "c/lhtllo . ,-" I I IIt\'4 ,
Ja,., 14)() ~lh .
247
Di al-! nosis Readil y idenlified lin Ihe ba~is of liS greell hod y which conlraSIS ~harply I" ilh a grey lail; max . IOta I length of2-W eill. bUllIsllally only 160-IXO Cill.
Fil!. JJIi
(;()/II 'O,'O IllIi 0\ \( <'I'/Ill/11111
Ih"ll
1'0rtll1,;.
600
DCSCI'iptioll Onc large prcocular; 2 pnslOculars : 7- 1 I ~upra labial~, the 5''' and 6'" or the (,'" and 7'" in t:ontaet w ilh Ihe t:yt:; 12- 1~ sublabial~; dnr~:.tb ill 23. 25. rardy 27. rO\\~ : 229-263 \cnlral:-. :--harpl y helll Oil Iheir side,,; :Ilia I scull.: dil'ilkd ; 120-15 7 palrt:d ~ub c:a udals. divided . Lighl green d r~a ll y. "11l11e ~c lk~ bordercd \1 ilh hlnck. olh(;l", Ivith ye llow. Ihe numher nr ye llowhortkred scalc~ increasing posleriorly forming a vaguc striped lalcral patlern ; upper ~ Ide 01' head !:trce ni~h yelkl\\; upper ~Idc or lail grey or bro\\'n ; lenlral sidc~ li (!hler: bt:lly hnghl or pale ye llo\\' : .Il1v(;nik~ altogelher ~omcwhal darkn (Ihis t:o lour pattern rcfer~ 10 Borncall ~pCCilllt:lb onl y ). /\ clUlch typically consists of 5- 12 egg~ (5X-70 ' '25-27 111 III ). Ilat ch l ing~ cmcrge ancr X7 Il' 125 days and Il1ca'LlI'l: -lO-)) em in lotal length , E(''' I ''I-! ~ I!:: t h Cl l u~y This s pccie~ is fOllnd in low land arca~ I,'om ~C;1 level Il' eiL:valions up to ahoul L,OO III \\hne it inhabits primary and sccondnry Iropi cal I l)re~ t s. tropical wei IlHllltane rore~I;.. mangro\.c f()rcsl~ . s\\am ps and marshe~. \\CI ~crllhlands. alld plantatl(ln~ . It is rather rarely ellcountered close 10 \. IlIage~ (;11/11 '0,\'0/11(/ (11:1'('(,/)1/(//11/11 IS usually. hlll not c\c1wmcl y. found close to rivers. on riler banb. near p(lnd~ and other
Ill ,
.13 9
( 'oluhridac
1l1~~il' area~
C(l\'erc,, \\,Ith tnnglcd \'eget:Jtlon ; it I~ at honle In trec!'> ami bll~he!'> <)vcrhanging water. It i~ an agile. ~\\' t fl. and \'ery acll\'e :-.pecies which 1', alll1n~t elltlrcly arhmeal . hcin!l partial to trectnps. tall bu s he~, and olher thick \'egetali(ln ;juveniles ma y be nwre tCITC~tJ'J:l1. Thl~ strictly diurnal sna"L: l'eeds 1Il'lillly <111 hirds. bUI ab(l readil y hunts arboreallllammal, and bat~ : .Iu\cnllcs ha\'~ been reporteLllo prey upon li/.ard ~. Thi, '1)L' cie~ has all lllicrestillg Ihrcat
Cientls lIydl'llh/lIbes B()t lll
\1(;1
Ie I H9 1
Thi~
gClIlIS I!'> clldcllIlC l(l Bornell. ~ Illall. 1101 di~lillcl !'rom Ileck ; eyes I-mall . pup ils rolllld ; :til oblique ~lItllJ'e betwcclI thc Iwo Ila :-,al ~hield~ : :\ pair~ of chin ~hiclds ; body long and roulld III cro:-,~- ~ eclinll ; dor~al s :"mnoth. withoul apical pih. III I 5- 17 r!lW~ ; \'clllnJl~ rounded : tail ll10derakly lon g: sulll:audais in two rows. '1hesc arc tcrrcstrial ~na"e~ with li)s"orial habib. They arc cOII~idcrL:d rarc . alld a~ a consequem: ~ hardl y an) thin g i~ kllnwn abnutlheir hiology. They I lead
tonI. alld Ihc hri!!ht hhlc - ~Iripcd IOllg uc e.\ tended and relraclcd in an c!>.lrcmcly slow. dcliberale nl ~ hioll . Vlating. occurs usuall y nl night 011 Ircc!': or !'>hrllh~ wilh a copulali
(j(lIlI '(J.I'III11(/
appear to li\e almost c\c lll"i\'cly ncar 10 water. ~o l "e specilllell~ wcre IlHlIld ~willllllin g in rall1 foresl sircaill s. T\\() species arc ""0WII. hOlh Ii,,; on Hornell. onc of which Oil :VII . Kinabalu .
Key to the Bornean species of Ilydrablabes 2 prcfronlals ....... .. ........ .................. ......... IJI!riIJfls • /\ single prcf'ronlnl ......... .. ...... .. ..... prueli'olltillis
Saha h
11 J'''!'(I/JI,d ,,,,, 111'1'1 0/1.\ (II "" "'(. Ix n . ) " .I'dl'llhllllll'.\ Jll'tlljl'fJlIllI /i.\ (f'.l< " '!' \N Il. I X()O)
I frdrllblab es prae.frolllalis (Mo('<)lI ·\lw. 1890) ' hluh/", IJi no/" \. 11 jJnw /rrw /lIh, (-do( ,)\ -\ 1(1 ), I X90 .. I '7 pI 1\ , li g. T~ Ill' lura )il y : ~·II K"Ii,b"llI
I",
IM"I>", 11<"'1"1'.' val' I'I
1( " ,," 1' /1 7
I n , '\1 11 11 I'n l
12
\1 "
1111\
1'))01 1 ' 2 S:
1' 111 ~ ' ''N '' <'I.:. ~ ""'" ' '" l 'I X..j · ~ ..j ; S Il II II( , 19 91 : 14'1 :
f\1
\1 "" 11 ,
It' lJ-lh .:! . .t7 .
' I I I !t IN I, L~ " t il It
I,)q()· 1T!..
Ta xollomic CO IIIIII C III ~: SJlIJ 'III M, & I'J(;J'J( ( Il)l)l)) helie\ eli I ~\ 'dmh/III/(' 1 !,me/nll/llIlis ICl bc a ~ y n() -
J~()
•
K",nh'llu
•
S"' ~ Wll k.
ilrllnc ,
•
1
•
I Kalim .. lllall
•
nym of I ~l'dJ'l/h1l1hL'., 11/'I'ioll.l . They did. howcver. not really ~ynonYlllise Ihe two :-.p~eie~ . /\s long a~ thcre is no comprehensive rcview oj' thc gcnlls. we..: cUllsider the separalion of Ihe pre!'rontais 10 warrani ~pee i cs slatu~ for bOlh laxa . M()e ()i ,. \ Il !) pub lished Ihe de~criplioll of' Ihi~ 'pecics Iwice , i.e. ollce III Le NalLJralislc. Paris: l-t-t 145. and again in ' OUV. /\rch . Mus . lI is!. Nat. . Paris. 3 (2 ): 11 5- 1Me 1\()('()i.;\I(I) 'S papcr In Ihe "Naluralislc" was published on .luly I" . IX90 . 111 the lir~t pnrl or this 1()IIf'-parl !'>erie~. M()l'I.H I,\IW wwte (in JJ1ICrpreialion) : " ... The descriplion of this collectioll l Mr. Whitehead 's eolleelionl will be Slllln pub lished ill Ihe 'ouvcllcs Archives du Mu~cul1l:
( 'oluhri dae
hu t we be lieve Ihat readers orthe Na turaliste will apprecialc to tind d iag n (lse~ o r the new s pec i e~ included in thi s w llcclion in this int ere~ tin g publi cation , whi ch, at the same time , all ows us to sct a date for these diagnoses ." In the '·No u\·el les ;\rchivcs", there is 11 0 date of pu bli eation . It is clear Ihal the first part oC the to ur-pa rI series in the " Nalurali sle" was pu b li ~ h e d he fore the " Nou velles Archive. " , and most probably the second lXIIi, too. It canl1ot. howeve r. be clari tied here whi ch ruhli cat ion serves as original description. Unforlunately M()(,()IA I(f) used two di Cferent ~ p e " ings for Ihi s spec ies in these two puhlicali ons. If Ihe ru blicalion in the " Nat lll'al iste" pro\'e ~ to havc becn published tirsl, Ihe correct name should be Ilrc/rah/((h('s prelimllll/is.
Diagnosis Pre rro nt a l ~ fused to form onc shi eld : three pairs or ehin shield s: m3 X. tntal length 43.6 cm.
DesCI'ipl inn Prcrront als ru sed to rorm a si ngle shield : I lorea l: 2 pre o c ul a r~ : 2 p o~ t oe ul a r,, : 2-3 s uboe ul ars: 1+ 2 temporal s: X-,) sup ra la bials, none or whidl louch the eye: 15 dorsal scale 1'01·\ " at mid-body: 17X202 ve nlrals: 72 suheaudals. di\ ided. Oli ve brown dorsall y: a dark ve rtebral stripe. ~ om e times hardl y visibl e: Iwo bl ac k d tll '~o l a t l: ra l stripes.
(icllus II/lemalils Y 1\ )\( , & R,\o. 1992 The gl: nus 11l1 ('m {/ lll ~ ran ges rrom I'v\a l3y Penin s ula 10 Sumatra , BoI'll l:O. and Ja va. Iso l:'tl ed locali ti e~ 3re knl1wn rrom eent ral Thai land. Small : hl:ad d is linct : chin "hiel d::. arrangcd a~y m metr ic all y: 2'" ur 3''' pairs ofslI olahial s enl arged . in co ntac l wilh eae h olh er: I rre-chin 'hi l: ld
int ernl r ted tll fllrm a ~l: ri e~ of' po inl s 111 Ihe po::.\('ri or parl or the hody; lo\\'er "idt:" yclftm , Ih e ou ler margin" u r the \·entrals dark brnwn: s llb c alldal~ with brown margin,,: a da rk mediall line along Ihe lai L
,
Di sl.-ihulion T his ~pec i l:s is endcmi c to Mt. Kin abalu . bUI nll prec ise loca lit y dal a art: h.now n.
Uein g crepusc ul ar and nllc tum al co lubricl s Ihey are "pecia li:lt:d in a diet o r ~ Iu g ~ and ~ n a d s . The latter arc e.'( tricaled rro l11 Ihl: ir ~ h l: ll s wllh the aid of' thl: anteriorly posil ionl:d ta n g~ of the narn>w. tinn lower j a\\'. II the species reproduce by layi ng eggs. I'our species have been described ( D \\ II) & V ()( d I 1'1(6), two tIl' \\ h Ic h occur Il n Hornell. bOIh on vlt. Ki naba lu.
Key to the Bomean species of ["Ier"alus 5 or 6 upper lahiab ............... ....... ..... .. .. . hl/'I';S • 7 upper lahial s ............................... lI1ala(,(,(IIIII.\·
3-1 1
( 'ulllhridac
S"bait IlIle /'l/I/lII ,' Ill /,,.i,'
• •
( 1-1' )11 , 1l'i27 )
IlIler lllllll.\' IIIlIllI ( ' / 'lIIIU ,' (I'I II R' ,
I X('4)
I lIfe rJl(llu.\' /ael ·i.\' ( H OII :, Jm M,'c,'p/Illi1,,' ,,,, ,,,, 11,,11 IX27 ,~ (I fmhh, I plldh" lilt" 1\, H( II I I \ 1, J It I XC '(h
I X27)
1)' 111' Illca lif ): .I :,.'" '
' -II . ~III
t 1111( 1)
6X , III 1<""11 1'/1 7 ' 7 () ~ "" 111"'1 11 ,.1: 'm hlrrt'l,halll\ lrT/t 'hra/n I t) \ I I{ lIlIoI Il));": - -L' :
I e)O 1.1
;-\11'1 It l" ,
11I1l1111 11 \ 1(11 ~\;. "' '' ,cl ln , 1()X4 . .::!..j S l l l 'HI '" 1"e' I' 1'l'J2" l 'I'. I ')'I·lh 2--l 7, 1'1')()c:2'J1 ; SII'IIlI N', 8. 1""1,11< 11)IN ' X7 I,,'erna/ll s I'/{ ' I'I\, I{ \( 1 & 't \ I , I\)l'~ - 1..t2
" ,II I'tI.\ IUt 1'/'. I 'HI
;;o, ~I \I"""
l>iaJ,!IIHsis Idellt lljcd by (he cOlllbina tion oj' as), llllllci rica l ehill~"icld~ and 'i - c) ~lIpralahi;tls, The lung lon.:a l lum:hcs the eye , 111fL'1'I/1/l1/\ IlI,' \ 'i\ call reach abo llt (,(J e m in tota l kn g(h .
Fil!. JJ 7
~
(; L
-<: iii,
J--t2
11I{<'I'II(/{I1,\ IlIl"'i ,\
['mill
thl' Kaillborangoh Road ,
I
Killaballl
• •
8rllllci
S:lra\vak
• •
I
KJllInantan
• •
Colubridac
Oesc ripti on One loreal, in contact with the eye: 110 prem:ular: 1-2 postocu lar ': 2 12 temporals; 5-6 (7) supra labials. thc )",1 and 4'10 or the 3,,1 tn 5'h touching the c:,c, the 6 11 ' vcry long: 15 dor~al scale row~ at midbody : smooth: vertebral s enlarged : I-IR-1 76 I'entrab ; anal seutc entirc: 3-1-69 suhcaudals , in pairs . S~IITII ( 1925b) rcported about two specimcns li'om 'vlt . Peri ssen with unusll3l1y high ventral (196) and subcaudal (71- 7X) eOllnl~ . One or his speci mcns had 7 supralabials. \ ,1cdium to dark brown dorsally, with irregular b,lrk bamb which do not cro~s over mid-body: head usually slightl y darker than hod : lower side~ hro\\n or yellow with e\ception or the dark spots which mark the ends or the dorsal rings. EC (llo gy/E thol o g ~'
Fig. 339
This arLllIreal species inhabib tropical wet f'orcsb and tropical wet montanc I'orests. If is act ive during the cvening and night hours . In Pcninsular \t1 ;t1aysia, a speeimcn was seen in a sinall slrealll (KII II 19X 7). I'\othing e1~e i~ kno\\'n or the biology ()rthis r,m; spccies. It is presumed to bc ~irnilar 10 other 11I1('rll(lllls and Porco' species. M.\I I-- ~II IS (1')9('c) round a ~pecimen during rainy \\'eather onthc Kamborangoh Road at 10.00 p.llI. Another one was discovered ca . I m LIp in a bush al about <) .00 p.l11 . (M ,\l I-- \ Il -; 1,)92:1). I}istrihutioll This specie~ is distributed rrom Lao~ , Ihrough Thai land and Pl:ninsular Malay~ia , to .I'l\a, SUlllatra. and Romco . On Borneo. it has been recorded from Sabah as well as rrom Saral\;)k and Kalil11antall . i\o rceords are available Illr L3rulll;i . VII. Kinahalu : rVlarei I'arei. Kamborangoh Road (1600 Ill), Sg. Langanan ('1150 m).
III/I'/'IIU/II.I //Jalu cHIIIII.1
from I'Dring.
inf'ralahials in contact wilh one another and 7- '11 supralabials . It allains a m ~l\inl\111I length or llf' ahoul 600 111111 . Description One lorcal. in contact \\ ilh thc eye: no preocltlar; 2 postoculars: 2 2 lempnral~ : 7-'11 "Ipralahial~ . the ." .1 and -1'1, or the .1'.1 to 5'1, touching the eye. Ihl: las t largc: 15 dorsa l scale row~ at mid-body, \cry Jilinl keeled ; vertebrals cnlan!.e
In tern allis
malllCC(III11S ( PI t ) ' R~,
. 1.1/, ' 111""1 )\(/ < ,l/oli/( CO II (/
t" ca til ~ : " 111
Jcr Niihc
1864)
PI II R\. ) 1'6·1. 27.1. r)al ~ ti~ . .\ r ) )W vlal :u:c,," Il'cll'lb uJ.1t Mal:,," ;:lJ
B \1 t III LOR ( 1951-:) reported about a ~pccimcll found in a rubber plantation . Nuthinl!. cl,e is kn )11'11 about the hiology of thi~ forc~l-d"el1ing ,pccie~ . It is presumed to be s imilar to PurcC/.I corilltillis .
\'till
l)jstribulion J)ia gllClsis This 'pccics is idcntified by the combinalion of as ymllletrical chinshield~ lI'ilh the ~ccond and third
lives Oil . umatra . L3nrneo. alld in the Mala y Penin,;ula . ~II. Kinahalu : PorIII/Crill/illS 11/1I/lIl 'I 'CII/I/\
II1g.
( 'u lllhnda-:
Gt:l1l1S
L('ptllrop";.\' Bot
II · (il R,
1900
'I h-: ge nLl ~ 1'<'II/I//'I)I I/r;1 rallgL', frolll 11i(; Malay Pcninsliia ~tlllih 10 Bornco anti Sumatra. L~traordln ari l ~ , lim, elon!!atc. cyl illdri c; h..:atl tli,ti nct froll1 Ilic hod y. \ (;1')0 tkpn.:~sed; e)'c moderai el ) large ; pupil vertical-el liptica l ; en larg-:d nlllg" In the upp-:r ;;1\\ ; II(htrib v-: ry largc. cach wcdged ht:lIVCCIi t\\O Ilasab anti the first :--u pralabial shield : dor~a l " with di :-.tlllct kc e l ~. li nel y ~c rr:1lcd . lanc":lIlail.:. :-- li ghtl y imbrica te . with api ca l pit s. in 17 ro w ~ at mid-hod y, velil rab ~eek:d laterally and notched: lail ntrclllely long and ~I im : :-- lIbcalidal s ill Iwo
'()od IS frll!!S and li /,Ird" anti tlll:y rcprlldlll': c by lay111 !!.
eggs.
rhi , genus i:-. ll1onotyp ic. The onl) "p..:ci..:" lias h(;cn '(lund al"o lin Bornco allti'v' l. K inaba lu .
Leplllrop";s albo.lilsClis (DI I\ lI RIL , BlI lR()r-- & 1)1 1\ lI' RI I, 1854) \I,hl '4odc', 411ho lils(,l1\ 1'1~ 'I\ Vr lu,alil~ :
i\1,,, ,.,
()/ ,IIi/('\
/,nl)( /tm alhojll\'If.\_ ') \ 11111 I~) ~I.
1·11'.; ... . 21.
1:!:
1>1 \ 11 lUI _ HII HtflN
& 1) 111'.11 1< 11 , I S') , 1a
'>ulllaira
\1 \" 111 '
~Il I HI 'c . (1)1>1
I WIII,I. 1·1·1
\ 1\11
\o)llIllllk ll
I (Hlia .
I·'X,
"') . IU RIII)! 1
~X
I IiINI I\I~ I.
191 7 110 :
&
S J( lCiHI '
"lJ ,1 .... I~N~h , ~~ 7
row ~.
'I 'r/tII0I ,IN\ u'''ol((\I 'II\ \ 1\ .... \ 11
Thc"c ,naKc, arc nllclLI rnal and c rep u ~c lliar. living 111 :--econdar), :llld pnll1ary lim;"t, rhcy are probahl y largc ly ;Irbol'l.::d. bUI ha\-c mosll y been ~cc n acl l\'\.: lIll the gro und . The :-.n:lKes arc olh;n I<wlltl :illlng ~ trcam s (I I' ill "tr(;;1I 1l bcd~ . Th..:ir I~lvo urit c
Taxullumic comJ1ll'IlI: For a dl'CLlS:-.ioll nC lhe taxollomy of thi s SI cc ics ~cc [) \\ II) & VOC;I 'I ( I \)\)(,). 1,1.'fI/lfI'(lldll\ hO I'/l/' ('III' ;,\' II, It I 1·t\1 d R, 1900 is a synono 111 llr I . Cllho/il\cl/s .
liil!, .' ,10 Adult 1.{,/JlII/ ·(}/,/ti.\ ,,1""/11\'4'11\' from Pei1"","ar i\'1ala ) "la. ~~
/>
c: c:
~
o
<5 ~
.1 -14
~~, ~l~~~~~
7
_ _~~
( 'ollllmd
Di ag nosis Slim body with all ~x Ir"llle l), IOllg lail: pupi l \'crtical-e1liplical : \'enlrab ~ l1ld s ub c:llIda l ~ ~~clcd :
Illa\. knglh 20 7 CIll, bill al' l.:rag~ S p~CiIll C II~ l1l~as urI.:
120-160 CII1 .
f)cSlTipt inn Siloul vcry hllll1l: I prco~Lllar: 2 po sl(l~ lIl ar~: I klr..:al : 2~2 tClllporak X 17) s Lipralahial~, Ihc .\' ,' to 5'10 or Ihe -I' and 5'" in COIllaLl \\ ilh Ih..: ..:ye: dursa l ~c ak~ III 17 !",I\\'S at midhod y. \..eelcd: 220-259 \,'ntrab : an~ d ~eLlk di \'ided : I 13-206 ~ lIhcau dai s. ill pair~ . \"~~kd . l)orsally UIlIl'Ol'lll dar\.. g re ~'. g l\: yi~ h bl\)\\ 11. gre)'i ~ h h lad, or blac\.. : \'CIItrod ~i d ~s y~ II O\\ ,~h \\ hil ,' 10 \\'hit..: : JlI\~lIilc~ dar\.. 1\ ilh ,10--10 whitish balld ... Oil Ih~ b(ldv all d a hr,ladel halld Oil Ih,' ncc\.. . Ihe balld~ elus..:r 10),:elhcl pu"teri\)J'I\ ~(llhal Ihe "' lIa~,' ~ app 'a r lbr k ~r
e l1ll ) 11\\1 (),'nll Oil .1 :1\<\ \It. h:in a h a lu : Illorl' preCI~l' lura"l\ dala .
EtolugylE t Iwl ng~
illh ahi l" lowland Iml i,'al 1l\l\:"I~ , \\'here II I~ rcpurted III II \e cio,c 10 '11',' :1111" ill hu ~ h,' ~ alld Ih,,' \.. \'q~l· lalion . \\11 ~Ir,'alll hallb alld g r;nel h 'lb , Thi , .';Ila~c I, pnlb:thl) lar~'e1~ :trh()lcal. hul II ha~ IIH),II) h,'cn ~eell arillc on Ihl' g round III 1')l'cSlo.,. SOllll'lilllC~ il \\' a~ ob,,'!'\ cd "II inl Illillg. II i~ nol'l lIrna I an d k"lb Ull I'ro:'" and ILIai'l l..,. pl'~y1l1g c .~pcc l ;t1 I ) 011 Ihc o.;cllllaqualll' "elllLl d gl' llU " 7h 'I lidlll !I'III'IIS '1hi.., ~ pe,'I"" I'" rc~anled :I~ 1ll1culllllHlil Ihmugh(HII
(;~11 11 ~
I ('III/II'tllilIIS (//hll/III('/(\
II .... range . I)i ~ t .-ihllt iun
I hi'; s lla\..c·s r;lI1g,' \,\Ielld ... Irnlll Ih,' \bl ;l) I'elllll';lI b h) \lorn,',I. SllIlIalr:l. and la..,. II dnc .. appal -
L.reot/oJl FII II
(.I 1( ,
\I
ItlHlIII
I X2h
I he g,'IlU" / . I'('f}(ltllI " dl'll'Ihutcd Irolll Ih,' kill (11' till' !'onll..:r l iS')\{ lilroll g h 1 ; lr~ l' palh oi l" dia ;lIld ('11111 :1 ,outh III SII!a\\ ,"I ,1I1d Ihc I'hill ppille 1, I"ill"Small 10 IlIc(hlllll-'L/l'd. ~klldcr ~11 : lh·~ . hc,HI de prc'"cd d()r" ()\l'lllrall~ . ,h "dllly ,,'1,111 Ilnlll hod \ , l'\' ~ r,' l alllcl~ ,""tll , pupil I l'lli t:: tI -m ,t1 . il lhll" lal'g,,: 'lIllCl'lor 111:1\11\;11 IL'clh ,·ul'lc'd . 111111 :1 ,l!:lp helll,"'11 the \el ~ 1:Il !!c al1k'll(11 aild lhe , "h"'qUl'lll ()IlC,. UOI'al " c: li,' ~ \..cclcd III lilli, II Ilh "PI" ,i1 plh. ill 17, 19. (lr ~ I IP\\ ~ allllld- hpl" . I t:1I11;d , nlulllk-d . hCIII. ,Ir II ilh blL'ral 1..,','1". "lIilGllld ;iI , ,mangcd II) a "Ilgle or IIIIl rnll ~. hH'Y
I h":~l' lTl'PlN:IIJ.1I t, Illll'tllrJI;d l'illlIhnd~ Inh"b lt " II'lde lal ll'l) (I t hahll:II,. ranging rrom l'lI lti\:iled :1rt:'I~. Ihrough gra,~ alld \andy plai ll '. 10 sl'..:,llIdar;- :II''': Il'rrl' ~ lrlal I hl'~ ked 111:11111) llil li/ar.!\. rrogs. alld '1I<1i-.l'~. 1IIIh Ihe \11I:Ill..:r 'p T Il''' a\-;ll prl'VIII!!, upon ill'>":l·h . .IUIl'lIlk\ pll"l'~~ 11111rC 11I11:n. l' cll lour" and murc C\'lli I'CI,I illg palkrJI'> ..\ II "jll'CIl'" III Ih l\ gl'I1\I'> ar..: 01 IP' II'IHI\ . III Illial 2(, ' IWl'll' ,
Kcy to thc Borncan
speci(.~s
2 ,call's l1<.:l\\ecn Ihe po"I<.Tior
of Lycot/oll
lIa~a l
and orhil .... ('/III/willl/,\'
• I ,1.:;11..: hrlwcrn Ihl' poslcriur 11:1,:1 1 and nrhit .. 2 2
0 prl'ul'lIlar, :lnl' lun!!ate lorcal and tlie prefroll ial rll!.:r 111l.' orhit. unc an!.:rtpr tempora l .....
............ ...... .......................... .... ,,,beillu,,s s<'lIl,,; • I pn:ocli lar: lorcal absL:lll; prl'fmnlal in ":OIl I:l1.:1lI'ith Ihc supra lahiab: 2-:1 allll'rior IClliporals ... I'I/m""i,'
·Iah. -17 1)1'1111)1111(\11 (\1 I n"d""-' pel'll" ,In Borne" .
'''''.
L.I'('ot/oll SlIbdllCll1S s(,lIfe; /Itnt!""
\uh(.'/II(/(I, HI/h,t II \'IH~'.
II (IN ,
1055
I \)" 1~' JJl' PIIlI1PIHl1l':-'
[1) :;;-'
i l ~: Plll'rl~\ PIIIh..:l.' ...... I. P ,dil\\ ; lrI 1... l.llId.
I.\·,nc/ull \lIh'lfIl III' ~II·IIII"\'
1.1' \
:-'"h"h
K",ah:du
IlrUllcl
•
•
•
•
I .. l'lllclllll I "I"" /II/(' H,,", 11'1 ::7 I\'( "dllll 1'1/1'111 '11/ ,1 ( '\/, I x·17 I.ynllf/J II ,wl" 'j" cTII,' ,\ I'lIll'i I 1 \110'-.:. I') ';'
I (j(':t I -
II.NI~\J . \I \I}..\ II ."'; 1 ~)l) lh . ~
17
J)ia~nu s is 111;'('1111'/11' arl' n;adi ly rc..:ogni/ahlc on lile hit, IS of' 11'1..: usually IhrL'l' I\hih.: rillg~ unlhc all l..:rltlr pnrtll)11 1'1' llie hody. I .. I, l('tll('/ is lbtingUI~hL'd f'nlllllhl' olher "ub"p":clc" by a 100Ier el.l\llIt of' ~uhcaud~d~ II illeh , :1cL'(>nlinl,!. \0 I I I' 11l'l ( IlI55). lIumher k~" Ihan ()'J . It dilTcrs frolll Ihe other I3l'rI1L'an ~pcl'ie~ Ill' II 'c(!(!rJII in Iha l il lacb a prc(lcular 1>hl l' ld and ha1> a long loreal wh ic h touches Ihe l'yc. T hc Ipngc"l "pCl.:ll11e n kllO\\,1I ml'a~ urcd I () l'1ll. Wllh awragl' \pel'illlen, heing hl'lwc..:n X() and 100 Cill IOIll,!..
:\du ll/:,'('udull
Dl'srripliun Onc lo rea l: 110 prelll'ular; 2-,\ p o~ t nell lar~ ; I 12 ICIIlporal..;: X ~lI p ra l ahl [ i1" , the J'" 10 5'''. 4,1, a nd 5'" or J'" 10 (,'" In eontal'l 1\ lilt tltc l'Yl': dor,a l, In 17 rows at mid-hod y, i-.l'l'lcd : 19!i-21 1 vcnlra ls: ;I lI a l ~eulc IllllTn~l ll v dll idc!! ; 60-6!i ~ u hea u dab. divided . rhe pliolido llc l'haraclcn~tic~ arc adoplt:d rrom I 1\ IIO\J ( 1955) and app ly 10 lit is subspee il's o nl y. I)al'" hroW Il Ill' hlaei-. dorsa ll y: napc regio ll whilc : I\hile ring~ l>rlanahle Width all a long the bodv :
I
~ill.J\\ak
Kdl'Illl1lllan
• •
• •
hdll1g rl'l'lll thc lail cnd willi IIIl'rca<; lllg age: grcyis h III hnm ni,h bl~lc" \I ilil 2 whllt: han d ~ or II'hilish spcl'b in Ihe ,I nlt:rior jl('rli()n ur till' hody, lhe latlrr hcinl,!. rell l;lins or w h ilL' ball d ~. I cry ,lid ~pccil\lcn\ With hardly :lny 'lic li IlIar"lll g, al all : Il'lIlrUtll dirty III grcyi,h while or hnm 11 . 'il'l'l'ral c1l1lche~ per vcar or 5- 1 I cggs (3 1,5-)(, / 12,5-1.\.5111111. a\erage ,\cighl .\.(,5 g): ilatc hlings I11casure ahulil 24() mm in knglh, ~Idulh
b' C)lu:.:~'/E tholog .\'
This ~Ila"c i~ Illlllld !i'OIll ~ca leve l lip to elcl al illll, ah{>\'c 1500 Ill . It inhahll~ 1<1\\ la lld t r~Jjl l e:t1 wei a nd dry foresls. lro pica l ;tlld sublropica l wcl tIlo nt alie 1 ()re~ls, ,hrub lallds. p lalllali( 11 ~. rice padd ies . cu llilHlnl areas, alld Ihl' cll\'iroll~ ol\'il lage". This SWill ~ lI a"l' is Inrl'slrial . n:trl'ating un dcr rocks, Slumps, pie..:es of wood alld Icgetalion, and arborea l. It IS lh)clUrnal an d rorage~ ac ll vc ly in the tiark llL'SS III lillt! ils prey, that in es~enee c(1 n si~ls or li/ards. especia ll y gce " n, and diu rmtl skillks, IIhic h arc ~tarled in thl'ir ,hcltt:r~ . 1~ 1'(,lIdlil/ SlIhcil/( 111.1' i ~ ra ther a~g rl'S"II'l' alld b i t r~ rea d i Iy whcn lllokSlcd . Distrihution Thi, subspccil's i~ nat l\C tn BurneD alit! l'al;l\\';t 1l (Phi lippines). :\1t. Kinahalu: !l undu l uhan ,
( 'nluhndal'
GelltiS MlIcropislllOc/o/l Boul I' j\( 01
I g93
R.
'I he g ellu ~ ,:I /(I('/'()l li.l'l lwt/o /l has a II iseo nt inuolls d ist ribution ran ge \\ ith rn: llrd~ be ing kll(mll (rolll ( ·hilla . Indl :l, Sri Lan ka . and rak i~ tan , as well a~ rrolll ' l hailallli ~u uth to SUllwtra. Horneo, and .lava, It cOlllpri :-.es Illcdilllll-'I/Cd :-.pecic~ : With very ~ll)Ut hodie~ . I lead di :-tin ·t I'rolll the h,)d y: eye Illode ratel , 1;lrge to large. pupil round : llne pair 01' very large 1:lI1gs: dor:-.al s ~t)'()ngl: keeled. \\ ilh apical pit s. ill 1<)-27 Hm:-.. \elllral ~ I'IHllldcll ; \ai l moderate l) lon g: ~u h ca ud als ill two I'(l\\ s, rhe IIlCllided \ p L:c ic~ Illay he diurna l \l'ell a:-. nocturnal and have a Illarkcd predilectIOn 1'01' the \ iClnit y or \\'at~r. Their delc nec hchm iour rc~em bl ' ~ thai ol'thL: cohr;!' III that Ih ey llaltL:11 the an tc-
;I,
rio r purtion ur the blld ~ ~ t re t cllI lI g It latl'rall~ and lilting It in\(l a \ l'rtlcal po:-.lll11n oil' the ground . I hClr diet c OIl ~ i,t~ li)r a major part o( Illlg:-. and toads, their Iarv;le. and II 'h . i\1I1opeclc, Ill' 1111 :-. gellu , arc o\'iparOils. I'our s peeie~ arc kllnwll, 1\\0 "I' \\ llI ch ,'CClIr Oil Bornl'o. onc nn Mt. Kinaba lu .
Key an the Ilornean species of Jl1acropisl/toc/OIl Lsua ll y X upper lahlal :-.: iliad, or \\ 1111 iliad, halllb: Ilcck panl y red ........ /111\ '/ ('( '11,1 • Us uall y 7 upper lahlal s; a da rk \'l'rtchral ~ Irllll:: lIt:ck parll y hille ............................. rlwt/lIl11ellls trall ~ \l: rse
-;" h" h
& 1)\ ,II (" ·/'IIpl,i.1I"tloll rltm/',I11,'/1J \ (11' ,,1 IN H" II . IX27) \/(/(. rtJ/Jitn/fldollll(1\ 1(.
(,/1\
j f) l l\ III
1'1.11 101 •
•
lvlacropisl/tm/Oll r/toc/ollle/as (ROil ' in R Oil ' . I X27) 1111/11"111,,,111, "/r,,d"I1/,/'" I" ca l il~ : '/,1\"
II
IlI," III
I·
11"11. 1X~ 7' .'3.'
KII';lh:d ..
HnlllC I
"'~Ir;l \\,oill..
t\ .IIIIlwnl.II'
•
•
I X'i·1)
T) Ill'
l)ia~lIosis
• I
•
hp ~. \hroal , allli he ll y UIlII;,nll plilk til :<.:11<1\\ I ~ h , with a row nl' dots Oil Illl' \cnlrab; lUI l'n ll e, deep r.:ddl ~ h Oil Ih e lr upper :-'I d e~ . i\ ~i n g lc <.:Iutch c\lIllpn sed 25 eggs N(IIIlOre ill (;,, Illation is ava llabic .
Fig. eXI\
\I,,, 1'I1/"\·,It,,t/,,1/
I'j,"c/II/I/('/((.\
I:alrl)' , lender, hotl y lalerally cO lllpl'c~:-.ed : easi ly Idenlll'ied b) li S hlu c Ihroa t a nd IhL' di s linctl y mark.:d dark \ eriebral ~ I ripe : m:l \ . icngth abou l 75 c m. Ikscri "fiu II Onc loreal: 1-2 prl'ol'ul ars: ,\--1 pos loc ulars: (l 'cas ionall y I ~ uh(lcular : 2 · 2. 213 or 313 tcmporal !'>: 7- '11. slIpr;lIabi a ls , Ille ~ "I and <-1 '10 , 11I0I"e ran.: ly thL: <-I'" and :,, '10 touching the cye: 19 )'(l\\ ~ or dorsab at midhod y, ' l!'Ilug ly k.:.:lL:d ; 124 - 1-1<-1 \emrals ; rill a " V" : nape hlul ~ h grey: upper ~I dl' of head 11111 fill'lll in Ihe groulld co loll r orthe hud y: lIec k hlll i~ h : upper
/I
I-. " ~
,
14 7
( ' olllhlld
El' olC)g .\' / E llwl og~ " " C'm/lil';UU/OII
I'it"d"lI/!'Ia.\ I... 1(""ld ill 10\\ 1;lI1d
al1d 111l1111;\11,' 1l'!.!IIIII ... \\ Irer..: il I1lI1ahll~ IHlplcal \\el Ii \I c~b, Iloplcd 1\ ,'I 11l11l1lalle 10Ie,;h . '\"""p'. IlIal~lrc~, plalll ,1I101", II,'L' paddi~ ~, an d gardL' Il ~ III ~lIhUlhall arc
rhe ,l!C I1I1 ~ (}/Igfl( /Oll" \el~ II iud, distrihllle:d 1\ nlr pll p" lallolis III Paki ~tall. Ilan
I hl~ lwautirul ';llahL' I~ C(1I1I1I1(\I1 , ,,:' p c~la ll y III thL' 10\\ lands \\ hCI1 arulI ~ ..:d . It r;II"'C' the all tenor part or ih h()d~ . Ilalt ell, lis Ill.'l' " Irk e: a cuhr". alld Illay bltc \ 1(,lously. I> hl r ihu ti o ll I'/IfIc/olll(' /a.\ I... dl;.lrihlltcd rmlll th~ south tl) J!(lII1L'Cl .. Iava . ;lI1d SUlllatra. \ I t. h: inaha lu : I kadqllarle'r, (15()()-1(1()() Ill) II \()( olle:ct 10111 . ,\fW'J'(I/'J.I//r'Il/OIl
\
1aI:l ~ 1'~1l11l , "la
wa;o; a I~o IllClit Illlle:d 1(\1' Il llrlll'O by III I{,,, III I I ') I 7) IIl1de:r the na Ille (}/I,!!,(lciull \'ill/flC '( ·/IS . A cUlrd Illg 1(> ':i II I IU 'II(; 8.:. I'II( 01 II ( I'JI)'}) Ihi ... i, a III i';ld~nt i l i~at i()ll. (}/io,:IJt/c1ll (,(1I"I't'/(\ I' a :-,pL'cic., 11\ il1g lIortlr llr Ihc 1\1.r1a) I'cllllhula. 1'01' \\ Iltcir re:a~on \\'L' ha\1.' c\eluded tlri:-, ... p~ciL" i'rom Ih,' l:llIn
or
Key to th e Rornean species of OIiK O(/OIl Budy \\Ith !-tripcs .......................... . _.. __ ... _... _..... 2 • 130dy wi thou t 't rip e~ ..................... .. ............. J 2 Dark slripc;s \\ ilholl l ligh l ,pllh ....
.. .. .... ....... 8 • Snwll red or )'ellll\\ llr while ~P(]ts ill a dar" VLTIchral siripe ................. _.... ___ ............................. 7
J Vc nt rals barr..:d \\l th dar" paltc;rn .................. .. ........................... __ .. fi ll rp 1If'11.\ ('(' liS • Vc ntra ls II l1 ll1ar"ed. ,potkJ llil Ihe: Iluler cd!!e~ tIl' mOlllcd ....................... __ ..... .. ...... .. ......... 4 " 19 dorsals at Illid-body..... s(,lIIicill('lIlS • 15-17 dorsal!> at mid-body .. _.. _...................... .. .. 5 :" 17 • 15
tlorsal~
dor~nb
al mid-bod~ :11 Ill id-hod:
__ ... __ ..... .\'II/)c({rilll/l/IS _..... _........................... (I
~5 black rill!! .... each with :111 \>\a l ye ll"w i~ h >pot .............................. flllIlltll'j(-r • Narron\ bro\\ nish ur reddish nCls~bars on bar" ..
(, Ilod y with abollt
.._ ...................................... __ ........ _._ .... cf.
.Iigl/a/lls
7 Anal ~Cll l c; divided ......... l ·e,.tehf'a/i.~ "ertebl'll /is • Anal scu te 1I0t tlil idnl ... _......... _ evere!ti ~Iripes arc ~ I early visible ...... . _.... o('{ol inelltll,\ • Only Ihe dorsal ~ I ript' i~ ckarl) \ isiblc, the ot hc.:r six ... tripes arc \\alltln!,!. or ICI) lilillt l), vis ible .....
H 7
1I1(,I'L'I'illliii
Co ilibrida..:
Tah . ...) Di ,triblltioll "i" (}f£gu"(I/I - SPl' cic~
Oil
Il(lnlc('. Sahah
Ofigllt/Oll (' ,'{'r('lIi H. II I I M,j
R.
I XI)}
O/igllr/OIl/IIl'\'(,I'IIIAII (<'; III~II\ . II~ I
K.
1X') I)
Ofigllt/OIIO('wfill (,uw,' (S. 'I~ 'II " 'C I ~III) Ofigot/oll {J1I1'(llIrtI.'·('('IIS (S'"II' .o I . 11'.1 7) ()//gIJlfOIl .: r..I/,~ II
1X(,2) ()/ig".loll .I II loco 1'/1/11/11 " ((;1 " ,' 11 K. IN 72 , Ofigot/OII ,'er(ebrafi." "('''(l'fJrafis (( ,I 'r- 1111 K. I X(,:,) O//g',,/Oll
,\t'/IIICII/ C (II .'
(PI
B Ol I I' (,1 ' 1{,
()/i,a.:, Od(lJ/ c. TlJ't ' tll HI H II '( ,II~. IX('~ . I. .:! 1
\It .
",":oh"l" (}I/ ,I~("I()//
,'I , 'J't 11/, II, " I " , ,I I< I ~ 'l-I l' 1" ) 1'1 I X" ", 4' s 2. "illl I II )I{I! 19 01a 6 1: III
B \U I I I I I
11(1 ,
:-'\1 1111 l'.l .l l .12:
~'P( .10 N
-'1 "
"11
~I "
'\ I ri g I J{ (t.q l
.)
n,un",
<;anl\\:ok
•
•
•
+
"
Oial!llosis
I g93
· 1 ~p('I
K,"ahalu
"
II K- .
Oligot/oll el'ereffi
I
• • I • • • • • • • • • •
()/igl}(/olllllllluhf('1' I Ii "t t I'''',IIL 1X')3)
.
• "
K:ol'l11ulll:on
• • • • "
• I • or
Rcadil) r..:ell!,!n i /..:d lln Ihe ba~l~ Ihl ' Cll lolll' pal Icrn dC~(I'ibed be Ill\\" Ventral ~I d c Ullililrtll coral red : allal ~ ulc ":Illir..:: J:'i },(JII ~ oJ' dorsals al 11mlhod y. ' Iol al k11glh k~~ than
50 CIll ,
I t)I ?-
1111 \ I
I~) ~ -t . 2-+ : ~ 1111 !l INt. PN I: J.q : i\1 \ 1 ... \I I '''; II)l) · lh ~ 2-1'7 . ) & (i !«j,,'\ I \'''J 1l),) 7 H )l) . " 11 I IHN I " ,\: I Nl dl< 11)1)l)
10·1
Ocscrilltioll On..: prem:ular: 2 postocular.; : I ~Illal ll ()rcal : 7 ~II p ra labi als, Ihe 3'.1 all d ..\11, III cO lll acl \1'1111 Ill..: ..:y..::
H I!. J.JZ ()hg utl,J11 ('1'('1'('1// li" '11l 1),IIlllill Valk y. Sabah .
u !~"
( 'olllb n dae
I I:: IL:ll lp or;Ii~ , dm d~ 111 I" Il.1\ " at 1l 1ill -hod): I 12- 1.:'-1 l":ll tla ls, allal ~l' lIll' ':Iltlr.:: -Ir.-72 d ll I(kd ' lI hca ud :ti\. l tppl'r sld.:\ .l!r"\'I~h hrllll I1I"h . Ill th , hload hla rl- I"h hl\lII 11 ,IIIP':", tilL' Illiddk \111 ' II Itil y..: lI \lII I~il al1d n:dd i"h . hlad-border..:d ~;ll ld k blotch.:,>_ th l." lateral lII1C, 1\ Itil 1111l'1 \ 11I:r\.:d 11 ).!lll ~ca k :< ilead bllll\l1 . II IIh a dark hl"\l\Ill, \'-:--il aped lllarl-il1g tilat OPL' Il S p\lstenllrh , ;1I1\It l1\'r dark hl"lml1 hal1d .:\te lldIll!; fll llll lilc UPllL' l sid.: IIf lli' h.:ad ule r the c)'..:,: 101\.:1 "I de cOlal I.:d , I ':Ilt ra ls latera ll) witil darl111;11 !-,11l~
I I"': all ' Pl'l'IL'S l.fllliS ).!Clllh 11111-11(111 11.
0\
Iparo\ls: otilerll I"l'
OligOl/OIl octO/iIlC(f/II S ( S( 11 .... 1 11>l 1{, I ROI ) t /1111,\
(1(/,,1111('1111/\ \ ( It" 11)11'.
~fl\(· 11. "1I1I"""\III\.'lllh
I XUI
(.k·. . ' .!.! I1 .lll.:d h\
~t)()
. ,\ IH'
IU f;._'Ii I ~ : 'lIt
1\111:11 ''i ~ )')(ll)
~...JO).I ...
pll,h.lhh' hl'llI ;.! J il\~ 1 \UIIOfl '\ ,../O/tlll"!II\, "I"llil \I~I)
I ~, ()!J ,'!Or!,," I')X~
(1/
/O/IUCOltt\, "~J
I'
1 ~ ()n ;l
I ~).~ I . ,~. I
I
~ /~ III
\I'"
1{" \ltl I\)I ?
\11 11
I~: "
f
H ,HI '\
'-l 1\ 1"1.,,," 1'1'14h ':4:
Ilia glwsis Fasi ly 1'l:'C()t!" I/.:tI h ~ iI " dl~ lI nc t ,tri pc:d p:ltll' rn . I)orsa l \c;Ilc~ III 17 1"1'\\" at Illi d - h od~ : 2 :Inte ri or te m porals : IL'n tra ls ,Iig ltll ~ all!;!u latcd: :l nal sculL' emi re. 1\1 :1\ k llg lil about 70 elll, :tlL'I'age ~ p L-C I II le ll , 51l-(1() Cill .
Fl' ''ICI :.! ~ / E t Iwln:.!,\'
I, a grolllld-d l\l'l llllg spu:i.:" nr the IOlllal1t1 li>l..:'as . ()IlL' ~ P L'Clllll'1l h:ld a Sl'1I1C ld Ii;a rd (II tilL' !-,CI II I" .\jJ/I('/I111110I,I/III\ III Ib stolllaL"i1. 1111 " rL' l' r':"l' lll " :tl ltil..: Il1lorlllatioll ;lI a li il hk 1111 ih hi o lng.' . ()/Ig(lt/O/l ""( 't'elli
I)j ~ t
rihu. iUIl II" " :-,pl'C les I' a IlnrilL'alll' lldl'llllte 1\ 11. Kinahalu : no prec i,e I('ca lll ) data ;lla liabk
f)c\uipl io n One lore-a l: I preoL' lI l:!r: ~ po:o.lllclI lar:-.: (, s upraoculars. lite 3,,1 "lid -l't. In contacl \\ !t il tile L'Yl' : 2 I 2 tL'l llpor:t1 s: tl nr:--a'" In 17 rn ll ~ "I 1I 11 d-hody: l(,I I X3 weak ly anglilated IC lll rals: alla l ,c ut ..: ,'nt ll"l:': U-6 1 suhe;ll llla ls. dll·i(kd . Til ..:~e \:liuL's rck r III , p ecillle ll ~ rro lll I\oml'ol lnl y. i)or,,
or
1969) .
C lule hes o f up to .5 l'ggs. E c olo:.!yW tholo ~y
' I hi s s na kc is rou nd in low land reg l()ns \\ here it
"
CL
-'5 0
( 'nlubnda..:
inhabits tropical wet and dry I ~)rc~h, but it I'" obmainly in cicaring", llpen sc rub lands and grasslalld~, along roads, in plantation;" rice paddies , cultivated arca". gardell~ , alld in the \'ieinity ( 1'\ illa gc;, . II is ~ecreti\'e. terrestrial and se mi f"o""orial, retreating under roc"",,, logs. pilc~ or wood, and in \c!;!ctatioll . U/i,!!,()t/IJII oct()/illl'aIIlS is nocturnal alltl li.:ed~ on li n lrds, frog:" tadpole ... . and olher snakes. but secms c"pe\:ially pariialill the eggs or li/ards alld bird ~ a, \\cll a" frog "pa\\'n. I his "peele" i" Illiid-Illan lll'red alld "cldol11 allempt~ to bite . III .·ingaporl'. a clutch \\,};, (II"co\cred in a Iree III the borings of" ,,0 111 e longic()rn beetle, a" hi gh a" 7 III abo,"c Ilic !;!rtlllnd (Rlllll) 19(6). ~erved
Ll istri huliu ..
li\e" in renill","l ar \l ala y"ia as well as on Ihe I nd\ll,c .~ lan ,,,Iantls of l3angh.a. Belitun J2. . .Ia\ a. ia~, th e Rlau l\rchipela g(l, Sllla\\'e~i , Sumatra and on Bornco . 1\11. h: inab a lu : no precise localil Y dala 'I\a ila l lc.
O/igllt/IIII IIClo/lI/l'aIIlS
Olig ot/oll plII,/JilraSCellS
(Slltl I (il I,
I X3 7)
Llt's crip ti o n pr..:ot:lIlar~: ~ or 3 J1()"toculars : I or 2 I Imcal : 2 I 2 01 ~ 'icl1lporab: !< "upralabial~ . Ihe -1 '10 alld 5 '10 (II' only Ih\.' 5 '10 III CO i1lacl \Iilh the eye: 19-21 ro\\'~ ,)rd(\r~al sc ak ~ al Illld body: 16()-2 1() lelltr"l ... \\ IIh ~ ll g hl al1gle~ : al1al s\:utl' l'lIlirc: -I()-(I() ~ubcalldal~. dl\ Ided . Adlilis redd"h b(,(\\\I1. ~re~ i ~ h hr(\\\'II, or d ar~ grey dorsallv. or wilh a hknd of"the Ill..:nlluned colo ur~ : hody \\ ilh wid..: bantl~ I~addle hlolcht:~) . ol'tcll \\llh undulated, darh. (i.e. hrown , g rey, or bl~l(h.ish) borlkrs, the illlCr~ paccs with 2-,\ da rh.L' r IlIle ~: a dar~ h,1I1d li'olll the
One to Ihr..:c sub(leular~ :
\' >J1tJc/UIIIJltrl'U /"{I\( '(,f/ \ :--" Itill.! I .
I X'I ')11 1'1 :1 lig. U c\. 14 T) 1Il'lol'alil) : i\11 "Mang. J,I\;) SWIII/('\ P'U',nlld \( 111.\ , IU
Fi:,:. J.t.t Ofi!!,odoll 1"lIl"lnl\! ', ' 11\'
frOI1l
Hundu ruilan. l.\:--tJ
111 .
RlJ( 111
1'11 7: 12(. U/tg,OI/UII /'III/lIIll1\ (' I ' II\
'-' \ 11111
Ilnl 1~ \1\ '1 11" 191" J .:!X : 1; 111 N II ·\ lt l)
~ -L
&
~JI II H I ' \
~ 11(,)(
~hl """ 1'»)·lh
I()')hc:
,
1t ")1
\
()S -t 1,1,--l .
2 11 . ~.I 7.
2\)~
Diag nll s is 19- 2 1 I'llw~ nr dnrsals al
Illid-body : allal ~\:ule cntin: : 1\ln anteri,lr tt:l11pOrak Ihe \ elltral "ea lcs 11Ia\ or Illay nol b..:ar blaeh. hl(\lche~ (If lip tn hal I' the individllal "cak~' ~ i/ e~: 11Ia\ . Illial kn gth allllo~t 100 elll .
.151
\\ :Itn and hid hl.' l1l'a t I a IUlhl." rnl."i" ( )1 hl.T ~ pl.'l.'l n1l.'n ' \\1.'11.' dl ~l' O\\.' ll.'llln plll11al\ 1:lIn li H'c, t :1I1d In ~Ct' ()lldal~ I(llc" t IlI1 Iitc ed gc' 0 1' pOl1d, al1d :1 ri\c r, rl"pl'I.'II\I.'I~ lS11 IIU'J(, s: l 'JloIl< 1l)l) X) It I~ a rare ' pel.' lc, Ilh ll.'lt h:h hcclI 11.'1."01 .11.'11 f'I'l)lll , tltllll(k~ hl.' I\l'l'l'l1 '1.': 1- 11.-\\"1 ;111t1 'IOU III a,"' ,1. (KI :ttl).
I hI.' r 'prodlll'til l.' hllllll g~' and I.·";( ll(l g~· are I.' nllll.' l\ lIn klHl\\ n. 01'1 '(JI'r1 11111 11/\ i' ;1 IlHlIlllt ) pieal gellu, \\illt I" nllly ~ P l.'l.'ll· ~ al~(l (ll'\.'lIlTll1g (HI 1\1 1. Kinaballl ,
Oreo('ulalllllS /ullfilsC/,i
131 It ) I.N(,J I{ .
( ) n '11~ ,t/rlllul \
1S"·'
I';;; ~
I QOl)a
.'0
I X9l)
Di,. I'ihu I ittn "I.'lllkllll l.' III Ilornl.'ll and 1'01' a 111111.' \1," 1."\1.'11 U lll' ldL'l l.'d I.'nticllll c It) 1\11. Kllla haiti II \\, :1' (llll~ rcn: IIII~' Iltal il 1\:1', :iI , ,' rCl.'llrd l.'d rrnlll (lllll' r rl.'g l(llh (Ii" Sahah, S :lJ':I\\a~ , and i'nllll Hnllll.'l (tl l "l
()I,i",It"'I'/l/J/ \ 11/'/1 II'
il .1 : ~
11.11
'I"{ 11(","ClOIf/\ 1'11I1"
1Ir11l1h. III I ~, )\ I I ' • •1 I'
1.\ fW 10,' ,,1 .
1..111.11>, <111 ~'Oll ,,",'1 1111111/\,
I" It \ ,\ 11
l II
:' ~
~III I 1.1,,'11
I{"" I I 'll!
1-11 , I ( \I 1'1 i ll :--''' ' 111 I'n I II, ~f\. '~:I)! II " ... 11)50. "hiL 1"1111' II)~~ . 'O.hU"'dl\IH' ~""" 'ItH. R! '\ 1"X ~l .2-l. "">IIIIII IN t, PN) qq \hl t..~lI" 1')1) 1h ~, 1 7. j "loiK 8. 1 \, I')"hh h'J \ 1 \'\ III' A (.1 :''':-,·.·\'',,'\ 11)1) 7 .:\7~. \ lllliI I,-X: h •. 11 11 ,1.,'1 1::\ hi , '"
J)iagno~b
(ienll" ()/'C'oca/alltlls B()t II
I '\I(,J R.
I Xl)()
I'ltl.' gl.' nll' ()I'('IJ('( i/U/IJ//' I, l it , lrIhlll ed 111 I' l.'n ll1 ~ lI lar 1\1a1 :l\',w and lin I\ornl.'ll. :-'mall. k' lll·'lIla!. Il:IIml\.'" ... , nah" ci"'l.' l v I'I.'SI.' I1I hllll1:' 1111,,1.' of'lla: ~'I.' IIl: ra ( 'II!rIll/II/'/'/ :ll1d ;\!(/CI'U(,II I kad nul d " tlnct f'rulll bndy : eye" ,Illa ll , pupil , roulld , I I~ e III \ / " ('{'()('It!II II/II., tlt e I'lrsl ,u pralahl al tllu l.' he" thl' no" tl'll : dl,tin gui shed rrolll ( '"lul//l/I'/II alHI ,1/11 , ' /'1)(,1111111111,\ hy tlte pn.:"el1l.'e of' 1\\ II I 11 I I.' rn :J':ti , al1d h ~ I" 17 I'll\\' , (II' dor'ab.
I""/IIS
Fil!. ( ' \ \ (h "I" 1:111.
I
'''"11111\ hllllll\(
""l! 1\:1.. 11 , 11 72 Ill )
/111 S \11 77'11 J. 1...:11 1111,111
SI.'I.' gt: ncril.' dl ag n(l ~" .\ , lllallll"sUl'lal ~na h' \\ illl :1 11I: I,i llllllll il'llglli n.:t:lIl'd 01' "'7 elll n~ scr iJltinn
(jilt: Inrt::ti : I rrl.'lIl.'ular: 1 rO ~ llll.'lIlar: 7-X "lI pra labiaL.;, Ihe -t'" and Y" touchin g tht: t:\'t:, Ihl' 7'" " Iht: largest : X inl'ralahlab . 011\ 11II.' h thl.' lil', t ..t arc III cOllla 'I willi Ih· :lIllerior l' llIIl~llIdlk 112 kill ' poral ~: 17 dnr ~ al sca ic 1'1)\\ s along thl' cntirc hildy, ' 1IIIlotli : 1_"i- I 32 Il.'l1tr; tI,: ;llIal ''';lIk 1.'11IIrl.' : 20- J 2 ~ lIh ,,; alldab . In pair~ . Side:, alld hack hlac~I~1t hl'lm Il 1" hla c ~i ~ li : dOI'~al l} \\'illi tb r~I.T ,t': tl t:~ whiclt forlll SIIIIlI.' ~ ind ,Ira lig/agpattl'rn and an allllll~t t:ontlllllOll, stripc Oil tltl.' lo\\,I.'~ I dors:tI ~l'alc I'I)\\ ~ : a lliill 11111.' Ilirollllli tlic I.'yc lil a) I.'\tcnd onto the' b(ld~, for it ~ h(lrt dl ~ l a n,,;t: : a V-shaped 1ll:I1 ' ~ill g Oil thl.' oceipllt and Ilapl' : \1.'1111'<11 si dc ye ll()w l o.;1i wliik. ,(lIII(.:Iill1c~ \\ ith :t kIt hrown :..pots: latcralillarglll , \'cillrais the e()lour ul thl' dOl ,lim : I",ve:r :..idc "I' t;1I1 hn1\\1l \\'illi tlirl.'e d:lrl..cr ~ tripl.' ~,
or
Ecn log y / l<:. hol() g~
I'hi ~ :..p '.: il.'o.; Ita, hcc lI Ii.llllld al altitmks betwccn 1000 and I XOO Ill . I. I ~ I (I
J)ist ri hu tion I'elllll:..ular 1\'1;da j~i a (I':tilallg) alld lloJ'llcn (Kali Illantan, Lon g 1\;lah, II T2 Ill: Sabah, 1\'11. Ltllnakll : Sa J'il\\ak. Kcl ahil Pl;lteall) , \1 .. h:inaha lu : M arci Part:i , LlIlllll I 1I1ll1l ( 1 1\1 I <)]O), I\"ic'silali (I XOO Ill). at ..t 20() il'1'1 (II \ I'" 'II 1900a),
.15-1
l'o lubl'lliae
licnus Parells Wi\( a I IC 1830 The genll~ /'I/r('(/, range, frolll eastern Indi a, through the south of( 'hina and .lapan, ~()lIlh through Indoch ina to J;l\a and Horneo, Some islands of the Philippine~ arc also inhabited. Sma ll. slender, harmlcs~ slIa!..es . Budy Ibtillclly c()llI pn: ~~cd lalnal ly: hcad ,hort and wide, tbtinctl y sct off rrom the hody: eyes large, pupib vertical: selilel ialinn of'chin asymll1clricallading ;1 menial groove . In eontrasl to the genu, A,,/o/,cIfill'll there arc 15 (rarely 17) rn\\s ordlw"als allllid hody, in con trasl to /1I1!'m(/II/,\' t\Hl palr~ of chinshields, hUI no pair or pre-ehin~hlclds. and Illl.: 2..1 and 3,,1slIh lahi a ls arc en larged . Slibealidab dl\'ided, anal ~cutc entire . The cre pu ~eular and noetuntal slIa!..e:-. arc highly specialised predators of ~Iugs and snaik The latter arc e;>..trieated from their she ll s hy means or f:lng" which arc positiolled ill the li\1I1t ora lIaITl\\\. finn lower jaw.
/\11 species nrthis genus rcp ludllce hy la yillg eggs . T he ta\onolllY within Ihis genII' i... hOlly tkbatcd . I'o llo\\'ing I{;\I) & Y \t\!o ( 11)92), 15 :-.peeies an~ 111 eludcd, onc or 1\\0 or\\hieh llCCllr Oil Ilornco. one on ,11. Kinabalu .
Ta:\o llullli e COlli IlIl'll I : I he e\l~tence 01 /'urt' a,\' on Borneo" dl\lIbll' d In S II I HI " <'<:. h(oIl( (11)1)1)). I.lnrnrlllnalcly no "pecillh:lh II ere n:e'\amined by Ihe,c aulhor" . ;\Ithough till' pre,ent aut hor shares thi~ I)pinion . a !..ey 10 dl"tinglli,h holh "pccle" IS pn\\'ided hen: .
(' arilli/lii'
Key to the Bornean species of Pareas 195-21 J \ l'lItrais ; 105-1 I X slIhc<JlId<Jis .. ... .... ..... ... .. . .... ..... . .......... .................... .... IIlIelllI/i.\·
• 161-1 Xl) \·clllra1s; 5J-')<)
( ', Irllla/II' ('UI"IIIIIIII"
Pal'I'II.' IIl1dlll/i.' 11\(0' ,
( 1\" 11 . 11'1 2:-:1
''l,'' H.
I III
nlfl
holl' HI H
•
I')()O)
II '\ 101 '.
1 ~ IJl' I"l'alil~ : ~1 . II ,1I1 ~ .
/'(/ I't 'tI' III IC/lt l /r, .
\1
\ 1 h \ III "- )l)" .... h
I l)t)O.I ' I N"i .
pi \ \ II - fi ;.!\
""iI \l lSlr',·1. "i1r,,,, ,,k ~ -lO . ~· r·,. . It)t)hc
"
Diagnosis ( ' hin"hicld~
a": lllllll'tncal : high cOlin" of\elllral;-;
( 195- 2 13) and subcaudab (IO)-IIX): slipralahlal"
and \ elriarl!ed ; IOllgesl (1\-111
I, \ 11 "
~pecimcll
Killilh,tI"
Htllll!:1
~ ,"d" ,'~
Kalllnanf ~JIl
•
•
"
PlIreas IIlI('/W/iS (R<) l 1 t r--< 01 R, I L)OO) 'mhh'l Ilt/wl",
sllbcalldal~ ('(/1'/1/11111"
<;:1 \1ah
/Jt/rcll.'
0
recorded 7 1.)
~
, ' Ill
19l)~h).
J)r~criptiull
Prcf'rnlll:ti ~ltidd ill C(llll
•
.,
19" - ~ 13 \clllrak <111<11 ,,;cllie ellilre; JI)~ - II X 'lIbc;llIda I'i. ill I\\'(\ W\\ ... . D(\I'~al "ide or bod\ ~and-clIllHlred " \\ I1h J1)1d dllr"
.1 55
l 'uluhlldae
pralahl;d ~, '\n uh lll, Iel y arran ~ e d . di~lally lapt:rIIlg. I.e , llll>rn -~ hal'cti . dL:II ~ e c hl "lL:r ui' spL:c k ks I'ruIJl IU\\L: I L:d gL: lll' l'yC' III lip; lIl\\l'r si des or hcad
Efu l u:..:~ l Et 1H) l u!!~ ;\ snake ul IhL: hl\\ 1;llllb, \lllh IhL: highe:-.I record sll:lllIllin g I'r0111 X50 111 ahOlc ~,' a-kll'l. Iii" nucIurnal and arh(1rc,II , pl," l1lllahl) l-':etii llg UIl s lll g~ and ,,11:11 "\ I II J., \11 , ( 1911-tb) 1(IlJJ1d a "pl'ei nl,' n ~ Jl1el1'l:, III' 111 a bu sh in a halllh(1(l 1'01e,1 al aroul1d 9 .00 P,111 , 'Yl1 lpalric;dl y 1I'IIh 11I1{'/'/IUII/I IlIt' l 'I I ' I Ie' repolied ahoul :1Jl Inlcresling re~lIn g posilion n r lhis parli l: ular ~ 1lL'eIIlICJl , T hc ~l1ake \\'ol1 ld lic curkd lip in ilS shL:llt:r il1 4-" ,' 11\:UI :I1' eo ll :-. whieh arc arran)!ed panl y ahole eaeh olher, hlilll
liil!, ( ' \ \, I
",II ('i/\ 1111<
llU l" IWIll \ g
Lall g all>lll ,
:-I'iO Ill .
Fil!, J-t7
{'(/ /'('al 1/ /1(
IliIli,l
Iinl1l
Sg. I
.111 ':;111 ;111 ,
;-; 5()
111
I he "Ilks ol' lh.: hl1dy \Ie rc ,troll)! l), cll l1 l prc~scd . '111l' tail \\,)uld li)J'JlJ lhl' 1!1\IL:nnusl co il. and Ih,' hl'ad :llId neck lI't:rc concealed he low Ihe IIppa or IJ1id-kve l body co li s. The "Il a "-; IJ)ailll ,linl'd Ihl s peculiar pnsllIre filr allulhe:r Illil1ule elen aner il had hl'CIl lakel1 oul oi' II » ~he ll cr. '1hi" behaviol1r 111;IY Ind lealL' lhal 111leJ'Jlo(k, or l1a111bo\1 sene as primar) da\ ~1lL'ItL' rs , Di ~ IT i h lll io n
1' lldL'llllC OJ) L\ llllll' O , ,It, Kin a ha lu : ca. 3(1) IIllh)J)1 Ih,' Sg , Langan:1n (X:"O In :u.I. ),
Genus
P,WllllfllO{~l'lIl1.\'fe.\'
(ie 1\ Itlll{, I X5H
'I hi" genu~ I ~ di slribuled frum Nepa l and ;\ ss;nl1 (1llllia IlhJ'llugh C hina In lhe norlh. 10 Sulawcsi and 1 : lore~ III the ,·a~1. II ahn inhahlh nlan), orlhe I'hil IppinL' i,land", S111a ll. ,,It:nder, rear-ranged, 13ndy e longa te an d di~ Inlel from body; "nonl \\ illl angl1l ar edge,,; eye» large . pupd " \l'J'li cal: i1 "lIlgle na~:tI sh iel d: fron tal n :ry narnm ; d\lr~ al ~cales ~Illoo lh . II ithOlIl
( 'olllhndac
Ta ll. 51 ))i, lllhllll"ll or
/'w /lIIJIII"h
IIO.I I, ·.I- S I'':C I': \ O il
HOrtll·O. Subah
/ ' miIJlllot/I"iIll.II ('I·
/ ';("(/1.1 (il'I\ " " ", I X5)oi
P.WlJ/lIIIOI~rlllls(('s {llIh 'l'r"li'IIII1I'
(H, >II
l'l
13, >II. I X27)
or
Il/ard,. but rmg.s arc Their diet C()!lsi,t s chldly a l ~o takell (lCC
Key to the Bornean species of PS(lIIIIIIOc/Yllllstes 3'" ini"ralabin l Ilot horde'rillg Ihc Illcn tal gWO\"l:, .\
pa ir, orchin , hiclds: dnr,al sid.: (I I' hcad \\'ilh darJ... and light longi llldin al "trcaJ...s ...... {I/lI"I'I'/III' II/II ., • 3 ,1illrralahi al hnrdcrtllg Ihe 1l1l:ll tal gromc : 2 1);"llr\ or chill shil'lds: d tl1' ~al ,ide or he'ad not slrcaJ...cd ........ .. ...................... .. ............ P;dl/I'
P.mlll",(}(~I 'Il(lstes pIIII'l'rIl/elltlls
(BOil / ),\ lIl11l11 tJ/ ,hn/Jll h \
in
B Oil ' ,
ru/l uta II
1827)
I~C llt 111 1, illlil
1 :-< ~7
5-i /
' 1 ~' lll'
Illralil~ : J ~t\ i.1
I'll/11 f t" . ol'n \I clc III
1"III11I11"dl'lI(/\lc,\ \1I1 11 ( 1I~j)
J 2:
I<JOl a.
\ I \N I I !! '
~ll ,II U 'c .
\l)l) 1
(1' . II I
1')Xl, ~~"' ,
I{I )I)I
19 1:'
I X'JO. I I 11, l en l 10, 11 1(01(/' 1 ()~ 4 ' .4 :
\1 \1 ... \ 1'" (l)l) Ih ::! 17
/ "l l llllll tU hllo ' (t', ' ,h.'C . \ \ """ 11 " ! l )l}(ll
.I
Br""~1
• •
• •
Dorsal clliuralinn ne it re . reddi sh bl"\)II'I1 , darJ... brown, greyish broWIl , grey, or alllw, t hl :lcJ.. . I:cIllaic, II ilh a ~ene s nr large ol'al rurou s or bl'OlI"l1 ,pOl S 011 eaeh ,Ilk 01" Ihe spllle, Ihese , pOI " , Ol11elimes cnnlluelll In lill"lll eros,hars ol'er lite ~plne : helly dellse ly peppered I\I llh hnl\\n . and \\lIh darJ... brOIl n spots or 10llg itudinal Illle,: Ilanb one n ornallll'llIed 1\ ilh whilc strcab _alld n ·t.: ll i anll'riorl ) . "laic, II ith a more nr ic" di , lil1cl , darJ... . II ide :-Iripe (l n Ihc hack. illl·o ll"ing the mediall li vt.: I"lIW" and hall' urlhe s l'\th nm ortht.: dor,ak SImil ar ,tnj1L's Oil tht.: nanb-, s ' parakd rmm the rOrtll l'1 hy a narrow 1\ hIll' ~treaJ... : lllarJ...ings In Iht.: 111l'dlan ' tripe V-,ba pcd 1\'ll h Indi ,iinci t.: d ~'e'< hell y nOI ~ o d(;l1 ~ e l y peppered ;1, in I(;lllaic ~, Ihl' , pec b oneil rnnlllng 2-4 IOI11-.:lIlldina l lIne,, : 0( ,' 111 nol a" dl slln cl as III [cma les. Ilead lI"ilh sYllllnt.:lnca l Inngitlldil1a l l11ark ing' in hUlh .·C\C', nil each , ide a dark s treak litroug h thl' eye, sOlllclilllCS II ith a II hlle slreak belml . P .,·uI/IJI/IUh ·JI({I!<' I /'rrh ·L'J"lI/!'JlIII\ is lile-be'anng Illth st.:lcr:tl ill te rs p{) ~.~ ihk ill a yt.:;lr.;\ lillc l Illa y cun la m up to III hahie ~ wllll:h ml'asun: 14 X- 17X Illlll ill kn glh al bIrth .
\ ~II
202 ~ 't \ 1I 11I
'\ 0 , I I U r\ II \ ~) l\.
J. I\.111ahahl
• • • I • I
"l lP
El'ulogy/ Flhulllg) Th i, snake I~ round li'om , l'a Il'l cI lip In abtlul 21J()() 111 . 1\ ith a lllarJ...ed prt.:dilcc llllll I'll I' htll ~ areas ((bllW
Di;tgl1()~is
Sllla ll I\ith a di ~lil1cll\"c hl'ad pallL'rl1 (sec heIO\\'); Pllptl, le1"llc;1I s lih: 3"1 suh lahla i llol touching Ihe I1lelllal grOl\\e : 3 pall'" orchlll"hlcld,. ·la\ . kl1 gth (,.\ CIll , ;m .:ra ge abOlIl 50 CIll .
H:,:. J-tll
1'.11/11/11/ ," /\"1111 .11",
pungil I. 'il':(J
/, ,,It ''I",,I' 'IIII/\ Imlll "t! . KI -
Ill .
Iksnil)ti4)11 I R 1~ ~ ' ", SI 1\ 19 75 )' One lorea l (occa,i llllall y , plit illio ~e\era l ~ h l eld,,) : 1- .1 pl'Cocular,; 1-4 po s tocular:- : 1-3 ;lIllerior IClllpural s: 7_1) su pra labial ,. Ilurtnall y Ih e 3,,110 5'" ill 'ol1laCI lI'ith the eye: 7-X sub labI a Is: 17 (rare ly 19) n1\\ , (l I" dlll'''1b al mId-bOth : I .N - 17X I ellt ra k ana l sculL' l1onnall > entIre: .\X -74 s llbcaudals.
-'57
III
~ilIIlUdl' .
11I\\ lalld Impleal \wl alld and slIhtropical \\ellll(llllal1\: li)re ~ ls , halllbo(l filre ~ b . 1l1l1lSI :-OlTlIbfand:--. Illarshl's and swallll):,. rice paddies. hl'd ge~ , alld gardcn~ In suhlII'h,lI1 ,lre,IS , 1111'; ag ilL.. :-0\1 111 ~n , lkt: 1:-0 u:-oually I(Hllld on 11I\\ legdallun III Ihl' illlllledialL' Vlcinlly or fi lre~t <,Ireanl~ , thlehe<" pOllds. and ulher wall:r budlcs. II IS hOlh dlllrnal alld IHll'llirnal. SOIllClIllles lerrestnaL hilt llI:lInly ;lIh"real . f()lIlld rrolll rllily e\po~cd areas 10 Ihick \ cgl'latlllll . II keds chiefly Oil lii'ards. !lUI abll hunt:- 101 J'n.gs. 7 ()()
dry
It In ,ablb
1()J·e~I~ . lropicil
(iCllllS
1'.ltllIlII/()(~1'I1lI1· 1('S 1III/n.,.,t!{'IIIII.1
IS II eli-knOll Il li)r ils a!!!!re~~ile di "posili()il , "trikillg lierc.:! y alld hil In g readily and I Iriou,ly witell handled . Uislrihutinll
occurs Ihroughoul Ihe l'lItire rall!!l' Ol'lltl' genus. I'rolll c p;1I ,llId '\ :-0sam ( India) 10 China in Ihl' l1()rlh, and frLlIll Thailand 10 Sulal\c"i and n ore~ and Ihc Phtllppllle~ in Ihe Mllilh . ML Killllhalu : Ki;III, Sg. KiplIn g lt I 1' ,I'{fIllIJ/(}(fI '/I(IIfL' l 1'1111 '('1'111< '11111.1
(5XO 111).
Ps(!udora/u/;oll .J ,\ ,. I X():?
Key to the Bornean species of Pselldorabd;olt
'I he gellu~ l's(' l/(lol'II/,diuli I~ di:-.lributed i'WIll Thal i:lI1d in Ihe nurth Ihll)lIgh the rv1ala~, Peninsula to Ihe Phillppilles. SllIllalra. Horneo. :tlld SlIl;I\l eSI ill Ihe soulh . Slllall. harllllc~s ,nakcs; hody "lightly ePlllprcsscJ dorsll\l'lIlrally. hcad 1101 dl"lincl rmlll body : eycs sillali. pupils round : eharaetcri:tl'd by the absence or a prelleulal "hicld (present in I)selldora/,t/ioll IOIl;!, I(' ('/),1 I Sll Ihal (h..: preli'olltal borders lhe eye , 5-6 supr:tlahial s: anlcrior Icmpurals \I allllng: 15 1'(1\\" or SlIlool it dor"al scalt:s Ihrllughllullhe It:nglll of thl' hod y:
Tab. 52 J)I"lnoullllll of l\ cIlllom"dlO/I-'pl'ell'S on
1',\{' //{llIrflhdll!l/lIlh(}/lI11 IlIIli.1 ( i , '\, I III «.IXl)(,)
I',\ ellt/orulltlioll ('ol/u ri,' (\ll r" 1'.I L'/IlI"I'II""ill/l 1(1//,1:,11'<'1"
({' \
' ) 1 I~ II,
l (Ok,
IXlJ2)
I X~ 7 )
I'" ('IIr1"ruhd,IIJI ,'ural 'if(" c'''''' ,' (<';, 11 II ( ' ~ Il . I t)1) I)
Lnreal ~itie ld ahsenl : prcfrontal in con Iact wilh upper lahial s ............. .. .. ................. 2 • Loreal shl.:!d presenl: prefronlal nlll in con tat:1 wilh upper lahials ................ .. ..... -' 2 I'rcOl:ular presl'nl: inll'rna"ai;. nol ill COlli act with
upper labials ........................................ IOllgICl'I),1 • I'rcocular ahM:IlI : iItIl:rtIa"aI" in con lact wilit uppcr labiab .. ,............................... ...... ... ...... .... wl/llris -' \ 'lore thall 1::>5 \'l'nlrab-: ItIOIT litall 40 suhcalldal " ..... .. .... ... ... ... ........... ...... .... ...... ,..... ... 1I1/1(1IIIIclllIlis
• Lc's" litalt 12() I,enlrals; les" Ihan 30 subcallda l" . . .. . ... . .. . .... . .. . . . . . . ............ . .... . . . ... . .... . .. Wrlll 'lICCI/.I'IS
PSl'lIdorahtl;oll co/lar;s (MO(,{)1 ,\RI). I X92) IdfU/d"dl' n}lIar" \h )<1I11 \1<1 I ~l' hrllall ~
Rl\ l' l .
l }-i l) :'
T~lw I n~'alit) :
I t})
H \l rll l...·o
1'1111":. 17
1' '' ''I '/lOti, ,'1'''1', '1/1 S".I I (ll' ''. S~I\\.Ih . :'\101111 B(llll e ~.
!'.HHdoruhtfunI ,·"lIrII l\
h \ 1l .1n er INt d I( 8... 1 I l ~t,l li
& 11 \ 11 1"
1.11'1' I,,~alit.l : Il) (,(, , IOl 110. li t!. 2
\ I hI '
I~'hh .
Hortl~ll ,
Subah
Klllahalu
IInll'CI
S;Jnl\\'aJ...
K,llilll;ll1latl
• •
•
• •
• • • •
• •
Colubridae
Tax onomic c() mm c nts: The
author i~ not qllik convin 'cd that l's{,lIdor(fhdio/l cI'{'re!li is indecd a ~y non ym of PW' /I domhdic!ll cul/(/ri\ , 'I his probkm can. hOIl,c\cr, on ly hc ~olved hy the examinatioll or ilion: llIatcrial ~o that the nonll,;nelalural conccpt ~lIg !,!e~k d hy IM,HI <' 1.1 \ liON (I %(,), which slill rcpre~en l s lhe lalcst rc\ ic\\ nl'the sy"telllatics of these ~I)(:cies, sha ll Ix: rnllmved herc. L>ia !.: no ~ i s
A small , I()ssnrial snakc: lIIax , 20.4 Cill in length: di~tin gu i"hed from other species or it;, !,!ClIllS hy thl.: rolillwing c(1l1l bination : nolnreal shield. no preoclliar ~hidd. li'ontal ill C(1l1taCt with cye.
Dcsc ript inn Supraocular~,
Inreal", prenclIand anterior 1l.:lllp(1ral~ allscll t: l.:ye sU IT(1ul1lkd by I l' ig. 349 !'s('I/(!(I/,al>difll/ , 'v/larls from 'Vt :1I1I1I1(, Sah:lh lIr 2 Pll"tllculars. lhc frontal. I prcl'rnlltal , alld lill.: 1,,1 and 411 ' ""pralabial ,,: 5-(, "u pralabials, the 5. the la rgcsl. Genus Pfy Cls Filii (d R. I X-L\ scparaling the ~(lle posterior Iclllpora I Ii-< 11 11 the po"tlleulars: 15 dor'i;d "c; dc 1'<1\\'S along thc cnl llT kngth I'he g.en u ~ I'll'll.\' IIlhahll~ lar!,!e parts or ( 'Illna alld lIf the body ; II (,-13-1 \entrab ; ana l "CIIlC enlin:: 25-II sllheauda b (BR""( 01 '1/, \1 \ 1\)-1(,). u'iually 111 pairs. India. Tn the "ollth it rea c h e~ Sul:1\\ e~1. III addllilln to I)lIrsally blacki"h brO\\ 11. bladi~h grey, or hlack, iri thl~ . it occurs on 'l'\.eral Island" llr the rhilippin e~ . dcscent: a ycllnwi,h, \\ Ilitish , or red(h..,h c(1l1ar heLarge. slender ~nake~ : hudy 1I11"L'lIlar. ~ Ii ghtl) COIllpr.::",.,ed laterally: head long , di,tlllct rWIll body: hind the paril.:tab, Illay hc 1\';lIlling: n:ntral 'illk: lighter. e~e.., larg\: . pllpil~ round: d()r~al ,cales "l11omh or i\ Il:lllak: contained thrcl' cggs . h.l'l'led \\ith grll(l\CS, a di"linet \endx:llw\\' Illa! be prc;.en t or abs..:nl. \ 'l:nt ral" 1'()LIIllled ; tall long: Ec() l n~~ l EI hulu :.:~' Thcsc '!nall co luhrids f'a\'our primary and ~ce(lnd subc:llIlbls III twn ro\\~ . ary lim.:"ts where th\:y li\'\: in th\: laycr or kaf IItlL:r I he ~rl'cie~ of'thi" gellll" aI" di urna l :111(.1 truly ag.or lind ~h.: ltcr b\:llcath rocks and 1:lllcn tree trllllf"" . ill: . I hl'y arc l:llcoullterl:d In a I11l1liltude or hahi tats and dn nnt nl'Cc""arily a\'oid ellli l\'atl.:d arl:a". Oi strihutiull ;\11 int l.:resti llg. hcha\ IOUI' can bl' ohsl'rwd \\'hen the ,nakc" eross through high gr:b". In (lrder tIl obtaill This spccil:s IS l'lKkmie to Borm.:n. Vlt. Kin a halll : Kamhnrallgoh Road (II()-( (II ll:ct Illll) , a heller ovel'\, iew they' lili Ilklr h..:ads alld allll:rIlII' Iar~ ,
( '"Iuhl id
pllr!l\ln:-. '1I'Ihl' hlld ~ hit! h aho\'l' lhe 1!r(\lInd , ,\1 Ihl' III11C thl' ~ III~~ IO lldl y hy prcs~ llI g (lUi air. I hc s lla" c~ ui"11H: gl' nu ~ 1'/ 1'0\ kl'd on rrog~. Slwkcs. li:lard~. hird s, a nd p
Ta xu null1i c t:o ll1l1ll' nl : rh l' gl'n ll s /.UOI'IS {'OI'I', I X6 1. ", a ~ ~yllil il y IIli ~c d II ilh I'mll hy 'vV \I I ( 1')2] h) alld I 1\ I' 'I ( (111(,S) 1111" ;IITa ngl'll1l'nl has Ilot bCl'1l cOllllllllnly an'l'pll'li. I. \/1 I I l't al (Il)') I) , ba~l'd Uptlll a "tud y "rhcll1lpl'nl:t\lllorpholug}, placcd hOlh gcncra ill 111l' 'yllOllylll) ur a grea tl y c" panded gcIllb ( ·olllJ,el . 1\ l'l'llIdlng til \" \111( '11 (pers . cullllIl.). data 1'1'(1111 th l' ill lcl'lla l an:lllIlllY supP'1I1 Ihe Illl'rgmg "r/un( 1'1 l\'Ith 1'I\ 'lI.1, hUlntl t II Ilh ( 'oll/hcl: Wl,therc-
P/l '(/\ c(ll'in(lIIl.\
I'IJ'I'" l i l ,l l'l/ .1 1'11 '(/ 1'
("I,
I( j"
Ronl(, ~"l
.. , ... , ... .. , ... , .. , ....... . ............... .. ..... .. .. .. ...... , "
' ,I I .
COIT/ ,h"t!iJ lI / ,, \.( 11 \ (, I .... !l 11 k. t X:jX. II~
IX5X) I ~ IH.'
Ittl"a l ih :
1l1l11h: P / OUln/II\'('In " 1\1! 1 1l 1t) ~ I ~
l o .. t.~. \.1 7. \ 1"
l l tlNI I H IJt\;.."\ I« II, H I ' I ( 'X · ~
"I,., I 'P) II>
~· 17
Ta \()I1I11I1il' ClllIIlII l' 1I1 : II i~ olJ\' iou ~ Ihat SI1'1 '1I1 (, & IN(dl{ (1999) ~(l nlll s l' J PI.I'U\ lil,li ·II.1 \lilh I ~ , ·uriI/Ollls. sil ihat Ihc illlell'111aliun pn\\'ided by Ihcm is Icrt :!sltk hcrc.
"nl'I'n,1
• Alle\ellllUlllhel' ordol'sal scait: 1'O\\.s
• Le" Ihan I:W slIbcalidak dorsal !-o on Ihc l11edian ~-4 r()\\'~ kcckd ................................ .. ('ol'il/olll,1
•
IX 5X) IX, 7)
species of PO'as
An "dd Ilumhel' "I' dllrsal sca lc rows at Illid-\ll)d y
S.,bah
P~rttsfil s(,l/s ((it 1\1 1111{.
2 ~ . ",, ' ''' ' , I')'J I
K('y to t h('
"'",,,oaIH
ilrun e,
•
•
111 1 K. I X~X)
' Ill K,
Ao/'l'''' (S. IU,
iell'e I'e 'oglli /l' I '/I '(/ ,\' :\S including Ih l' gl' nu ~ /.({IWI',\' hel'c (Sl'l' a btl I) II ") & " ( I( ,I I I IJll(,) . Slll 'BI\I( ; & 11\101 R I 1')')1)) dllllhied Ihal/'/I 'o,\' ""I'm,\' II'lIild occllr Ilil 1l11I'l1CU :IS Ih i~ "PCl'1 l'S h:ld m:vl'l' bl'l'n con finlled I(ll"ahllut 100 ycars . '\,llwilhslalld ing, 1'/rIII/,III'm,\' is cUl1sitlcrl'd III Ih l' lull()\1 ing "l'\'.
1
<;""'" , '~
•
•
KaliJllanl ~ ln
• • •
(,OC, III COlllacl with the eye: 16 rnw~ or d,)r~al ~ea l es al 111Id-hody; dorsa l" slllooth : I X'\-19K ventra ls; allal SUlk di"ltkd ; 155-1 79 sLlbc alllbl ~ III III,) rows. Olive hnl\\ 11 to bn)\\ n dorsa ll y. ()ccasiollally 1\ ilh a rcd, black -bonILTl'd \'l'rkh ral 'Iripe and oblique bar~ : a blat''' lall' ral stnpe i'rolll thc Illld-pOrllon or Ihl' bod) III the tai l \\hich touchLS Ihc Illargi lls or the vC1l1 ra Is ~ Illd ::>u bcauda Is and widl'n~ posteriori y so thaI the la il Illa) he c lllire ly hlad ; 1Illd e r~i dcs pall.' ).: \1(1\\ . ( ) \, Ipartlll ~ .
J)ia g lll)si~
I';c ulu g) l EI hulu g ~
Most c;t:-; t! v re~,)glli7cd hy its latcral slnpes lin the poslerior portioll or Ihe hod y which arc 111 ,tark c()ntra~1 with thl' li g ht IClltral .,Itk. !'l1'll,l'ji l\( ·11.1 l!-o Ihc ollly Unl'lle:l ll ~ Pl'C I CS II· ith 16 1'll\V~ Ill' dorsals alll1id-hod y. lis l11axilllul11 kl1 g lh is about 300 Clll .
ThIS ~ n akc I ~ found in 11m and hill y areas wilerl' il inhahih \\c1 tropical re ll'es t ~, ,hrublalltb. and planlatilln~, II is usuall y found in den~c , tallgled vegct3tion in the illlnwd iate vielllll y or water. SOll1e illd iv idllal~ were t(lllllt! s\1'llllllling alld other, lip in trecs as hig h as (, l11 e tre ~ above thc gro und . Llttk is known ailout ib hiology. exccpt Ihal It ked s mainl y on amphibians: H,1I1 'IIH OR ( IlJ5X ) rep,111ed a specilllen feedin g on a sllla ll chidcn.
Orse ripl iun prcoc ular; 1-2 o;mall slI\)ol'ul ars; 2. pnstoculars; 3 lor~al s : '2 I 2. tl'lllpnrals: ') ~ l1pralahia l s. Ihe ,'i'''
( 'nlllhndac
F ig . .130 /'/y"" '/11," '11\ 1"10111
P Ori " ).: ,
Di stributi()Jl T hi s s pccic ~ li\c~ III I'cn111 ~ ular 'l alay~la and on Ilol'llco, Sumatra. a~ Ilcll a" on so n]l' of "IIITtlIlllding "mall cr is land ,,_ ,\ 11. h:illahalu: Klau, Poring,
C:;Cllll~
Thc
RlllIbtlophis
FII'I I (i l R.
I X-D
gc nu ~
Rlwhdo/lltil i" di ,,11'Ihlll cd frlllll Ihc parI:-, of SIhc1'la and Japan, ~(l ulh Ihwugh C hina . III 130I'llCO, I lorc~, Sulallc"l , and Ihl' Philippinc 1"la nds , 'lodcr'III.:I), hirg..:. slend..:r. aquallc Clllubl-id,,: dnr~al~ kccled. In 17. 14. or 21 I'\m s. Wilholll apical pil ": hcad slig llll ~ set off li'OIIl hody; cycs 1;llrI), 1001'gc. pupll:rOlilld; Iwo "ubslalllially cnlargcd l;lI1gs in thc rcar Ih..: upper jaw. which may lralls ll:r \CnllmOUS sa lil a inllllhc blolld slr":;1I1I whcll allowcd 10 hih:, Ikpcnding (In Ih..: "p..:ci..:s Ihc vcnolll \'aric ~ in I(l\ ici lj and Illay hal'(' a scriou"l y dan1,!l'l'OlI" clli:et Oil 11l1111ans. Inlcmasals hroad anlcriorl y. no~lnl ~ p(l ~ IIi(ln c d 1011\:1'ally; !:Iii long; allal scule and "ulll.:alldal" dll idcd , Thc d i- and Iloclllrn a l s n ah.c~ occa"l(lnall), ..:,h ibl l amphibious habil" , 1\11 ),PCC IC" arc ICITcstnal and reed I() r a maiM parI lin rrog~. Ihc lr tad pole" and fi"h . Till') rcprlldlll:c by la YI Ig cgg" , "O\lIh\\c~kr1l
or
Col ll b nda ~
~abah
IXJ7) IlIl11bdlll,h;v l'fIlI,' p ;I'iIlIlIl/,\' ((;, ~ ,", 1<. IXn) Uh llhi/ol ,h;v ..llI:r.\ I,rgll.' (S, 11 1 11 "
N!III"'/"I,!JI"
, .
'11/1111;11/(///1,1 I II/JIll/III II III ,.
Ulw"dlll ,h;\' /11 l1 r ll lll'IIS;,\
(S,
III'
", I. IX~7 )
(S ~ III1I . 1')2:')
At ka~t 15 ~p~l' I ~S arc c llrr~lltl y know n. Illllr will ch !t\ ~ Ull B nl'll~u. tllP un rvll. Klll ahalli .
or
Taxonomic cc)llIll1 l' nt : \Vh et h ~r Nftl/lldo/lft;, sl/hIIIlIliol/l.I' oeCl lrs on Bortle(l i~ at presclll lInd~ar. It Ila~ li slL'd by SI I' II II~(, (1991). but omitted by S II ,11 111\\ , & 11\, 01 R ( 11)1)1) II ithollt any Cll ll1ll1 ~ nl. It IS inc luded in the key bel ow,
I
KlI1ab,lili
2 No blm:k ~ tn.:a k • 1\ black ~ treak
the upper lip ~ .... c!,,:r .\'lIrgos 1i-<1I1l the eye acros~ the upper lip
" (1 1110:1111:111
• • •
• •.,
Rlwhdophis dlly.wI/X0.\' (S(,tl l (0( ;(
I.
l R37 )
/rO/lidnn Ull t' (hrL\r " ~rJ\ '\( JIll ~d I . I X-3 7: J 12.1' 1 I :! . li g (, &. 7 T~ p,' lu'''
B IlgllL
n 'P/Ur/OIlIlIll\ mrll'lI/fI/II' \;11
T)I.~ I ,,~a l ity : /i'O/IIr!rlHrll(t\ I
2 anteriur tempurals ......................... ................ 2 • I anterior tempora l ....................... .......... .......... J
'i .lra\\a~
• • I • • - • •
1%-1 .
Itn '.,':'<.! II\
III
' k
I Xt)Oa
1")
1:-('111> ~'i~ 1
J{nuu 1(1)7 , l)
1931 : 10 ,12, I
HINII \ lt p "-':. SII X, I{I'
~ .j .
Nairn (hl't ',\(II ,t~rI din \fI(~t1
"""h""I'"'' (I"T"").!,(, I'J~:
.Jd\,I)
lorc/IWIII'- ~hH. ·f.ll \UI>,
KII1:' 11:11111"" " Ikrll''''''''
.\:O/rIl .Iln'CI/).!,
Key to the Bornean species of RlllIhdophis
Brune i
lO. 1\1 ,,1-.'",
1)1
II \ \:-..
1 9~9 :
79, XX
I " ,II' \ ,I{ Ll' \/ ' k tl} ~2 . 1·1. \I " 1111 \ tIJ:-- ~
9.191'7 2X7. I')X<) I<)x . I'NI..-
.U. 1 <)1)2~ 1,1. 1'1')-1[, 2.l
s.. I" '" I 'N'I I ~ O H!JII"""/'/'" d')'!'.IIII,I:".I , 1\1\I
I-. 'It ,
I '),)-1 11
~ .j 7 ,
I <)<)IIl" 2'12
(lll
........ .. ........... . . , . . ,......... .1111>111;11;(/11(\ .1'1111111;11/(/1111
J Upper lips bri ght vl:I"lllililln ............ IIIlIl"IItieIlSi ,\ • 1\ ye lluw . hlack-edged ~ treak on the lIppl'r lip ...
Oi a/.: lws is Eas tl y Id.:nt i lied by li s particular paltcrn on the ncck described helmv. 2 anlc rior t e ll1poral ~ : m
... ,", ... ,.... .... ,........ ,....... .......... ,..... ('oll.\ picil/lItll ,\
,.. ~~..t
'"
£
~ If)
£,c.
Orsui plio n One lo r..:al: I (rard 2) prl'nclllar~ ; J (rar..:l y 2 or..J) Pllst Ol:lIl ars: X-9 supralab ials. Ihe 3'" to 5'10 or the 4'" to 6'10 III con tact I\ith the l'y~ ; 2 12 or 2 13 Icl11poril ls: Il) dorsa l ~('alc I"\m ~ at mid-body: dorsa b strongly ke~le d cxccpt for thosc in the lirsl row : 13.'\ - 1X..j l'clHral s (S It 11\ ' (jl "~ 1<)22): anal sc utc di vided : 3~- 1 0 I slIhc
Colubndae
i.c. yellow turn~ into brown . or red turns into brown . or g reen bccol11cs n:ddish brown: ve nt ra l ~ide whitish. margins o r \l: lltral~ ~11tllted With black. Juvenile specimen s exhibit the same markings as the adults, but more di stinctl y so. In addition . they possess three light spot s un the occiput: head dark. bright ye llow nr white stripe i'rom the lab ia l shields up ont( the nape, where the ~tripe ru ses \\'ith its counterpart i'rom the other ~idl' to i'orm a wide V. According to III I I '\A~ ( I ()-I9). spce imen~ rro m Mt. Kinaballl have a temkm:y toward s s ubstantiall y darkcr co l our~. ( 'Iuteh cs or .1-1 () cl;P (I () .5-3J .5 . I ::! -::! I 111111 : 2.22--1 .0(, g) ~e\ c ral times pCI' year. Inc ubat illn takes 5 1-6 I days, hat chlings measllrc I-IR-::!::!O 111111 ill Iell g th . ;\ ca~e (,I' /\mphi go nia retardata is known . Ecology!!:: t hul ug ~' RIIiI/Ji/o/lliis ('liIT.WlI gO,\' is Illllild li'om sea kvel lip
to abollt 160() 111 altitllde. but is morc cOl11mon in hili I'. rugged areas and m{)untainou~ rCl!,ion~ abme 5()O m. It inhahit~ lowlalld tropical wet ro rests. Iwp ica l and subt ro pical we t montane I;,) rc~ t s, bam boo 1;'1I'e~ I~ . I()rested mesic ~crllblands. marshes ;lIId swall1p~ . and occ urs .dso ncar human habllation ~. This snake I ~ often round in Ihe immcdiate vicinity or ~tream~ and olhn \\akr bodil's. II i~ hulh dilllll,d and Illicturn.d. mainl y terreslrial. bUI G ill sonh:times also he ~een ba~kin g o n loll' vegetal ion . It i ~ IIsuall y 1(1111111 in thic k. ripari'lil vegl'talion close to water. keding on amphibians. ~mall mammab. li J:ards and hird ~. T his ~ nake is widespread. ge n~ra l l y ~o lllm o n . and i~ onl' of' Ihe m ()~t common s nak c~ on M1. Kinabalil. II i~ eon<;llk red h"rllllc ~ ~ . but hil es ora l I line olh er ~ r ecil's or Rlwhtloll /til (i.e . R. /igrilllll) havc eall~cd ra tal envc nomatio ns in hllm a n ~ ~o that the prese lll spec ics II1l1st he con~ idl'r~d potent l
Ie,,,
-
....
------~
(M II ""l iS IYX 7 ). One s nakc wa~ di ~c\lVe rl' d in primary rainforest ( I'. . hl " ~ I (lS Il)X(») . threl' other~ alon!;! Ihl' Sg. Silall -Silau in Ihe Il ll'enllOn h(lur~, an d ,lIlolher lim: was observed ill dipt erncarp li)re~1 bet \\ een Sg. K1J1LlI1git I I and Sg. l.alll;an'lIl in thc afternoon ( 1 \1 ""I ~ 11l1l2a) . Dis tri huti on rhi~ s pecie ~ ranges from Chllw In Ihc nOl1h. sou th tn Ihe rhilippint:~ and Sldall'e<;i , 1\ 11. " inabal.. : K iau. Sg. LlIlganan, Lobang (II \ \s IlJ-IlJ), IlcadLJu art cr~. ballk or the . g. L,wagll. ea~ t or Ihl' Sg. I.iw
RlllIht/oplzis colispid //all/ s (Gl ; 1'111 ' 1{ , I xn) IS 7.'!
Tn'I"do/loIU\,COII\/JiclllaIU\ til '111It{.
",t.)(l. ri ~
t\ lat.lI1g. S~lra\\.Ik. HorlH. .'~} In JllldoJl(ltll\ (Of/.'fN( 1I1al,,' _ I I " I '4 II I ()OOa 70 N/whdol'ln\ t OIl\l'tC'll/alll\ :--, 1I I1 HI'4 I ')q I 1"0 , I <)<)-Ib : "47
4
I)(ll'
IIIl'a l il~ :
I
~ 1 \1 ... \ 11 ...
dangerou~ .
The ~ n a k e \\'a~ rOllnd on rog!;!y day~ on hl'dl!,l'-like Ihidet s or IJil l/eris and (j/('ic//('1I111 krns wherl' il prol'l.:d to he a ~kilrlll climber (M \! " ~ Il l , 1() X5) . Several spec illlen ~ were disco\ ered lin roadside~ with a vcgdatinn or grasse~ and rcrn~ (I-Icadquartel's). (Ill grassy. llIo rl' s UI1 - e\pll~ed spots in Ihe rainrore~t !lOI 1'; 11' i'rolll Ih e Sg. Li\\agll, and on a fore sted ~Iopc rar i'rom II ater easl 01' the Sg. I .i\\ ago
Di ag nos is I ~ asily idclllilied hv ih particul ar head p
a: ~)
y ,l lhroligh :;"', ,Ir Ill< :I ' :lIHI-I" , III conlacl 11'lllt lite cye; I'> rim ~ ur keekd dorsab al mid-hody, IOll'cst .. 0\\ Ilia), bc S1l1oot lt ; I J(,-152 IClllrais: anal ~cll ie t!i\'id c d ;J(, - ('()~ub(alldal~, ill palr~ 11l1 R' "111 1')1 7, III 11\ " 1<)-1'1, '-;11 1'" 1<01 1( 1922). 111\)\\.n or rl'ddish hnmn ahml', lI'ith a dark Ill't\\ork and IwO IUllg iludlllal ~l'nc" OrYl'II"W ~pots or Irall,ler"l' linc,: Ill:ad II Ilil ~m;ll l )'e llll\l, dark-edged , p(ll ~< a ydhm , blad-edged streak lin the IIpper lip. alluther i'nl1l1lile cye 10 nlrllerorlhc Ill(lulh: usual ly a pail: Iran" \: I ~e mark 011 thl' Ileek. n
Thi " sll :lke 1'; UIlClll11llHlIl in ,omc an.:,!". and COI11111011 in lither .... It i, reg:lrded a" Itannk,", hut it s hite IS act uall y po icnt ia ll y d,lll!,!lTOU", Di,t ributio ll specie" Il\ e" III I'e llill'iul ar I\-bla ys la . on SlIIlIalra. l3o rneu , ;11lt! '(line ,,111;lIler Indon es iall 1;. land " ,\11. Kin a ha lu : ll e;l r '1III 1')OOaL Sg. I\i
Thl~
Rlwhdopilis IItllrtu/l!lIsis \1110 \ 1IIU1
Thl" s l1;lke i" Illlllld at I(m del atiol1s II here it Inhabit'i tl'opieal Il l't lilll,,,t~ . I iltk i" kl1l1l1'n or It " hiulogy. One "peCl lll ell \\,;1;" co llecled rwm thl' dry it::lI' lltler III the 1\lre ~ 1 Ilun r ill S,lrallak (D.II 1118:. V'II" I il)l)(» ~tl It i... proh'lhl y a lerresl rial Ilx est\h\t:lkr. Nltllhdtlf!his (tl l/\/" cil/(/l// \ i" act il c at ni ght. There i, 110 illformatiull ahout it ~ diet.
'\\11 111 . )9"::"
IllIlIlI
,\(flf'/\ f1Surlldl ' , I\'I\,
~21 1) -),
E CHl u g~ ' / Et Iwl ng)
ftel.,,,,,,,
i\"nl"". -;-;()[) I
HI \I I
\I{J)
I) 1)\' IIII'"lil.' : \tl f\ l lIIlItl.
Itn( 10. 12. I (' X..L .2
~ 1'1( . lq \
~(I . ,2,1>1
11 ,\,' 19--'l)
~
UIUl"d"l ,hl\(1111H Jr!~II . \1 "1111\ & 1)1\/11< 11)~<'
f\ 1 \N lilt \ I') t"~' ~~ Nllllhll"/ )/I1\ IIIl1rlld(.'II\/' (\ 1 \1 t...\!I1 .... 1I)t)-lh - 2-P': " I '96h. h" , :-"11 IU ,\, ...\:.. 1r-.i1 d II 11'1)1) 1>-:-4 \ '0'1'1 \ (Ill"
1(25)
leT I
'-'\11111
&
(Sf\llrJl.
1 2 1i~ ..
II
......... d. } 1It11l "lld" " " , .
( ,I I< ..\:
1 ,'\
l)ia)! lIosis Easil~ recog lli /ed by it~ hright rcd upper lahi;t\ ,hic:ld~ , ()Ill: antenor temj111r;t\ , Max . kll gllt ;Ibou t I 00
CIll .
I)"suipliun Olle lureal: I-~ preocul:lr,: .\ pn~t() clI lars; I 1~ tC11lporal~; 'i ~lIpralabi ab, lit..: -+ 10 ttl (l'lo tUliching Ihe l')'C ; II) dor,al ,ca lc: nllls al mid-hl)dy. dor,ai, , trol1glv kelcd c\cept in thc 1011'e"l roll' ; 17()- 1X7 \entral :..: ana l ~c nll' di\ ideLl: h3-() 7 subcaudal s in two row~ .
Dorsilill grey \\'ilh ,hOI1, illliislin ·t, blal'k crossbar~. a ro\\' of light spots a~S()ci;IICd wuh the ends llrthcsl' dark Illarking" appcars at mid-hod y and rlln~ dOl\ 11 Ihe back, convergi ng 10 a singil: roll' :tntninr to the vent alld eXlcnd,ng to the end or the lai l; head lind nape broIl 11 , LIp per lip bright \-erIni linn: eye brown, with a light grecn segmcnt in lhe uppc:r portion ; pal1 of the IH.'ck bright redd ish orange WIlh n;gularl y arranged black spots; hclly
( 'n luhridae
grcyi~h
yelll)w, with Itll1gltLllllnal ~crie~ of ~ma l l hlack spot~, b~st marked to\\'ard~ the oilier 1I1argin , urthe ven tral ~: tail dark grey bellm , I<:c ulo g ~ / 1::1 holol-,)'
Di strihution
1/II[/'I/{ / (,lIIis ha~
bel'n fo und at elclations hetween I ()()() and 2500 111 ahm.: ,ca k\'<:I . It thu~ app.:ars 10 be ~ tri ctly slIb1l1ontanc. On~ ~peci111cn was kept al iv.: o n a diet or rrngs . Thi~ i:- the only s pecie~ of snake whieh ,I,ccmb up into thl' rog fi1rest reglllil of IVII. Kinabalu . ;-.Jo further detail , ar.: avai lable on it:- bIOlogy. R/whdophi.\
Genus SibYl10phis
FIII
I (;)
I{,
,\'ibYIlOph is meilll1ocl'phll/fiS lIIl'ltllloceplw/fis (Gle\)'. 1R35) 11 COr!fJllfIII1I1f1Ut(ph"hl\'( I H\'t. l ~ .~ ) llll
~I '" " IIlJl a"
11\
I. 12h-IN1-)) 1"
'T~ IJl" IUl' alit~ : '\\1I l!l' ~n: 'iUb"'L!ljlh.·I1I1~
h)
(!l IN tl ll
~,
d"'· ... Ig.lldh..·d
K (IX"Xii 2X. III ",'rr~l l" alld " 'illlgdpOfl"J"
il"" , " " I< II , ')1"
IlIlIrtU/t'IISi\ i ~ all elldelllilt: of H01'l1eo. lIere, it lias di~elllercd (lull' nn Mt s. I\lurud alld Kill uhalu . :\11. h:inuh"lu : Ki au, l1c a dquaner ~ (M \NIIII \ pcrs . COIllIll .) Kalllhorau goh Road, SU111 111it Trai l (250() 111) (K t'\ ()\\ II ' ~' per:-. COIllIll .. at tlie ,al11e tilllC r.:pr.:sc ntin g the l11a:--illlulll altitudinal record for reptilc~ on MI. Kiuaha lll ).
N/ltlhd(ll'lJi,
pupil round, I lorea l s hield , aud 17 rows oJ'dorsals at IIlld-hody. Its co lour pallern , dc,cribcd helnl\, I, al ~o e hara · Icri~tic. MOl" , le ngth <)l) Cill .
1843
I h.: genu, Si/~\ ·/l of ,hi.\ ha s a \\ Ilk-rangi ng di:;tribution, ex tc nding fi'o1l1 Sri I.anka and thc Indi an slIhclllltinen t, north to Nepa l. ce11lral, easkrn and ~o uth nn China , Japan. through Suuthcast i\~ia. and thc Philippine,. Slllall. ,knlkr: head moderately di~tll1l:t rrom body: pupil s rnund ; 17 ro\\~ of do rsa l ~ a t m id -hody , snloot h. II ith tubercks, blll \\ ithollt apical pit:-. XI() supralabial s, the 4,10 10 (" I, touching the eye. Ana l ~cllte and ~ lIh calidals dil'ided ; hl'mipeni~ entir.:. (irllllnd-dwe lling. diurnal ~ nak e~, tlll:ir di et cOl1 sisb prilllaril y or li 7a nls, hilt al~(l in<:ludes fro gs and in~e c t s . Th.: specics or thi ~ gc nu ~ an: o\'iparoll'. Altllgcthcr six ,pecies arc kllO\\ II. (111e occur~ lin Uorneo and nn I'vll. Kin ahalu .
fi r.2
I're'lllllably ol'iparou<; like the otlier <;PCCICS ofthi, gcnus. nolhing preel~t: ha , a, yct bCC0111C knll\\ n ahout tlie species' reproductil c hlllillgy.
Description (['Vl {)I(I, ,\ t'\ IlJ7 J) Nas,d ,)(llible. no,tril in -hetl\cell : I prcoClil ,lr; 2 plls((l e ular~; I loreal ; X-IO :- upralabial ~ . normall y the 4,1, to 6'" touching the orbit ; 2 I 2 telllpllral ,: xI() sublabia ls; 17 1'1)\\'<; of dorsa l, throughoul kn gth of Ill)dy, ~ 11l()oth; 12X-ln lentrab ; anal scutc di \ ided: X7- 140 subeaudal s. I \cad re ddi~h hrllll'u . oll\'c hro\\ n. or )'e ll\l\\ i, 1I bnm n. 1\ itll well -dcfined, ,1I hci rcular , pot, nrJight pig1l1en t rando1l1ly di~tr i hutcd o ve r tli.: ccphalic scales: lahial~ with Ill' lI'ithout darkcr 1l1ottling: nllchal arca reddi~h brown. nl' :-ello\\ isll hroll'n , lI'ilh li ght. lalcral hlnl c h e~ or a ~ trcak : dor,al a~ pect ofhody :lnd tad rcdd"h bnllln or olil e bnlll 11 , lI'ith light ~Iripe:- o n the uppe r hal f o f dors,d ~calc 1'01\' 7; uSliall v with a scries ofal tcrnatlng dark bars, taper ing tn ~poh on thc tail. c:--tending int n the latemllight stripl's, Chin. thwat . l e lltnJl ~. and ~ lIbc au dais vc llo\'\': a ~erie~ of dark ,pots at thc edges of the velllrais alld ~ uh c al1dal ~; a mil' nr mid-I ~ntral Ileck ~ prescnt (I I' not. (h iparnu, .
I XII)
\,h I '''''1''11 \ ,t:. I 'IIiIlUIIII\ III R ( Iit!! It) I ": ) 4. lig t! : 'i \ 1I 1I 1 I In I HI. 2:'. ")2. 1\1 \'J IIII " 1l)~J ~(I .\tln '''illill\ nI( ·/( lI lnc ·
SllIall sk ndcr: head rnund . hardl ) di ~t in ct; be~ t Identifi~d by the I'nllowillg c0111binatlon of traib :
I::rulug~-J [thC)lu l-! ~
1 he ,nakc i~ fi)l111d ill low land area~, but it I~ pnncip,dl y an inhabila11l of hill y and nllllllltain()u~ regIOn " atl'lclatillnsupto IO()O 111 . It llccur~ In 1011 !and tropical wet f i.l rc ~ t ~. twpica l and ~ ubtropical \\'etlllOl1tanc f i.l r~'t s. bamboo ti)re~ t ~ , plal1tatiou~ , and in cullll
( 'olllhridal'
lerre ~ lnal snakc, prc : rril1~ lan).!bl n:gClatlOn along nwrs, nH\I~1 IHI\he .~ alld dll111p;" of bamhoo. II i~ (hllrnal ami "ecm, 1(\ kcd un rllre~t floor ~kil1k,. frogs and In~cl'h. c"pceially g ra~"h(lpp c r~ .
(iClllIS [) 11\1I lUI
Stegollo!flS
& J3l1ll{()N, 1X53
I he !,!e11l1" Sl<:~OIlfJ/II\ IS nati\(, to Ihe 1~lal1d~ ~lr lhe Il1dlIhcalidals in I- ~ 1'0\\'''.
NIll'turnal li }:--~orial.
Ii i:.:. -'55
lern:~lnal "lakc", tllev arc partly SC1111 They arl' lillll1d in a mllititllde llfhabital:--
S" :I!,Ol/olW h",.l/( '( ·l/,,'
rnll11 the Ikad4l1ancr".
Ilistrihuli un II!. 1I/1'/11/1f)('('/ J//(//1/1 I~ di'arihllll:d li'0111 thc Malay J\; nll1~l1la III the north tll "umalra and Bllrnco. "II. h:inahalu : K iau .
.)'ihl·J/o/lhis
ranging ('fIllll raill I 'orc~ts to cultllaled lalld . J he ~ Ilakl':-- nl' lhl' gc nus SI<'gIJII(J/1/1 ked Oil rcptile eggs, s111all rrn~~, li/ank ~Jlakl's. and ~mall 111;1111111al" The)' are (l\ ipamu~ . prodUCing ellllt:hes of (, 10 12
eggs. ~pc(ic~
''''n (111':--
hmc heCIl described, onl' Oil Horncll ;lIld lin 1\;11 . Kinah:llli.
Stegollotus hOI'lll'l!lI s;S 1 \/t','.!.tilllllll').OJll j,U \I\ h,C .lt~
,pccic~ 01.:-
(" ' 1(,
1967
19()7 X2 I ~ P l· I<)(.·ali.): '\bllga Ilmd 1)1\ ,,,,,,,. <",11 ,1\\", \/' ",!!,O IlIJlt/\ /'0"'1('( ' 1/ ,\/,\ " HI 'II\PII ,\:. ~I(I(.I~ I \ \l)X..t : 24 : ~I \I " , , , I ' I 'Is) III. I'iS 7 2>:1< . I'nN I')~ , 1')')1" 1-1. 1')'12" I '.f 1994b 2YJ.2 . .17 . 1 {"J(\\.' . 2"2.~II II II \(, 1l}l}I ·J.f 7: JNt, IH & I " 199(lh 70 , \1\' 1111 \ & (11( 11"...; r..l \" 1')97: 3 ') . ~
"'k.lllI . """,I
I
),,1, Ill.
( ·o lilbrida..:
Dia gnosis Uni colourcd , allllost dark brol.\1I1O black: pupil l ertical -elliptical : Eac h I,entral sca k \\ it h an obtusc, latera l ang le : vcnebrals cnlarged . ~ lax . totall cngth 130 em. O CS lTiption
Two prcocu lars: :! pos\llcular~: I lorea!: 9 supralabiab, the -l" and 5' h in cont act \\ ith the eye: 9- 10 s ub l ahia l ~: 2+3 tcm poral s: 17 d llr~ ab at mid-body: vcrtebra l ~ ..:nlarged: 194-233 v..:ntrals : ana l ~CLltc ..:ntir..:, (,O-7IJ subcaudab in pai r". Head and hod y greyish black dorsa ll y and late r;lIl y: s upralahiais grcy with a pinkish till ),!.":: I.: hill g r..:y with a hint or brown: wntrals lig ht grcy, ..:al.:h with a hhldi sh band anteriorly, ( )vi parou s.
(iCJ1US SI()lic~kill
.It wo!\., 1870
The gcnus 5;/o/i('::kill is native to Sabah on Borneo and the Kha si Il ili s In Indi a. \~ o dcrat c l y large. hannlc s~ colllhrids. I kad dist inct fi'Olll budy: bod y slender, I.:ompres~etl latcrall y: eye" small. sometimcs prominent. pupi l rOlllid ur vertical-cHal: no s trib pointed I'llrward : tai l lo ng: Dorsa lb ~Irongly keeled. c llipti cal. juxtapo"cd . increasing in "i/c 101l'an!'" thl.: ICIlIcr. !\ :-.ingle na"a l "cal\:. Velli rais \\ el l-developed: subca uda l" entire . These arc terrcst ri al snake" wl1l)s,,: hio logy is cntin.: ly lInknown . i'wo species haw been described . nfwhil.:h lIllC lil es on Borneo and Oil /VII. Killaballl , 'I':nol1omil' ('OI11I1U' II. : The gencri c l1amc S/o/ic::/, III .
F cnlog) / i:: t IwhlJ-:Y T hi s ~pcc i es li,es in the ralll I('re"t~ and i" pr..:dominantl y tCIT\;~trial. \11 \I "'~llIS (1994h) found a sp..:ci ll1cn II'hich lIas climhing conlidl.: lltl y ill branchwork, Vv' hcn it felt thrcalcncd it stru ck out quick ly. hut did nnt att e lilpt to hitc . During it s tl'nll sport it str;lIlg kd an 1'.5 Clll largc (l'I'/(}(/lIclr/{I,\ hll/II(,lIsi.\ II hich call~cd the ":" pu lsilln or two ..:ggs in th..: lalter. /\nother, ~e mi - adult "PCCiIllCIl Ila" dl sclll..:red at around 1'.I)() p.lI1 . as ii crep t \\ IIh I cr J ~ I ()\\ 1ll0ti llll ~ over thc forest Ilnor (~ I II "'~III'" i')~N) . When it wa~ pic ked lip, it contrackd 1 1rm l ~' around the halld alld produ ced a sccn t reminiscc nt or a hill y goa t. One Spel.: illlell wa" located lx'ncath a "un -c ''!lO"ed ~ I ab or roc\.; on a gra"sy rnadsi d..: ahm'c;1 tricklc o r spri llg wakr (1\1.\1 I-- \11 ,., 199 I a, 1992a). Whl'll uncllIcn:d. it hid Il~ head unde r its body coil~ . It I\a~ due to "hcd \ erv suun . ,'not hcr specililcli 11';1''; roulld \\:ry actlle at about 'I .O() p.l11 . during a heal ~ tll)l\ npour (1\;1 \I I--~Il l " 1l)l)6c ).
dcseri hed b~, JH
SlOlic~ ki(f
bomeellsis (B( H 1 1·1\( tI
\ lllll, -~UI(/ /'tHllt't 'Il"\
Di s. rihution Thi~ spcc ic!> h;I" to da tc on l) bel'n rouilli in Sa hah and Sarawa \.; . :\11. Killaha lu : Hundll Tlilian ( 13 7() m), Sg. Si lall -Si lall ( 1-l50-15()() III L Il eadqu ,1I1er" e;\. it (151)1) Ill). Kamllll rangoh Road ( 1500- 175 0 Ill). goll' C(lur~e . Mesi lau ( 1750 m) .
~tl
H, """,h.,t". t!"rlll'''.
\/lJht ~ h(fttl
1')(11.1
11 \\,
p>x..t " I
"01'11(', '11\1\
~7.
1~) 5 1)
,,,
R ( III. I
\..\.
PH :
I' t
199)
I'~
I :-.:tJiJ. 1'\2
~~IJII Ie 'C 11 \\1 1 .... , 11 IqOO"
:'-'0, f\ 1\\j IlIl \
~ . L S II I B I NI,
I IU "I,jt,
II II \;1 i ll{,
..t " h g.
ItJ X' \(I . '29 . \1
Ill'
-:'0,7 1.
"'1(, . '\ \ 11111
1H9())
R,
I Hl'alil~ :
""!! I
I,n l
II)I{!)
,~. III
II II "\~ I'\\ :--'IIII, IH ' \ 1 ""1("
1t)·J
'h
~
I It Il)l)l} : ... -.
\'0111_"-/.,1 '0111( '"",,, \1 "'111\ ,I\.:
\I <" ,,"-I\NN
11)1) 7 ,l}-l
17
( 'olllhridae
Sloh,.-k'll hOJ'//( ·, ' II.I;.1 Ii i:.:. -'56 (abll\c) I'w ill ( ·r"cl..cl Rall ~ 'c N I'. S'lh"h Fi:.:. J57 (len, tiOll 1 Ihc tlc
Dhlgnos is Readil y recog ni/l'li hy ih l'xposl'li ~kin bl'lwCl'n ~ "IllC lillrsal sc:dcs . ,\ row o['small sca ics heIWl'l'1l the prcl'rollt , d ~ and the frnnlal~ alld sllpr"lll'ld"rs. Max . tutal length 75, Cill .
pi inn \l' ry narrow ; prdi'ontab large, scparatcd by ~ Illallcr sc:t lc~ rrolll the fronlal and the slipra ()clll;lr~ : 2 PI' J prl'(IClllars: anolher 7-X sillall scaic~ Illay (\ccasi(\lIally ~ lIrr(\lInd thc eye: noslri ls large:: I IOlcal: 10- 1 I ~ lIpralablak nOlle oj' which. or thc (1 ,10
and 7 ,10 • inl'onlac t wilh th..: ..:ye: : klllporai s very small : dorsab ill 30-35 rows. parll y sepa rated I'rolll c;I '11 olher by III1Cll\'cred ~kin : 20-"-~ I(J \ clilnds: an,,1 ~cllll' l:lItirc: 124 undi\'ided , uhe
J) r~{'I'i
IlIl e rnasal~
Oistrihutiull This spec lcs is cndcmie to Sahah alld has on ly been round on Mts . Trus Madi and Killabalu and the (·rod.L'r Ran gl'. 1 \,11. Killllbalu : 4200 fCcl (I h t\ II ~(,1 1 I()()Oa). Vic:\\' Irail (1600 Ill) .
( ' (llllhrid~e
Gen us Xelleillplris Gi'l\ 1111
I{,
I g64
The di~trihlliion of Ihe gCllll" .\ ·(,II ('/aphis eXlcnds from Myanmar and the Ma lay Peninsu la to Sumatra, Borneo. and J~va . I.argc. slclltlcr. harmlcss ; body muscular, round : head dislincl from nape ; eyc fairl y largc, pupi l round ; dor~a l s smoolh. in 17 nw,,~
/\s f~r a~ is I-.nown 10 date . the ~pCl: i es or thi s genils are ()viparoll~ . Two s p ecic~ have heen deseri hcd. hoth inhahil Borneo. and nne ()cellr~ Oil MI. Kina ba ill .
Key to t he Bornean species of Xell elaplris 140- 179 slIbcaudals, brown abovc with 1110n; or less disti nct hlack tra nsverse bands anteriorly .... ..........................................
streaks, separated by cream-coloured interspace!> .............................. ...................... ......... . ellipslfer
S3hah
•
XI'II l'illl'it i.,' I'/li" ., ifel' H," It 'h,1 R. I'JOn . \ "III'/II/ J/II' "('II/g m/IJIII., (C\~I<'R. IX'l 7)
1/(' \{/K() /I 0 1IlS
• 134 subcauda ls. body with I X black-edged brown
Kin aba lu
•
Brunei
Sara\\(lk
• •
Kailinalllan
•
Ta x()lwmi c CUIllIlH' llt : SJ( I III (, & I (,I \( (I'J')'» dotthted Ih e occurrence or this species on Vl t. Ki nabalu. but failed to pn)\'ilie :1 rca~()n ror their doubts .
wilh the eye: 212 temporal s; 17 rows of dor~al sca les; I X(,-20) \·entral s; anal sc ule divided ; 12<)1)-1 slIhc:1udab. ('rcalll-co loured ahove, wilh I X 10 20 large. dlipti calor round . blac k-bordered . brown markings; side~ cream-coloured. ~ Ii g htl y ligltl cr Ihan the dorsum . spotkd or marblcd with black ; head and nape pale brown. sidc~ of nee I-. with interrupted. hlack. dongatc marl-.ings; ttpper lahials ye llow ; pO~lcrior portion brown wilh a black latera l siripe ; lower sidc uni forml y ye llow or crcatll -whilc . Prohabl y OVIparous,
I)i ag nus is Dor,al sea Jc ~ smooth . ill 17 rows at mId-body. /\ large species tllOSI casil y rccngni/cd by thc eo lnttr pal tern described below. Ma" . lenglh n.:corded "II 1;11' 232 Clll , hul it Illa y cven g ro\\ larger according to I WI' I IItt · ( 191\)).
Ecol ogy/ F.I hol og ~ T\\I 1·llIt (I<JXJ) Slated that onl y ~I" SpeCltllCn, wcre known . Vlo~1 ofthetll ~ee m to ha\'e hcen eallght In m()Utltainou~ fi lrest an:as ht:twet:n XOO and I 100 III above sca levd . Nothing else IS I-.nown ahnut the bIology or this apparcntly vt:ry rare spccies.
lkscriptio n One prellCttJar: I largc ~tthoctt lar : 2 postocu lars; I Imcai: X_C) ~lIpralahials . either .;eparaled from the eye by (he subocular. (It' Ihe ,10 or :'i'" in contact
I)i sl ribu tio n ThIS s pcci e~ l i\'c~ on SUIll:!lra. Illlrn eo. and In Pl'nillsular Malays ia . :\11. Kin a balu : 1I'llhout Jlrcci~c localitl cs .
X elle/aphis e1lipsUer HOI JI.I' I\(OI ·IC I lJOO 1',,"e/lIflIIII ,,1111,,11/"1 11111 '" ~(d K. l'IOOa . IX·1. 1'1. '\\ 1
'1'.\1'"
IlI l'a lil\' : '-;ar,,,,,,, nvcr \ ('lIdlll,III .\ 1'1111"1/('" IJ! R. H III 11)1 7 11),.1. 2·1. SIIIIII", IIJIli ' .12'1 ' " " "
8:
t)'\
I 111'" \I{P &
\I \1~M"
" I( 'i
.1< 1 ,
1'1'1-1[, ' _-1 7.
h HoI k 1'1'1'1' 79
t 'oluhndae:
(;~llliS
XelloclIroplIis
I X()4
Gl 'l\'l lll ' R,
I he: g': llll ' \ '('IIO"/II'IIIJhi, i, nativc tn the rq;ioll bd\\'cen Pakl~t ; 11l In Ih.: WC'I . China 111 the north alld ea~t, ~'Iuth 10 Sula\\'.:sl. MOt!.:l-al.:!) large:, hal'lllle:-:~ \Vat.:r :o.lIak.:~ . Bod y I1ll1seular, roulld 111 crO ':o.-'.:ction: heau IOl1g, l1ari'(m. di~lllh.: t from hod y: l'YC, large, pupils round : 110 ~lI h~tal1l1all y e:nlarge:d t<':l'lh : dun,al "ca les ill 11,)17 r(l\V~ at Illld-hlld y. 17- 15 a Single head-length anlerlor tll the anal scull' . I)or~al,; I.cckd cxccpt for the IllII est row~: I clllral, rolilldcd, lai I long: allal ,c llic alld ~lIhc;ludal~ dll'ltktL the latter at least pal'l 1<11 I). I\ith pits: hemipeni .. lilrked_ The diurnal. highl) aquallc ~nakes arc rel10w lled for hlling n:adlly. I'hcll- diet l'lll1SIQh of li sh alld li-ogs and their tadpoles_ The:y reprodlll:<': hy layillg eggs, parll y ill lar!-:e IIllinher, .
I .\'el/i!ch,."phi.\ 1II1II'l/llIl/n' I I
1>lIIM.
I X(,-l)
.\ 'I'I/(}(·III·OIJhil· (rial/gillig I'm .\' (15011" I Xn)
XCI/od/rap"i.\'
IIU1CulafliS
1I'rIINdllll" fII ,\ 11111 1 II/fltlf' \1.lrtaplII,J
I
/"UI'I"Ollulll ,\ lilli e Ulfllll\ ,
Xofl" V(l11
n
lIIelt (!lIl
IXld.t : 0'
1)l'l' luca lil ):
1l ,lIIdl'.:t 111,1'.111 H OIIIl't.l
IH':. II
1'1I11:r '\1\.
IIII t \(:.
(EDt· t.tM I, I R(4 )
\h w ()II.\IHI I X90a
IJ }:.
SIIlIII )K I~
Roo, )! 1'1 17 1/(1 ,,/a/lt,\, ~Mtlll IlJil . '2 1'1
.... l ·t! )lHl/r IIlulo, '" \N 1111 \
Vllll(\ 111(/( "Iulu
I
UI NI I \ I~ 1t l'\: '-: 1C u, l< 1 '\
\'1 ' 110(' /"01'/11\ IIItllU/,III(\ , '" I t I 1\1\ \ I
pnn 1X It):-.t... · 24.
I ()41 ') 1. ~ 1 \1
t\f\H .'
1()i'· ~h .
2·17 r)ja~lIo~i~
Moderalel y large: I11l1sclilar: dorsal ~calc~ keeled . I. ye VCI') largc: ::! anlerlor temporals prese nt ; no .. ub~lanliall y cnlarged tceth . Mosl easil y idenlilicd hy thc ~o lollr palll'rn de ~enhe d helow. Ma'( . kngth about 100 elil . Dt'scription I·.yc wry large: I loreal : I preocular; 3 (rarcly 2 ,II' -tJ po,toculars: ~ - 3 tClllpora l:o. (rarcly I 13): I,)
370
T'n .. pecle~ arc kllOll' IL 1\\11 ol'which liH' on Bornco , both also Oil ML Killahalu . TaxCllHlllIil' fUIlln1l'l1t: X(,III!c/1I11IJ/ii.1 flisr'lI/(}/' was Iisted by S II I Illto-' , ( 1')') I) as a IllIrd specie, or this !,!CIlII, li)r Borneo. but \\as thell again rCllloved from the list of nat II'(' sna ke ~ 11: S II I 111M, &. I (01 H ( 19 N) . The lattcr eon li1rllls with the author's opinion .
Key to the Borncan species of Xellochrophis All ventra ls wilh dark margins: a dnrk lateral poillt Iht' ventra ls; upper side with no black Iriangk~ and no red laterally ............... ............ IIIfll'lI/lItllS • Ventrab laterally with altcrnating blaeJ.. margills. rorlll~d by the black lalcra l I riang l e~: upper ~idc with lateral black Iriangics intcrspaccd by red ... 011
................................................... (,.itlll~lI/i~I'/'IIS
<;abah
Killabalu
BnlOel
"iara\\ a"
Kalillllllliall
• •
• •
• •
• •
• •
(rarely K 01 10) s llpralallial ~, Ihe -t ,l, 10 (, '1, (rarel y Ihe .\,,1 10 5'" or the 5'" to 7'" ) bordering the eye ; I I) dorsal ,calc ro\\" al Illid - body. all dllrsal~ s ir,)n gly kee led: 131<- 156 \'entral:-. : anal scute di \ Ided : X4 - 1 17 suheauda Is (h ,," ,\1).\ I ~()-l). in 11\0 rows . Olive 10 brown dllr"dly : -l rol\~ of' small. square, blal'k spots, Ihe interspaec~ lilled wi th two longitudina l r()\vs of ycllllw spot s: supralabial s y cllo~" with blad margins : head oark above. wi lh black spot s: lower sidc1' yellow. thl' \,Clilra l, hordercd with b l a~k anti a blaek dOl lalerally. Oviparous . F:cnlo~.vlF.. hology 1//(/('11/0/111 IS a snake or Ihe pla ins, inhabiting lowhnd Iropic.d wet rores t;;, marshes, ,wal1lp~, plantalions, and rice paddies . II is invnri ably round in the vicinity or I.v akr or in wet areas. It i~ dillrnal. h::rrest rial. and semiaquatie. Little is
.\ '('I/(}c/im{l/iis
Coluhridae
~
Fil! . .lSI!
\"J/{whrll/' /II.\ /II(/( 'II/a/IIS
frolll SUlllatra .
known about il~ bio logy. other than thai il fi.:eds llil frogs, toad~ , and li sh. Thi~ snake i~ mild-mannered ami docile, very se ldom attempting to hil e.
Xellochropltis
triallgllli~erus (BOil ,
li 'u/ 'Il/Of/ O!!" l ltjlll,~ II!tg.I·III ' I II III
.
1};:!7 "i,'i
.1:1", 7i-oplI/Wlflllf \ Irillll,t!.lIft t:./'nl,\,
Dis I rihu lion
1)1 R.'IHI
II)
17
111(/ /"1/ 11111/\
i~
I X27)
I) 'pc JII l'alify :
x-t
'valll\ (nrmg, II".!!..t'ld. I ,u Nti\I{ ,1 & '-.; H II ,l-t ' , I{)X-l
nati\'e III the f>enln~lI lar Malaysia , iJorneo. Suma tra. and ~ome sma ll er I ndon e~ia n i ~ land s . 'II. Kinahalu : wit ho111 prec ise Incal it ic~.
.\ <'Il lwltm/lltis
_ _ _ _
:!-I
,\00 '1\ tnwl).! II" !.! / 'I'II,\, ~ "' 1I111
P); I L~. .\"II()( Iw(}phl' fll lm~ II",t:.. ( ·ltI S I I I B "lt II)·}I 1';;' I \ "' f/ W 1m II /it " (nOIl,t!, lIltg C'rif.\, 1,\ 1... ""'''' 199--1h :' ,", 7 .\ ' III1JlIllfn\ In tlllg u!t>-!f''-O , 1\ 1\ II... M ' ..... ICJ/J )
37 1
( 'ulUhrllhll'
,..
o c
C' ..., :2 ::l 'oj
1aS~:" f iJ!, 359
Oi
Muderatel y largl:: mu~c ul ar: dor~al ;,cale" keeled . I- YI: or Ilormal "i/e : ~ antcrior tl:l1lporal~ presl:l1 t: no sub"lantiall y cn la rged tcclh . \llost eas il y reeognl/ed by Ihe lalcra l, dark-co lollred Iri ang les and Ihe red interspace;,. Rccord Iellg th 1_,5 cm. but ave ra ge specimcn ;, II1Ca~ un.: o nly XO-90 cm in Iell glh .
9 !' upnalabia I;" Ihc -I'h to (," tllilc hilig Ihe l'yC dorsals III 19 rm\"; at mid-hud y (17 I\cre qllolcd by MAl 1-.\11" I 9()2a), kee led exce pt li)r Ihc luwcs t row: 13 1- ISO vcnl rab (FNIII 'I( IXX5l:a nal ;,e llte di \'ided ; 57-1 (J'i "lIlx:alldal s, in pair;,. Dark o li\'(.: aho\'c, wit h "Iliall black ;,pots: a hllcral ;.eri e;. or lar),!c, tria ng ular, black spots. til..: points reachi ng the Icn tra);', alld s(l1I1l'1 illles forming hands a<.:ross the bclly, sepa rated by rcd int e r" pacc s: M)l1lct ilil es red s pots in:--ide Ihe black tri ;1I1 g lcs ; nn Ille po;.terior part or thl: body I hl: ;,pots arc indi stillct: head dark: uppl'r labia Is yd I< \\ or II hit l:, \1 ilh black ;,lItures: lower "lIri;ICI: ycl lo\\l. uniform . o r Ihe ve nt rals l:dgl:d wilh hlack. Young an imals are more vivid ly coloured . 'i-X l'ggi' (21)-. -I ' 15- 17 mm : -I .J9-'i.2') g ). Incubal i(ln lakl':-- S'1-60 da y;,. t::cul ogy/ t:: llw lu:,')' . \ i.'I1(/('limfllii~ Iri(lllg ll/iJ!,('r/ls i~
Ol'scri ptio ll
Eye lIlode ralely large: I lorea l: I (rarcij ~) preoclllar:-.; .1--1 PI P,loclllar:--: lI;,u
Fig. Jlln
.\ "'IIII(
t\lalaY\I;I.
/tnlfllll.' Irlllllx"iI).!.crw
frolll rCllill ~ lIl a r
flllllld li'om sea lel'c l tn altitlldes ahlll'c 13 50 m. It Inhah its reg it'lls coven.:d by In\\'laml tro pica l 1\l:1 fllre ~b. tropical lVet muntalle fllrl'sb. and pl anta ti on;" whcre it OCC llri' 11111~tl y III ril'l'r~, streams. lllar;,laes. swam pi', Ilooded ricl' padd ies, a nd pond s . .\'('I/I/ cliroplii., Iri(l llgllfige/'ll.\' is invariabl y fOUlld in Ihc thic k vege tati on imml'diatci y adjaccnt to water, or submerged in thc water amo ng aquatic plants . It is diurnal. and largely aquallc, a llho ug h it is also at laol11 t: on lalld in the riparian legetation . Tili ' ~ nakl' ll:cOS on amphihian ~ alld li ~h . Thi~ ~ p L'eic~ i ~ ~o metime" rat hL' r aggrl:ss ivl', st rikinl.! and biting \ iClllus ly when molested, bllt it i ~ hanllles~ .
Di strihuti o n .\'(,lIuclirop/ti.\ IrwlIg ll/igerlls i ~ distributed from
Laos, Cambodia . Myanmar and Vietnam in the 1l(lrth . to Su lawesi in the ;,ou th . \11. Kinaba lu : l3undu Tlihan ( 13 70 111).
3n
Crolalidae
Crotalidae /\ s far as L3orneo i~ concerlllxL this 1~l ll1il)' compri~es ~Illalllllllledium-~ii'cd snake~ . u~u;lIl y lI'ith ~h(lrt or moderate ly elongalL:d bodies and taib. The allllnst triangular-shaped head i~ di,lilH:tl y ~et on' fmm thc hody and has sma ll eye~ with vertical pupils. The upper ja\\ is f\lrni~hed \\ilh so len() g lyphllu~ 1 ; lI1g~ . These arc long. hinged. ho llow teeth which arc ro lded hack parallel 10 the skull roof intll protective ~kin pockets whenlhe mouth is shut. On opening il. Ihey arc erected up to a right angle by individualmll~cle~ . The venom is injected through the duc ts orlhe teeth . The potency of the venollls \arie~ with the speC1e~ ; some require illll1lcdiak medical attcntion to avert 1;ltalities. but the venolll of several species remain ~ unknown . The Illll1ily is marked lor the presence or a sensory organ \\ hose pre~cncc is rccogni/able e"\krnall y by lorea l pi ts. rhi organ is able to di"ting uish minute dilTerenees in telllpera tlll"e" and so enablcs thc snake to locale prl'y, predator~. alld sex ual partners. Pit vipers Illay he oviparous or OVO\ i\ iparou" .
Key to thr Bornean genera of the famil y C rotalidae (iular ,lIld inlersllpraocular ~eak~ Sll100th ....... 2 • (iular and inter~upraocli l ar sca les ki.:e led (;\) .... . ....................... ............ ...... ... .......
Tropido/{/ f! /I///\
2 More than -to slIbcaudab .......... ... Tri/l/ pr f!s /II·/I .I • Lcss than .15 slibeaudais ....................... 0 1'0l'"j,
Bl
R(oI :i{ III
Genus Omp"is H()(iI' & Ro 1,\ () Ilm il' , 19H I
The genu s Om/illi, i'i di st ributed frolll cenlral Nepal and Ilangladesh in the Ilcst. ea'twards to eenlral China and Japall ill Ih.: ea~t. and China in the north, ~outhwanb , through the Indochinese Pellillsul a and thc Malay Penilbula h) Sumalra ;lIld II lIrn el) , Small to m.:diulll -si:tcd; lTIu'l:ular: body stout; tail short; hL'ad triangular. distinci rnlill neck. thi ck . 1I1kn swollen. abovc I\ilh small. a~ymmetricall y arranged sca le'i. Two to four i'lm s of ~l1\all scales iletwL'en subocular~ and supra l"hia l' ; ..;ubcaudals diV ided or entire . The~e arc gru und-dwellcr~ which prefer hi ghe r clevali ,l11s , Dl'spite illl:ir apparcnt plumpn.:~s they call st ri ke in a flash. 'I heir \'enolll s ma y he I;wd filr hUIll;]I"' .
Snake~ of the gC IIll ~ Om/ );';, may lay cgg~ or gin: hirth tll full} <.b eloped you llg. It i" kllm\'l1 llf ~() me ~p.:ci.:s thatthc klllak" guard their clutche~ (or egg"). Fivc speciL's arc cu rrently reeoglli/cd. PilL' of which inhabits L30r1lCO and ~vlt. Kin abaili .
Ol'opilis c/wsl'lli 1,."I1c.'/'{'SIII'1I\ (1It1\I'1/1 '-' \11111
t "nll i l ~' :
",.It"
l i"/IIlIft'.\/trll\ .
, 1<1 1\11 '~\I
1931)
(S\IIIII .
I'n I
2l).
pi II
M I "Ill.tlx tlu ItO\l 'UI L o\l IH I"" I 'nx. ·1';;,
tl~
\h ~ IIII '
I
I YIlt'
19X." 30:
8.: :-,,,. ,Ill' I ()K-I . 2·'
OI'''I,hi, ('110\( '111.
'-; 11 1 B INI. , C)') I
'" IIIIIN',&' h e,ll(
'':;7,
1' "
\ 11 "
11)\'4 0 2· IX,
P)qll 22,
f)i a::! lIflsis I)i ~t inglli ~ hed
frolll Ihe cl(l~ely rL'latcd 0. I/ulllli hv Inwer Illlmher of dorsa l sca lc nl\\ s ami sllh lahial~ . :Iax . total length Ilcarl y (,:; cm .
c(JllI.
v
o"
Fig. 36 1 ()'''I ,III' (/wll'lIi. paratype lrom Kiau (/'IU ' 2._17.11.
Desc ripli o n
His I ribuli u n
!Jody stoul ; head llat. \'ery di sti nct rron l hod y: eye ~ Illall : ,hicld, llnllppcr ~ ide orhead rclallvcly large, ~ubll nlm ca le . \\ cak Iy k..:c1cd : s upran c ular ~ ve ry large: (> 't lpr:rl ab ial ", tlK' 2nd helow, but 110t bordenng. the loreal pll , the J rd the hi ghe~t one, separa ll'd rrom eye hy 2 , mall ,ca lc,: I0- 11 ~uhlahials: 19. 17- 1(), or 15 rows (II' d o r~als, Ille mt:dlan Dnes on th e anlenor body ~ tron g l y kecled. tho ~e on the p o~tt:rior body wcakly kt:ded : 130-14J \'t:ntrais: anal q ;Ule entin:: 20-JO "lIbcalldal s. in lXIiI'S. B rnw ni ~h above. with Irreg ular b lackish , Ilght l11argint:d ,po t, which ru:>c to form bamh in tht: postt:rior portion o r the hody. Lower side ye llo\\'I"h. \\i lh a 1I1ulliludt: or 'i 1l1ll11 grey spot s, An obliq ue, black stripe WIth a whik lo\\>'er border behind 11K eYl' .
Thi~ s pet:ic~
E c o ICl :.:~'/ E llwl(l :':y
s uhmonlan c "peC IC", recorded from altiludes hetwee l1 l) 15 and about 1550 metres ( II Q-{'olleeli on) ,
37-1
is endemic til ROnleo, it ha ~ (lill y heell round in Sahah ( Kg. I'a go don. Maliau l3a ~in, Sin~urnn, Crocker Range, and It. Kinahalu) . Mt. Kina ha lu : Kiau , I k a dqll arll:r~ .
C;Ct1L1S
T,.imeresu rlls
L ACl-f' f 1)1,
I K04
Thc di~trihuti()11 range or till: gcnu" 'fi-i/J /('I'('\ /I/'11 S e'(t..:nds from nOl1h..:rn Indi a in th..: wes t, eastwa rd" through large part s orc 'hilla to Tal\\'all. sOll lhwards through the Malay Pt:ninslIl a, tll thl' Ph ilippin..: Islands and to th..: Indones ian i:-,Iands of' Wetar and Timor, S11lall tu lll ediulll -·m·ed venOJllOUS snakes, l lead uSlla ll y \.,;rv di st inct rro m hody. it" tipper side covered \ il h slIpr;locll lar". illterll
C'rntalidac
st.: nsory pit with onc nasa l po re; hody round or ova l in t:ross-seetion : tail short to l1111tkratdy long, prehens ik ; anal se utt.: entin:: subea udals most o rt cn dividt.:d . The crepu. cu lar and nocturnal. bush- and trecdwdlers s peeie~ live in forests as well as ill opcn. bushy areas, gm. slands. plantati ons and gardens. Thcy fced (Ill sma ller mammals. birds. lizard. , and frogs . ~I ost species are ov(n iviparoLlS, but aft.:\\' are oviparous. Thirty spec it.:s art.: kn own at rre~cllt, of which li)lIr or five live on Borneo. Thc nccllrrt.:nct.: on this island of Trilll<'r<'SIII"II.1 a/ho/l/hri.\". based on an old rt.:<.:ord. has not bcen con firm ed . Thrcc spt.:<.:it:s art: fo und on Mt. K inab,t1 u.
rah. S7 DISlnhUIIllil or 7'·;II/(,/"l'.\I1/"I/s-spccics
Key to the Bornean species of Trim er esllr/ls Snout strongly projecting. bilobed or ~ p a tulate ; 1-3 supraocular plates. nat or sl ightl y elevateu .. . .... .. .. .. ... ......... .... ... .... ................ ......... bom eellsi.\·
noutnot strongly projecting, hilnhed or ~pa tul ate : I nat supraoe ular scute ........................... ........... 2 2 19 dorsal scalt; rows at mid-body .... .. . III a/co/III i • 2 1 dorsa l sca le rows at mid-bod y .... .. .... .. .. .. ..... 3
3 148- 157 ventrals .................. p opei ortllll Mlbah i • 182- 191 vcntrals .. ........ ...... .. ......... .. .. .I'll/lillII'll/illS
0 11 IlMIlCO .
Kinabalu
Sabah rr;III (,/,(,.w/'ll.~ ho,." el'lI.\;.~
(1'1
rr;lII f'rI'SI//'IIS IIIII /co/m; L, II
II' ~S ,
I '72 )
I kll. d ,
--
I \).'\}(
• • • •- • • + +
TI';III l'r('S/I/'III pope;oYIIIII .\·{JIIII /,i ({' (i ' ,.\" & Tnll/f'n ' \lIru \
\lIl11e/ln/IlU '
tR II II I'"
K HIMIR . I') ~ I
IX22)
Trim er eSlI r fiS horll eell si s (Pl"ll' I{S, 1872) f tJ'o !,o/dJl.\ / ' f} /,lIc ' I ' f} \ ' Sill ,twak , Uonh:O
\'
PI II k". 1 1' 7 2 ~ ~ I
T!"I m T ',\lI nl' IN olln'lI ,\', '\ 11 I HINI t 14)1' I
1" 7
TYPl' Illc~ l it y : ~ pal t
l
Di ag nos is Lasil reeogn i/ahk by ih projeclt:d. spatulate. uptllmed tip of th <.: '.;nout. Max . kngth X3 em: averagt:-sizl' -1 5-55 cm. I> rsc ripti ull Si<.:ndt:r: head di~tin ctly set o fT frOIll body: tip of snoLlt sr atu lar: 1-3 supraocu lars. flat or sl igh tl y butl on-shaped. sl' para tcd by ')- 1 I ~ca ks : 3 preoc ulars: I sub, eu lar: 1)- 1 1 ~u pral abi; lls . the 2nd high. hord n in g ante ri or margin or lorea l pit en t irely or a lmost entirely, thc 1st separa ted from the nasa l shi eld : I C)_2 1 rows Llf smoot h dorsals at mid- bl el y: 152-1 XU vent rals: anal ~cu t c entire : 45-5, subcalldal~. di vi ded . Dorsa ll y li ght , ll1'dium, to rcd di ~h broll'n . more rarely greenish, \\ ith an irregular pattt:111 of dark
IIrunci Sarawak Kallmanla n
•
• •
•
-
•
•
brown spots and grecnis h hroll'n to reddish brown markings ; urpcr ~ic k o f tht: head dark with a lig ht hrown lillt: li'om th e posterior margin of eye to bchind anglc ort he mouth : nUIlIt:rolis li ght hrll\\n dob on labia Is: \Cllter li ght to greY I ~ h brown, line ly dotted with dark: 10ll'er >.idc of tail dark hfll\\ll. .Iu\·cniks more cont ra>.till g II ith ye ll ll\\ tail. 7- 14 c!;!.gs in a clu tch .
~
a; <J
--...-~-~;..;;;:::.:--- ~
( 'nltalidOl":
EculoKV /EI h o lu ~~' IS l(lUnd bl'tl\ CCn ~ca Iewl al1d a llttudc ~ or at lea~t 12()() Ill , with a pn:dd L:l.:tillll It)r hill y rq~lon~ at I.:lc\'al i(H1ShcllVccn ."i O() and I()O() m. It inhahits low land troplGll wct Illrl'sh, trupica l I\ct 1110111 , IIH': lorcsh , ..;walnp~ , I\Ct ,hruhlal1d~ , and plalllatlllt1S, It (ll~ tl.:n lives In damp pla cc~ or c1 o~c In waler, This noctUl'llal ~pL:cics is hnth arhorc;t1 , [rL:qllenting logs, \\\Iod plli:~ , thi ck bu ~ he s and trce 1()liagc up to 20 III ahove the ground, and telTc~trial. 1~c1ill g at hOllle among Ical L:S and planh on the 1(lrest Iloor. Ju ve nib; SOIl1Ctil11c~ climb into Im\ vegctatiun holding "" to twi g~ 1\ ith their pn: hen~ik tai ls. In tlw, position thL:Y onen pu ll tk body into a !lal coil. I\ ~ a rulc, thc snake i~ 1110stly cneouillercd in tangled wct vegcl,ltion, r /!o l'll{ 'C'lIsis lCed~ m;lIn ly Oil ~ mallll1al11mals , lizanb alld rrt>gs , It I ~ an aggres~ivc snal..c. stril..ing PI\)ll1ptl y at lillie prmoc;]tiun and biting rcadi ly, Nollllng i~ I..l1llll n abuut it..; \'elltllll , 13 analogy with r plll/in'lIs. thi s sl1al.. c should bc n:)!a nkd as potcntlall y d a ngcrou~. 11.111 I (1963) rep0l1L'd Oil a bite or a 7hIJ/('/,('s/II 'lIs e hl/l('11i III Hnillci . but it I"; almo..;t ce rl ain that r hlJl 'IIlTllli,\ \\a~ Il1Iolvcd . TllI~ bite ncarly I..ilk:d a middk-aged 111<\11 III cxcellellt hl:twld coil up around Iheir clutches and I..eep thi ~ po, itlllll lInt il their yOllll g c\elltuall y hatl'h , Th i~ ~ n a l..l'
T/,;mt!/,£!surus malcolm; l ,m 'l niwcn.',\lIrr/\ ,'''111(1/1(1111/' 11/(1"
tuc.d; I) · Sungl;
illuJ/ I .)\ I
I{l\~r.n~"r I-.Ia(l.
li 'JIIH'I'f ' Ufnl\ ,\ItNla /raltu,
tl
f{t()(
KIII( ,I . Il)
ii,
193R
\X . ~1 5
'I ) Ih'
""';II>.iI(I,<'<1 n llll ICc!
mil/co/1I11
f\ l \Nlfll\
ItHU· .1 0 .
"flO'
M \I I'JX"
&
" HUiiU\
II)X--I
S\1I1111')1.1 II .
2~ ( p .ll l ) ~I t "IIt\I,
""II)
(pdll L 1 '(1
(9 1)1
J . . 7 Iparl
(I.!ITt, )r
"-lIJ\f{P
I
Uia !!l1l1sis
Max, tota l length 1:\.1 Clll: clu~cly rclated to TrilllC/'( '.\I/I 'IIS IlIlIlIIlmllllS, btlt difTering by l'ewer rnw ~ or dor"al scaks. and lowe r vC lltral and s ll bca lldal eoullts.
Dl'sniptillil Thrce prc() c Lllar~ : 3 postoclllar~ (inelu :i ivl' lllle ~u bol'u l ar): 2" 2 tcmporal.;: X-9 ~upralabi:Jls. thc I '" :ieparatc rmll1 n a~a l : I pu~t s llh() c lll a r : I I) row ~ or keeled dorsal ~a tll11(l - hod y: 1(»)- 17-1 \enlra i'i: anal ~cu tc cntirc ; (, I-X I , ubca udal s. Upper ~i dc s I I' head and hod y hlac\., with Im g ht grecll "pols elln:-.ist ing or triangular grcen are a~ I1n indi\ iUlial ~ca l es: a tcndcncy tu 1(>rI1l illdistinel transvcrse linc ~: \clli ra l, dark laterall y and un their anterior l1larg in~ , uark or Iighl grecn on their posOislTihutioll tcrior eugcs; tail patl eJ'llcd wilh ~crie~ or unilill'ln . /'rilll (' r ('s lIrlis hlll"l/t'{,IIIis inhahit s the r e llill~lIl ar parallely :l1Tangcd. red ~ poh between dar\.. grecn caul'v1a laY,Ja. BtH'Ill'() alld Sumatra, It i~ di~tribllt e d ill dal sealc~ : sli heaudab light grcen, ,()nll.:tillles li'al1led all parts (II' Hornco , M t. Kinahalu : Poring , \\ ith hlack . Juveniles gra,s-grccn above. often wi th hlacl.. rral11L:S beFil!, :\63 li'1I111'/'( 'I' /lf'II,I /llld ,'ulllli, JL1 vc mk i'rom Sg, S liau-Silau, 1(,00 Ill . tweenlhc sca !cs: tai lt n ~ Iark eontmst . i.c, li),!ht brown wi th imlislim:t. dark. tra nsve rse bar~. postcnnr Ihird or tail dar\.. browll , l,alcra l s llr l ; ICc ~ ye ll owi~h with reddi s h stripe: vcntcr grcc ni sh yel low ill the anterior quarter. othcl'\\ Ise erea m -yc llow: supra- and s llhlahia l ~ bri ght y cllowi~h g recn : lalcral stripc 1\ hitc abovc and rcddi ~ h he low, ()II bottol11 rows or dorsal s,
Crotalidae
... '
".. .f~
"
.." "
~
(
f
w
1 _
@
Fig. 3 (,~ Recc ntl y wc rcec lI'cd th i, pholO
~
or a sl1ak~ rrolll
to be 71"11111"1"<,.1111'11.\ SlI lI/ilI/ WII I.I . Duc 'Ilpalicially. \\C hc,' tlall' III In cl lld ~ /: til Ihe hcrpdor:lllila "I' \.11. Kll1abalu lIlItil ~ pcl'ill1l'ns arc' at hand. III chcd. Ihl' pholidn~is . Por ing. Thi,;
10 Ih ~ I;ICI. Ihal Tril//('rc.I/I/"/I' wlI/illrilll/lS IS Icry '> Imllar 10 1/11111111'1111/1'
Ec ol ogy/F: tlwlog ~'
Di stributiun T h is :; pccies i~ e nd c l1li c til 1\11. h:in a b a lu : Sg . Kc nokuk (at Kiau) , Kiau , Blindu Tuhall , 1.1111111 I.um ll , 1\(;;ldqu:trt.:r s. Kamb ora ng o h Road ( 16()O Ill) , Kunda ,a ng ( 1525 Ill) . Sg . S i l::Iu-Si la u ( I S()() Ill) ,
';':C I11 ';
11111/, '0/1111
Trim ere.m rlls popeiortlm saha"i R Hd :. ,\ so.; & KIC\ t-. II 'i{ , I ()X I
Trillf('['('.I'ffl'll.I' IIw/co /llfi ha~
been recnrded frolll altitlldc~ bct\\ cc n 915 and I (-,00 II I. r-\il11~t or the spcci mc ns IIL:r': co ll ected llnlh.: forest fl oor or as l' i Clim ~ of road traffi c. [\,1 II ... III "> (I ()X7) li)lInd a speei mcn on a rn::ld~ldc ICl,:t.:Ialed wilh Ii:ms and l,: ra~~ al around l) .OO p,m. du ri ng a ra ln shol\ cr. Thi~ ~pecics i~ nocturn al. It:- dict is 1I 11(,. 11 0\\ 11, but adult s p cc imcll ~ fed Oil rat s and gro und ">t(uirn:l, in ca ptivit y (, ' 111 ' 1111\(; & I'HoI 'l{ 1l)l)X) , /\ j UICn ilc aen :ph.:d frogs ::Ind t3tipul c"> frol1l li1rcc ps ( ,1 \I 10.~Ill:; 1%7).
T
I
(I(}1( " /\
HolhrrJI',\
,t:.rllIWIIt ' U,\,
'>11111 "" 1'
l'IOla
1o( ',n \1< 11 I ~'NO,I
ClX
'It \
1x02 J, (p,lrl"l. III ({ """
gr"JllIllc.'II\ «nol ('(I/lIht.'I ,t.!nIlIlIlHU'
IIINI I" II 1900,1 ' 711: 191 7' 2~5 Iparl..
1'1
/hlUe 'n'\I I/TI.\ ,!!,/HIIIIIWtl,\ (nol ( 'UIIl/I( 'I ',t!,ro mIl1l'1I.\, II \\\
I'n!. 10, ,0, '\2, 1 ,,11"11.,1 I'nx of '. \1\"'11 11 I
I Xll~) ,
I%~
Tnlllln ',\ Urll\' /Ji)!1t'iJrllt1I \(lho/II R lc,1 "'\ ...... '-': ~ K"II K.
""1111
.111 19NI I')()
T~' I)~ l" c;lIi t~ : \toulIl KIlIahdlu . Kulap" RII ~r.
IInl l
llrth HUI"I1l.'o li 'llIh "'t.',\ lIIl1 ,\ fJO/'(,OI'lWI,
I 'HI 'It \Kll
4\.
~t tll, IU N
14) , I 2 1.
SIIIIU", I ()')I "iI Tl'l nlf'1'4 \111'11'
,\ 11111011'01111\ 1IIe//olI lIlI
I/Ilt/udnll I tl\l l~IIK.t. 19.1:\ ). \1 " t
T,.imt.'I't'\urus
/,11"t 'O/, /II11
,aholll. \\
(IIO(
\I t t...,
\Il\ \1\
1,1111 ('/,('\1 11'11\
\'
1l}l}2a, 13(, ... Il)l)-lh 2 I X
Taxo nomic comJlle n' : SII II'III!\(, 8:..
I Md ' \{ (19l)l)) agilin 1I ,<:d Ih t: old name "I; 'il//('['('IIII'f(1 1I(I/I(' [)[,II f11 and rcfcrred 10 thc corrcction ny thc uriginal author.
377
( rOl alidae
11 th<:n.: l~m; ,I ppears Ipprnprial<: to qllotc the COIll ment givcn hy D ,WIIl & V
com mon along streams. bllt Ilothing i" known of Its biology. The nominatt: [()rm inhabits lowland tropical wei forests. but is mostly f(lLlIlci in tropical and slIhtropical wetmoniane li1re"ts. b,lInboll fort:sts. wet montane shruh lallds, ~\\'alllp" and plantations. II olien oculrs along densely vegetated banks lll' rivers and stream s or c!o;.,e 10 watcr. This noctlll'llal ...nake i" largel y arboreal and li\'e~ in bushes. tea ;lIld coffce plant:-.. shrub". and low tret: f\.lli
Dia)!lI {)s i~
i)i lTering f'rom the other "lIb"pecie" by the combination or the rollowing trait s: _ I dorsal ,COlic rows at Illid-h\ld y. l11all: ' with ft: \\ t:r than 15X \'clltrals. i'cllwle" With fc\\·er than 150 \cnlra ls. :vlax . total lengt h 7X l·m . Ucsr riptiol1 Supmlahials (i-II. the I" ,'parate rrom the nasal "hick!: illlt:rnasa is eillarl.!.ed. 'iepar"tl: li'olll each lItht:r by 1-2 "ca le s: ()- 14 "cales between the nalTo\\ "lIpralH.:ulars: temporals weakly or not kc<.:led in adult ~ pe C lmen., : 21 dllrsa l "ca lc rllW, at mid-bod y: dor:.a l' kech.:d e:-.ct:pt for th()~t: in the lir~t row: 10- 13 ~ lIblahiab : 141'-159 \'l:ntrals: 59-74 slIhcaudals. Upper :-' Ides of head and body unifill'll1l y green : tip 01' tail hrown : s idc~ or head and ve nt ra l sides slight ly light<.:r than body ahove; adult specimens without postocular stri pe. Males with distinct lateral ~tript: on iiI'S! row nf' dorsals I'rol11 ncck to su bca lldal,,: Stich st ripe i~ Illi~ s in g or hardly recogni/ahh.: ill females . l lemipeni s bilohat e. ·mooth. without hoo" , or ~p lke'i . Tri/lwre.'·lIms / /oJldol'/ lIlI .\lIh(/!ii is Ii\'e-hearing. The 1l1a ll il and form produ ce~ ahout 10 yo ung in a littcr which me'hlll'e I X-20 Cill In length. E c o lu ~~'/E t hnlu ~~'
Trill/eJ'('Slims p"pciomlll .\lIhll"i is found bel\.\t~ell
"ea h.:vel and ele\'ati(llls of' at least 1475111. It
37X
i~
Di stribution Till" sub"pccil:s is a Borncanl'ndemite with record ~ being a, yl't limitcd to Sabah and Sarawak . 1\1 1. Kinabalu : Sayap, Kiau . Kellokok . without locality 640 III ( II \ 'III ~ ( ' II 1900a), Sg. Takobang at Kiall . Sg. Kadamaian at Kiau . Bundll Tlihan , Sg. Luidan at Allndu Tlihan. I kadqu ar ter ~ . R hlol' 'l ,\ S " &. KI('\\ll R 191' I mentioned a specimen or I'dt. Kinaba lu at an elevation or4175 Ill . Thi" is obviousl y a lIlistake , i l ' rvlt. Kinaba lu i ~ only 4095 m hi gh. Perhaps the high was 41 75 reet (1 270 Ill) or 1475 III due tu all error.
(jcnLls Tropit/o/aI!IIII1S
W i\(;LI 'R. 1830
The genlls TIY/IJic/II/lle/1I11\ shows a disjunct di~ t ribll tion pattelll. II occurs from the M;lIay Pl:llinSlila and Indonesia to the rhilippint: ~. but there i, also a specic:.. Known from ~outhcrn India . Characterized by the absent:e urllas,,1 pores, st rong ly keeled. <,111 a II sca l c~ Oil head. and Keeled gular ~eale". SCl;llnd slIpralabial docs not border the anterior Illargin o f the loreal pit . and is burdered abovc by a prei 11VeaI shield (I) .WlLl & VO( ,I 'I 1l)<)X). Tht: species of the genlls Tmflirilll((('l/l/ls arc crepuscu lar and nocturnal and live predominantl y on bus he, and s hrub ". The y arc \ e ry lethargic
Crotalida.:
aJllbu~her~
preyin g upon small malllma l ~. li;tartis. fro gs. and birds. TI"IJ/Jic/o/{I(' I11I1S arc Iivc-bearing. Three species arc known . on.: or which occllrs Oil Borneo.
. TroP;I/o/(J ellllls wagler;
rf}"I'id"/," 'III1I' 11It/>:/<'1"I \\ " ,II K.
Wi\( iLFR ,
IlOO. 175
I R30
TYll<' 1 ",' '' l i l ~ :
.\"" Ta \onolll ic CO llllll c nl : Thc ta xo nom y of thi~ speei.:s or sp.:cies complex is high Iy lInsal i Sr~lclOry and cllrrcntly undcr investi ga tion (Kl (II pcrs. comm .). Thi s form might have the lIan1<.: TI"IJ/lic/oIIlI'IIIIIS lI'(Ig/eri sc!JIcgdii UI I I 1--1 IL I XS7 in futur.: (bl-- \\JIl.\1l & COl 1.1 in prep.). For thc authorship of this ~p<.:cie~ ~ee l),\\II) & VOId L (1l)96). IH a :':lI osis
Largc, ~touL max. Icng lh aboul 130 cm. averag<.: XO em in f"cm a ic:-. male:- are ~ma l lcr and thinner. Tail knglh/ tolal kngth ratio is bclow 20"0. I> cs rripliun (valid tl)r Ilonlcan sp.:cim.:ns only lDA\l ll& V()(dl 1l)l)XJ) 10-13 strongly J..ccled scales hctwccn thc su praoculars; J pn.:ocular:< 1 enlarg.:d ~lIpratlclilar; 1-3 postoe ulars: I suboc ular : 0-6 s mall lorea ls: 7-1 I supralahials (Fll-- \IH 1')6<1). none in contact with eyc: 10- 12 sublabi ab: dorsa l ::-ca k s kC<.:kd. in 21-27 rol\~. outer row s moolh : 130- ISO \' entnll~ : anal scut<.: enlire; suhcaudals ill rair~. 46-5<1 . Coloration va riabk . a Jllixtur.: of various shad<.:s tIl' gr<.:ell alld y.: llow. limning a bandcd pattern dorsa lly whit.:h i~ olkn not c learl y ddim:d and C:\ILl1d~ onto Ih<.: tail : uppcr s id<.: of head g rLen wilh :v dlowi~h SpOIS: lahial shields yel lO\v or grcen : \cntrals gree n with ye llo\\ margins or ye llo\\ With dark gn:en margi ns . .Ill\ cnilc~ g r<.:ell with r<.:ddi ::- h wh ile d()r~olatcra l S P()I~: a whitish :-tripe, bordered wilh reddi sh beIllW. rnllll th.: lorea l pil to th.: ()t.:ciplll ; tail with a r<.:ddi~h or blat.:k tip. (hovi viparoll:-. ~i\ ing hirth to abOUI 15 yo un g, with a maxilllllill r<.:t.:ord llr4 1 jll\'en il e,; in caplivilY ( 1 . I~t 1l)90) .
Eco lo~y/"= th o l ()~y
r
lI·agle,.i is round in Ihe I (\w land ~ up tn elevations or at least 1300 Ill. but is most f"equent in Ihe well-walered lowlands. It inhabit:- lowland lropieal wet for.:sts, tropical wd Illontan<.: rorcsl~. c lo~ed or open ~wa mp s ancl ma rs hcs. l1lan gro\'e~ . and t.:oas ta l s\\" Jlnps. It ollen oct.:urs close 10 watL:r. alonJ,! hanJ..~ offorest rivers and ~trcaJlls. in foresl.:d ~wal1lps. wh<.:re il lives 011 slruclurcs overhanc.ing water, in hushe:dose to rieL: paddies. and in ditches in vi II a J,!':S. 1hi ~ very :-Iuggish. crepuscular and nocturnal '-pet.: les rLIreats durillg the da In hollow trec trunb or in tang l.:d vegcla lion. where it can remain t.:oi led motionle~s li)r a long time. and becomes ae ti\ e after du ~k . J\dult~ arc strictl y arbor<.:al. prdcrn ng low .:levat itln ~ ollCrecl by bushes and s hrub~ . but .Iu\cnik!-- olten 1'01'agc on Ihe grollnd and in lall gl~ I :-S. Thi~ snake Iced:mainl y on mamlllais. li/;II"{I ~ and "·og:-.. but al~o take ~ birds; juveniles prcy upon !I'ogs and li:lards. l hl/Jido/!I(,IIIIIS lI'Ilg/e!"i is a vcry do.:ile ~lIake. rarc:ly slriking or alleillpting to bile unless handled roughly. Il owevcr. some ~pet.:iJllcn,; arc: Jllore irrilnble Ihall others. and Ih<.: \·enolll is pot<.:nt. Sever<.: case~ of .:nve nl1malion hav~ been repoJ1ed. whieh lead 10 both lot.:al and sySI<.:IllIC sympt lll lls. ' 1Ill ~ ~ p ec i<.:s sholdd be regarded ns potLlllially dang<.:J"Ous. I>is iriblliinn
This snake rangc~ fro m Ihe Mala y I'enlll~ula and Ihe (,realer SlIl1da Is land:- to Ihe rhilippillc~ . 011 Uorneo. Ihis spet.:i<.:s is \\ld<.:-ranging . ·1t. Ki naha hl : lI eadq llart cr~ (I I()-Collcct ion) .
Fi!!.365 TI1Jf/u/u/llclI/l/.' "'"j!. /" I"I trolll J),,..;lIl1ahlll, S"bdh .
(
y l in d rll p h i l d ; l ~
----
C ylindrophiidae Thi'i 1 : 1I 1l 1l ~ () \ll pr i ~c'i 'illi a ll 10 m()(krmely larg:( ~ nak cs The l)(ldy i'i cy li ndrica l ill crllS~ :-.ecti on, thc head not di stillc t. A mc nla l groO\c i ~ pr(:o.( Ilt. Or the () sll pra la bia ls. the 3"1 III ..t 't. arc in cont act I I ilh thc cyc . A Io\"(:a I i ~ absent. T hcn: arc 19-23 rows or d llr~al 'ica le at mi d-hody; thcy are ~ I lh lo l h . k t: lh a r~ prc~c lIl nn 111\; palatt ne and pl<:rygoi d: the max illary ted h count 1) - 13 . 'I h( e(\opte rygoid i'i robust a nd O\·t:rlap:o. 1ll:lx llla a ll d ptc rygo id exte ns i,·c!y. I'hc prcmaxi llary lI a~ al proc(~:-- i ~ lo ng and (x kll t!:-- hetwl'( n Ihe 1I:1 <;;II 'i (('1''111,\11 c t a l. 19(3).
(,el1l1S
Cylilll/mphi.\' W '\( iI
I.R,
(eis and other
I R2X
T he !;:t:nu~ C:l'lim lmll!lIs I ~ d islrlbu ted rro lll 1 :10 '. Mya nmar. and Vil:tnam ~(l uth tn the ,\ rll blands; it a Isn ucc llrs O il Sri La nh.a . Sllw ll ~ p ec l e'..: budy cylilld rical ; head c()\'crcd with large. sy lllllll:t rl cal shlcl ds, \ ery smal l and not di sti nct rro lll budy: mcnta l !;:l'Illl\ c prese nt ; eye'i \ l:fy "1\01 11 . pupil rollnd o r \'e rtica l-e ll'l)\J e; no loreal: no pn:ocu lar. 5-6 ~lI pral ab l als ; nost ril 'i itllat ed in a ~i n g lc na'ia l; 17-21 SllIooth dnr,,,1 ~ca le rows at m idhody; v( ntrals ha rdl y llr llllt at a ll widelled: tai l very shorl, with a blunt tip . T hc:st: sma ll 'ina h. e:-. or 'icmi rossorial h ab it ~ li "l: bl:IH.:a lh k a r ht ter. IOll'iC harh. . or othl:r ma te ria ls cOv cring th( ground . but the) arc more ra rely fuu nd in tilt: sllilltse lC Th( y pr( y upon Iruly ros~ori a l sna kes,
Kry to the Borncan species of Cylilldrophis 17 dorsal ~cal e row ~ at l1IiLl-hody .. . ('l/g kllri
2
longi tudinal II 11Itlsh or black stripes on the back; head and tail )cIIO\\ ish or red. lower part of ta il wh ite .......................................... ... .... Ii I/eli Ills • No longitlldinal l>lripes 0 11 the bac k. head and tail abo \ e l:n lmm.:d I ike the body. lowe r part or tail vc rm ilion .... ..... .. ... .... .. ........... ... .... . mfJils /'IIflils
Cyli/ldrop/ri.' ruffil.1 ru./lils
SI I I I · III~I •. 11)1) .. H.j{ P.
' 111'0
(L ,\ I 1( 1 " II .
Cylilltlr0l'ltis ruffils
17bX)
I •
rt~llil.\'
111//11
t
\I 1<1 Nil .
t7r.x: 7t ·tYpe tOl' alit~ :
Unlll,i
•
" Sllnnam~ "
Dia gnosis Small ; sli brossorial ; read ily identified by red unde rs ide lI r ta il tip . Lo r(a l and preoc ul ar shields abse nt : 1l1;I X. tola l len gth Ill:arl y t) () c m.
Kali n,alllllil
• • I • •
p o~toc lll a r ; (, supra la the )',1 and ..t ,t. in contac l w ith eye ; dorsa ls in I t) o r 2 1 rows a t m id -hody, SlIlllo th : I ~5-2 4 5 le ntrnls: a na l SCli tc di vi ded; 5- I U ( tll ire :-- ubcall dOl i , Dot'sal l irid e~ce nt blach.. da rk hro wn. o r blacki sh hrow n; irreg u larly s paced w hiti l' h. re ddi sh. o r be ig.:-coloured hands. 11 0 1 e n eo l1l pas~i n g the body emire ly. tlie las t. or th.: two pllsk ri ormost o nes , 011 hia l ~ .
1111 , ITOII
S;Jrawak
Descri ptio n No lore;" : no prcoc ul a r: I
( L \\ .tON II . 17 6R) II/)! IIII
Ki nahat"
- l-
I XX I
and il1\ t:lkbra t e~ such
T he ~ n a kcs g il.e birlh 10 fi.l ll v deve loped yo un g. Ten spec ies are known (S ~ 1I111 & SI IlI" I t)t)~l. th ree or I I hil:h live o n Ro rneo . on( o n MI. Kinaba ili.
Sabah ( :I'/iIllI"/lI,III ,1 l' IJg /'II/ ' / (' IJ I I I ( 'I·/il/drlll,IIl .1 III/l ' (/(II ' I~ I \
li~ h . " / a rd~.
a~ ea rt h\\'tlllll~ .
Cylindrophiidae
Ahoh': Fi:,:. 366 Dorsal "iew of ( :l'!illllm/I/'/1,\ r mIllis from Penins1IIal vtabysla.
I{i:,:ht : Fi:,:. -'67 Ventral "iew of Cl11I1t1lV/I/'/I,\ ti'OIl) Thail '1I1d.
I:
millis
the wil reddish or bright rcd. the firs t one on the nc<.:k likewise: these bands may. hll\\'\;"er. be absent ; throat with almost entirely bla<.:k scalcs ",. . ith white tip~: ventra l side alternating blackish hrown and whitish ; underside of tail tip red . .Juveniles arc <.:oloured more vividly with the collar being particularly more distinct. I'emales give hi rth to Iit ters of 5-13 babies of abou t 20 em in lengt h. [culug~,/E I holo~.y
m/fi/\ rulllls is found at low ele\ ati(lnsand inhabits non-Ill (lied !tmland tropi<.:al wet fore~ts . plantation~ . cu lti\'ated areas and garden". hilt is mon.: frequently found in swamps. in the vicinity of rice paddies. and in ~uburban and urban ditches. cana ls. and ga rdens. It i~ sublilssorial. living <.:1ose to the surli1ec in loose. wet soil or in muddy areas wI1l.:n.: it can burrow with easc. It i~ al~o found in rodent bllrrow~ or other holcs. under decaying wood or vegetation. piles of wood. boards. rocks, stllmps. or in forest litter. ~octurna1. it i~ ( :,-tillr/I'III,lJi,
-
9
often seen loraging at night on the ground during the rainy season. and it takes to sha ll llw water when.: it ~wims effortlessl y. Thi~ ~pe<.:ie~ ti.:ed~ chiellv on I ther terrestrial snakes. but also tal..es ~IO\\ - mO\ ing burrowing li/ards. eels and other lish : ne';tling mammals and im ertehratcs (ill<.:ludillg worlll~ and myriapncl s) h,l\e abo been reported . Whcn it i~ surpriscd or threatened. it raises ils tail and curl s it ovcr its hacl.. , so that the red ventral colour Ilashc~ a warning. pos~ibly as a mimicry 1)1' cubras . Dis trihutiou C:l'lilldm"hi,' /: ru/III.I' inhabits a regioll rallging li'om \~yanmar an d China 10 the 1~lands of Bornell . . IIlllatra. Java . and Sllla\\c~i . 'It. "iuah .. lu : llcadquarters ( II Q-( ·ol leetinn).
:I XI
Elapidae Elapid ~l1ake~ ;Ire char;lckri i'cd by havi ng. pr(llcrogl ypholl~ teeth . They posses. two !!rOll vt:d, largel y syril1 ge- like teelh 111 Ihe anl erior portiun of th e upper jaw which are conneckd 10 wllom-prndllcil1g gl:1I1(.b . Th e~c ~Cr\e to injlT I \ ' (,110111 Ihrough Ihe dlll:ls and gro~lVes . The I~\l llil y is tlll1hcrl11ore typified by il~ l11el11b e r ~ havll1g. round pupil s and 110 loreal ~hieltls. It co mpri scs ~ l1Iall "pt:cies or ~o me 50 CI11 (ClI /!/IIjlIIIS). a~ \\ ell a, the l:1rgcst Vel10mOll" snake, Iht: k il1g cobra I ()Jlhi(}IJ//lI~IIS 1/lIlil/lIlt) . which ma y grow lip t(\ 5X5 Clll ill tolal kn gth . rhelr venom, can be very potent ';0 that a hite Illay he ha /.ardous. orten even lilla l. fo r human s. Th ese snakes reproduce hy lay ing eggs.
Key to the Bo.-n ea n genera of the fam ily Elap idae Tail Ilallened; in habi ting Ihe ()l'I.~a ll .. Seasllakes • Tail mo re or k~ s round .. .. ........... .. ..... ......... .... . 2 2 Vel1ebrab Illllch larg.er thall d()r~ab (A) .. .... .. .. .. . ... .... ...... ... .... .... ... .... .. .. .. .. .... ...... .... ... ... . 8ul//.:aru.\
• Vel1cbrab not Illuch largcr than
dnr~al ..,
........ .. 3
3 I J dorsal seak rows at mid-body.. .. Cal/iopllis • More than IJ dorsal sca le rows at mid-body... 4 4 2 large m;cipital shields (B) ... ... .... OpiJiol'haglls • No large occipital shicld ... .. .. ... ............... Na;11
Ge llll S
BlIllgal"ll.\· D AtJl) I , I X03
The di stribution of thc gcnus HIIIlj.',(/ /'II.\ extends frolll A f'ghallislan in the: \Vest, through large pal1s Mehina inlhc easl. to Ihe so uth through the l'vlalay P c nin~ula , Ihe (ireater Sunda I s l and~ and Su lawesi. 10 Java and Ilali . Moderatel y largc tl large: Il1lN:lllar; bod round or triangu lar ill noss-~ection; tai l shurl: head littl e dislinci from body ; cycs small , pupil s round : 1(lreal shield ab se nt ; dnrsah s l11ooth. in lJ- 17 rows:
vertebral.; di~lin c tl y enlarged; ~ u bca udab enl ire. or intennix..:d with 'lIlgk div ided ~ubcaudals . The~c nociurnal ground-dwel lers nl1cn prcscnl th 'Illsches .. , innlfc n. IVC and even too I
Tab. S9 I)l slnhllll o n IIi' 1I1111}:lIrw-SpCl:le, 011 Burneo. Sab.1h BIlIlj:/1I'11.\·./il sciaflls (S,
II ~ I 11>1 K.
I XO I )
I lJ.lX I X43
HIIIIK lII'IIS .fl(l I';Cl'PS ')(I/II l'IIS;'" 1.0\1 Kill(" , IIl1l1g{//,W 1/1/1 ·let,/ ).\· //(/\ 'IC<,/I.I
]X2
R I·I 'III.\lI III ,
• •
Kinahalu
• •
-
Brune i
•
-
Sarawak
• •
Kalimantan
• •
Key to the Bornean species of
BlIIrg artis fasciat" .\'
(S CII NI 11)1 R,
I ROI )
BUlIgarlls 15 (rarely 17) dorsal sca le rows at mid-body: all ubea udals en tire, with the excepti on of the last pai r. 130dy norm ally e l1(;ircled with co mpl ete. alternati ngly blac k and yellow-co loured rings ... .. ...... ... ........ ...... .. ... .... .. ......... ........ .... .... . jfl.\"d alliS
• 13 do rsa l sc al e row ' at mid-body: pos terio r
subcauda ls paired; do rsa l surface no t enc ircled wi th blac k and yell ow rings ....... ........ .. ........ ..... 2
2 Dorsa l ~ urfac e wi th dark . white and red bands in the posteri or part of the body: tai l banded ... .... ... ............... .. ..... ....... ....... .. ....... j1(1 viceps b(l/lIl'/1sis • Dorsa l sur face un iforml y dark, tai l red without bands .. ..... ... ........................... pm'iccps jlOI·ice/I.1
{'I('w/" h" " fll.IC" l/lI 'il ",,·'D ,·k. 1XOI lX' (d c'~ nrl"\11 I""cd (1 11 R",,, I I 17')1>. 1. pi ' I ""' tH' toc .. til) : 'vlall\()()f ( "" Iah . 1IcJlt:a 1 IJIIIJ ~c / "' '' / fl \uafll' .
I IH Ni l \1<1' & . jO< lKI N I t) X4 24 . I
Dia g nos is Eas il y idenlified olllhe basis of'i ts ye llow and blat: k banded pa ttern. and by having a usuall y hlllill tail tip: head ha rdl y di sti nel li'o m bod : max . le ngth _25 em. ave rage s pc c i m en~ ahou l 150 ell l. h ;1l1a les s hOlicr th:l ll maks . l>Csc ri plio II O ne prcneu lar; 2 pllsioe lli ars: 7 slIpral abi a ls. Ihe ] ,.1 alld ~ II' bonkr Ih..: 'y..:, Ih ~ 6'" largl'st: I 12 tempo-
ra ls : 15 dorsa l scale ro\\ ;. at mid-bod y:
Fig. J 61! /JIIIIg tll"ll\· /a.' cia /II ' fro m
rCl1il1 ~ u l a r
~ I t 1 1\1'< ,
dLlr~a l s
Ma lays ia .
.1x .)
'1l11I(llh : ~()()- 2'\..j I I nlra l, :
Th i, ~lI al-. e i, 10llnd 111 I(m lan d alld I ill y rcg iom up III altlludc, or al least ~.\()O Ill , but II ilh a predilectio n li1r the lo\\'l ;l1ld ;.. . It Inhahih Ihl' l11argllis or 1(111 lall d lropl cal Ilel and dry ron:sts. tro pical and ~ u b l rop l ca l montanc fill'CSI:-- and sLrllhlands. and e;.. pecla ll ) ",an,p all d ll1ar:-hc~, planlalions. :lnd clllt l\ aled area:- lik e rice paddle ... , /JIIII,I!.(/J'I/\ /lI,\cilll/ll 1:- round In dry. o pcn area;., ill "cruh land, and fi el d, or on road;... ;" well", ill Ih icl-. I), l'Cgcla lcd dalll P pl al:\:;" In Ihc , icinit y o f lI'a ler, II here il i~ quite I'rL'qllc nt. Thi, snake IS a ,een.: llle. I..:rn.:;.1 1'1 a I ~ p c 'IC', whi c h IS prcdolllinantly nocturnal. cmerging. a ncr du ~ I-. from It~ da y- li"Ie rct reat in Ihe )! rUlIllt! or lIndcr vcgclal wn . logs and r()ck~ , II ulkn OCC lIr" In the I icinilY of human habitati o n;.. and IS fOllnd lInder pdl:;' ll f lIood , on palhs alld rO:l tk II may en te r water Tim, :--pceies fee ds c hie fl y nn ()ther ~nah:~. but also 1:lI-.e ~ li/ :lrd ~ :l lId alllphihialls ; li ~1i :l lId eggs ha\'l' al~() hecn reporlcd in the dil:1. !JIII/gllrlls /liscilll//,I' I~ a dallgcroll~l) \e IHlIlHlU ~ ~1l<1 k ~. lI'ith it highly dli.:ctil'e lenom ; howel er, rel\ human fa tal aec ltknt ~ hav e bee n re portl:d . \tlAI"\'Il~ (jI)I)2a) i'
'lX4
RIIII~a/'ll '" fllll 'h'ep s ba/Ilt!II ...,'is
Lo\ I Rlll( 01 " 193X tn /" hel lllnl'" l. ,)\ I It Jt ll d . \1) ~x. 1 ·~ - "~' I1{' IIt .. alil,l : "'CIlII~II~ RI I CI . Ill';" "',III ~ I I. "' 1I1"h"I". , ', 1 " 1111
/llI lI gOl'ln (/Ol
j ~ L:1
IJllng,fUII \ / 1t1l /(4'1", Il( II III " ,II'
PHII .I h 7 (1',111
1.
III
I H I "I I:\ I~I . ...\ \ 1( 1( dH ' \1 11 ~'" , 1IIIllh . ~·I " JJII II.t~ CII (" / 141\ / 4 II" II) ~,
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11)1)1)
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11 K! It":. ,llh,..)·\
nfl.,;' 11}1 >7 11, 11 Hit. I ) (tOO I q
11 4111
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I S96 ,7:! (rPIIHa r) ) ~ItI II I I K Il
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I C)· IX 'Y 7. f\ 1\, ~ H 1111'\1.
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li lt ,
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'I';l\unumil' f Unll11l' nl : The :Iulhnr IS COlli inn:d Ihal tlli~ taxoll I ~ Illuch rath er a good specie;.. Ihall a sub:-pecies , For a detailed dl~CUSSL(l1l ~ec VI)(,II 8:. 111I1'1\l ," 'J (I ()'n) . Di ag n os is
13 1'\)\\ s ()r dnr;.a ls at Illid -hody : n:adil > di slill gLlIsllcd 1'1'(1111 all t)lher ;"Ilal-.c~ or Ihe rt'g lllil by Ih colour patte rn lkscl'lbed below : max . lotal k'lIglh I .n em . DCSHip t ioll
Olle prcocular: ~ postocula rs; I 12 tl:ll1porals: 7 supra labi al~ . J " and 4 11' touching l'ye: 1,\ dorsal sca le roilS at 1ll1!1-hod y; ~()6-~~5 \'cntra ls : ;III a I st' ute l:n llre; -t ~ - 52 ;.ubea uda ls. l'ntire anteriorly. di, ided morc p()~ t e l'ln r l y I k ad alld lIape rcd, morc rarely ye llt1ll i~h bl'(lI\ n t,) ycf ll1\\: an terior p0l110n orupper hod, black 1,IIIh a lighl \ ertebra l line: dor~a l ~ ol'lirst IWo rOl'" whlll: l\'Ith blacl-. cc nl re~ ; po ~tcrior half or body banded hlacl-. and cora l-red. thl: eigh t hlael-. hands split by \\'hite halld ~ , graduall y il1 cre a ~ in g rrom one II'hite band on the tall to four white bands at m id-body, Anolher group o f\\'h ile hands 0 11 th e alll crwr hal l' () r lll ' Iwdy is lim ited III an indicat ion , The bel ly is white ante riorl y, with bl ack margins at the ventrals and coral red postl:riuriy 1,\ ilh rCl1la inlkrs of thl' bands (V O(;H & 11 01 I ~I""t\ 1'1'17, KI.< 11 & (,( 11 / "1 20(0) , I Ae all I::lapid~, thiS laxo n shou ld bl: {1\ ' iparou ~ , F:eo In~~,/ F. t h o l n~y 'I his krai t is fo und in low land a nd hilly rel..tinns, I\l ilh a prl'ciileclioll filr Ihl: laller. whl:re il inhabi ls
Elapid;u; -------
Fig. 369 IJlIlIg llrl/Sjlill 'IN'I },I' h(/Illel/I·j,· li'ol1l K undasang,
Iropical w<.:1 ('on:SIS . Th e ~nake is terrestri;]l, perh ; lp ~ serniaquali c, and nocturnlI gh a hi ~ h -ca ll o p y diplerocarp tiomillaled pri rmlry 1'01'L:st at Poring 1-1 01 , prill gs (SS O Ill) . Th e air l elll perailln: W; IS about 2S"( ', and Ihe air wa~ saluraled wilh \I,atci' duc III I·cg.lilar ,howcrs late in th c afl cJ'Il()(lll . BUlh snak cs we rc ahoul 1<;0 ClI1. Ilill thc l11ale was a little hit IUll gL:r than Ih<.: femail' , Thl' two snakes were IlI'i , tcd around cae h olher. Di sllllh: d by th e li gh l l) r a 10llrch Ihe ~ pe c i 111<':11, tricd to Ike, but did 1)(11 illl<':l'l'upl Ihe copulal lon , I'm at least 20 m i nut es, Ihe Id't hel11ipeni ~
COl ,
1-100
111 .
or the nwic cO lild be traced In thc t'cmal es cluOica hy Ihe , \ve ll ing ,
l)iSll'ihut ion
'I'lli ,
~ubspccies was f"ound :11 MI. Kill ahalu and Ml.
Tru~
Madi 11 550 111) in Sabah . 1\11. h:inah al u : Kunda san1.! (ca , 1-100 Ill). K cnokn ~ Ri\L~ r ncar K I
Uenlls Calliophis
(iR i\Y,
I H35
' lIl;dll o Illcdiulll-si7e d: Vel)' ~1c 11(lc r: head , mall , nol d istin cI rl'Olll hod y: eye~ ~mall, pupil round ; hod y rou nd in cross-seclion: sc alc~ ~ l11all I J Hm', dorsa l s at mid - bod y: \'<.:l1lral:- l'llul1ded : lail sill 1'1 : Sllhc\ 8:. L.\\\'S(I\J ~O ()I ).
or
l' lap ldal'
\VIl en Ill.:: Il:d Ih .::alenl·d, IIll' groun d-dwc lling alld IHH.: lllrnal ( ' (\r ~tI ' 11 01 1...: ;. e" llIbil Illkn:,, 111I1:! dl'l ...·llcl' bd1a\ lo ur ill Ihal Ihc)' curl up Ihc tall lal c rall ) lo lornl a llal ~ pll a 1. \\hic h IS the n prese nted , to Ihl' PO k llllal ;ill ;ld c r 1\ 1111 il s colourrul IIlltk r~ Idl' lip /\11 ~ p cc i e;. "f Ihi ~ 1:! ':11l1 ~ arc III iparoll", l' lg ht "pec ll'" an: 1..Jl(1\\1l . tllll o r whICh occ ur lUI 13url1l;o . and hoth a lso l)Jl 1\11. Kinahalu .
Key to the Bornean species of Calliophis Ventral "lIrl ~lCe or body and tail ullifnrllll y bright red or coral-red: more thall 34 subcaudal s ."".,."
..... " ..... ,.. ,"",., .. ' .. ''', .. '' .. ,.. ,' ;1If1' sl;IIal;s 111/'1,/1,\'.1';
Sat>a ll
• •
CIIII;"!'It;\ hi>';rgllllllelraflle";11 1131111-.1 R, I X5 ')) ('ull;"I,It;,\ ;111(',\ (;11111;\ 1111'1""'\ ; I Hi li ~I ". I X5 9)
IIItl
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I
:-.i""
20 I
I) Ill' In r:l li( ~:
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\ \ \·,,1 I lUl IH'll ( 'O IlIIl/I/,1\ /l1\I ' )!,t I/O 1 /.'/role/('II(t/)
I \\\ ...1 IN "'001
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IllJl
I
KlIlabalu
13rllm.: I
• •
• •
I
Sdrawak
• •
"-allrna lltan
• •
li)I 'L"~t s,
Culliophis bh'irJ,:ata fl!lraflll'lIia (131 I I "I R, 1X59) Flath 11'11 alII I
b;II;,.~ara (('(rowel/;lI
." ." ......... ,,.".".,"",.,"', .. ,' "
• VClltral smtilce \lrblld y alld tai l wil h hlack .:ros,,bar;.: k"s than 35 ;'lIb..:audal s ..... ' ...... " .. ... ." ........ '
'JII\
I~
)1 q
l)ia:.:IIH~i~
l ' a ~ lI y
Itlcnlilicd by Ih led head and ta il: body with ratlnll (ill cOlllra"t IOllIe IIlher sllhspc:c les ); r-- Ia \ , to ta l kn glh 150 e lll , :I\ erag..: U()- I-IO CIll : klllak ~ s llh ~ l a lltl a ll y snlalk'r than Illa le .... rt:
n('~crilltiufl 11l1«. ( 01 I(S\l \ IlJ4 X): (Jile large prcocular: 2 ~ Illall pos lo c llla r~ ; (, ( 7 ) "upralabi a l.... y ,l and -I " IOllc hln g e). . e: Ii 2 l e lllp()ral~ I ra rel~ I 1 I or I I I 1 I ); I \ dorsal ~ c al c J'(lW~ :llillidbody, tll))'"al " "I1O(lth : 23 LJ -J O(1 \ ..:nlral s: anal SCUll' L' lltirc ; .16--1') ~ lIh c alidah . di, idcd , Htld y dark (11 ' hladl ~ h blue ; a pair ofli ghllongilu dl mtililll's both dor... o lalerall y and late r:'lIl y. the lal e ra l one wldc r an d "Ilualcd untllc bollOlll twu ro\\ " of dnr~ al s ; h..:ad . la il , alld a ll Imvcr side" ( )f" hod y alld lail 1I1lilill"llll y hri ght red. I'I II ~ snake I... O\,lparOlls. hilt III deta ils (lrll~ rerrLldUCli\'.: biology arc ~n n wn .
trtlpiea ll\ l'lllHllllanc li)rl'"h , alltl p1
Cal/iophis i"ti}sli""lis fltl!passi (RII ·H I' R. 1X59) 1-. /'/1" 1/"'1,,,,,,"1 Hili ~I H. IX"), 201 - I) I'C Inr alil .\ : !'.. I"l""g. I ~orn~t) 1),,!t/}/~Ir{\ IIIlt"'lIIr"l\ "ill
/) /I/io/}iI/\
11I/( ' \'III1(lh\,
-I hi S ~Il a ~l' IS liHllld from "ca level lip 10 elevat ions oral least 1750 III II illhabil ~ lowland lropical wet
,.1'/"
i( I Xtj(' .... . 404 67: IH 1{.t')1 1l) 17' )"J
B Olli I NI,I
,\/OlH '''' d IJlt...'~/I1W'" 1II,V.1"olclc.'molll.\', '-:'11111 19.' , - 10. 1N ,\I,ul('oro 1JI(c.'\fillcllt, t.: \ '('/'c.'III. \1 "1111' 19X.' - 30. \1 \1 ~ML'" I 'iX .~.
Ill. 1')1\7 ' lX'I. I'/XX :
I 'i";. M " " " "
Eful"1!~' / EI hulug~'
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'\ 111 I IIIJ
,\/tlllI " II"t1 111/1',1.1/11011"
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17 ~ :M\I~\I" &:-' \II' ! ') ')\
1'l'J. lh
SIIII""' ,
2-11 ,217. I') ') ('c , 2')1 S\1I11 1 19' I 12. " HI N!I\I{/) &.
1'1'1 1
~I(I"'< I ''\
-,~2
( altto/ dllx i ll lt' \llIw!i, (/ht"IW,\,\·il. SIII\" 11\ ...... ' . H')l 200 ! - 2.1'/
'In
&. L w.,q \
Ijlap idae
Diag nos is Small. slen der, max . leng[h ahnu[ 72 cm. Dis[ingu ished tj'om related rorl1l ~ by the co lour paHem d..:scribed below. I)csc l"il'l io n One: preocular; ? Pos[oc ulars: I+2 temporals: 6 supralabials, 3'.1 and 4'1. hHlching eyc: 13-15 rows or ;;lI lootl1 dorsa l sca les a[ mid-budy: ~OJ-2(l3 ven tra ls: anal seu[e entire: 16-2R su bcauda ls, divided . Upper su rlilet:: alld !lank:; irid ..:~ee nl black : two par,II Ie!. pale grey stripe:s f1'om Ihe cyes over the parie[::II" and pnSl1elllporal" right 10 [he tail tip: a re:d :Ire:a on Ihe su pralabial "hiclds. interrupted only by a blad. bar hel ow eye: "uhla hial s anri prc in rrama xi llar:; wilh som..: black SpOlS. mol''': Llfthosc 011 lcmporal. paric:tal and frolltal shicld s: a reddi~h hrnwilmedial stripc 1)11 aillli il 5 sl l1all sea lcs )lns[erior 10 tilc p:lridal nLllch which may e:x tcnd up to the lail in somc individllal~..; prd'rontal sh icld wilh a ligh[ b.5 scales ill wid[h alld spaced by J to -I scak,,; occas IOnally thc underside may he unirormly red: upper si de ur lail wilh n:ddish brown. dllubl)' in Icrrupkd medial stripe; In",,"c!' side of rail red with black bands . This sl la\..e is uviparnll', layi llg tip lu4 eggs at a time:. Ec ulu j.!~· / E lh o ln gy
( ·!llliolllii.\' 11I1(,81;1/(/li.' (liep(/l ~i is fllllllli J'rom sea
level up to allitude~ of' at lea s[ 1750 Ill. with ::I prcdileetion J'or I(lr..:, ted hill y :In.:as. II inhabils kl\\lalld tropical wet f"li'CSt-;, tropical and sub[rop ica l wet Illonlanc lixes ts, and plallta[iolls. II is a IIOCturnal. secretive. ; lIt t! selll ifllssorial snake which li ws bencalh or in the fore:st litleI'. under rocks. l'Oot~ and fallen logs alH l oliter dehris, and IIl1de:r vegeta tilln clumps . Thi s ~nake feeds oilly on othn small. f'ossorial snakc". I[ ha~:1n dislinel dclCnsivc di ~ p\a y. whclI IIHll ested. il !latte:rt~ the body and rai ' '':~ it s laillO show the red lInder~ide nf it. SOIllC[imcs i[ lIlay Ilip ovcr ex posillg [ite hright pal1ern ol' the hell y. (Jne specimen was dise:overed cross in g a road ::I[ an e leva[ion of 16tH) III at arollnd X.OO (1 . 111 . ( '1 \1 KMi lS I ')')4b). r-,·1 \1 K\ It ~ & S,\l ll f{ (199]) re:porkd
tI.
g
Fig. J7U ( 'ollio(J/II',· ill/n'/il/a/is //II'/Jllsvi Ii" 1111 St! .. i lau -
:ilau
\)fl [itt: dkcts oJ'a bite inlli c1cd by this subspecics. Ii resu lt ed in ~e\c re local s\\'el ling which, hOlVcver. had tii sappeared ;lncr tWll days .
Ui stri bu rion Titi~ snh~ pccies
is ellliL:mie I(ll\orne(\ and has bce:n round in all parIS (If the: island . M I. h:in abalu : Kiall (')O() Ill), Blintill Tu itan. herween I k adq llarl crs and Kundasang ( I-IOO-J 550 m), Kal11htll'
CJcnus Naja
L At ·RI ·1\ J t,
176X
Thc genus ;\'uill inhabit s a va~ t area incilidili g large par" or i\ I'rica and i\ ~ia . Moderaldy large:: slender: max . Icnglh ahout 2 nl : hodv mll~clliar. round in cru"s-s..:c ti oll: laililloder
or
3~7
Tit.: roud ~ pcctrul1l ,r th.: c()hr"~ compri ses smallcr rod.:nh. hillis. li~lt . alll ph ihian';,li Lards. Jild ~ Ila h.e: ~. /\ II the IIlC ludcd spcc ics arc ll\. ipOl rnll~ . /\ ltog":l her I X ,pec i e~ or NI/jll arc k nOl\'n. o nl v o nc or which nl' ellr~ l)ll 1~ (lrl1eo a nd nn \ Ilt. Kinabalu .
Najll
.\'IIIJIlIINIlUI
\ClIII II I /J/hltllll\ \ .: 11
tunliit) : ~" I " I, ,VCl/t l ll'IIJ((titWI\
\1 1 II
1 :-\~7
IIC
277
I)pt'
'\ lIlll.ll l
SOltllf"I/'lIt lr,lln \.11
IUl'alil ):
,\lIlUclfl'rllleI
Me I I.I·R, I XX7
IqO I ,l : h7
I~ III-il '(.I I~. l };ll( I~" .1:<:' .tfh.'1 \\' 1 -.11 K --\: I II
·un)
1 9J I · I O. ~X. ~2 , 1\.1 \N I IH\
I l)X~ ,
ml,d'l'n,
~.lliI\\ ;\k l~\1l
\ (1/(11111'11111(1/1\ \, tl.
'\'O/ cl' h l / OIll IUlt '/lI\ , '\\1 1 II
I )pt'
Dcscr ipt inn One prcoclilar: 3 pll~lll 'lIlar,: 7 sllpra lahl ak .\',1and -III, luuching cyc : X .;ublabiab; I. -25 dur ~al scaic rows al mid-body, dorsal~ ';l'11oolh ; 111 2-206 Icnlrab; anal Setlle entire: -10-55 suhc:llldais. tltc anicrillr oncs cnllre , thl po.;tcrior (IlK S di\·ided . <' ;n.llind c(lloration yc ll ow 10 yellowi sh greell , 0 11 Borneo also black upper s ide or the head hrow lli~h yellol\' or olive ; lips and Illmat yellowish : hood II It lh)U I pattcl'l1 : e~· es dark ; tongue pink ,'enter usually li ght yclkm , pale yellow. or black: individual specimens Wllh irregu larl y s hape:d brown ,pnb nn the underside or Ihe hood . Juwnilcs occas iollall y \I ith lighl Iran"erse l ine~ . i\ clutch (lrthis speclc~ ma y comprise up to 2.1 eggs.
H) \LlII' milO,
I
Iii N It ,\~ ll
oL\ . \ In,
KI N
'~1 .
\,
\1 1\'1\ ...
l<J t )4h
Ecn lng~,/ Et holo~~'
II)X ·' 2· '
\,{/.{ .\ 1111), 111 '11111. \\ ' " II ~ '" I IIIIk l'l I') X')
.)1).1 : ""
I III
I,
1')') I .
2~17
DiagJln~is
Distingul';hed rrom all olh..:r ,nake~ OcculTing nn IVII. Kinabaili. e'l.cept the clilubrid .II({ c l'O/ilsllwdo/l 1'/lOdoIlW/".I. by Its abilit y to Illl'll1 a " hood " . No lar:'e sh i eld~ hehlnd the parielal~ . \la'l( . to tal icn!!tl1 150 Cill . av.:rage I (lO- I_O cm .
Fig.37 1 JlII I:llik
.VoIi ll IIIII/{I/I'U//(I\ro ill
SOlllh Thailand .
is l arc abo wkcll . I'lli:-. spec ies i~ I ery cOlll mun l'I'c ry1\ here 1\ ithin Its rallge. It i:,- nol aggrLsslve. alt hough it display~ it ~ defensl\ c po~lurc at the: leasl prll\'ocalioll and dcfl;llds ibclf vigol'OlI.; ly when cornercd. T hi ~ snake can spil il'; ve nom accurately Inwards an aggrcs,or. It i:; a dangerously I Cl1(lIlltlllS animal , wit h a vcr)' potent vcnom . , lIlll':roUS human l ~llalili l.'s have heen rcplll1ed from its hi tes . .YlIill SIIII/{//IW/u
Distrihution
The ,pecie~ ranges rrom the Malay I'cllill suia and l3ollle() to Palawan 1 ~ 1 ;) lld ill tile Philippines . II is fUI1herll1ol'c 11<11 ive I(J Su matra an d Ihe , 1Irroundillg i~lands . II IS present ill all pari s or 13l1rneo. IVIt. Kinahalll: Ki all . BlIlld\! Tuha n ( 1370 m) .
JXX
- - -- -
Py Iho n idal:
Pythonidae T h l:~c arc Illcdi nl11 -sin:d to largc ~ Il ak l: s fo und in Ihc lro p le~ and ~ ubt rop i l:s . T hcy inha bit nl:arly a ll type~ of hiotop..:s. Ihl: rl' an: dcse rt d wdkr~. arbll rca l l i)rc~ t (\1\(: 11 ..:1";, a nd s p el: i e~ wh ich ar..: high ly aqua ti c. ll li; ir skl: kto ns illclJl(k rudim l: nl ~ o f a pelvil: gi rdk . and their j a\V~ cil rry largl: numh er~ or tt:..: th. Their lab ial shi el ds arl: pll tl:d scrYing a~ hi ghl y tkvdo pcd se nsory organs wh i( h arc ahl e In p..:rl:t:i\( minu te difll:n: n(es in tcmpe ra lure. Pytho ns reproduce hy layi ng eggs \\' hich, ill so me :- p e(le~ , arc aeli\t: ly ill( uha kd by the re m;) lt: snak..: .
Gellu s
Py tholl D AUIl II\ , I ( 01
Thl: l i":IlLIS P,·t/UIII i ~ \\idely di~ l ribu l ed In larg..: parb or A fr it:a and ,\ sia a nd . wil h e n/lOll r(, l iclIIIIIIIS. conta ins o nc of the Iwo 1,Irgesi specics o f ~ na kcs O il carth . Somc spec il:s llfl his gl:nu s are \ ery "kntk r, lllhc r,.; arc ralh..: r plump. I-Icn d always disti nci fro m hody: distingui shed fro l11 relatcd gcnera by a vt:ry lk cp first lab i,i\ pit and th c a hsc n(..: or a p ica l p ib (t.;NI)1 )l\\'f)(»)l
& .
1'IMS()\J
\\'alc r bodi es. I hC lr dici con~ISlS pri J1l arily ofma J11ma b and b ird ~. more ra rc ly o rn:pl iks . /\ 11 ~ pec i cs or this genu s arl: tl \' i p aro u ~ \\'il h the rc J1l ak~ activc ly incuba ling thclr eggs by co iling around th..: cl ut ch a nd rais ing Iheir hody tl: lllperalllre. I he gcnu s co lllpr i~es len ~ p ecie!\ . Iwo or whi c h li vc O il RtlrllCO, both Oil II. Kinahalu .
Key to the Bornean species of Py tholl
1l)l)O ).
l\tlost of tht: sp..:cics Ill' / '1'1/11)// arc pri mari Iy terrcstrial snakes and nnl y a few coul d b..: lel'11lcd :-cllliarblln;a l. Thcy inhab il a \\ ide ra ngc or hiotopes. and ca n ()cca~io n a ll y be 1\Hnld in Ihe \ ic init y o f human sellkllll' nh . They arc oliell eneountcrcd nt:a r
Tah. 6 1 D isl n h ll ll ll l1 " I /'\lhol1 -s p':,·I<.:s
lln H \lrn~(l.
I P)'lh ol/ br";I l'IISfl';II ; S 11 '1'1' \1 11"1 K. IXXO Pylh ol/ rl'l;..,t/(I{/I.\ ('1 ' 11 '11 1111 H. IXO I )
Py thol1 breitellsfeilli S II
Rostral and li rSI 2 ~ lIpra labi als with a pit ":3c h; less than ::!RO vc ntrab .................... bre;ft'llMe;lI; • Rostral and firsI4 slIprn lahials wilh pi ts; more than 295 ventra ls ...... ..... ................... ......... r ('(iclI/(I{IIS
11\f)"Cl I I Ie
I RRO
Sahah
• •
KlIldballi
• •
Brlln"l
K"ilmOlnlan
Sa l ;l\\a~
• • I •
I
• •
bd ow the ..:ye to the na~al: dilr..:ring fro m Pl'llto// by the l ~l C t Ih al Ih..: all ie rl or pair 0 1' parie ta\:' arc ill hroad conl ac t al the ml:di al ~ lIt ure. w h c rca~ Ihey arc not or oll ly \\ cakly in cOlltacl at f>l'Ilto// Cltr/ IIS ( KHl( .JI, B \)(J..I ·)( & Sllt NI 2(0 1). Ma, . total Ic ngth 170 CI11 . Cll rt ll.\
1)1 '111011 hrc,tl i II\J('im ') I l ' I) '\ l IINI 1< .
il ): I L"\\ ...·h.
I. XO ."76
I ) p'-' Im.'al-
B tlfl ll'\l
"1'111,,11 hn'I fc..'I1'/(' (III , KI, )tdl, B \ IH"'~
&
"'111'1
~ ool _
122
nia ~ n os i s
A snakc ort he low lands. 8od} \cry slock: lad shon : di sting ui shed from PI·t/lII// hmllgel'lll/ui bv a singil: supraoc ul ar shi ..:l d abo\c thl' Cy'C alld a n m or ~ m a ll or g ran ular scales e,wcring the arca fro m
Dcsc riptio n
Very 111 11 ~c l1 la r, stoc"y hody: h..:ad d i~ llJl ct: tad short , 2 prcocula rs. 1-2 pllstoc ul ars' I ~ upra()c ul ar : a s..:rie:- o r "lI1a ll , gra nular subocu lar ~cal<.:-, e'(knd illg
l'llilihat dallces ha \ e occasionally hel: n l)hsel'\'ed (L \ I "I 19l)7. K II('>IIl '\ II( 8.:.W IIIIM\ N !')I)X) , ;\ner
a peril)!1 (11' gra\ idily or ahoul I,() to 90 da y" up to 3:! e gg~ Ilia ), he laid in a clutch . in nnture u ~ uall y at
I'il!. 372
" l 't/UIII "/"{ ' I/< ' I/\/("II//
!null
I'urlllg,
up 10 Ihl: na ~ ill : lI- l:I ~upralahialli ; 4X -6 1 dor~al~ at mid-bod y: 150-1 (,9 \'l:lltrals: anal SC Ull: I:lllirc: ::!::!32 ~Llhc a udab lA l'tl\ ,\ & Alltl 1999) , Uasic co loralitlll brown or y dlowi~h hrown . laterally \\,ith lighl , darh-hordcred spots: a row of lighl ~pllis Ihe spine. hroadl:nillg towards lail and fu s ing 10 form an unintl:rrllptnJ line frallll:d w!lh darh. Head li ghl ahow. \,ilh nn indica tion ofa linl: which begins on th e r(l~tral and s plils lip into -l lines postl'!'Iorly; a darh ~t reak from in front of Ihe eye 10 onlu the hody; all ohliqlll: poslocular stripe fro m beneath Ihe I:ye ha ckwank Up 10 J::! eggs (50-70 .' 'i-50 mm on average lEI{111 ' I 1' ''1 1.'1 al. Il)(n l) ma y bl: laid in a elu leh , Neonales measu re -,-,--IlJ Clll and weigh around (, I g,
~n lllc tilllL hel\\'l:c n Jul y and '\ tl \'l:lllber. The eggs arl: depu~iled ill heap ~ or gras~. aCClIllllllation~ or plant mallcr. ~ llIl1lp S of dL:;ld COCOlllil palills. t>r old rt)lting. hnril.ontallrec Il'unh" "I 'IIi"/I hrei/(,lIs/dlll I'clllaic s incubale Iheir egp aclivd ) hy coiling. mnl'l' li g hl l ~ l'l' mOI'L 100:-'1.' 1)" a, the s ilu;Jli('n may reqllire. around thl: LIlItch and rai,ing the bod y tempL:rallll'e IhwIIg h I1Il1sc ularcontracliulb il'and when nccl'ssary, It i~ 11(Issib lc Ihal the remaic eWIl wlkct s \\,all'!' Oil her hod y tn suppl y IllOi SllIrl: to hcr eggs , Spoilt or Inkrlik eggs arl: sorwd (luI and pu shcd aside with thc head, ;\t a tcmperaturl: or ::!lJ-3 1"(' llil: nC(lnaics hatch aner 5<)-61 da ys . One to two days ma y pa s, belwecn Ihc clillilig open of Ihe eg!! shell and thc ac tu al hatching of a bahy, I'vlaks malure al an agl: or Ilm:e years. reillalc~ at four ycar~ of agc,
Uislrilmliun This spcc ies i~ a Bnrnean cndelllik rang ing over large parl~ or the island. 11. Killaha lu : Pnrin!! (500-6()O III a,s,I.).
Pylholl r elicli/alliS (Sl'))'\JlIDI ' ({, I gO I ) <;, IINl " ' 1 ~ . I)\() I 26-1 T) p ~ Iota Iii, : ,,"0' !,:'H' n: ha'l'd Ull .... pC ..' lIl)l·,I~ <.:ih,' J hy '-;,1 1:\ f't'lIl " ()EIl; II( ' (171·1 pI (,2" " moll ilia Ill ... 01 Japan" , 17J5 ' pI. 79 : ,1\ l"TOI). and " " tl\;1 11 ,,1''''"'''' 11 7 I' . pI. KU, 111 L' ITn!'). h " pCCII\TI) 1'1"/11111 r, ' /fI'UI,III1', \1 \1 .... \ 11 "i ll)l)2.L I l~ , Il)lJ..th. ~4h IJ""I'(' //(III(l1
I)ia g nu~i' E colu g~-j E lhul ogy
Thi s ~ pl:eic~ i~ parlicularl y cOllllllon in Lulti\alCd areas , lI ne it Ji \'es in !'ICC paddies, rubher plilnlaIlon~, coconut gro\es, a nd othl:r Iypes of 'ieClllldary \egelalwil. II ~ pend~ l1loSI (lr i l~ lillle in rodenl s' burrows. pile~ of g ra ~~ , or eanhworb, Rm\l:nl s limll Ihe ha~l~ of il s diel (Elml I I N 1.'1 al. IIN7 ). bUI larger Illalllmais. c hidens and dueb ha ve also hee n recorded as prey. Allugl:l her. thL: biu l(lgy of thi s sulbpL:eies in the wild t~ still poml y hnown ,
390
Very lal');c, l1\a:\ . length ()gO Cill. but ndull specill1en ~ rrolll K.alimanlan lIlL:aSllrl: al'llulld ~()() Cill (R I<,)I II R I <)t)X). The rl'liculall'd pallcrn \\ hich covLT~ Ihe L:nlirc hod y I ~ a characlcri,>lic Irail of Ihi~ snahe . Dl'SlTiplio n lonl y animals from nl)IllCO, AI
1 1\ ,\
& AIlI '1 2()OOa)
or muscu lar.
althollgh slender hnild : lail dl~tin e t frolll body: head vel): di:-;llIlct : eyl: with a \'ertica l pllpi l: rn~lra l and IiI'S) Il1ur suprn lnhia ls ",ilh dCl'P
Pythonlda.:
pits: 12- 1() ~upralahla l !>. 6'" to X'" III contact \\'llh cyc: 3-5 It'real:..: 2 prem:ulars: J postm:ular ': 61-X2 dorsals at mid-hody: 271-.-,-"n \,clllrals; anal .;;eute cntire: 55-9X prcdominanlly <11\ ided ~u h caudab , Ilolh se'(es posse~s ana l spu rs, but those of thc males a re larger. Rasic coloratiOIl or body vary11lg from pale ye ll ow 10 dark or o live brown: back covereu with l'Ou ndi!>h . oval. or rhombic nlark11l~~. fi)rming a rcticulalcd pallern th rough ru~i 11l: Ihesc spob arc yellowish 10 brown and bordered I\'lIh black and YCIIO\\: a black lonl:!itudin al linc I"ro111 the lip of thi: ~nout \(l Ih.: nape cro~s1llg through n:ntr' of head ; sides of heau with a black tcmporal ' tnpe hd\leen eye and neck: helly yellowish, sparsely speck led with black . When exposed to bright light , the body . hows a metallic gleam . 12 10 IOJ eggs (10.3-11.(, , (1.3-7 ,5 em): h ~ll ch lings measure 6O-XO cm an d wcigh 12()-170 g. L l' ulll ~y/E l hlll()gy
i!> 1(1I111t1 rr011l sea klel III up to ahove IJOO m alt ilude ami inhabits low land PrimaI' and !>econdar~ lroplcal \lct li)rc!>t~, tropical open dry ror.:st~ . tropical \\'.:t montane forest~. rocky ~hrllblands. SII amps, mar!>hcs. plantations. and eulti\,ated areas, a!> Ilcll a!> suburban or e\'en urban area!>. II is most 011l:n IlHlnd close to water, along river banks and canals, and is frequently di covcrcd in or around hU11lan settkments, ncar or inside hOll ses , I\ctive at dusk and main ly al night, this Python is telTc!>trial and se1l1iarborcaL orten conceal inl:! it sc l r in dense I'll Iiage or Itlll tree!> o\'erhanging water or In large hollol\' tree trunks . It IS also a \ery good ~II immer. I l~ di d eon~ist!> ehidl) of' 111;I1II111als ~lIch ;'" larg.: rat~, rabbil~ , small deer and dLlme~l1e anllllals (cab, dng~ and pigs), hut it <1 1,0 takc~ birds, e~pecit he considered potentially dangerolls ani1l1al~. (ir'l\. idlty in /'I'l/WI/ r('licIII(/IIIS i~ ~aid to take (,) tn 10) day~ , 0\ ipn,ition takes place mainly during thc 1llonth, or February and March, \\Ilh clutch.:!> heing deposiled In ea\'ities in Iree trunb. The female coils around hcr eilltch . but lhe nlll!>clilar Prl/f(1I/ Fl'lil'lt/flIU.\
Fil.!.373
f'I'II/o1I rcliclIllIl/lS
rrol11
I kral11a~\lI,
Sahah .
eom ulsion, kllO\\'11 li'om olher ~pee le, or /'\ tllIJlI arc rarel y l,bsened . The youllg emerge aller a period nfabou t three lIl()nlh~ . 1aks 11Iature ,tikI' _-3 year~ . female, aboul olle year lall' r (arter \ " I}II( :'11", K M(}l1K 1926, TKJl li' \1 IlJXO, \VII( II( 19XI. R()<" & M \lUI (' 1l)9 ~ , I: KIl1 I1 d ai , 1997), Di s trihu lion
Thl:
1',t/lIJ11 /'( 'IICIIllIIII \
Itllllll'ollllg, Sg K'I'''''gtlll.
WI
Typh lopidae
Typhlopidac I'his 1:1Il1 d~ 'ol11p ri,es ~mall , wUrllI- hke. ((),soria l ·;nak~ ~ \\hosc bodies mallllain ahou l thc same di:lIl1 eter throu ghou t thc enlire leng th , The head i" nol di~tlll ci frolll the body. the t a i l l ~ short with a hhll1t tip, 1 he eycs arc hidden h e n~ath sca le" alld il1 visib ic in some s pecic~ . I'he sca les or the hod y arc 1:lirl y untfi,rlll II ith 110 modtfll'u Icntrals or ~ ubC: ludal " . Thc) ked on anI:-> a nd !cllllites. their larvae and ct:t:s. nthcr sma ll Ilbl'l'l S. and \\'orll1s. Most 1) 1' thc rcprcscnlatl\ es or thi s lilillil y reproduce by laying eggs, bUI a lew givc hirth to fully dc\'cl opcd young. The two ge nera pn;~ent 111 the Sun
Typh/ops sp. A ~ pe ( le s urlhe gl' llli S T'Jih/IJ/" Was reeellily ph,,tl\graphed on Mt. Kinahalu . Its wide ro"tral shic kl and thc hi gh ,'l\llnt ()I dor~a l sca lc rl\\-\ ~ , uggest that thi s Ini g hl be TI '/Ili/O/)S hIJ('/,!,O('h' U R( ',\I: I I(S \I \ . 193-1 (W \1 I 'I( II pcr~ , CO IllIll . ). TI,/Ili/II/II' k"ekl.O('hi
Fig, .175 / 1'/11110/", sp. Irnlll 1',\1 II1 g .
392
has so I:tr b ~e n knol\n on ly li'om a sin,!.!ic ~ p eci men co ll ected 1)11 Ilunju I, land ulrthe easl coast or Bornell. Ihe pholog rap h d()c~ ul1l() rtun :llcly not pcrmll lin uilquestillilahic ideilltfi ~aii()n . Ir II is not li,/Ili/U/I," ko('klm('/" , II IS most certainly an as ye t unde"cnheu s r el' l l'~ (W \1 I \( II r ~ r ' . (l)l nnL I. Loca lity : rvtt. Kinaba lu : Poring,
Xenopcllidae -----
Xenopeltidae These arc ~nakes ofillcdium Icn g th~ with cy lilldri cal bodies. The boncs orlhe skull arc \\cll-united . Till: prciliaxillary bonc touches the llIa xiliary. bearing well-developcd ledh in line \., ith max ill 'lr), tcdh . The ectopt ergoid is loo ~ L:i y attached to the ma xi ll a. The prelhllltal is sutured h ) thc lIa,al. The (knlary is very loose ly attached allt crio rl y 10 Ihe anlerior elld of th c articular. but fre c bL'illlld alld Illovable . TllI:re is no corOlloid
Genus Xel10peltis
ROil.
1827
The ge nus X ello/wl/is inhabit s a large portion of" ~ia, ranging from the ~(l ulh oj'( 'hina 10 Mya nma r, and SUllih 10 Ihe Indo- , u~trallan /\rc hlpclago. It al~o inhabils ~nllle or the I)hilippine i s l an d~ . '!ax . kll glh nearl y I metre; head large. nol sci o il frolll body, cUliered abovc wilh large. sy mmelrica l
sca lcs. tapering 10 a \ ery Ilal. round. broad ,noul ; cve ~ Illall. pup il \ertica l-e lllJ)lic: hud y round, CO\ercd wil h large, very SlllllOlh ,GIles \\.Ith an iridcscelli shecn ill bri g ht , unil ght. 15 dorsal ~ca le ro\\', ; lad very short. abolll 10"" nflotallenglh: anal ~ClllC d"'lded: s ubcaudal ~ cn tlrc. The ge nlls \ '(' IWI }('IIIS compri ses on ly t\\'o s pel' ie~, une of \\hl ch occurs on Hornell and lIll MI. Kinaba lu.
Fig. .176 \'''"111'''/11\ IIII{(u/"r from SOlllh Thallallli
, ~, .'~
~,
...
,.
: ; /. '
,
, . :' ... 'Y' .'
.
... " "
'.
,
.. . ,-
'.
,........ .
:-i
"
.'
_•• :':, 'J," ,- :: .. • .,;,. -
..... f
• 0'
..
•• :::>
Xellopeltis
II II icolo I'
Bcm, I X27
F:cn ln l!~ W t Ilfllu l! ~
In hab it s 10\\ land an d hilly regions c()le red \I·ilh Iropi ca l \\'ct and dry ro res t~ . hilt i ~ chi clly found close to wa ter in upcn \'cgclat ion ~ul'h a~ Ill ar~ hc s, ~c rllhl a nd~ , hedges, plantali ons. cu lti va ted i1rea~. ri ce paddies, in gardens alld olhcr 'o ubu rban arl'a~ , and along ditche s and ~ trea lll ~ . It I" semil()ssllrIa l. IiI IIlg lo r 1l10 ~ t nf the time lInder Ihe ~ url~\(x In \\cl. loose or sandy ;.oi l. It Illay dig It s O\ln bllrf'(l\\,~, hlllnwre oltell lil,cs in h olc~ In the ),!ro lilid and <.lcc upies burrows ol'lginall y excavaled by rodents, is lo und lI lldn rock:;, Iree ~t ump :-,. clumps o f !!-ra~.~, logs. <1I1d alllong rtll,b and dcca" inl; \'cgc talioll . l\etl\c alnight and al da y, it Ilia), o lie 11 hi: seen foraginl; anll)llg \cgetation on the gro und . Thi~ ~pecic" li.:ed~ on lizard s, nlher "lake,.,. "'lllallll1
.\ eIlIJ/ I<'Ii;, IIIIICU/U/'
l) ia l!lIusis
M a\ total kllgth
nltioll
~ec
<)()
g,eIH.:nc
elll . tai llcll),!t h · 10 CIll. Ddi-
dia gl1tl ~i~ .
lll'sr r ipfiun Ilead Ilall ellcd ; ~ 110ut rounlkd : Iln >. tril ~ hetwccillwo lI a~a l s; li-onta l a~ long a~ it>. lbtal1ct.: frollllhe rostral; pre,' e lllar~ large; 2 p\l ~ lo cu lar~. the :lIlt cri ,' r UIlC largcl'; I ~ 1I1all ~u praocular pn.:~e nt . slllall cr lhnll Ihc upper po ~tocli lar : a large. ~epta gullal interparietal ill thc ecntre of,,) pari e lal~ : X supralabi :, ls. the lir,,1 in eUlllacl \\·Ith Int c rn a~ab, ..) '10 and 5'" in cOllla 'l wilh eye: I pair "I' ~mall c h i ll~hleld s. in 'ontaet with the three anterior ~lI hlahiaJ:., : 15 dorsal ~calc ruws tilroligholll: dursa l ~ ~ nlll () th : If,")-I<) .~ ve ntral .. : ,lila I ~c ute entlrc; 26-32 dl\ idcd subcauda k Lpper :-,ide hl'(>I\'1I or blackish. Indc>.cenl In bnght :-' lInli ght : ~calc~ with li g ht margi ns: s upra labl al sca k s pale yelluw: lowcr "i de \\ hll c; young \\ ith a li ght co lla r. This species is 0\ ip:lmu~ with a cl utch comp ris ing hetwccn f, and 17 eggs .
I)islrihlltiun The dl sinblltlon or thi s spec ies rangcs rrom Cambodia in th e cast and Chilla in the nort h, sO lllh thnlligh Myanmar. '1ha iland . :1I1d the Pen in,.,u lar Malaysia. 10 large part s or I nd o ne ~ ia io thl' Phi lippine Island~ . It i~ also wide- rangi ng on Romeo . .VI I. Kinabalu : Poring (I IQ-('ollcction).
'1est udincs
Testudines
(turtles, tOl1oises, and terrapins)
There arc about 300 species of't urtlcs* dis tributed wn rldv,ide ill tile entire warm and 1l10st of the temperate lone~ . They compri,e terrestrial as \\ell as aq uati c reptiles marked for their bony ~ h e ll s \\hich :1I'e ill most cases ri gi d, but furn ished with hinges and therefore movable in some gl:nera . Their venehral \:OIUI11I1' arc large ly firm ly fused to their shelb al1<1 . as a eonsequenCl:, an: freely mmable onl y in thc neck and tail portions. The head, lill1 b~. and the tai l ca n bc retracted more or less (kep ly into the openings of 1111': shell. ;\1 1 species inhabiting the Sunda region belong to the suborder Crypiodira whi ch retract the head and neek into the shell by verti ca ll y bendin g the vertebra l column int o an "S". Their Jaws arc rurni ~ hed with hurn -covered shear- like structures and lack teeth . Depending on their w ay~ o r life, these c helo nian, have distinctl y flatten ed to roundish l imh~ . Fredy 111()\ahle toes arc not found in all fa l1ltii es . The ~e anima ls ean live tll ,·ery o ld age~ with some s peeie ~ reaching 150 yea rs. They rep roduce by layil1 ).! egg~ \\ Jlh clutch , i/es va rying between 1-2 and up to 100 eggs. Many s peeie~ are known 10 po sse~~ the abi lity for a delayed tCnili/atiOI1 (Al11phigonia retardata). Juvcn ile specimens o ft en ex hi bit ~hell shapes and eolnrations di ncrent li'om those of adults and usual ly require several years to mature . Uorneo is home to II species whic h re present 3 li:llnilies and 10 genera . Ob 'ervalion'i on Mt. Kinabalu arc limit ed to 2 spec ies rcprc ~e l1ting 2 families . IhlTl'U'l'd "1".I(lr l(lrtlll\~' ;tn.! l~rraplll 'l
Distin guishing features in turtles nuchal
gu lar
keel Illarginal
hum.:ral ..J
l!
(;l.sw i
, p:
axi ll ar
g
pectoral III rrarnaregina I
'';
U
"r: "u '-
c..
c
celli ral
-f
t.IJ C
~
"
abdomina l
0'-
<:> .:=
IIlgulila I
~J
femoral
-'2
anal ":
or carapace (CW)
Fi/.:. eX\, 1II Seutes of the shell or a turtle .
Key to th e fa milies of Bor-nea n turtles (aner Limbs elongate, ,ub~ t antially ilaltened , Ilipperlil..e (A) .. ....... .. ........ ......................... Cheloniidae • Limbs short. morc ur less round ................ 2
M ANT III'
& (iIH)\~\ l i'"
IC)9 7)
B
2 Shel l so li , lea ther-like, ilat , without horny scult:s (n) ................................................. Tr io Jl ~l c hi d
• Shell hart!' wit h horny
~e ute ~ (e) ...................... .\
c
J Peet with di stinct
toe~ (D) ............. Bataguridac • Peel without free toe~ , "elephant r()ul -li ke" (1-.) ... ............. ...... .. .... ... ...... ..... ......... ...... ... i'estudinidal:
,(»)
Ilal agllmiae
Bataguridae I ill: 1 ~lInll y Halagllrl lh\,' c01l1 pri ~e, ahuIII 70 s p l'cil:~ whicil are: distriblllnlill We~1 and Suut heasl l:ilrope, Nonh 1\ 1'1'1 ca. Wl' ~ t. Cl'll ll'"I, S(IIIII1 , and SOlilhea I As ia . I he gl'11US Rliillo c /<'III1II1 ·.\ I, Ihl' \)1lly re pre,e llta tive (If' 1111 , l:l1nily 10 occllr In Cen tral alld South A me ri ca . Sa\l.' Il lr a 1<"\\ cxeeptions. I\alagurid" arc lI ig hl ~ ' adapkd to a li ll: al and In waleI'. l'hl.' l1· bon y shell " a rc lilirl y variabl e ill ~ lIape. but u"uall v ralher nal . and :II\\ays covered \\" 1111 lIorny scu ll:s . Both lIah·es Or the ~he ll are u~ually ri gi d , hili ill ~nnl,' spl.'l'ie, IIIl:Y arc united lIy COll1lecl l\'C li"slIe Ilial allow ~ for ll exdl llll Y. Duc W Ihe,e h in~e~ parh (lr the ve ili ral , he ll Gill he II10vcd Ie .g. ( '/fortl spe:cies). I'he 11111hs bear rre:c wcs wllll'h are: illt.:rllllkcd hy 1ll':llIhr:l1les. l"lIall~ \1 ith :; ,'1:1\\ " Oil Ihl' rrolll and..J c laws \HI 111\; 11Ind i'eet. ;\si:11l \()nOI,e~ an: III ge llera l om l1l vun;". which con"umc alli111:11 :" well :b vegclaria ll rood. hutlhere arc a ls\l ,(lI1le: ~pcc l ai l /cd k,'dcr, . '1lIey rep ro du cc by ia yillg eggs 1.\ ilh ~ol i. p:m.; h- Ilke shell , 111 lar)' 111); 11II11Iber~ .
Oil Lhll"lh.' O. IlIi " 1: 111111 )' I ~ repn;~e llkd hy tn 1\11. Kllla h: llu .
I) s p l:l"i.:~
III X ~e ll\.: ra with \\111 1' nile gelllls alld on,' "pcci es lIali v.:
Key to the Born ean ge nera of Bataguridac
(in pan aft.:r M.·\'IIII \ & (JI("SS~IM.. 1\ 19lJ7 )
6 or 7 Ix nlral-, (A) ... ...... .....................Vo/(1("/W/.I ·S • )ccntrnl " ............................... ..... .. ................. .... 2
2 Fl"llnl tCl:1 witll 4 claws ( 11) ....... ............. 1311/l1,!.!1II" • I'm nl tCd with:; claws .... .. ..... .... ... ................... 3 ~": lI1C " presl:nt (( ' ) ............. .. • Axillar and inglllnal ,ClItcs liny Dr abscnl . nn tl:rim ilnd pll"terior ~l:d i'l n ~ of p ln~ tl"\lll joinl:d by a hingl: (I)) alld m()vabk ... .......................... ('lIora
.1 A ,illar alld ingUinal
c
I)
.. Anlerior S,'c!101l Df pl astron with hinge (D) :tnd 1ll11\"able ... .... ................... ... Cl'dl'lIIl's • I'laS11"01l without hillgc~ ........... ................ ......... ::; 5 I'la'[l"On \\llh bnmn radial pattem (1:.) ............... . ................... . . ... ....... . ........... . ................ Hl'IIS('lIIy.~
• Plaslron withoul radial pattcrn ........... .. ............. (, 6 Carapace oli \'c 10 brownish. males with thrce black stripl:s and ~ h011 nasa l Irunk ............... .. C(/I!{/.~ I11 · • Carapa\:c dark brown to blackish. ma les will1Dllt lIasal tmll k and nl) ~ t ri pcd pallern ..................... 7 7 Whili sh ye llow spot~ on l:ycs. angles or IIlouth. and oc..:iput ( r). pl :lstron blad,ish ................. ... .. . .......... ....... ....... ... ..........................•\k IWI1/"l1cAid/1I
• No whllish ye ll ow "POl S on the head. plaslron unilllrllll y ligh t ... .... ........ ..... ..................... Or/ifil/
396
F
nat ag llr i d a ~
- - - -- - -
(fCllllS
Heosem)'s
S TU I\H oI- R,
1902
inhabit Ih ~ soulh wes t of' India. SOlll hAs ia , the S undn rcu ion. and Ih<: Ph ilip pinc b lanel!>, Scmi aquatic It: rrapins \\ ith a di slin<:1 v<:rt<: bral k~~1 on a n a lt e n ~d ca rapace: ;lIlk no r margi nals usua lly sillooth , po!>tcrillr o nt:s serrated: pl aslro n joi n<:d ri gidl y wilh carapacc; toes or the fro nt Ct:d with :'i. Ihose o Cthe hind Icc I with -l claws. Only om: sp<:c i<:s o f thi s genu s is rc pre 'cn t<:d (lll H o rn~ o and also occlirs on MI. Kinabal u.
1I{,() ,I'i'II/_I 'S
<:a~ 1
Heosemys spinosa
(G RA Y,
183 1)
Fill'" '1'"1"'" (;"" . .1 t ( I XJ t) 20 '1I'Pl' tuctllil \' : (' c Ol aOl g. h'r,'! 11I 11' l(lIW "", J \\ , 'I IS" ~ 1'17<) ' .\.\. fi i!. 1201 {1t 'U( 'II/I 't/a ''I'f,t1 l!.lel"l 'l'(,l1,1!, /('r;. (nO l /" .\ 111110 "!,c'ng/cn (j \ I H IN. I ?X'») 'v\." 11 11 I I'IX 2: .>1 ( il '/ J('IIH 'dll "/ '(,II,l!,/( 'J'I . (lh)1 7, '\ 111'/" \'/ ', np.lcJ'l (j \H I I"'~ 17Xt) } M \ 1 ~ \It " I <)<)-1 : 2~(, Dia~nosis
la rge (C L li p 10 22 cm ). bl'(1\\ n tn brown I<: rra pin with a vcrte bra l I-<:c l and
F ig.377 Dn l',,11 "ic ll'
Vt.: nlra l v i.:w o f a j uvcn ll e III ,c l\\l ad lilt s pcci o f J" '/ls, 'lI n 's I / JIII/lI'(/
distil1l:tly ~e r ra t e d III a rg III a I SCllt~ s (limit ed l(l lht: ant crior ;\lld post~ n (l r (l nc, in adult s p~c illl t: n s ) . Description
111 0 d ~ ra t el y
rc dd i~ h
ITIt' n
nl' a JlI\t' ni k to s ~ l1l ia dlih
C' L lip to :n C Ill , PI. li p to I X <.: 111 : ~ h ~ 11 n at, \'<:ry wide: :'i ;,tronl;tly kcc kd <.:~n t rab. all wider thall klllg:
~ pCt: lI ll e n o f J" ' (1\1'1111 " " I , i/HIS i i ,
39 7
i3ataguJ'ltiac
en~ t a l s , each wit' a ~ pike in young, smooth in nld SpCClll lells: I I lIlar-ginals, terlll inating ill a poi nt ill juvenik~. lip, or Ih e ant.:rior and poste rio r marginals ~I i gh tl ) r~l1~ed. lateral lllargl11a l ~ of ol ll srceiJlIl~ JI~ i dJlIll ~t , ll1ll0th : I Jlll(;hal longer than \\ Ide. 2 ,up ra cau dals: I axillar, I inguinal. no infralllar)!ina h..; pla , tron large wi th wi lle bridge . ( 'arapacL: or i UI cni k~ a 11110,t ro und and \'ely ~ ri k y. 1110re elllllgate 1\ 1111 increa~ing age and progre~ ~ilely ~1ll()(l ther. I illlbs with largL: horny scak:-;:
-I
front lnes \\ebbed over hall'tl1l: lr kllgths. hilld toes only with ba,alillcillbranc!': inguinal region and ha:-.e ()rtat! with pointed tul1L'rcle~ : tai l lery ~ hnrl. Cnrapace n:ddish to n:ddi"h li ght or d;lI-k hrown . I'ertebra l "eel light browil throu gho ut in you ng s p ec il11 e n ~ : plastron yellOWish \\ ith a hnl\\ II ra dial pattelll Oil e;l(: h ,c ut e in jUl'e niks. In various sha dl!s or brllwn \\ithout rad ial pattern in nld ~pCClnH:n~ : head and lilllb~ brOlV1I 10 gn:y ish brown . Side or neck With a sma ll vcl lll\\'is h. reddis h. or li ght brnwJI ~P()1. Ecu l ug ~'/ I::t holu:.::v .ll/il/lI\'lI iJlhahit s ~ lll a li . "hallo \\'. ra~ l - nll\Ving. ekar st rea illS in the 1()J'ests eovel'l li g the hill" ;IJld mounlains. but is a ralher PO()l' swimme r. It o rt en kaves the wa!..:r IlH' longer periods of time to frequent shaded moist local iOIl ~ or to blll)' ihclr ill the laye r or lear litl eI'. It keds primarily nn plants alld fruits. but ~ nail s and worms are also con;"lI mcd. Acti vi ty IS limil ed to the coo ler hOllrs or Ihe mornings and cve-
II
nlngs .
1-3 large. el ongate eggs. CL or hatehlin!,!s 5 1(i .) 111111 .
Distrihu ti o n Blinna. Thailand , Peninsu lar Ma lays ia , S in gapore . . lImatra illc\. 1'1Iiali l3angka . l3alu Isla nds. and Plii all l'\atlilla Besar, Borneo. M t. Kinab ll lu : wit houl precise loca lity.
f mm top Co bas~: I h 'lise/III'.\ (adillt. CL _0 CII1) fig.379 Dorsal \·i.:w. Fig.3!!O VClIlral vic\\'.
\}liIlO.\'1I
39X
1 ri()llydlldac
Trionychidae Thi s f;ll11il y orcryp1 odirans includes abollt 30 sp.:ci .:s. Their distribution range ext.:nds fi'\lIll (lrlh Amaica, through Africa and Asia east to ew Uuin.:". They arc eharac teri/ed by 'o ft shells o f ro llndi ~ h to elonga te-oval shapes. The ca rapace and plastron are no1 cnve n.:d by horny seutes but a lea the ry skin. Wi1 h marginal bOlle:- bei ng absent. 1he hah es o r the s hell s are joined by li ga lllcn1 '. The s u r t~le e o f the plastra l bo nes is d isti nctly structu red, lI ~ lI a l l y ka ving their depn;ssi olls (call os i1i .:s ) well visible th rou gh the skin . Th.: elonga te h e ad ~ o r ~on-:- h e ll e d turtl es tcrminate in 1run k- li ke noses or variou s kngths. Their horny heab arc Cllven:d wi th tl e:-hy lips. The indi vidual genera are mainl y di s1ingu ished by anatorni caltra its. such as the ahsenu; ofpn;s.:ncl' of bony plates, their numb.:rs and si tu a ti o n ~ . Roth the fro nt and hind fect arc r1l1'll ish.:d wi1h 1I1I\;e free el aws eac h. So l1- 'helled turt les ha\'e a reput atio n ro r bit ing more or less r.:adily. They feed mai nl ) on other animals and reprod\lcc by lay ing eg.g.s in larger num hers. On Borneo, 1his family is re pn:s 'nled by 3 species in J gc ncra wi th onl y one ge nus and nnc s p ccic~ na ti ve to ML Kinabalu .
Key to the Bornean genera of Trion ychidac (after M " 1111 \ &
U J«()S~M \ N
1(97)
I.ong nasa l trunk present (A ) ...... .. ......... .. .. .. .. .. . 2 • I.ong n a~a l trunk absent ( Il ) ............ P('/(Jche/I'.I' 2 Carapace wit h vertebral stripes (e), -l ca llosit ies f)°K{/l/ia • Carapace wi thout ve rtebral ~ tripc s, :I ca llositi es .... .. .. .. ... ..... .. .. .. ... .... ......... .......................... "I//I) 'da
Ge nu s DOX{llli{l
G RAY,
1844
i~ lli st ri bUl ed from \.1 ya nm ar a nd lli c Mal ay Peninsula to Java. BOrllen and the Phil ippines. Relativ el y small s(ln-~ h c ll ed 1U11 1es ((,I. up to 35 cm) with an oval carapace and a dark. often indi sti nct. vertebral stri pe. I.argc clonga te head \\ ilh long na~ al trunk . T he genu s is monotypica l.
f) ogall io
Dogllllia slIbphlll{l
(GI.OFFROY,
I riolln \' lIh, " OIlIlS (; Hl I IKtI' . I X()9
1X09)
11. pi 5, fi g. 2
T~l)l'
IUI", ..di,y: 1I 1l kll rm 11
()og {/I//filtll/{/
1) 1
Ro" "
t'l1 5'
326:
\1", ""
l )c/ochdl'S /I,hn1111( flol 1',.,011\.\ hihroJltl 0 \\ I I 'JKil. 1'1. -1 1
\I ,
_
I"X I : 10.
1\.1" " .. , ~ I ')l) Ia ' 2..\ 6 I ~:'ll). J \t "O I" .. 1 f\
Fi!-:. JR I
P O l'tl'~ ,t \ ) t' / )".<'; (111/(/
.1/lhp/(//JlI
_
"":'___
...J g
1'1'\)111 PC llIIl SIIi ;! r
rvl a la y~ia .
399
I rlnllydllll:ll'
1) ~~rr ipti4)l1
(' L li p to _') elll, l'arap;lee \'er~ 11 <1t.o\'
age .
3-7 perreclly rO llnd eggs (2_3 1 nlill III diamcter).
Ecn lng. /F. tllUl f)g ~' I>. S/lhlllll/Ill ~hows a preference for clean , COllI. fa s t- flow ing slre;lIl1S III l(lIvland and hill y country. II climbs vcry \\cll and onen see ks ~ ilellcr all 1011 ); rocks oiliside t he water. O. -,"/lIIIJ/mlll I ~ crepllscular to 1l0ctllrnal, spend ing 1l1Osi oflh.: day in hidin g. II reeds pr.:dol11 inani ly 0 11 ollH:r animals. L>i ~ t rihllti oll
Myan ma r in c!. 1L:rgui Island s. Thailal1d . l't.:ninsular Mala y~ia , S in gapo re. Borllell, SlIllialra in c !. l'ul;1lI ,' ingh:p and Pul au Na luna 13L:~ ar, Ja va. Philippin!.: I s land ~ . I\lt. h:ill ahalll : Sayap.
-too
Difli:n:nl l.:OI(Htr pallCIll i"l"Ilm PCl1l1lslIlnl
-1a lays 1:1
I
j"
Ihe ca rapace III
nogili/ill
\'lIh/II
(; lossary
Glossary ahioHc: inanimatc: ph y~ l cal and chemica l as pcc ts o f the cnvironmc nt o r an or~alllsm (con tra ry In hioti c) adapl i\ e radial ion: the c\"()lutlOnary tk\'elopl11 cll t Ill' nc\\ spccics fi'lllll an cxisting spcc ics as a rcs ult or adaptatIOns to \ariolls cClllogiea l ni c h c~ wilhin a eOl11mon geographica l di ~ l n hut ion ran l,tc adult: scxua ll : matu rc. grim n-up allopatric: li\ ing in dilfcrcllt gengra ph ica l l\:gion~ (collt rar) to s~ mpatri c} allIphigonia n 'tardata: I : delaycd fl:nili7atilln. 2: thc ~lOragc of \ iab lc :-.pcrlll by a fcmale amplcxus: the matillg pu:-.i lioll of L1nura n :l mplllhian~. th c em hracill g or the I "':malc'~ hody Il\ Ihe male'" fo rclimh ~ annuli: ri llg" or ring-lih: mnr"ln~s apical : at thc tip or a :-.Iructurc lHluatic : li\ ing in thc w:ller arboreal : living in :l lim:slcd l'l1\ IHlnmcn t arlm.-icul: Ii, illg 0 11 trecs aututOlIl~': tIll: ca pab ility to hn.:a" o lT the tail. or part" nf it. at predesigna tcd places as a dckncc Il1 cchal1 l"m hcnthic COJlHlIUllil~' : plan" :IIHI an im a l ~ \\ hic h ill habit thc bot to m of aquatic eco~y~tenb hiodiversity (hiological di\er"i t) : a ll aspect~ tIl' thc dl\ cr"e animatcd wllrid (c.g. "peCles di \er"it). geneti c dl vc rs ll y, di\Cr"l ) o f ecosystenb) canlhus 1"O~II'ali~: ,kln-covercd, bllny ridge Ixt\\een cyc and nostri l carnivure: mcat-con'llll1ing "pecIc" caudal: referring. to the tail con~pl'l'ilic: belollgi ng to the ' :lIl1 e ' 1)C": le ~ t'unn..-gelll'l' : C\O llltl tlnar) pmu:s" th rough whi 'h unn:latcd . gcographi ca ll y o rt cn \\ id cly ~L' pa rall.!d ~IJI.!cie, de\cl\'p "il1li lar ,Irllcllln:~ (habi tus, bch;1\ iouI') due III ~illli l ar e..:o logical cha llengcs lTt'plIst' ular : a..:ri\ e al 1\\ ili ght hOllrs (da\\ nand or dll"") di slal : mure distant rromt hc centrc ol' lhc b()d~ or a de linl.!d ponll or rekrcncL' than \Ithcr pmb or the bod:- or an organ
diuntal : ac tl\C dunng thc da) dorsal: referring tll thc had dorso lat{' ral : rekrring to the "Ide of the b ac~ duno\'{'nlral: in a ve rti cal direction cndl'lIIir: hClIlg e,clu"ive ((l a certain gl.!ogra phl cal reg io n: II' a species ha" de\l.!lllped \\'lI hin thai rcgllll1 it is called autnchlh()nou~ l' ndolruphir: Il:cdlllg on Int crnal resourl'C~: here : l a dp \lle~ \\ hid! de\dop \\ ithoul a free- ranging ~ ta !.!e and arc therefill'e Indcpendent from an cx tcrnal food :-.upply : they de\elop dm:ctly rroll1 thc yo l" 1Il:1~~ . fUl>so rial : li\ ing ~uh t erra n ean l ) (linderground) !-(t'nulypc: all genetic trall~ \\' Ili ch tletCrIlllne lite ,trllcturl.! and I'unction of an organi~llI ~nl\ ill : can") In !.! egg". belllg " pregna nt" ha hilat : Ihc nat ural cm iron mcnt an anlll1al Ii\' e~ In hahitus: gcncr,iI appcarance, bud) build hClIlipcllis: pall'Cll matilig llrgan o r mail: rept ile ~ in c. \l'd.: Incertal' ~C dI S, prelllllillary taxllnOl11 lC ass ign mcnt illcuhation: the dc\elupment ("hruodlllg") of'eggs infra-: (pn.:lix) !tmer inguinal : being ~ itu a t ed in ur Ilcar tlte groin i~olatilJn Illcchanis l1I : ;111) ge ncllca ll y induced d if~ I"': rence bClwecn ~pecie, \\ hi ..:h pre\enl~ thc~e frolll matlllg ullda natural t'lrl"lllll ~ t a ll ":C" jU\ cnill' : nol yl.!l 'c\llall) lIlatlll'l.! , nul rull) gnl\\ II lat enll : rderrillg to the s ilk mandihll' : the lower .la\\ Illa~illa : Ihe upper .1a\\· median: rCf'erring ((l tlte Illiddle 01 ccn tlc llIil'ruhllhilal : a portion url lic manuhahitat \\ hleh olTers the CI1\ lronmental cir..:u nb t:lIlce" an nrg;lIl1 "m rcqlllres ror il s ~u n ' l val lIlilll e~ i~ : the adaplallon of'cnloralion , hody "hape, and or hella\ iour to th e 1Illllledi all' CII \'lrO I1Illcnt hy I1llllli..:klllg re~pcdl\e ,tru..:lure" or th c lat ter 111 ordn to cscape thc at lentlPIl of' pntc ntl al prellatllrs (c .g. the Illlll:ltlon 01 plan t "trlleture,) mnnut) pic: cOllt aln ing nn ly d ne IInlt pn superior ") ~kl1latic rail"
-101
t j I ()~sary
or
nalural ~ clcel i4111 : .11": L'lHllrihllllOll \'arlOlIS gC J10I yp..:~ In lh..: n ll, prin1,' or the sub~cq ll c nt gcneratl on: \1\"er tlll1C lhi, re~lI lt ~ ill t: hallg..::- W the ge ncllc cO l1lpn ~ilio l1 01' popll lali\ln s lIielll' partiliuning: the l,'nden,'y or cn-cx istin!.: spccics l(l occu py dilkrel1t ..:cological ni t:hcs wilhill 111(.: ir t:lllllllllll1 ly inhab ited habitat nidll': Ihe ~ pet:llit: hahltal within all CCO,.;yslClilt ypicall y inhahll.:d hy ;1 'pct:i.:, lIurlllrllal : actl\'e at I1 lght nmni\'nre : a ~pec i e, thai t:lllbll lll':S hOlh plant and all I11 1<1 I lIl:ttcn:1I 0\ ipusil inn : the lYi ng or cgg~ papilla: a ~ lIlall . fl eshy proj.:elloll paravcrll'ilnll : bein 1,' s iluated ne.\ltolhe \ c l1chral <':(11 L11ll 11
phenot ype : the uh violl>. lrail s oranllrg;lIlislll whi ch are Ihe re~l1lt orlhe inter:Ictio ll bct\vccn gc notype and envirol1mcnt phylngl'nc ~ is: lh.: l:\ollitillilary history of a la.'(o1I011 11r gro up pupulaliun : gl'l) up printli\. i du ; tI ~ \)I'a spet:ie:-lI'hi ch inha hll:t t:crla in gCllgrapllll;al regioll at a (cr1:11 11 poinl or IlIlle I'l'pruducti\'l' iso latiulI : spatl ;d aml lelll poral sepaIhc sa m,' spcc ies: :111 mliol1 \)f' popul ation, lI11pllrtant prccondilion lor the dcvl:lopll1cnt of' lIew S pl.'C "':~ ITMHIITe : eve rythin g an orga ni ~m rcqui rcs lo r its e~ iSlenc e (c , ~ . ruud, ~ p a\\'ning s ilc ~. etc.) ~exllal dimnrphis m : thc dilTercnccs in hnd y ~ h 3pC andior coloration bc!\\eell Illales and tClllilles or the ,a llle ~ p e!;ie~ sic!: re:ll ly ,,,! (rl'al ly ~ \l \\'rillcn) sil -and-wail : rUra),!l ll!;! ,lrakgy in whit:h a prcdalor rCII1;I1IIS illlm ohi Ie until a prcy COIllCS wit hin st nk ing I'angl' speciali()n: till: prO\;es>. of' Ihe developmcn t of spce les ~tratificalion : th l.: ti ..:rill!;! or ;,torey- lik e, \,I.:rtit:il ily org
or
or
or
syntopic: occurrill),! togeth..:r. illh ahllin g the ,alll,' bioillpe taxon (pi. til.\ il) : any sC lenti lit:all y n;lIlI..:d ca tegmy or planb 411' allill ia ls (e .g, s p eei..:~, ge nus. 1~11ll il y) taxol1ol1l~' : the ~cie llt:e or clii>.sii'v ill g o rganis 111 s terminal : "1;11' rrulll the '..:n t1'\; " , bclong ing to the 1110,1 dista nt end ty pe loealit~· : Inells lypicu , : the loca lit y at which the ho llllype or a ,peries has bcc n cu llct:lCd type spl'rilllen: 1110"t holotype : rcli:rellc": ~ p ec illl en upnn \\·hlch
CO ll ve rsion table e111 (ccntimctrc) ~ o.,,() in . (in ch) In (Illeire 1 = 3.2XOX n. (f'ect) g (g ram) (U)4 0/.. (ounces) I (litre) - O.2 h4 gill. (I\lllclic;lIl gal lon>. ) I ''C (degree Cebill ~1 2.63 " F (deg rees Failrenheit)
Bibliograph y
Bibliog raph y Literature used in the in troductory chJPICr~ and furth er readings are marked by an astcri~k (*). AlII, 1' . (1')27): lur Sysl"'l1lalik lkr asialisch..:n ;\11..:n ,h:r Frnschgauling 1(/{({( ()I,IIIJrIl.I SII/. Her. 1I'!llIrr. FrcUlilk Hedin : 1.' -17. (1')'>1) : Li,ll' dl:r rl'7cnkn i\l1lphlbicn lind Rcpldll;n Anura III PoIYl'cd:nidac. 1);t~Ti.:rr.:ich. Hcrllll. 55: ~I' I + -1 77 PI'. All \1,\ , A .t' . & W.( '. Il H()\1 11 9X2): Rcpmdll~II\C biology of '()I1l~ ~ pCCI-:S or 1'11I/alllll,I (Rhacophoriua.:) and olher philippinc anurans. Ph ilipl' . .I . Hint.. I I: 2()\-
22() ;\ 11"1,:, R. (19 70): i\ ~cy Il) Ihl' ladpl)il's of Iill' cllI1linenlal Unill's Stalcs ,lIld ( ·:lIlada. I krl'clnlogica, .i"llIboll Clly, 26: IXO-20 7. ;\ 111(" It 8;. (j . F. j""I\" I"'oj (19W»): (iuilds nfallu ranl;lIvae: n:ialinnshil's among ,kl'cloJllllcnlal Illodcs, 111mph()lo g i e~. ;tlld habilals. I krpctolngical vlonogr. J : X 1-1 II') /\ 1\111K'''J'oj. 1..( i. ( 1921 I' SOIlIC r":JlI iI.:, ;tnd balral'hians frnlll (elllr,iI-H..)mco. \kud . 7001. I'"hh .. Knsllana, (>1 (7): 1 19- 12. . ;\ 111\ \ , vI .I(: I' 1\ llI 'I, (1'1')'»: '\cuc L-rk<.'nl1ll1l'~": 7U 1', ,Ihull CI/lm.I (T'IX/" '/l IJ /I/(I/ ·/'/III,I) rl<:. ;-..J..:w Engl. Zool. C lub. l1o, tlln. 7: X7-XlJ. - ( Ill.IX): Nnks UII " ,V"cI(/l'lIn '/I "", Proe. Hiul. So .... W;lshlllglllll, 5 1' I') I- I'!'\ . I.hlOlIIl R. r. '-" (i ,K. I\"1 1I1 (1'116): Ncw an'p lllb,an, and a IlCW replile fll lll1 S: II'a w,l~ Pro,', N..:\\ Engl.l(1ul. ( 'Iub, 1l11'IllIl. (.: 11)-22.
11 11("'1 ,1(. I). & T. B,\llKI I( ( 11)9-1): Th.: KI\)()d -l'ylhon and oll,,:r ' lIh'pce h':s of Ih..: SlwrI-Talied 1')II1\\n (/" '11",/1 c llrll/l' ). Thl' Vivariulll. () (.1): 30-.1 I , ') 11 \1((1111. 1', ( IX9 :'i~) : I h<: cr5c): Nnl..:, 011 Ih e sna~..:s of 13 11n1":o and adjaccnl islands. Pan II. Sar,l\Ia~ 'loll' 1 1,.o~ , 1111. I: 7X-X5 . Ihl ( 1111 0)1{. i).r,,1. 119:'iX): Snllle nole, 011 Ihe ~nakc, of i\~aliall . M,dacc,1. Mala y. Nat. .I ., Kuala I.umpur. 12: 1()3 - 1 I I. - HI U( " , .1. 11. (" R.S. HI \~(" (1 1)')0)" I)I\l'r'" ' and disInbu1tllll pattcrns ,n Ih .., flora of \1(lUIH "In:lhaili . 111 : Ih ", P., "- . K \I M \1\ 8:. R. (ill " I'" (his )' rhc plant dl\<:r~l1y ()f I\laksl<1 Dordrl'dH (Klu\\cl AcademiC Publ.i· 1-I 7- IW. - Ill \\1 \". R.S., .1 . \1. H I \1\''', (' \ lIH'i1i & 1'. W OOl)' ( I'}X5): Drollf!hl and fi)rc~( fire s in Sahah III 19K, Snbah Soc .I.. Il)X5: 10-.'(). *1\1·( ( \HI, () ( I ')()-I): Walldenllf!' III Ihl' grl'al fi)r~ ' IS of B Ol11~o Lnndoll 1311(1 ,M \1\. I<-A.M. ( 1956): 'I1,C :UlallllllY II I /In ' 'I,I//.I IIIl/SIIIII.I . I. I'roc. "Oil 1\ 'd Akad \-Vet , S~r. C. SI}: 2('.' -::!71. n'lon , P. Y. (19 72 ): l lnd~,crihcd and lil1k-kIlO\VII tadpok' li'llIll \Vc~t - Malilysl;1. I krpl'loluglL"a , .lohnson ( 'ity. 2!-i( <1): :13X-3-1( •. (1'17S )' Th~ amphihlan lOll 111.1 III' Pc II I11 , "1,11 I\I ; da Y~la Kual a I (II11Jlur (I rtlp,.::!1 Pres, ). 13() pp HI HK\, P.Y & J. III ' 1l1(1( ... ",' (1<)(, I) . I "I,I";',,,u//I/I'" /li}.!,rlJIJv. a ne\\ JlciohalHf frog lilllllilie 1\1 :liay PCIIIII '> llla. Wlih I ~en'ar~' Oil Ih .., (il'lIll' / ,"/Jllllml<'/IIIIIII In SOlllhca, 1 -" , ia. Copcla. Washln)Olon. (<1) (>-II -h-l X 1111 1"'1 1(, 1'. (I X.'iX 59): Repldll:1l ell \ " 'l·h..'11 lan"'ill~alVang ("cSI~lI'l vall lh)rn ~{)) . Nat. Ilidsc hr. cd . Ind .. l3ala\ ia, II>: I XX-I X9. (I X5')): R<.:ptdli.'n vall S111iallf!. \el1amcld do,lI A. If. TIII·I''',. Nat. Tijdschr. N..:d Indic. I-I . Jl:lrl 11). 20: 200:'()2. (1~6()a) :
R..:pldl i.'11 \all Agal1l '\al. Tildschr.. cd Ind., Bala\ la, 20: 32S - .~::!{) . - ( I XhOh,: (her dc replllli.'n -I;llIlloi \ all SUl1lalra. Nat. TlJd'l'lir. ·l'd . Ind., BataVia. 21 : 2X-I-29X. 13", n(,II(, O . (I X92) . K:l1;llog d" I lalra c h, ~r-Sa1l111l111ng im \Iii Ise ll 11 I d..:, S (: lle~ 'lIb<.:rg.I ,,· Ii,,'1l naturfilr'chl'ndcn ..:hr. (ic-scllsc haft in I rank lllrt 1\1 . I rall~III'1 M
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(I ~i; ( I S'),I.I/: I )n Ihl' hl"lp<.:IOI()l,! ll"o ti 1;1"lIa (II· l'al :lI\an and 1I,llah;I<': . -\III1. \ I:lg '!al I list. , L,)l1d'lII. h. \l'1 1-1: 1-90. (I ~()-Ihl ' ,\ 11'1 Il l" rl'plill" <1l1d b;lIl :lclll,lI1\ ('()Ilc:l·k'd b~ I)r 1'.. I\'I"dig ll,11I1 nil " <': 1"1: 11111 ("'Il(lra). l\klll;1\I'l'II,bll(k ,\nll . 1\ 1" , . ,'1\ , Illr ,,,II (,l'll,na: (,13 -(,I X II ~N k I: Third rl'p,l!1 on :1I1""IOlll lll Ihl' h"Ir:ll'IlI:l1I ("(II k CIIOIi III Ihl.' 1\, "111:11 -11 1'1(1) I\'IU s~ lIl11 I'I"I\{"' /.,,,,1. So,· 1.'"ldOIl (>-IIl-I,·II. ( I S'!4d I: SI.'<:"lId I ~ poll "" addll ",", ,0 IIIl' II / an i n,ller111111 111 Ih,· ,,1111":11111\1(1) 'l. l usI"UIll. 1',,,, 1,,"1. "()t". lllllllllll" 722 7."\(,. II S')-II.' ): { ·,II:tI" J:'. " l" I,IIi1,· , ":tk", illlhl" Ilrllish .\111,(,11111 ( '- ,IIIII ;IIII I\\I 'ry) \01 II ( " 1lt;1I1111lJ:'. Ih,' ,'(IIlt lt",oII "I Ih,' l llluhrid.ll' ;\"d\ pha,'. I ,,"dol1. (Ta) lor & I·ralll.'i, ): \1 I 3S2 Pl'., 1'1. 1-211 I I X(15a) . I ll"l'ripll'Hl "I -I 111.'\1 l>all:l("III;II" ti,,,·,,,",'rl.'d ' \1111 M.lg. Nat. Il i, 1.. b~ lr ( 'harks I los,' 111 Il()"ll"" I Olilioll , 11. S,'I 1(, 1(," - 17 1. I I X' 6h) 1lL"l'rlplll l ll " I" 1\\0 '"' \\ "Iak", 01 ' 11,,: g , ' l1l1 ' ( '/lI;lIra;\1111 . Mag. dl1all IIllIll 1\11. Klila Il,tllI, \Jonh Hunll'" :11 . Il i, 1.. I.olldoll. 7 Sn -I: 151 --153 . 11900.1). lk, t:npllol1 (,llll'\\ rl'p,ile, ,11111 hal lOldll;tlls Ihllll Hnrnl'o. I'ro, 1 1101. ~oc. I .(lildoll I S2- 1H7. (1900hl: D":sl.'l1 (1 110 11 r~ pllil', Irol11l'.:t:lI,. 'vlalay 1'l"IlIIl, ula . ,\1111 . \lag. '\Iat. I I lSI. , 1.(J lllIon , 7. S~I". ) :
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,1I111UI,li cd dlCl""I,,1 alld "I'y II Ith lIallllal hl\tory lIull" l' ran"ful1 "Ill \lall1 (I ' dllion (1IIm,mal. I,,','olld Ull changcd clh tinn 19'171 1-2(,1. (IINX): Rt:lil'''c nplIllll (If 1i/lI/l'I!,II// '1I1 1111/1011 / " 111111, 194\) (Sc' rpclltt: ~, ( rotalilla<:). \\ IIh a d"cu '~ I"n ,,( Ih n:iallllllsllll'" ( (alllatirY,III , 22 12 ) 7,-)( 7 I) \1 Ill. 1'.. (i . Vex " I L\: () I' \111\1 I , I II)I) X) 11111"//('. \ /11(/ "I ' /(fIIl/1l (( Ii 8:. Dill) , 196(1) C" l'lpl'nt.:~. (1,llIhlldaL), an add,tioll Itl tl1<' , lla"I' I;llilla of VIC lila Ill . \\ It Ii a lI\t "llh,' ' pc't:IC ~ III tht: gcnu , , 111/1""1'\ 111<1 and a 1l0le 1)11 lIS 1 ~ lll' ' P '\'1(;' J '1;11\\:111 \Ius" - I (2)' X<-1)2. -t-D \\ls. D.D. I 1<)(,::' ): lallllllais "f thl' 11,,\ 1,IIld r,lIl11<'rl'sl Ill' 1'<,\1111 H"I1lt:II Hilil al. 1\-111.,. :-'1Il ~"PIlI ':: , '\ I 1- 129. D \1 l'lI'I. (0 , \\ II. ( I I)X4) ~o"I-lla 1-:g lll g tIl'l'la y III 11l>I"IIL,1I1 Irngs. Sar:mak t\ h h . L Kue hing, \3( ) 4}: 177- 171'. DI t\/ I K. \\ (1')<)4:1) ' T , ce hole IIICt:dlll g III Ihe load "d"I ,/tI'lI/" "r<'I '/I" " (PI I I I~~. I Xfl 7 ) Alllphlbl:I - RI'plilia, ( cldcn I.~ ' 224 - 2~(, (1'J')4h): /111 HIIllog le 'Oil /'''/''1''''-'''''' h,.."1'//"'" (1'1 II R,. 11'(,7 ). hl'rpctnl;lllna. \\ ('ill,tad!. 1(119.1)' 17 20, (11)96) : Replllielllll1d \111plllhlell dCI S:lIl1l1l)(>llg 11,lhlll'el. Sarawaf... I3lll"llcll. Sauna. Lkrlill. 1)(( I): ~ 5-1 2. 1), ' 1' \ \, t\ 1.1{ , 11912). Sill tilli s t'ollet'II(1I1' de rcptdc~ el dc ha lr,KJ-:II" pru\t:l1anl d ' 1",\ rcf II pel \1;,1 ,11, Hilil 'vIII ', I (isl. ",lI. Pari" I X. I')X-211<; I)RI ~ ' , . .I. III)~G :I ): I III ~ ' o( (he gcnll, 1.,,/,1,,/1/'11 1 {, .. IIII (Pdoballdac' ), AlIll'hlbia -Rq,tdla . L,'I,kll. --I : ~1) - IIl:' , 119l'i 3h): '-;1>111<: I1l' \\ Irng~ hl>m Sa,:l\I:I" , ,\mphlhla Re pti lia. I -:ldC:lI . 4 : 10,-11 \ II ')t-i 7) BI1I11<\Ul tr~\:II"\l&' III thc' gt:llll\ 1'/11/111/11/\ (I{h:l"llphnl1lbe). ,\ml'hihl:l-Replilia. Il'idcl1. K 11)-17 I), 1\1"S, A (I '»)(II): ' otc" sur la '),I" II1:lt Itlu,' c:t la rq';)rtlllllil dcs . \mpllibil'l1s Al1tlun:, dc ( hllll' ct dl's rt:g ill l1s .1101 'Il1alll<.;' IV. ( 1;"~ llie"llIlIl gC:Ilt:1"1"1.' c:t ",h 'CIlCrlquc: dc:~ PcltlbatKia,' kgoplir:- lilac Bull ~Il" L IIl1l 1 YOI1 . -11) 4(,<)-4X2. ( I I)~ I) , ( " Ie dc, gClln:s Ci \\1I1'-ge l1le ' 110111111,111\ ti c Ranollk-a (Alllplllhll'lb .\1l11Ur,'~ ) du 1I111mk. ;)IlT IdcnIlr,e;i1I011 dc Ieili s e ' pcle '- I ~ pt' S CllII Sl'qllt:ll ce , 1l11l11<:Ill"iaturak;, \1 111111 10111 lIal . 111 ' 1. ' lIppl I ~' 22 5-2l-.4 . (II):-; ~ I : II 'I ' /()I ' /WI//( ' I-II / I ""~, IX4 \, a 'Cllllll ')'111111)111 III «/( ',,/,/t'-"I<' D \\ls. 1')35 mulpl1lda,' ) J IlL-I'I' , II> 171 -1 74. ( I I)S ) I" Ntlle I'r,'llIlIlilall'" ' III Ie l!I' IIII' / I'I ,I"/III'1I ( ) I III ",, 11)1'11 (,\l11l'h'("l' Il ', .\11111'1") . .1\ t:t' dl.J),(IIIl\t: ,('ulle 1"I'C(l' 1I1l1"elk "" \',cllI,1I1l . \l Yit:" (';)11 '. 214) 14!- I.Q. (II)X(,) ' t\1,well ,III";) t;)\1111111111";) h; JIr~Il' It"III1-: IL' a I I ) \1 ) ICs, ('.lri ~. 51 1-2 ). 7_1).,. III)') ;! ,: ' ot e~ ' III 1:1 t'1: " , IIIL' all'''' dt: , R;lIll".I': I '\ l11phibit:lb ,\11\1111" ', ) 111111 mCII ' S" t' 111111 l~ lll1 , 111 (iO ): \0'\- 1'i 2. 1) , " ,, 1' ... \ 8:.-\ ()II IIK (I'I'IX) .\ 11 " \\ ' 1' I' t'Il" or
Illhliography
I ('I 'luhm, 1//11111 (11hl ,\I/I,h,)/"(II li'ClIll lIl'rlhcl'll \,ie'lnal11, ~I Ilh a Il'l IC\\ o llh 'la\ClI1<>l11)' ... 1Ihe: !;!.cnll~ 1."I'lohm, hll/lIl 11\;lah'llIdac. tl. lc'!!(ljlhl)'lllac'I, I)II111CldJa, Pan" ,I( I) :
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120(10) S)IClll.lIll" pi ""1<'11'111"'" 11111/111(11,,1'/\ ( ,K\I'I '1111111.. 1. 1l't2(») ('\l11pll1ll,1. \11111 :1. Railldac' l alld rl'iaL.1ll' r\ai 111,1 .. SPl'C I'IIhl. '0 2 1' III I 372 PI' 1)11111 \ 1\ '%, \\ I , c\. l 'I III I II II'IX(') : III"hl!!) nl" \IIII'IIII1I,ll" c' l~ ,,,II. I \1<:( il:111 I Iolll~"o\.. ( '''1111' I. (.7(1I'P r h l ~1I1<1I .:\.11." , ,Ii.: (i 1i1l1~11I\ (I X.1(,) : 1' 'l1clnlngll' gCII':r;lk (11 11 IoslCllrc' f\;o[ure'lk l'(llllpt.:Ll' dc, Ikpldc,. I'ans, Vol , 111.1\ :'IXpp IIX IllI ' l ' rI'0Iol"~ le' )!l'nl'1'ak llll Ili , l oilc Nalllrclk l'lIlIlp,,"lc' dc, Rcplile, . I'all', \,ill \', 1'11.1 I X55 PI' DI \llll li,A\I( .. (i HIIIKI,I\ & ,\ 11 ,\ [)I \ II RII (IX'i-la): I II1CI"loglc gCII,'I',de' nll I Io,(I,irl' ".ilurdle clllllpick (I.:, RCpldl'S. i'ans, \inl. "" II ). \ ~ I I 7XO PI' ( I X~-Ih) ' 1', 1p':I<,lo,l.:'" Oil II ,,[olre' 'Jalurdle c(lI11plcIC dc, Ih:pldc:, 1',IrIS, Vol "" (2). \il 7X-I- 1) 'ill. DI I"'. I· R (1'12 .11 , On " collc:CIIOIi 01 rcpllics I'm lll Sara\\:lI. . J. tl.la l:oy. Iii. R"y,d 1\ "at. '')O( I 1--1. I III I\ 1I \ ~I I, T .'\: I' S". 01 (1 'I II')X"). 1.11<: 011;0 lIl1p";alllluul1lain , 2. 'I he rcpllle, 01 \11. K'l1ab,,11I alurl' \1aI:oY,\I;oI1,I, 'I( ; I' 11>-27, I III I 11\'" A,('..I , IIX(,'i1 Rcchl'rchcs ,"I 1:1 f:IIII1C l''l),' (lI' Il'glqUl' dc Iloln':o, f\cd I'"dsl'hr, I llc'l'k " l.e' lde'n, 2: 21)11-211-1 I III ",1)1\ , S.B, ( 1')1) 11. I hI! celllll,lIvhlllo,l.:), Horncan Lrl'C rrn::s ICllndy Rhacophllmla.:) , /.001. J I .inn . Sill', I I) II -I ), 33 7-.15 7 ( I ')I)~) ' ( '('"I "hlJl "lid IIC" I·bllJldlll!,!. heh:" lor of a IInllll:,1I1 I'ml!. Rill/II "" '1111 ( 'OPl'''I, Wa,hlngloll. i-l); 11 2 3· 112 7 I \ 111('''1\, S.II 8:. I) I~ I '« I(I< , \'1 ( I I)l)J), S\"Il'lIlallC~ "f'Solllht:",1 ASIan R,JIlids ; tl.lultlpk ongllb "I' 1(lIC.:k,,"C'~ 111 Lhc suhgl'llu" I.IIIIIIIJ'" '' h ',1 II 'II / IN'" 1( 1. I krpl'lologica , -'"hll"llI ( '''Y, -1'111) ' 22-.1 I hlJ 1(''''1. s.n, 8:. R I,. I " ,I J( 119(2): I'I,C c'ulllparali,,; c'..:n lngy nl' 1nl.:.:d ;,,111 \olCcl .,' B"l'Ile'an Ii·0,l.:' , .I . I krp . 26(-1) ' ,IX2--I')(l hlll" II'I, S.Il, IU . I", 01 1< ,Ii.: I) , I, ,, I'-J)\I( (20011) ' Moil:<:ulal 'y,lclllall,' ;oml bHlgl:IIl;.I: lphy (If Ihe I':lngt:d I' rogs 01 SUlIlhca~1 ASia . \.Iolccular Phylngt:llclll" and F\". IUilon. 1<111 I: 111-1-1 2 1-\11 S,H, 8:. tl.1,,\ ,R, Kill III 11'1(10)' Thc 1I1.::r~cll(II" "f oehavi"ral and Illllrplollloglc,,1 challg\! In Ihc c\'olulion III a 11I)\'cl 1,\('OIlIl'(or Iypc: " Flying" frng~ . h'
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NCI/P) I 10011 1/" 1111111 r,'fI' 11/11111\) IIlld Illllipy lhllll (/ "'111111/ ( 111 '1//\) In Sumalra 111111 K:d'll,alll;oll . Imlolll'sil'n. ,\I"chlllllbclll'i'1/11l1l I' orsl'illingsprnjcki illl :\IIIl",g. dL" Hlllldl"Il)JIIISIlTIUIll~ fOr LJIll\\ cit , ,' allll ,,'Ioui/unu Rc'aklor~icherheit. (I Il\cliil1;;IIII,cilll 1)/ pp , h i 'I. I ' (14'17 ). Hlo()(1 I'yllo"" .. , 1'\'Ihllll ,'/lI'/i,, : ~lIb~pt:<:1<.:,. ':;11\: alld hret:dlll!! Lilt. 'icrpcllt., 171(,) 116-1.::>-1 . hi, I .. L'. '1'1 8:. \ . 111 "'1, : 119l!1) Kl.!v 10 CIoIIll"c ;\m plllhi:1. ( ·llOll g'!"I j.!. I dillon' 01' ~c' ICIlI'L'~ and I'cchIII,!UC', 1-1\ I 1-2 I 1-3(,-1 lin ('hllll'sel l 'ls( IIIIL J.(i . IIXX:'I. flbel ':llIc: K"lk\..lhlll 1'011 /\ml'hlhicn lind Ih:pllioclI al" "lidoSI - lhII11CO ,\reh f\allllg.::,ch .. Ikrllll' -11 · 72, h II I'-' ,I I(, L..I.11 1('6). t\t:lld '1a",lit'alJ(lIl dLT Rcpltlle'n nach IhlClI lIaliirlll'Iol'1I Vl'I'\\anul<;e' h;lftc' n N.:b'l c' IIIC'r \'L' rwalldtschalb- I al~1 lind c'lIlelll Vcr/clt:lllll~Sc' lkr I{Cpllliclb'1I1l1ll11"')'! dcs I.. t... lo>ologisl'ht:1l \.lw,c'lIm, III W'CII \Vlc'lIlJ. (,. Ilul'i)llcr), \iii I 6() PI', II X-I .'\): S~ .'\c'Ill:t Rcpllillllll !';N', I , ;\IIlbl)gln~sac \"lIclllholl;oc . \\',"" (Hrauniilll'r & Se' ldel), 11)6 I'p. 1·1< \11\1, J B .. \\' FI(I \ , II KI It" 'II 'll ~ (I::. I'v1. \ ,111\/11 (I 'NIl! ' tl.ltbSl'~ and Il\cmoils ofl\1i. Io:llIahalu Kola KIII:thalli I Sabah 1'~ lrb I'libi. 12) h « lSi . n ,R (U l ) II (IX)): Amphlhlnn Spt:I'ICS\11'1 h.: \\llriU. ,\ laxonOllllC and ,l.:Cog l~ lplllc' al rt'li:rl'lIct: , 1.•1\\ renec IAlkll I'rt:s, (\i. The th~OClaLilln of Sv,IClllalic ( '"Ilet:Illlll'). 7.12 pp. - f ~()(I()l : /\mph,h,,", ~pet' ICs ,)I' lhe' world : All olllillc Icl~ c'IC:III'C, \, 2,20 II . Sc'plc'lllix:r 2ll()O) I I " \1),1, I L (1<)(,-1) : t\ 'allall (;(,Ikcli,," "I' snak.:~ of lhl' lo:y,'ln I J,lI\c"'1ly IllII'IICI' Lxp.:dillOIl , 111111. Kyo>lo> (iakug.:i I 'Iliv , Sl'L II. 2~ : 71-77, (il ,," 1(1l) , ~ . II . II XIl'I) : Sur ks IOliul'~ Ilmlks, nOUVl.!:lU gcnrc , IIlIS k 110lll de Trifll/I'I . l'1 sIIr Ia f(mnallul1 lks Larapaces . /\nl1 . I\lu,,_11 1'.. 1 nal.. Paris. 14 : I ~O. * (,11\1\11\ . I. & L. 11I :--J (III'):'): Sayap - Io: lIlab,du P,lrk Sabah rd~ilillgJay 'l (l'd,lndlll. I'llbl.) (i\lllll\ , J. I . (1 7 ~<)) : (aroli a I 1IIIIl' Sy~ll'rna per Rcgn:t Ilia aLur,I':, (i ,F Ikl'r. I .:iJVig. I( ») : I(H \-1516. (il "' . I'. & ~1. VI M I S 11t)l!-I): /I I'lcldgulde In Lh.: Am · phihi ,lIls and I{cptolc., III' J\.1adagasc:ar. I C\ ~ I "",CIl (M , \'Cl1l'1':S & F. (; )a\\ \'crJags (ihR), -IXO PI'. * (;("\ 1, J.I'. 11972): FJ,"l.!~ nr K inaballl I\otllillnal Par\.. Hilil . 11l~1. SrI. Nat. Iklg .. -IX ('IJ. (;()"~ I 1(, I;, L ( I %()1. i\ sllllploflcd labk li'r SI
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(I XJ Ib): Synopsi' Rcptiliulll 01 ,h0l1 deSlTiptlOIl or Ihe spc'cicsof rcptilcs.l'al1 l. Cataphr;lcta. TOI1ol ~l's. CroClIdik,. and Fllal io~au ri am, . London (1 rcut1c!. \\ 11111 I>.:. Co) ran I : \'ii I X-' pp. - ( I XJ9 ): ( 'atalol,!lIc of the slcmkr-Ionguc'd saurians, 1\ ilh dl'~t:riplion s of ma ny nl'll gc ncra :tnd 'pCCICS. ,\111 1. Mag. 1\al. II isl., I.ondon, 2: '1 '1 1-'1" 7 II X" 2): Ikscriplillil or sOllle nCII ~pc' CIC' of rcplllc ~. rilic'fly Irlllll the Urilish t"IUSClIlll w ll ccllon. I he / (ltlIng ical tvll 'c.:!"",y: 57-5'l. II X4S) : Cal;il lI)!uc' or Ihl' Spl'C lIll Cn~ III' I i l.a ld ~ 111 the collection (II' Ihc Ilril ish ,itlSelllll. I,Olldoll (I3ntish Mlisellill Nalu",1 IlislOl,) . \ XVII I 2X 'J pp, (il~I\. ,1 , E. 8:. TI\.II(llIIHto.l ' (IX-,O-IXI2): IIIII s t ra llllll ~ lIr Ind ia ll Illnkl!,;y: dli c t1) sd cc·tcd Irom Ihc' c(llkc ll on nl 1\,1;I,illr-( iCllcral Ilard wickl', I.R.S.. I .'., M.lL \ . ' .. \1.~ . 1. \., &c" Volulllc I (pans 1-."<), Illildoll (Irl'utll'!. \\Url/ , II Clillcl, JlIll. 8: Richt er) ' II 2 ). pi 1- 100. - 111-I32h- 1XJ5 1: IlIuSl r,lll11ns or Indi an Inn lngy : c: lllcil y sckctcd Ii'olll the cllllcell(\l1 Ilr\ibior-( ICIICI'alllardll Id.c. I.R.S" I .'" \I ,ILI .S" ~I.K . 1. \" 8: (" \ 'olulllc II (par" XIXX) . London IAdo lplH I' Rldlh.;r ;lIld Co 8: !'arbury, A)lell , and Co): II 12 ), pI. 1- 102, (i HI I I(. A , I~, (I '>70): Th,' rdalion ~ hlp or thl.' s ~III~' rcfcm.:d 10 thl' gcnu s /J(l S/(/ , Blel IlIra, C;IIlIbrid),!e, 3.. X: 1-.10. - 11'!7"): I hc gell ert c rclallonships Ilr Ih,' scin,'1l1 li /ard gc'nu s /. ciollJl'im/tl and it s Iclatilc' Au"r. J. Zool.. SlIppl SC I., \lo. '1 I: 1-67 11')77): Thc' ~y~ lclllallc~ alld l'lolllti,,"af'V Idaltollslll)1S or thc' ~c incid It lan l !,;L:IHIS rl go'IJIIIII J. 11;11 , Il is1.. II ':; 15-) .. 0, (ilws. 1', III ' (I'J37): Eine Ill'U<: 1 ~ "l cd\Sc all~ Nonl- l1ll1'1 lco ( 'ill.,'I ',' Aillllf,IIIII"IISi, . /001 AliI LClp/ig. 117 : 13(, I,X , ' ( il( I~II( >I P. L \ , (llJ.N): l ip vlt. KIIWil.t\II , SCI. \1(111 '\.Y.. I III: ..01-41 .. , ~().. - SIX (ii ' 1111 ~, I\ (I X5Xa) ' l'at:tlll~lIc' o r Colllhri lll' 'Ila~cs "I' IhL: Brilish 1\,1II"CI1111 , I nndoll, \\ I ' 2X I pp. - II X)XiI): ( ':tt;ti ol,!lI<: orl he l3atradlia Salil'nlia ill thc l'" Ilceli(lIl or Ihc Urili ~ h 1\ hl"l.'I IIII. I.ondoll, ) flO pp (IX(I.~) : Third alUlIlIl1 "rIlC\\ S)1I.!C'ICS ,II' snakl's In th,' L:ol lccl 11111 Ilflll\: Ikill ~ It :VIUSl'UIll , Ann. Mag. :--Jal. I list., I olldon . .I. Ser. 12: J.+X-«,5 . - I )X(I"): I'he rep ltics or Hnt"lt India , Lond on. x\lii I .. 52 pp .. :!() pb, I ) X(!. ): I'ourth accolln t 01 IIC\\ S)1\:l'Il'S or '"a ~l' ~ III Ihe l'"llccll(HI " r th\: I1rtt l h \lusc ullI. Ann , ;\lag. al. I list , LOlldon ..t S,'I'. 1:-: XlJ-'IX. II X72 ): ()nthclc!'tlks, lIId ;lIl1phlblallso J'BlIrtl" o, PnlC Ion\. Suc , L('lIdon: 5Sh-(,00. I \.\ IS. ( '.1"..1, III \ 19.. 9): I'hc gClIlIS NlIl1'1r III IItc l'llllcl' II ,11I nCthc Rank, VIU S(;11I 1I alld II , dl slrubu li on ill thl' I\l ala y 1'<:II 11I' lIla, Bul\. Ranks 1\111 ~ .. SlIlga!,olc. I <J : 7X-97 11950): ( ' hec ~li s t "I' tItc s lIa~ e' or lite I lIdo-Au~lralian Aldll!,l'I;I).!" (R,'pti lla. Ophidia) , rrl'ubia. Ilog')1 , 2()'
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.I .. : .I')-!<~ , - ( 1900h) : NIlles JIIgml'ilI111i111/.1 .
O il lit ,' IlYIIl~ I'rllg N/iil(,fll,/i"'-1I.1 J. Str. 13r Roy. f\ ' , Soc. SlIlgap"re, <-1 :
<)(,-117 , ( IIJOlk): ( ' ltc,: ~ I lSI or Ihl' Rcptlks alld i\mplll hl:tIlS III lite Ranks :VIlls,'um , Sillgapur<:. SlIlg: lp",e, 6 PI' II IRIII"-'" K,A. ()lJX2), ( \ 11II1sItIP display ill a 13(\\'IIC\1I\ il lIg. I'I'Il' Hil,\' Slit:. \\ ashinglnn. ')-'1.'): (,~ 1-62 .. 11 \I(II\\II 'to.l, I', R. "" .I) ' (i R\\ (IX27)-'\ sy ll()ps\ ~ nflhc ' PCLIC" III' saunall r-:ptlks. co lkl·t,'d In Ind Ia by \1 :1.1"1(1\:1I<:ral Il ard\\ id,c. /."0 1. J. I.IIIHh'Il, <: 2 1.. -229 II \1 S( 111111. ,\ . ( 19'lX): I \;t It illig Iliid '\,Iclt/lld il des I}[IIIl,'rti \\'ar an" ' ,1/'(/1/11,1 c!III1I1'I'/I" (S, ' II""I, I, .~l)). h<: rp<: I(\I ~t llna. 20( 11 5): 27-1 .. , III ~ ", '. w. ( I'Xl6): RCI'i'loll tkr ( ialtllll)! I )/'{/( () I Aga lllldal') , TClllllllllcKia. Ll'ltll' II. I: 15-'-220. IIII{I(~I" \I . ILl . ~ I. l lHlll (1'1')2) ' Ghl'r Ii al lung s(I\\il' \clltaltcll HlIl , / //1'/11111/ 1111//1/1/'111 1\11,,11. IW,II (/\11111 ;1, Ilulllll lda,' ) in 13inh'p lind ' \c" ,I"IIIII , Salina, Ikrltll, I.+e l: I-,-IX, I II) lie \V, R, (lin I): I\l all ng ca ll , (II SIIIlIC !1lI!-:' Iroll IThailand , I icldi:llla /001, ( IIIC I: II, "X(6 )' (,I-X2 , ) 1"..1\)\. 1'. (1l)~21 ' ,\ 11"\\ III1Ibkss Hmc/iI 'IIIt'I. " (Sallll:l: Sl'illl'idal') li','1ll 1\11."in.lh;lhl, :--' .-llllfnco. ( ol'l' la. Wa~hil\g t "n, I.. ): X"O-l'I-+.f . (I <)I)(») , Bo rn .:a n (jd~,ltlld II/ ;trd , of tltl: gC IIII ' ( ., noe/(1('[1 /11.1 (Lacert tI, a (' l' ~~ (\ llld ; I <:) \I lilt (\c,,:npllllll' of tlHCC nl.'1\ SPl:C'I-:, .11'".1 I krpl'l(I\. 13(.,): l) I- 1m ) 111."1\, I. 8.: II. 0 1\ (I'J')")· ,\ i,/i""()lIIfl1l,hll.l (/(I"III/( "' 1\ 1\I to. III s, 1'l'J I. a .1 11111'" S) 11(11) III o r 11))/IIc/fll,IWI'II,1 11('('( '(In! (I'I II K' , 11-;71) I R-:p ltll a "qllantata: ""lIIcldac) B(I\III. zoo!. Bl'ilr. "5( I ): :"7-/111 . 11 ~1 / 1)\J . .1 120(11): N\:IIl' h~CIIIIIII" s,' l ur h'rlpflall /,lIngs hlologle 1'1.111 , l/wt'II/I/(/ 11/'(/'//111 1110 11 . I ~~71. Sauri;I, Berlin, 2~ (2): .. 1-4.. II IIII "I '~', 1>. 1199.. ). I IIl1d 1Il'1 scl tc'lIcn 1«(/1111 1,(/1(/1 '(/1/, '/111\ alii 1\'11 Klllahalu aliI \3nrtlco. Salamandra lOt 1): 22,\22.. .
(1\)'I5a) : \ IlItcrsucltulI).!,'n lur ..\ I\\lrC' nl alllla dcs 1\11. Kina ba lll (11 01'1\<:0) let! I: l-tnfiihlllll g, 1-a'11l1' l:n I'dohatidae,lIl1l(lI\1(!ac, i\llclohylld;le S;luna. [krllll, 17( I): :--1 0 , (1')I) 5b) : l IItc'rsll c \tlln gcn 1111 Alllll clI!a\llla d,'s r-..It " inahalu (HOI II CO) Ic, t! )1 ' 1';lnlll,,'n R,lnida c . Rh;lwpltorid;t<:. Sallria. I krlln. 17(.\1. 'J-17, (l'l'J5c): /.ur " C IHIIIII ~ dc, 11I1'I 1I;lIllliu'ch 'S C/iIlI','I'///li II "e'll 1, " ')1 1~ll , Il'I'i2 \(1111 \It "'lIaba lll . herpctll-
-HI 7
Bibliography
blllla, \\ '<:III\I"dl , I 1'1(.)' 27-2') (l'iI)Xa): HrlllJ1lll'~\.'lltk \\'assLO rlri),c ltl' ,illS HOIlILOO ()i\1 /., (il'l'"'II\..II.:il':II, ~ I (h) : lX4-JX5, (11)'iXh): 1\1,,.11" 1o;'lIal1;llu Ski//,Cl'II1':~ I .LOhl'I1":II1111":', 1'1;I)1h.:, (,( 2 ): X.' '10 (I')!)).,e) !\lllllel\..IIII!!CIIIIII I'oripllan/ungsbiologic lind 111111 Rill 11\11 Nlllal/fII' 1"'/( '1'\1 (HclIll ,( .! 1<, 19(0). Ih:rpdoJ:illlla . \.\ ell"\.,,II, 20( 1161 ' l.'i-IX (lO()O)' :l/c '/III'It l'l'IIdlli "llIIdallll (1'1111l ', IX(7 ) ':111 I 'n>,dl rllii 1111 i\mpk\lI s. Salina. Ikillll, 21(2): 37--lO, 11" " ,,\. 1\1.. 1\1 1o; ()11 \Y ,,111, II ,-S. '1'0%, II () I\ & 'I I IIKIP\ (11)\)')): 1."(111(11111': rc-cl ula(""1 ul'(h.: SI"(II,.,r /)r(/('o ('r1/'1I1111/1 <.1 '1\ 1I11~ , I X(,4 (R.:pllha :\ !!;lIl1lliac) I\m phlbla-l{cpl11la _() : I 'I'<~ I 0 11""\1, R, (I XX3)' (llllle\\ alltllitllc kll()\\'IIIi"g' Irl'" I II1\.' Vlalayall ArdllJlclagtl, Nnil:' I L:yucil 1\ lu, .. Ll'itlcli. :i: :!Yi -24-1 II III~I' III " i\ \'\' (I,'XII. ()II" 11..:\\ gCIIIIS "lid '1',',""01 ,1\g;lInida.: fllllli SIIIII:III:t NOlc' I l'ytlell \Ius.. LCldl'II, , . '" \-52 , 1.( './.. " (1IIIl'l'II;Ill\lllal ( 111111111'S"'11 "" IOlllogleal N,'ml.'l1dalurl.') (I '1,XI ()1'"11011 " 2,l lnl.:rprclalinn ,)l'lhc nllmi11:11 ' PITI':S ( '"llIh(,1 1 -27(, (I I)K:'). I nl~rIlal ",",11 ,,,(Ie "I' 1('ol(lgic:! 1 Ih)I11Cnd:l1l1rLO i'hlld ('dillon , l.ond"". IIl1crnalIC,lIal i'ru'l Ilu' 10lllngl 1';11 '\Clllll'lIelallllc. \ x , 13Xpp, 1" ,,1/(' lei ', (I<) ' -Ia) ' ()11 a l'"II.:t:lion (II' :l1111'1t")I:II!' liwil MI Io;inah:llu, NI>llh 11,,"1\,<' .I. \Va sh, :\l'ad. Sci .. -I-I . 2:-0 -_.:' I . (11154bl: SV'lcm:lIie' :lnd loogL:ll!!raplt) "I' Pl1IlIppilll' alllph")I:I I-Idcll.lll:l lon!. , Chicago. .n(4}. IXIS,'- (I 'J"6) : ~'>lIlI' :ll11l'llIl1lan' l'r'lI11 Ih.: 1<'1\1 lands of' North 11(,1'11.:0, 1-ic: ldl:lI1:t I,nol., ( 'hlca!!o, 3.:1(3(,)' .\X'!...t24. (I '15K:.): IllI c\' I1C\\' '\.. III\..S r.:!all't1 I" ''>I,/t' 'IIf1l1l11rl)/tIl' I " II'/C:~(/1I/1 (PI II K ~ L I i\'ldl:ln:t Illlll .. ( 'In 'ago, 3')(241: 25 7- 2(,X. (I I).'i:':h)· i\ I1CII (1l'C\..ll ol lh.: g.:nu" ( 'IT/"c!(I<,/rlu, II lilt a \..cy (", Ihc ' lk'CI':' 1".111 HOl1lcn :lmllhl' PhiliPPlllC ,,lal1lb S:lIaw;t\" 1\111", .J.. KlIChlllg, X' 2(,(-2(.4 (1<)5X,,): i\ I1l'W Inad lI'oll) S:tr:lw:lk . S;lra"a\.. Mu s. J" KUl'hin g. X( II): -l 76-17X (ItJ:;Xdl : 1\OIC' olllh,' n"I'l1.:an <11:1.' " SlIa\..c. Sarall'a\.. \Ill ' , J.. J..lh:hlllg, X( I I) 4 7tJ-4X I ( I tJ(,Ii:l) , '\"I es <>II (h.: Illali- 01' th.: gl'IIU' 1" ''')1''11'1'1/( '. 1' lddlall;1 1001., ChICago. 39( .19): -II <;-11 X ( l'I(.!)h). .II, rCI iCII "I lit,' -"1" ( 1')(,0,): '\ IL:IIL:\\ ollilc agamid li/.ards ()I Ihe gelllls "/tol"l}hIT' I II 111(1' III ( OPl'IJ, W:tsllll1gl'"I, (}): 22 1225
-lOX
( 1')62-1 ')(,3 I: R"\,,I sOCll'll' C,\p.:cllIllIl1 III "II111h Ilnnll'(1 1'1(, I : Rl'pnn X. !\111Jlll1ll1:1 . I'm<: , I 11111, Suc. l.ondon, 17.:" 4:;-16 ( 1'1(,,'): T\\o n.:\\ sp,-,cics 01 ' rrogs fro II 1 il,lI'nco , hcl,ilana 1,,"1.. ('JIII:ago. -11120)' 151 15') (19661 ' Th,' 'ysk'malics and I.llc.geogl':tphy (If' lltl' all1 pl1l111a ,)1' Ik,l'lll'o. I Ici,li:lIta/oClI " UIIL:ag". ."2: I --I() ~ . - ( I")(, 7 )' A IICII L:olubnd 'n:l\..c lll' Ihl' ~en ll s ,'i/('gOJl"'II.I rlllln Homen, 1' ll'Idiana 1.0,,1 .. ('hlcago, 51 : 77-X.1 , ( I 'nx): I h..: li'''gs :" Id loads, Chap!. I ., 111 : Kiliahaill . SIII1111111 <,I' Hol'llco K"la Kinab"III (S,d):liI SOl'll'(Yl. :111 320. - ( llJX()): Relah\c abundanl'" or li'og, and li/.ll ds III I(!rcs" of SuUlhl'asl ASia . Iliolmplca, 12( I): 14-2,' , (19'/1..1<1): \lnrpholo~ 1 'al alld cl'lI lt'~i cal \ 'arialilln in llil' Ilyill~ I 1/,:II'l!S (gellils Ih'(/IO) , Ilcldi:IIl:l loClI., l'IIIL:a!!o, IIl'U': SCI'. I X' 1-.15. (1IIX.1h) : I.alla..: III' Soulh,'a,1 Asian specie s of' I ',/}/OhJ'IICIIlIlIll alld '-"I,/olin/( Itdl(l (i\nllla : 1\:I"hallda': l Ill' RII"I1", A . &. \ 11\ \ 1I, K (cd,). ,\
<.:1;""
Hihliograph y
l,c,1 R, R.I-'. &. P./\ . (;RIII, (19Xl): \Ian:ili(llllnl~onl~all frog~ of Ihe IlIIollIl,s/C'l'hll(/ sp~ci~s I,!wup, with (,,"scriplion of two new species. I' leidiana ZooL, ( 'hicagn, lIew SCI. 19: 1-1 2. I ~',I R. R.I'. 8:. N.S. 1\ \III· (1'l5 1»): T\\o new frogs from Sara\\,:1l-.. Sa r:lw:1 k MI" . .I.. Koehing. 9:270-276. I'CoI'R, I t~ . &. W. II. Mll K (I %5): e\\' SIX'CIl'S of sc incid li/ard o rthc gl'nus ,\,1111'111)/111111'"11.1 li'mll Sarawak . '-'rael J. /.()oL. l-t : 11-t-1-tt). "',I 'R, It I-' 8:. \\,.'. KIM, (1%1): /\ nc\\ GI\'e-th\'elling li7.ard of Ihe genus ( " T/",/tU ' /l'/t,,' 110m lah , Sarawak \11 1" . .I., Kuching, II ' 27-t-27h. ", ,1 '11, R. F.. M. L \ I-.I~I , /\ . HI 1'1 & P. Y '~ IIII , ( 1497): /\ nl'\\ s pe cic~ of 1'<'1'/111,,/111 (/\nura : I\1 cgophryldae) frOIl1 Horneo. ASlat. I krp. Rl'Sl'a rch 7: -tX-50. "<:1 '11, RT 8:. /\ .E. LI \ 11 0' (14(,(,,: rhe la.\OnOnllC ~IrlIlIS (If Bonl'an ~n : ll. cs or Iile gcuu~ I'\('uc/omhd/fll/ .I \~ and of Iile nOlllinal l,!l'nus Idinplloli., "·10(',11 I\1(1). Proc. Calif. /\cad. Sci .. .14: .107-.\ l-t . ", a'I(, RY 8:. I L M .\1\\ ( 1,>(,,5): The sy"leillalics and l'volu lion of Iill' (}ril,ntal co luhriLi snakes of thc ge nu s ( '<111111/(/1'111 . l'ic llilalla 1001.. Chicago. ·19: 1-311-t . 1"' 011<' R.F. & R.B. SIII"I"', (I')X')): I-rogs of Sahah Sahah Parks Puhllca lion o. 10- K(lla KIIl;lhalu (Sahah Pat ks 'l'rusll;es), 12') pp ( I')(H ): /\ nl'W Spl't: ll'S of li'ng 01 Ihl' ge nus I .,'p/ohm../11'//" S\IIIII (/\l1l11'a : l'e lc1halldae), wi lh a key ltl Illl: speCil" frolll H"rneo. Rafl k , Bull. Zoo l.. Singaporl' . .w( I): 'l')- IO:1. - ( 19'12): '1he IIIOlllall' :11I1plllb1:111 1;lulla of m>rlhwl'~ll'rn HOflleCl Malay. Nat. .I.. Kual a LUlllpur, 46: -t1 -5 1. (IC)\)-t). I·irsl Record o flhc 11I:11 e1 g~llIl~ 1',I'I'wlolal"/I ',' (1 .:1CCI111i;] : /\),!:l111idac) 111 Horneo. wil" lkscriplion "I' a n<:\\' speCll'S. Rallies UIlI L7.ool, Singapore, -t2( -tl: '161%) ,
( 1'>96): '1'\\\1 IIl'W SPl'C Il'S "I' rrllg~ from S,'ulhcastl'rn Sara\\'ak. Ra llks Bull. /.0"'-- Singapnle 44(21: 5-t 1-)-t9 - ( 1997) : A fielel g ilide to Iii, frllgs of Blll11ell Kllta K IlIab:1Iu ("alural 1\ 1~IOI); Puhlll'alions in ,ISS(ll'. \\ ilh Science and Ih:h nol ogy l lnil). 211" pp. IM,I R, IU -'.. It H, SII IIlIM, 8: I-.L '1' , (1'11)5): Nl'\\ SPl'c"s alld ne\\ n:cords orAnura ns fnllll Homen. Rallks UulL 1001.. 'Iingnpnre -t.l( I ): 11 5- 1~ I. (11)'16): I-'rogs:l11dloads III: W['I\(" K. 1. 8:. i\ . PIl11 II1'I'" (Leis.): Klilabalu SUllllllil Il l' Ilorne(l Kol a KlIlaha lli ISah;l" S..e.: Sahail Par~ s) : 352-367 IMoi le. R.F. & I'. L. 1 \'1 (I'NO): ({c'cl'ntl y dl,co\'cred a 11(1 newly ass ig ned li-rlg Iar~ae t ({ ;l1l11be alld Rhacopilnmiae) litl11l IIlll'llctl. Rallks Hull. Zoo l.. ~1I1),!apllrl' . .,~( I): ., ') .
- ( 19t)(':l): ( ' h l'dh ~ 1 .. rlhe I rog~Clr H"fllcCl . I ~amcs Hull IOllL. SlIlgapon:, ·j,I(2)' 55 1-57-t . (l'llJ()h): The nalur:l l hi ~hl ry .. I' :1 III pi II hl:lIlS :l1ld n:pll ks In Sabah. K..la Klllah;1I1l (N:Il11ralll1'lory Publica llons (Hlll'llL'll) SLin. Ilhd. I, \. I 10 I pp
,""11
IMoIl(, R.E _F L. T;\ ~, :vI . L 'I-.I\I & 1'. Y N (2001): 1\cw spl'cics I)frhl' h/:l1d (icnlls .\,'/" '/1(1111111'111111.1, (I accrl1lia : Se llleldae ), wllh nole, un .:colog lGI I and geographic distribulion of spec ies In Sahah. Mala y,,:!. R:lllles Ilull . 1.001., Si nga pore. 49( 2): I XI- I Xl) 1"1,I'K, R.F , F.L. T\ 'J & P. Y \\1111 1,., (2()1I0 ): 'I he frog 1:l1l11a of Ihrel' Parks In Sab:lh . Ma laYS ia K Inaballl 1\ 11'1. . ( 'roder R:l1lgC ParI-. :11 It! T:I\\'aul lills Park Sahah Parb at. Jour.. 3: 7-2x. - (200 I): A ne" Inad or Ihl' ( iell ll' 11I \[))I1(1 (Allura: Bu11.)l1llla.:) Irulll Borneo. Rallk, IlulL 1001., Slllgalwr.:, -t')1 I ): 35·.17. 1,,'1 R, R.I-.. H. &. /\ . I Will-. (I ,)%) .. ':W Spl'l:ll" l,f ranlt! rrogs (/\lllph ibia : Anura) Imlll ( 'c mral Kall111 al1lall _ ilorneo. Rallks HIIII 1.001.. SIIl!!a[l<,re. -t-t(2): 36,,':\(,'). I'll a R, R.I-'. 8:. II K. \IOKIS (I 'lXX): "iaxonoillic stalus :ll1d rl'produc:tiv.: billingy of'blll'lll':l n laclpllie - ca l 1)'111)( frogs. ('opel:1, Wasillngloll, (-t). II )(,O-IO();! (I ')')3) : A cOlllparisoll of am phihi ;1I1 eOlllllllll1 lll eS Ihrough lillle and frolll pl ac\.' III i1nrnl':l n 1(11.:,1, . .I lrop l,co L, 9: -tOl)-B1, 1"011(, RY.. I I. K. VOI{I ~ &. KJ. I·KI •• ~ I K (Il)Xh): Orgal1i/alion ofa COIllI ll llllll y o flat!po ks III rallll(lI·,,1 ,I rl'ams 111 80nlc.:O. .I . Trop. I:coL. 2: 1')3-20). i'I(,IR, ru ·" ILK . VOKIS &. 1', \,\-',\1 ~I R(19X6) ' I al'\;11 Ir;lIlport 111 a Hom':: l1l r:lll1d frog. Cl'pCla, Washlllg((lll, (2). )2.1 -525. I'J(:I·K. R.I ', & IU . \'V \~~I k" I, (199()): /\ t':lItrllicnld-lll-.l' !\lIl1ralilarva Ihllll Soulhl'asl ;\'Ia. 1001. ~c i c.: n . 7: 5575(11. lsI-. \ 'IJ \R, 1).'1'. ( Il!7X): i\ lIew speC I(;S of HlIl'hollrtllll : Fir~t record 01':1 lil 'c:og lclsslli anuran ill !!"rnell. ( 'ope!:I , \.\-'ashinl,! llln, (4): 564-566. h"'\~Il;\I{. D.T.. R.B. SII II'III'C, 8:.:V1 SII\IIIlI 111'1 II'NX): Rl:pcli1 onlh" lito I.10nl\.'() Illmll\clslly hpcdlllllll I')'n Iknluang Karillllll1 alllllli" 1':11-1- . \Vesl K.:11l1l1allta n I allpl. 1~ lIll1l1all W1Idlili: SantlUar\ , Sar<m ak Part I lel pc.:ltll;1l1na. Bentuang Karlllllln all ona l Pal~ 1'11I.led PI) 26 !J.1. Wordl \-\ Ilk hllid I(,r aillre - Indol1l'sia Progr:ll1111le PIIlllla11a l-. I')')X: 1- 12. .h((IIN) . S,K . 11 979): /\ glllde 10 i\1ol1111 KlIl ah;lill NaIional Park . Sabah all\,":11 Par", I'll hilca lilln '\ (1 ." Kllia K.ll1abalu (, ' ab:~l Nal10nal Park I Il.L,I':l" ).:". pp. - (IIJX(» : K.inabalu ParI. . Sahah 1';lIb l'"hil c: lIlIlll '\0.7 Kola I,,"ahahl (Sabah Na lllll1a l ParI. 11l.I~l l'e'). (,,\ Pl'. *JI 'I-.IN"'. D,\I. (1'I7X ): fhl' lirst hllndl ed YC: II s. II1 . l lll'l'Jc" I),M .. ( '. \VI''J III I..R DIMoIl \ (I tb ). KIII:l ha lll - ~lllll 11111 of' Bornco KOla Kllla ba lu (Sahah Soc Mllllllgl )' -t:' -N. .1[\\1",.1. K.1l1 (1')30): Nclleo.,llli '
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11')101 : l ' llll~ lIelie ,\( '1111/,/"'11/" - :\11 IIl1d al ld e lc 1\l11phd) ICIl \ lIlI Ikll (">III1l.llla J. al. Tlld 'cli r. ' ..:d. IlId " BaWl la. (lli ' 1 .'\-2 ~ ( I'ID I lit e alllphlllld 01 lite IlId 'l-/\II ~lra liall !\n:hlpt:iago. I cllk'l (I ' .I ilnll ), \II I J O~ pp. J.- \1 I' . I· (I ~251 I· Illi g..: Ik ll1nhull gl'lI 711 l\1crrcm~ Ii alldbuch hIS lall OkclI. IX . ."1"1-'92 . 1'1. III. (I X27) IOlllogl','h,' l\1ollPgraphlCr lsI' \!."/,,,., fIlii il"I,.1 haltl/'II.li" l.e.\ II(IIX,J. I L) I, (Scrp": IIIC:": I'h lp"la..:). Sallria.lkrlill 22(2): 19-22 . K, III . II. (I 1'2(1): Iklll~i ge l UI KCIllllilis dlil 1\lllphlhll.;1I Ill: KI III , II : Iklll iigc / lil 1""log ic ulld \'crgkichclldell ;\lIalll/l1ll', Frallhflll1 a.ilil. (I knnallllschc HuchhandIUllg ), 75 I ~2 . *KI In t.\. S. (1')76), Ncpcllthl" III' Muunl Klllab:liu Kula K illabalu (Sabalt Park., Puhl. 2). *Klllts('II" 1<, II (1 1)1)0: D, l' cpl phyll'CItCIl MUll ' gc,c1I -
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,;cilal"tcn alll Mount Kin aba lu (Nord Bortleo, Sabah, Malay,ia) NOla Ikd\\i gl:l. 5 111 '2): 1-75. \""'1, i.... I.H .. I'. Y" 1I11 'I I/.: S. !\ lIt '\~1I (1')'1') : /\ cOlllpara· ttl'l' ,t udy 01 Ihe amphlbiall I'opuialinll III Iclc lcncc 10 habital disturbancc :ICl'llsS an c1i:I'atl"llal gladll'lIl III I-: lIl.thaltl 1':lrk , Sabah, \Iala~ 'Ia . Sabah P:1rb Nat .ltllll'.. 2 27·4-1 1..\1 RI ' 11..1 .. (1 7Mi I: SP":l'IlilCIi Illi:thl·UIIi. i:\ llllwlls '),llllP'1I1 rcpfll,ulll Vll'lllla, 2 1,1 pp, L.VIII , .I.D .. J. L. KIIII\" &. (i .A. S\~llll\lJ ~ (1l}<)I).· 1he SIII:l\\c~i Hl:o l'h Raci:l. ( '"llIh,,/, (/""(/.1 I "'I )\'{f.I . •lIld d rClliarha l'lc "I'I(lpal','~l li c ag.gregatlon. I'a"ilic SCI.. -I ~ : .1:')-Jt> 1. I I ~'" . R.I'. (I X,O): ()hsCi V;1I101l' g~ II':rak~ 'lir k, reptl k~ rec ucllhs dan' Ie \'ll :ag.:. In: IJ I'1'11(1<1 \, I .. 1. : VIl) age' allhll' dll m()ntle. C\cculc: pal Ill'dn.: till RllI , sur Ia Cnl'\'dlt: tI..: S;I M;I)""". ia Coquille. pcndant les ' 1I11l1'~ I S22, I X2~ ,'t I X25 . /.olll"gle 2( I): 1 -6~. 1.1 \1" ", A. I.. , ( 19."'5): SyslCIII:lIIC ",,1..:, oll lh.: i\ ~ l all 'nak<' 1,'('(1'/01/ ,1"h('I11('/1/,1 Phil.l . Sci .. II/lamia. X-I : 1<):;-20.>. ( I 'J( •.\ I: Conlribulilln 10 a rCI ICI\ nl Ph ilipPIlI": ~ lIah cs . III. - I Ite genera ,1/,lIll'IJm ,lIld I'hd.l Sl'I .. Man ila. ')2 : 523-550. - ( I t)(,-l ): ( ·nlllrihuIIl.n, til a I C\ ICI\ "I' I'llIhpJ1l1lc sllake,:, 11. - Th c gCIICI:I I "'111'1" ,\ and Sihl'lII>phi.l . Phil..I . Sci .. Mall ila. ')2 II t)(,J I (J I: JIl 7-.~X I. - 11')hX): ( 'Ilnlrihulion III a J'l'vic\\ "I' I'llIhl'PIIlC: ~ nahcs , .x II - '1he.: ~nai. e' "I' Ihe gCllus /)('lIclr,."'l lltis (Scrp..:nles: Co lubnda..:). Phil..1 S..:i ., ~ilantl .l. '17 : n 1- 1<)6. 1.1\ lioN t'\ . Ii. &. W . ( '. Il RO\1 N ( 11)5')): i\ 1't:\'ICW ol'th..: ~ Ilak ..:, Ihe ge llus PI'(,lId(l{'(lhdil>1I witlt rc.:l1lal'i., Ihc sl:III1' Ill'lhc g..:ncra ..I.[!,l'fll,ltis and lil'ltfo,l!l'lIl llti l (SC:'1'CIlICS' ( 'olllhridae). I'rot: . ( .tlli'. ,\ ..:;!d . SCI ., I') (I -I): -1 7:i-50X . Llllll1ll1 J II Ill , T.W. I \N (IX')3): 011 rcptlle, Imm ol1hn"rnco. "te' I.cydc ll MII, .. Il'l(kll. 15(3): 250-257, (11)05) /t'lll
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.:al Prc s~) . 124 pp. I 1\1. K.K.P. 8: P.kl . N" (19'11): N<:pcnthlphllou" larvac and brccdin!! hablt ~ oflh.: slid.)" Irof!, }\al"plir\"lllls pi('/(mSlIg l/1lI TSlIIl IIlI (Amphibia : ivli.:rohyltdat.:) Rallk" Bull lolli.. SlIlgaplln.: . .w( I): 2(1)-2 14. 1. 1~~.l l r (I X07): Ik,chrcibllngder ·alll raltl'Il.Sallllllllln!! d<:r 1 1 11l\"c l,,1iiI / 1II{o~ t ock . 1.I\elleAbthciluJlg. Kostnc" (Adkrl- I:rb.:n): 51- IUO 1. 11\ ' .\1 ,' I ~. ( '. (1 75X): Sy"lcma natural' p..:r r..:gna tri a naturale. "t.:l:U lid UII I l: la"l:s. ord ulI;.,. ge nera. 'pcc les. <':11111 charal·t..: ribus. dilkr":llliis. S)11O n ym l ~. l oci~ . Tomus I.. edit. 10: X24 pp. LtD) Il, \tI .. R.F. IN(,I K 8:. F.W. KI \I' , (I'ihX); Oil thl: dl\'I:Is it)' or r..:ptile and amphibian spcci..:, in a l3orl1..:an 1<.1111 tlln':sl. A 111<.: •. I\ afl.ralist, 202; 497-:_ 15. L, "'11(11)( ,I'. A . ( I')JX); 'c'W ,na"..:' oftht.: !!t.:nt.:1<.1<. '1Ill1l/limll. //1111,1:01'11" and /iw",r<'.\III'lI.1 from 1\·11. Kinahalu. 0.1h HIlI'Il(;(l. Prol: . Bio I. Soc . Wa,llIl)glol). 51 : -1 1-10. "Loll. II. ( IX5.:!); Nott.:~ oran asecnt oflh..: mountain Kilia H;l low .I . Ind. Arch .. VI; 1· 17. *1 01\1(\ , .l.H .. D.\\- . I II ' & H.C. SI" I (1971): I· n-':ct o f drou ght on Moullt Ki nabal u \1 alay. :'\at. J.. Kuala 1.1I11lpm, 2() : 17X-179. *LII'IN(" D.M.. C. WI \I &. I:. R. 011\(,11 \ (197X): Klnabalu - SlIllllllit of I ~ornco Kot a Kin 3ba l1l ISahah Soc . MOllogr.). *M \( KINNON. K.. G. II \11 \, II. II \IIM &. A . M \f\(, \1 I~ (11)97 ):'1he eco logy of Kalima ntan London (Oxlilrd Ulli\'. Press). \1 \1 K'" I~ , R. II'IX5); Amphihicll und RCpl il i..:nl'llml\ lount Klnahalll (-IIOIIll), N"rdholnCIl . h cr p e tl) l ~lU n a. V...·t.:instadt . 7(35); ("U , - (11)~6) ; I.in ~t.:itcncr Rudcrli'lbCh \'I)m Mt.Kinaha lu . ord· HLlI'IIl:ll' NhC/ell/,hllr/ls ('\'( ' 1'..111 (HOIII IM,I~) Salina. Bcrlin. H( 1):.1 . - 119X7): Il crpctolngischc Ikoilac htun gcn alll ·It.Kina· ba lu BomcLl. r-.lrtt . 1 001. ~h" . Ikrlll). () 1(2): 26'1·2l)~ . ( I ')XXa); C'lrllle/mll'llIl 1>111111'11.'" (1\-1(~ ) 1 \({(), I XI)O). Sallria, Berlin , 10( I): 2'1. (I ()~Xb) : RII/" IIIHII'pcr ai, H":I\ohllcr (\(:r Kullllriand~c h aJl T;II11hllnans (Nord-Bnn1co). Sauria, Ikrlin, 10(4)' <)·1 I. (1<)~Xl') : Walldt.:rllllf!..:n 1111 1\1ount Killaba lu . N;II ,oll;rIpa r" . I\nrdbornco. . at. \ llIs .. i-ran" fllrt 1\ 1. II~«,) ; 16 1·1 S I, - ( 19XKd): I k rpctologi'chc Stlld ic'n an ..:in":lll Wa ldbaL"il :1111 Mt.Kin aha lll, Ilnrllcn. hcrpctot:ulna. \\'·cllIstadt. 10(5.1); (,· 11 - I 19X9): l1c rpct'llogi~chc Ik ohach tun gt.:n alll Mounl Klllaha lu, Ilornco II. Mltl. 1.001. ,111' . Ikrlln, 65(2): 17')-200 - (1l)9Ia): lur I krp..:t o l:lll11a dcs oht.:rcn Llwago. \til Kiliaha ili ' Nordblll'lll:o. hl: lpctofall na, WCl nsta dt , 1.3(72): 2(,-34 . - ( Il)l) I b): ,\1,1"'""'11017/111.1 'p.n. ISalin a: Scinl·ida..: J , 'Olll
\tIl. Kiliahalw Nord· HOIllt.:o S;llIlla.lkrllli 1'(3); 232X. (1l)91c); Dil: Ilogcnfi ngclgt.:C"lb d":l (iattllng (I.,.,,,r/(/('· 11'I1I,\' am I\lt Klnabalu Nurd HIll'lIt.:\) . h..:rpelillalilid. \\'..:i nstadl. 1.>(7-11: (,- IO. ( I 9'):!,,) Ilt.:rp e\(llol!" ' hl' l!cob.l c litllll gl'1I alii vlt Kinahalu. Nord-l3om..:o III , 'villi Illoi. r-. lus Ikrlill . 6~ (1 ) ' 101 -1 IX. ( 11)')2b); 1('/1/11111/,1\ l'It'III' "I'" (. \l) llIa . 1'l:lllhatrdacj \om :Vlouill Kinaba lu ord· l3ornco Sauna. Ikrlln 14(3); 3-6.
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(1')!)3) . Zur Zcicilliullf!"anabrlll ;il IIml IllIn I'Il"l..:ll· \crhaltclI dc~ I{lIdcrli·o~..:hc, NI,,",,/,hlll'll" (/I1,!.:"II/! I.I'Irt.1 I(x 1.II I \KIl, IXl)O. Salilla. lkrlill 15( 4): \~·,x ( 1994a): I:inlgc IkIllCr" UIl!!CII/U ":,,111<'11.111 KII \\, Il)X4. Salina. Ikrl ill 16(2). 9· 1\. ( I ')<)4h) ; I krp ..: t ologl~l'hc Ik obadllllll g": l1 alll Ml Klnahalu , 'md-Ilorncn. IV . \1 It I. I.()ol Mus . lkrlill. 70; 217-2-19. (11)<)4e): Vijf kikkcl'" bij dl' J\ksrlau-gnlt 01' 'viI. Klllaballi (nllordelij" Ilornc'o). Lar:cna 52(4 ). X6·I)O. (I ')').jd); Lc'll/dd/allll' ,bill! \all Mt Kimballl ; ':.ehlllldt!(irnlltb lang('allll/llIrill ,lc/rlllu/li). I accl1a 52( 5 ); 111 I D. ( I I)'>:;a); OWl' het \ool1plantll1gsblotllop van l'lir/alltll.1 UJl Mllunt Kinaila iulllt)()rdcllj" Ilon1c·Il). Lac':11a 53(~ I: 6X-7-1 . (I ')<)5h): \Vcr rid' hier; Frosc h odt.:r I kll ~L'hr..:c kc'! Salll ia. Ikrlrll. 17(2); ''i-.IX. ( I')')5e)' Di<: Krl'lt.:nl'rll,\:hc d":l (i atlullf! ,1it'g "p!rIT.I Kl III & \ ' \N II \'SII I. I X22 alll \1 oun t Kln aha lu, i'ord HIlI1ll:(). hCIPCtol:II IIl:t, Wc ill,tad!. 17(\)4 1: 6-10. (I'NSdl: K(lpl scllcn lkck,'n b":l h':lllI:lI (ic,r\lie('\li.;m VOII IU,III "/"101'11.1 UI/,!.:ulim,·I,., .1 Alii, 1<)2 7 11\lllphlhi a . Rha cu plrlln (~ I l:) , Sahlillillidra. Rhl'lIlh;ICh , \ I(-I ); 24524(,. ( Il)\))..:): Ilomcl)sc Ikrgagalll..:n (/'!r".WIIIII ·I.I) I an MIlLInt Klllab;r\u . I :Kl:rta. I \:1Iit:1l . 54(2). 65-6'1. (I ()')Ila); Voortplantlng van d..: J)w..: rgpad / ''''''/ili/ 'I 'I/(' JIIIS,'rt/ . i.accna, I eldcn, 54(-1 ): 12')- i.l.i ( I')()(,b) ' Ikmcr"ullf!l:n IIII' l ~ura" ll\ 11 :11 r: llll gcr I ' ro~chartcn alll \I ou nt Klnahalu. N(lrd - ll n ('(l c'll. hcrpcl11 t:lII n;I, I X( 101 ). _O-2(l . II ()l)(,l:I ' I krpe tlll()g l ~eh..: n 'obaeiltllll gt:1I :1111 \loullt I\ lltl. /.()ol. \Iu s. Ikrl. Kinaha lu, ord-Ilmnco. V. 12( 2); 277-21)5 . ( I')')C,d) ; I .c/illlhracllllllll gllllllll,~"II,II,1 'Jl 11 (t\ nura ; Pt.:io\latidac) ""ll Mlllint Kinahaill. Nllld-lloln c(I vIllI. 10lli I\'hl" Her! 7:!(.:! I 21)7- I() I 11l)I)(,c ). I lntt.:r, ul·lnmgc'n I llill A" \I\ Itiitsrhytlrlllu, 1'011 SlIIlImll HOI II (,I~ . 1l)IX alll MI. "-1I1;lhalli. S:III1I.1 , Ikrlln 11«4)' 1·16. (11)l)XI: lU I 1'lll1pllan7Un!!,I)(olllglc' dcr Ka..,"adenli·i\,r:Itc, lit:r (iattUl)g \1" 1'1""',1:< '11( ',1 \ ,,, •. lIN I. hCI pctolalllla, Weinstadl. 20( 115); 21 ·2(,.
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I Il)l)l) I: I )IC 1\.;lIilqllal" lcngcllll:ll1scilall c""'S Ikr)'.hadl~ s 1IIII.lIiII.!O. h<:lp.:lol:llllla . \VC"I\I;ldl , ~ ilI20I : IJ - I'). ( ~ Illlll) ' Illc' Sl'iil ,IIII,Jrii'l' ilc lin (ialtung /.<,/I/o/"/,,r 1)1 ' 1«liS. I,)X() ,,111 ,\1111"11 "-,,,:oh:olli . '\I"rd-n"lTl<:o. "lOl pcl"I :" ",,,. W,·""I :,dl . 22( 12,')' II - IS, \ 1\1 f..~11 .,. It. (' 1\1(11111 & 1. h II I Il)l)l)): /\llIflhl C:1I l'lI r~pllkn I': ,n l)e nlillak()1 (S:'0;il1 , r..,,1;,k" I"). - LaCLTI:l. 1.~ldcli. '7((,) ' I') I- I l)~ , \~ , II "'~II '. I{ &.1 1\., ," , 'II (1')')lJ): Ikscl llill Ill).'. 1';'11 hel "i\..".:rll'I" \:111 1( '11/111,,1,,\ 111'111 ',1/ vi \I'll, 1')!)7. l'<:n 1<1""<:1 1111 ,iL' blllillC l'v1t:gophryi,la.:. I :II;,' lIa. I cld,'I1, :-7( 5 1: I (>I ~ I 1>5. ( _1l01l) ' 11,'s"h,,"OUII)! \.'111\.'1' n,'u,' n ·I/I \('/lill· l. aJ'Vc ( . I/lIiIlI"(~III"<'I) Il'n Unr1lc'(). Salallland,a, Rhc",had, . .' (,el: 121-12'1. ~ 'I II HII s, I~ . J "- ',SI' II c\: J 1\.1(11 1 (11)1)1»). Die 1,:l1Yl' VOII S/IIII,.UI.I /lIh, 'r;IIII!',III ,1 BOlli 1\1:1 R. 191X , l· i1li: 1l~1I'" cC:lllroicn,dc:niilllil iell,· Kaulqllapp.: ails L\oll1cn (,\lIura: R:,,,idac) , I kqlel"/oa 12 (1 ·21. 17- 2_ . \l'l l ... ~II ,. Ie & M , M \1\1" (19 1)71: MI')!'''I,III'1 'S kohal'll I/lIi,l'lI\ n':lIn p"l"halldc!" F!"l\Sl'l1 1',1111 'h ,uI11 Kinahalu , S~1lI1 i:l. H... rhll 11)11): .j 1-,\ 7. M 'II f..11I", It &.. K. 1{ 1I111 (1')1)31 : Nal'lil r'l)'. III L('/J(olllla,\ I,i, 111,1 M.'\ I" "liS, 11)112, 1!\lIura : I'l'I (\balidac). S:!mia. Ikrhll 15(31: 7. 1). 11 1)<)(':1): 1)1l' llalllnrri;,,·Ii.: d.:r (ialillll)'. 1'11;1(///11/,1 \l)1ll I\lIIH IlIl K III:tlxdll '1','111 ' CJllt:rhlll'k lind dlc
M II "~Il '. It ,'" II. S·II II{ ( 1<)<)3 r J)ll' ~(ll !!l'n c'incs I3 I!>S<', lOll .\111111 //1'11 illl("'/I1"''''' ('1(,(,,,1/1. Salalll:!lIdra. 2l)(~ ) : I :;~- 1 5,1 - (11)9111 : RlIli;:, ll'lI l1ll!! 'Oil 1'111'<'11,1 I/Iwlwl;.1 ulld I:r,t11;1<.:1 1\1 \.:1<, dll'seT ;\ 11 illl Naliona lpar" MOllnl Killa baill l\1ala;' ~ia , S:l lan1a lld l·:t. 12( I): ~5-5g . VIAl 1\·111'. I;. V. (IW,lI): SyslclTlalic: dl\islon and cvollilinn I,rl hc' l'I'llIhrid sn;)k ... g.:nlls 'V,,/rl\ . willi <':'\lIIIII,' nl" 011 thl' ~lIhlalllllv Naln ·i llac Pro<.: l\l·ad. Nat. SCI. I'llilad<.:lpl1i'I , 11 2(' 1: -+1 -7 1. M 'I ~ 1111 '1'. l J. ( 1')X2): Dic ( ,all llll).'./'/I'('lIo://(/1/ "-11111 . I X22 I.:i l 2, Salln;l. Bn hn , <11.\): 5-12 . ( II)XJ): \. xkllrSIlHI alll viI. K IIlah,tllI 4100 III. Nord 13''1'11':" . Ieil 1: ( 'ht-c""sil' del' l IcTpeln(;,Ulla nhcrhalb hllOli1 (iNN . hl'qlclol;llll1a. Weinsla(h. 5(2,1): 20-.1 1. rv,l .I~ 1I11 I. l l, &. W. l)l 'V II< (ILJX 21: I- xkurs loll ;1111 Ml. Kin:lhalu -I I(10 III. I\'(\nl - n Of11l'(\, T\:il 2: I krpelolngi -
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,,<,iIe' I: IIlllriil'kc. hcrpcllll:iIlI1;I. \VcIII,I;cdl . -1(21): 11 Il). (11)<) I)' nl C 1·.l' IIl':1I \':lI1kc'lknpl;lga mcll del' (; ;1111111 .' (;II/weel,llltlll,l' "- \11" (S:IIIII:I : .l\1-"IIll1da(') Il l. (;11/11'('( ' I)/Iltlll\' gmlll/i.,' ( ; 1< II. I X-l51. S;luria. UCrilil 1.1(.\1: .1. 10.:; /\ bb. - ( I()I):') : I>I C I.L·llI':lI Wln"cl~IIpl ;I!-,:IIIICIl tlc- r (iallun).' (;I/IIl)('( 'I)holll" (Sauna : l\).!'lIllidac) V. I>ic' />< ' /lii-( "II Pl't: , Sauria , [kriin 1.1(.\): 7-20. - (~OllO) : Ikscnplillil "I' ;t II,'IV !-'CIIUS, /III)\'/('({Iu/('" gL'Il. 1101 (S;lltria : Agamidal') fnJm \'\ PII 11 I K il1:rb:d u, '-"rill [lOrtlC". \\' llh rCllln,.\", (\11 Ih e; )'.':IIC rI l' idcllll iy ,)f (;{)I/fJi'c'I,/ia/1I1 I'e/III/,;,'ll ('l frlllIl; URI \.\ . Il}l)I). IlaIIlalit y ad. I\ladra~ 25( I): 1.1-20. M.II\ IIII I. lI. & \'\ . (,KI N,\II ~ (I')l)-I): (iki llliegl'll bei HI1JJI.-Iu"c/" <'I'I'{OI('/l1I (I\.I 'III. I X~() I IS,lun;1: ;\ g:O IlH "lae). "auria. lkri lll Ih(31: 17-.\X. - 11<)<)7): ,\mplldlll''' , " Rcpltlil'lI Siidnst;I~ l ens . Me'IlS!l' l (NTVl. :;12 pp M . II\ 111I' \, l & ( '. S II Ii II (llJl)X): NiIllCIJI)IIIIi'If\' ('Willi IJIIIII'It"I'" "I'. 1\ . ('\I\unl : Rha C:(lpllOridac'). ei ll Ileuer Fl lI).!li·o,d l \(\ 11 (ic:r \'l;rI :l ii'l'h<:lIllalhin~c'l. SlIll l;tlla und 13nl'11co, SaurIa. Iktllll. 20(YI: 37-12, rvl IR~I\\111 . J.. 1\. J)1 1"OI\. i\ . ()'II'II, L. [' ''1)1 I I 8:. ;\ . 1111 IIIl (2001l1: Phyl,>g.endic rl' lali,)fbl1,p~ III lilt: R:lIlld;lc . IlI(k· pe'ndcnt origin "rdll CCI d",dop"lcnlllllhc' gCllera 1'llIillll//I,1 allli Ji lr/OI'(IJ/II, C.r.,\cad , Sci_Vic .. 32.1 : 2X7-2')7. M'\1el. 11. 8: R.I', 1",,1 It 1195:;): NIII.:s 011 111.: ~ 1I ; I"l: " ol ' ih l' g<:IH I' ('''/III1J(tr/II , F! eldl;lIla Lo(\l.. ( hicago ..~7 : 1(>7-
2( 1) . M 1"111, M. ( 1"71)): '\lnpllll"'I'" 1>1' Salxlh . I, SyslL'l1lalil' alilimiura l hlSl,>ry mIlt's. ( ',>nl L binI. Lah, Kyolo Lllil'" 2~ (4) : 30J -.\.t(, - (Il)X~a): /\I11Jllt ihI;' '' ~ "I' Sahal! , II. AnHlslic: charac:tl" ri~(ic~ til' Ilm:<: Clllllllllln Anurall ,pecic" ('01111'. bill!. Lab. "-YOI" l llll\ .. (_6): 12,·129, ( I LJIQhl: (all C1laral'lc'rislic:s 1>1 ~l'\'c ra l /\nlll'an spccit:s Irolll [a~t I\.alilllanian. ('01111'. biOI. I,,,h. Kyol" I " i\'" (261: I \ 1-11<1. ( 1')')5 I: Call produccd by a "voieeic!>s" fi·ug. I
( 'a ll c: llaracll'l'i,lic, an d SY"':llIa lic rel;llitlilshils of a 1\'1 ;11"":111 lrecl 'rug N"('/;.w/lI.I /';eltt.1 (!\nura : RiIanlphllrid ae). I IL',V , 1.. (): 6~ - M . - ( ll)lJ7): ( 'all charac:lcn Sl leS "i'M"laysi;1II Lt'/'((lIII/II.1' wilh a d<:s<.:rrplll)l) Oflll'() ni:1I specil's (Anura : Pc'lnhatithlc). t 'oPe'ia. Washington, II): I :;X-165. M,I I\I.I. M .. ·I. (·!I I'<.,IRIl & J. :-\.\I
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M \I"'I, M .. T. 111 1-.1111 & II . "J\ ~1Il11 (19X5): The amphihian,and reptile, co llected li'om Bomco. MalaYSia . Bul l. 'loya ll1<1 SCl.vllls. 1I0.X: 151 - 15'1. M \1 SIll, M .. T. SilO & 1. 1I1 SIlNll~1I1 I (1l}~6) : AeOIl.'IIC 34): Dil' S<:hlang,'ngaItLing {){'Iltlidlll,h,,' il lJi I 1''1,01 H in ~ysll'matl,chcl lind /Ilngc,'grap hl s,:hcr Hl"/ichlln!:!. Ardl . '\Jal.. I <.: 'IV i/! , 3: 1:\7-20·1. (I')(.~) : Die i\l1en lind I lnlcrancn d<:1 SCllll1l1c\..halllll,chlallgcII (( '111'1 '\1'1",1('(1) Sl'llck . bil'\" Frankfill1. M. 4'1(3 ,.-1): 1'>1 -2 17. (1%9) : i her die V:lll ahlilliil dcr A..:hlslrCIlt:llllall<:r. ()/rgfldo" ()( W/rIll'IIIII.I· ""Ill'\.. . bioi . h ank Iun 1\1. 50t5 6): }3'1-345. .\'11 1111 ~I ". M.II. II'I52) ;\ g<:lu:n<: ,)"oJlSl' "flhc II /ards "ftlu; subl;lIllliy I ygoSLl11llnal'. Smlihson. 1\11'(.'. (",11.. 117( 17): 1-.15 *I\1JIIIIII{( '. I .. (1l)2X I: I I rtlpll..c:rnas Vlilank Ursko!!ar Sloc\..h"hll *t I'no): hlr"l 111i: and :lIII':I1III1C' III Ih, ~ I al:lY . \r<:llIpclago I.Illldol1 « i<:Llq..'l' /\Ikll <'I:. UIl\l'II1 LltI.) M()t'I.'1 \1(1). I,. t I ll')Oa I: [{eeherd1':' ,I II la 1:11111': h<:rp<:I ..logiqlle dcs II.:, dc lIol'11co d d.: l'al'III'llI. Nou\ /\rdl. 'IllS lI i~1 al .. Pan , ..,(2): II S- I(,l' . (I ~')()b) : Diagnose" d 'e'pl'ec, 11I"I\ l'lIc., d<: ICpl"<.:' <:1 dc hatraclcns dc, ilc, Bornco cl 1',lIa\l'all. I t' Nailiralistc. Pans (Scr. 2). 12: 1-14 - 1'15. ( IX'12): "'nul'dle eonlrihllllnn a Ia 1:1IIn, h<:IP':llIllIgll\UC de: 1I11111l:1I. M<':I11. So<: 10111. Francc. 5: 1'10-20(,. M'"{(;I , F.C. ()'n}) ~nak<:" orthe sulll:11111ly SihYl1opll1lnae. LII:1YCttC, I.Oll isi;l11a. 1.llli\',rslly "I' Sl'lIlh\\·';'lcrn
I.OIIIStalla, I'h.l). ths,erlall,'n : 1-2(,0. MOKI, A. (I')')}): A note nn Ihe sldl'wliltilng IOl'011WIIOll1l1 111'0 cnlllhrid snakcs. (JI,llIslh"IHl/'I" /I'pl( t/ and 1'.'1'11 "m{'/I"do" lIIt/cml,.1 rhc Sm\....:. 251I ): (,7 70. *M,ml', J. & L L. DII I\\\N (IX~5) . (onllihuIIOII' lolh<.: natllral 11Islory oJ' 1.all'alll\ Vl'rh. Hala\'13 C'cllonhl'h .. 1(.: 9(,-'>7 . MI Sli R'. ('..I.M . ( 19K,): '1:IXOlltllll) ol'lIlc gellus /)I'lI('fI L. (Ag;lIl1ldao.:, I aO:l1ll1a. Reptilia). Ion\. \ hh .. I cllkn. 1'19: 1- 120. NIHl'\s. I'.M .. /\ S. 1"1'11,. 11.-1) \0 % 8.:.1. CIIIO"I,,,,,1)\1 ",-\,11(1) (I')I)~) : :Vloq)hologica\. heha\'lIlI:!I , alld g<.:IlCIIC Jl\'crgencl' of 'ympatJ It: Ilwrpholypcs of Ihe trct: fi'og I'tJi1YJ(,(/lI/I'., 11'1/( 01111 ,. ,111.\ III 1'c'llilNlJar MalaY'>I:I . I krpctologica. )4(2): 12l}- 142. NlJllll. (i.K (11)26): All allaly"" 01 Iho.: rl'l1larkahJ..: case' or di~l ribulioll al110llg Ihe ,unphlhI3. wllh dl'-,cnpIIOIl' of Ill' II gl~ncra. Ant rv1L1~. f\{)\ .. N~", ' Yt)1k (~12) : 1-2--1 . OIIII ·H.i\ . &/\ . 1)1110"( I'!'J'» ): fhl'idl'IlIYlIII:'IIl('/il'glllS\{/ J,!.l'idCII.II,,111t'1 A'\111 I(~I''J, I<) I('I /\mpillhia, R:1IIIda<.:) , wllh COllllnCl lis Oil SOIlIC ;1'peo.:ls 01' st;lllslical SUppl'rI 10 la\ OIlOII1Y. It'ologica So.:lpla , 2X : 2(,')- ~71) . 01 \. II. & T. II IKII1' (I'IXX) : /\ nCII ~JlCo.: l <:' 01' I <'I'ltlo"a( '.1 ,111." (S:lllll:l (ic\..\..,ullliat:) I'rn11l Sahah. Malv,ia Cop<: la. W'I',hln!:!toll. (.I) : (,I (,-(,21 . - (19'1 I): ra\OIlO1llIC le\(c\\ .. I Ih,' II/ani, nf Ihe gCIll" ( 'lIloil's (I \ II K I X17 (Aganmlao.: Squamata) Irnlll Sabah. I\ laJaysla. Tropieal/no\..4 . 179- 192. - (19961: I hl' s,:colld '(1l''':1I11CII til' ( 'III,,/c '. , ~1I111"""II " I'I,' de (iRIJ' (Squal11ala ' !\ ganudal') 1'1'1'111 Sahah. MalaY'la . II ith COI11I11<:IlI, ollth.: la\tlIlOl1l1t: , lalli' oflit<.: 'I'':<:I<:S J. l1crp .• .'0(2): 2X~-2L) I. (2000) ·1/,lulllio/I' 1111,'11111 (Ill JII M,. 1930). :111111101 SYIInnYl11nf..l.ol'Jlll/tll\'anIIlIlIl11l .III 1)/ . I~I)'\)(Squ'1IUal ' l' ·\ gal1l1da.:1. ( ·lIIT. I krp '101 I'l( I) ' 11 - 14 . 011, 11.. T. 1I 11-.1I1I. l\l Ko'\ LX 1. II I1)\KI119X'I). 111l1lSII," l1est ~itl' of a s.:illcid I,/:lld .\i,/i"II(){IIII1'1,IIII' !'1IIt/hlllli "/1.1'11 Ir,l1l1 . ·ab:lh. \i1alaY~I:1. Ill'll). Rl'l. 20(.n .\X-.N . o I I II.. S. SI 'J( ,OKII 8.: I 11 11-111 \ ( Illl)(» . II, n '\.:\1 SP":CIl" ,)\" 1.III"'Ir''tIlIl'lI'· (Reptl"'I ' (,o.:\..kollldal') lrom Ilorn.:n. '-'01'<:1;1. W,I'hlllgll'Il, (2 ): 4.1.1 - 13'). r IRld R. II. \\'. ( 11)2~) ' The bn:\ ICIP ll id hog, "i' Ih,' gClll1' ..til< m/ll ·11I ;\1111 . Mag. Nal 11 1,1.. I lllldl111. 10. ~l'r. 2. -173-11)'1, - 119 .1-+). A m'''lIIgi':tph 01' Ih e frogs III Iho.: I'a nilly 1\· l1 cr~,hy l!d ; l e . I ll11(loll (Hi'll Mu,~unl Trmtl'l"). \ iii I 20~ pp.
-113
Bibli() ~ raph y
*I' IKKI '. H.S , ItS HI ,~ I" 8.: .I. ,1. BI ,'~I ,\, (I'N2 ): '1hc' piant ' ,,1' \ ·ll.unt Klnahalu I I'crn, anti I'crn "lIlcs KL'w {1{,IY;llllnt"III C (;aldclI') *1' \\''11. J.. ( ' ,1 FI<\\1, ,, 8.: K 1'11111 11'1 " (19X5). i\ lickl guide II' lhe 111 a 111 Illa Is ,,1'1I,)IllCn KII:r 1:t 1.lImplil (Sahah Soc. 8< W,"ld \Vlidll k hllld Mal;I Y~ laJ , ' I'I \,DI I III I". 11M <'<: Il.! ' l'11 1'1''1 ( 1'n2 ): ;\ .I.lxll ngic,tl l'.\ IK'lilllllll to \1t. 1-:11I:lh.tll l .I , I cd. 1\1alay, SI. Mus .. Sillg.tPOIC. 17 I-)l'i I' IIII~\, LI, ( I 'IX I) ' i) rc i hL' mL"r~ c ll s\\'e ri c t\ac1l/lIchl cn: Nct/ pylhnn, Nash," n- I L'gllall II l1d (i u ph lT-Sdl i ltl~ r(i l e , 1)", I\qll:IrIUIII , 15 (1 3'1)' \.3 -37. PI II R'. \\ . I I X(.-1I: 1') hL'1 nL' lIc' .-\l11plllh icll ( 'i il ,hlllw l/ /( " ", 7if"""/", , I ,/II"II/Il II/ JI(I\ , ()g lll o d llll ). - t\ lh T . J.. prCII\11\~ ; ld WIS'. llL- dlli 27 1-27(•. I PI (I X(.7 ): Ilcrpclt.llIglscllL' ,Ili /CIL MllCr. Ki'illig i. " J..:ill. Wi" Ik IIIl I. 11- 17 ( I X7 1). i'lhl'r IIL'Ul' RL' pli hl'lI au, ()st ,iIi'i c~ lind Sa rawak (Humeo), 111I"/,iigllCh all' del' S;lInll1ll1ng dL" Ilrn Milrlilli' .I . 1)," "I / 11 (i 'lilia , Mbcr, Kiini g l Akad. W i,~ , Ikdill: 56(,-SXI. (I X72 1: i'lhL'rs icht del 1'0 11 dCIi I k rn: II , t\·h c. (i , [)"na IIl1d 1)1'. O. Ikl'l'al'l III Sarawak aliI' n Onl co ,'on I X(,) b l~ I X(.X gl'sall1 l11l'hL'1I ;\ll1phlbiell. I\nn. MilS, Cil , sl,lr. nat. (,L'II01a, 1. 2 7 ....j ~ , pk 2-6, ' 1'1111111'\,1\ & 1\ I I~II' ( I')XX): Pllcber-pl ant s o rl.a~1 Malay" a .lIld Iinl llL'1 ' a(lI re 'Vlalays iana. Kllala I Uillplir. 1.>(..\) : ~ · ,n 1'1" ". I & II ( i 1)1)\\11" , (2000): " ( '"//lh, '/''' lerSII ' ·· I )II 'fI \.'~· , \ IL'gl t.' U:lhk lallg h: or l1:lIll":S. I krp. R l '\ , . J I (.IL II(,- I ,X 1" >1'1. ( ,II 8:. S,II , 1" 11'1 (1'1.1.\): 1\ silidy of lhc grecl1 I"i I iper' ",' , ,,"lhl.:,,, tclll r\"" ;1I1l1 1.,1aiay\ ia . l'o1l1l11llnl y Itic:lltliicd ;1' 7i 'III /('H "'/l/ll,1 .!!. I'IIIII ; II I ·II \ I Shaw l, wilh dc\criplioll nfa Ill' W~pcc l e~ frllllll\:lI illsular Ilid ia, AlllcT, MilS 'Jo'llatc, . (, ~ O : 1- 12, R \ 11 , j) U' 8;, 1>, 1 Y I"', 119(12): Phylogcnct ic systl.'lIIall C' 'lf Parl' lnaL' (Serpelll ,) 01 SO IO"l.'''SICrn ASIa alld ad Fll'l.' III ,,1.llId, "',," re lallo n,hlp bl'II\C'l.'1l il and Ihl' gcn lllg\ cliall gl's. [In (hIl IC\l'. I' ngli sh '1IlI1m:!ry [. I\ cla /00 1. Sinic:I. IX( ~ I' 1 ,II_I '> 0 R \"III'. S,M,A 8:. ( ' II DII,f'-(, ( Il)t)l) : Ohscrva rllon, on J (1/ '(1 /1/ /\ ,(til'll/II /' kctling on ()/tgot!1I1/ "('/11 //1 /('11//1," 11;lIl1a lll),<1I1. t\1ad ra'. 2-1( I ): -1 X 'l, R \\~ II " 1'1 . .I .ll.. ( 1'17 '; ): (i l'og l~ lplll ca l variallon, IIlcillding :In l' I'olllilllllary Irl.:lld. III 1" {I/ l/l/lo t!1'IIa v/(,,1 /111/' '<'/'11 / ,'11111,1 (lIt III • I x27) ( IIOIglllal·. 111lIl1alop", ldac. Scrpc' nl es), \'ldC'll ' ~ Mcddl dan ~ k , n:ltllrh. 1·,)!'clI .. 13X : 39-().j , Ril d I\ \ S'. l. /It I·.. KI<'\\1I ~ (!')XI): Zur Sys ll'l11allk der !! riilll' ll ( ;ruhl' llott l' l'Il de l' (;aUlIn g '/1 '; /1/('/'( ' \'11/,11" (ScrpclliL" . ( 'rot ;"ltlal.: ), RCI Sui"c Z(lol .. (i cnl'vl.:. XXI I) : 11,1-211.\ RlI lIl \. II N (I')()(' )' ( '1111011' nL' ~ lln g pl acc of ,\ '/II/ flfl" ~)( '/(}/II/('tlfll" . .I. Sll , Hr, 1{(lY 1\\. Soc,'sillgaporc" 4(,: 263. RII 111 . 1-: , ( II)'):! ) n ioaJ..lIstlk der Rulgcl11cin'chaf1cn e illes
,,\14
Ill a la ys lsc li cn l'i c ll a lldrcgc l1\\ ;"dcs Verh , Dcut sc h. In,,1 (i e, .. SlUll g:1I1 , X(I{ I ). 2(,'). RU)III1 'IC :V1.I\ . ( I'),)X): Slatli s. pnplilatinn biol,)gy al1(lc'llllser\'atl () n of thc WallT Monito r ( ' (/I'{ II/II" " II!> 'a l"}'). lhe Rcll ~ ul:llcti Pylholl (/ ' I '/h u /I I'e/ic 11/11/11,1 ). anti the IIlood 1'}1hon (1'1 1/11 !II " III'IIIS ) in SlIll1all:! alilllP,). xi\' I 1,,\1 pp, (I X..\ XI: De,cripllnlls dl' ph"il'lIrs c, pcce, 1I01l1 d k , du genre / .II/Illy m s, 1-11.1 dr. Dlerk.. AII,,(ndalll. I' 4-<>, ( I XSX ), Ilandiciding Illl de bc'od i..'ning der dierkllndc. Brl.'da. KO ll i llJ..l~k e 'vl ilita lll' I\eaticlill c, X\ I 6JO pp, S. II~( II )I, I\ 1\ . (1'1 76): I-'rsl- 'Jach<:lIchl dc, l.ip fd fi'o' l'he, I/t';!ol'hn '" /1(/.\/1/(/ . . alam'1I1 dra. l'rallkfll11 'V1. 12(2 ): -;5-6)<, S'" 1\ IIIII~ . J (i , (I XOI): Il i\lO ri ac' i\mplllbiorlll1l natura"s <:1 lill'rariac , ra ~c i c ulll;' ~cc llndll ~ cl1lllilieliS l'1'lll'odilos, SCII IL:OS. ( 'h,lI11a C\
"1'
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Il ihliography
Lolkgil. dig.:ss JI. dcscrip' " , l't u.:pingl'ndul11 curavli. 'Iomus I. Amstclaetiami I /\lllstLrdam]. .Ia n, soI HO\Vaes he rg i o~. J. Wl' tstClllllIll & ('UI. Smith: (12). 1- 17X, pI. I- I II. - (I TI5) : Lut.:upielissilili rcr Ull1 natural 111m thc~allri accurala desc riptio. I:t irlllllhu ~ ~Ullli c lns i ssi llllS c\prl's"o, pCI' univers:lm physl':cs histuria ill. Opus. cui , III hoc re'rUIll gencre, 11l1l1ulll IXIi cX lstil. I·. x tnto Icrrantm llrbl' cnllegit, di gl" , JI , dl'scrips ii. ct d':J1 lngc nuulll curavil . TlllllUS II. /\m~tl'laed:lI11i I Amsleruam) . .Ianssllnlo\\'ae,hcrgios . .I. Wl't StClIlllill &. ( iu l. Smilh ' I ,::, I, I- IS-I. 1'1. 1- 11-1 . Sill I H lIm, R. (1'1(11 a I: A itst nr lhe reptiiL-" or i!orncll. .I . Str. Br. Roy. As. SOL.. SlIIga porc. JS : -I .l - ()~ . (19() I b): Onl wo new ~nakes fn)m Bunten. /\111 1. 1\'lag. Nal. Ilisl.. Lllncion. 7. SCI'. X: 51 (Vii 7. ( 19(5): ;\ nc\\ 11I.: lrd and :1 nl'\\' rrog from liornc'o. !\nn. !'vlag. :--.Jal. llisl., Londllli. 7. Scr. 15: 20X-2 III. - *( 1'116): A nalilralist III Ilnrneo Lonuon n. FI ,hcl 1:llwi11l .td .). Sill". C . K. (199-1) : OhSC rt':1I1 0m, in tltc cop ul atory 1)L'ha""IIi' o r C;OIl.1{}.\(lIIUI f/Ilc<'plltll/lll/ (UOII I. :VlaIayan Nallll~ i1i ~1. ·IX (2 .'):-1 -5. SI()I\l I\;-.KI. .l.B .. J. llm l\ ll\ k R. L.-w. ~(), (2 ()OII: rhc phylogem:llc rclatill11 ,hi p~ orA,Jan Coral Snakc, (I lapidac: e,i/lioIJIt;, a11d :\llIIic(}m) based on lllorpllt)logica l and nlllk<':l1lar c h:tr:u:: l~rs . 1krp.:tologlca :'7(2): 2J3-J ·15 S\lI 'I,11 \ , '. (I ')J I) ' otc' on the (il:tnt Frog. NIIIW 1II,,,.,.odoll . HIIII. Ranlc~ Mus .. Singapur.:, S: 5'J-6::'. *S~III1I, .l.M .I3. (l'IXIl!: Thc \cgctat ioll ort hc sU lllmit 10llC of MllUllt Kinahalu ':w Pity")1.. X-I : Q7-:,n. SMIIII,i..i\. & I. Slim, (l<)l)X): ()~slTlpt i on ofa nel\ ~rl\.:ci.:s III' Cr//l ulml,ili" (S':I'PCllt CS : ( yl ll1drnphilll:t .:) rWII1 Yamdcn:t 1, lal1(l, T:lIl11l1bar An:hipdagll. Indunesla . Rallks Illill. 1.0"1., SIIl!!a pl)I'l'. -16(2): -II'J-· I ~-I . S\IIIII, IVI .. -\ . ( 1'123):;\ r~\·i<.:\\ orthe'll/:mls or lit.: g~ nu s li"l'id"l,ilorlt.\ on Ihe Asiatic: vl;lInland. i'wt' . 1,',01. SOl'. 1.,)l1doll' 77:,-7XI. 11 92-1): ,.:\\ Ircl'- Ii'lIgs li'OIlI Inuo-CllIlla and lite Malay Pcnlllsula. I'roc Zool. Soc Lonuhn:)1.lon. Re'l'i.llndllll ( 1:lvIilia and Ailiphihla. Vol. III. Scrpcnlt', lor & I·ranci.,) "I I :iX3 pp. *S\I\ 11111 ', H.I.. ( I'16XI. rhc hilds O( HOIIII'O hhnhur).'1t
(Oll\'cr & Boyd!. *SI \1 ' 1' , (' .(j .G.J . \ \ 'l (1'16-1): Planl gco)1.ra phy 01 the mOllnlai" !lora or Mt Kinab:111I Pn><.: i{ov. SOl: .. SCI'. B, 161 : 7-3X. *S 1\1'1, O. ( I X'J-I): On the fl ora or viI. 1-.11I:tba lu, NOl1h 13 On1 l'tl Tr; lIb . Linn. Soc. Lllllcltln. [lot. , .j . (t'I-2(-, I. * ~IIII\, . (I 'n~) : ('ollllCtcn inl wC\llichL:1I tvttlall'chcn i\rchip.:1 '1hL: Ilagllt:. B"~h\ll. I 011 lion (JIIII~ I'lIhl. ) '> ILI N"\( 1I'lie I'. ( I XK I): Cibcr cinc IICUC I'ylhollart (l', Ilinl/ 8n 'I/('II.\I(,IIII) ;JUS BllrllCO. SIll . Ikr. Ka l' Akad. \V"s .. \\/Icn, /\bl.l, X2 : 2(,7-:~hX . S II J \II<" R, L. ( 19112): l'II 'cll": lI(l1/ h"ldl, a IIC\\ namc (01 I' 1I1111/l/ltW('I JIIlIItIIll. i'roc BI< II. SOl'., Washlll!!ton, I:;' 37. - (1')07): I k'l)e:tolngy 01' Japan alld adpl'cnt tC'ITllnry Hull. U. S. Nat. \1I1s .. SX' " I 577 ~ .. - (l'n2): Li,,1 of ,na ~ L:S co llect ed III Ilul u11gan, north<.:a,t BOIllCll hy larl l.ulllholv, 1'11-1 ~y t. tI.'lag. Natur\ . Kn.,lialla, 60: 77-X-I *St. JClII 'l. R.W. (IX(12) ' I Iii: 1I1lhc loresl' IIllhe hlr hl't. Vo l. I -I.ontioll (Smith 1·ldcr & Co.). SICltIC'I-\. r. (IX 70): ()Ih~r\alllil " ,111 'Ol1le Indlall ;lI1d Mabyan arnplllbHl :tnti reptiles. .1. i\"at. SIlL Ikll)1.:l1. ( ·aiculta. 19: I J-I-22X. SII II BIN." R. ( 1'19 1): i\ <.: h t:c~lis l "I' till' s lIa~ es of lhlrm:o. i{ a llle' 111111. LOllI.. Sing:lp,"l' 3'J(2 1: 12.' -362. (1'19-1): A chcci-, Ii ,t of thl' ' "a~ c, 01' Hlll1ll.W Add<.:na ami wrrigcllda . Rallk, Hull /1)('1. , SIII).'aporc. -12(-1): IJJ 1-'1.16. SIlIII"'I,. R. 8:. R.I . 11\(.1 ~ (III'IX,. Addilil1l1all<.:cord'ollt\\o rare "lakes rn Hll IJo rn cll. wilh Ih l' cO lllinn ;ltll1ll 0 1 I rillll'r('slIrm IIllIlclIlllli LCl\ 1~II' . I a, a d"tllKI spccie,. Ranks Llull. 1.001.. SllIgapnre ·lhI 21. 325- j2K (I')()'»): A lil' lti gllllie 10 IhL: , nakcs 01 BOrtlC'II. KIll a Klnabalu (Nal. lI isl. Pub!. ), 2~ -I pp. SII I HI I\( ., R.B & /\ . \VIII\(, (~(I()()I ' ,\ Ill'\\ ' PCC IC'II( I'm!!, I 'II il(/II I 11.\ ('ITlllml, IlI/Il""III.\ ( I ~ hacopllllndac) rrn l11 slllltll\wslcrn Sabah. MalaY'I:!. i{:lJ'lk, iJull 1.001.. Slligapnrc. -IK(2): 2').\-21)6. '1 ", F.L. (19').1): C h cc ~I"1 01 h/a rds of ,>abah. Kot,1 K lIlabaiu (Sahah Par~ ' I nNci.:s), I X PI' '1 \\IClI<. I ~ . II. (1920): Plllitppllll' :1I 11 phll1l:1 I'hil. J. SCI., r--,·Ian ila. 16(.\): 2 15-.\5 7. (1'I_2a) /\ddililln~ In thc he:rpctologll'al iaulla o l' lhc PhilippinL'lslalld,. II. I'hil.l . S<':I., Malllla, 2 1: 25711).\ . - (1922h): The ,nakc" o(' lhe I'hlhpPllll' 1, 1;111(1-. 'vI:llllla (Burc':tll nf i'nllllllg), 312 pp. (1960): ()l1lh.: cilc<.: lhan 'pc'CtC,,, II/h 1'01,1" \ II/III/II, 11I'1iJ/\' and !c'/l//mll,II/,' gllllil/lI\·11.\ \\ lilt d<':'Lriptll1l1, 01' rc I:! tcd ' p<.'c i.:,. Kans t llll\ . Sl'1. Hull., L: l\\r~IlCl' , .jl)(-I): .17120. - ( I'J(5): 1111(,5b I: I\cw 1\ 'I"tl<': and i\ nIl'an GI<:c illans \\'llh rcd.;,c rtI'IIIlIb orccll:lilllltitcr ' PI'I'I~' . Kans 1 /111\ . Sl'l Hull.. I.a\\' n.:n<.:l', -1(, (" I: 25J-JO::!.
-11 5
Illhllngraphy
II')/>XI: rhc' ( ', I~':lhall "l'lk \1 "rid "1.1\011<)1111,1':111:\\ , I :1\\1~lIl'l' II 1I11"~ lly "I K~UI";I~ i'1l"S), X-IX I'P I \\I<> K, 1', 11 & R I .. I' IIU I 119';X): ( IHllrihullon 10 Ihl: herl'elolog\ 01 I h:ld,lIId 1\..ln, \11111 :-'1'I Bull, I .l1\lI:IKI:, 1)0;\.1) ItlJJ - IIXl). i 111 \, 11.1). (I ')~()I I hI: SUlllia SII.:II: SOUlh l;a,,1 ASia . , (iI:IHlI()rphlll.. ~-I ' -10" .. ):'7 11<11 1"' \1 1. I (I')~O) : IlhcT dic I .:bl'I""o,;"..: uml Plkgl: dc, N.: I /p)'lh\lll~ 1'1 dlill/ 1<'11<'1111111/1 (S, "" 11>11<) 11\:11>1:10laulI:I. W"'"I,ladl. 2 (.) . 2(,·J I. 1' 1111 I>i. J.J (I X.\X) , (lb,dkall1>11 dl:r lIalr~.... hi.:r, 11111 Ikrl,,""achllgulI1,! d<':1 1,>"liell Thlelc dl"" "1 \llIhl:dlln~ 111:1 Rq>III";1I kill . SIl.... SCI. al \0,;1I1:1I,ill:1.2 : I 'N. I 111 11''', \I \\ I, 1111:'-1). '\ok' olll\ l.t1ayan replilc'" no . i . ilull Ram", Mu" . \1I1)!aporc. 2:'> . 107-11 7 (19X ~) 1hc' '1Iak.:, 1>1 MalaY:1 I Ird ,'(1111011) Si1lgaplll<.' (SlIIgaporc N:lllllllal 1'111111:,",1. I <>7 PI' I '< ll11<11 f 'lI>. (i 8:,\ I SII~I ,.,,, (Il)I)()I ..,\l'I;l 'S1lio,;a li ' l\llll l'ylhllllS (:-'cl")1':1I1",. I'vlhonlllae:). J 101>1. , l,>ndon. 221 : ';(>"(>0.1 . \ '1NI I', !'vi (1<)1)1) l'hyl,>)!,,"':lic SllI(h.:, ,III r:IIHlld li'o)!, 1/\l11l'llIl1la . /\lIl1la) \llIh :1 d"':lI,sll>1I (>1' Ihe: llngin and nolliliun oJ' 111<.' \ ":lkhl.lW l'Iad.:s III Mad.!ga"·:lr L:1II'lIhll,h.:d I'h1) Ihe,is, 1IIlI\'':ISI1)' 01' BOIIlI V," ,I I , (i (I II'))) I i! '/ICIt'l'l"/,h,, ,'/I '/cll/I' (t ·, 'I") 11':11 fiil die: I ,luna ilOIIlI:"'. 1\1111. l",,1. Ill" Ikl hll , 70(1): 1-17- 1-11) I ~()tl()) · /.1\1" Ik'"I11I11III11,! dn lInl.:r:lrt"n \':I die I cbl'nuliirhllll)! \,111 HI/ligan" 1[,,,'1, " 'I'" "a/l/el/l'I,1 I. Il\ IRllk;I ·. 19.1i\ ISI''l1c:nICs, I I"pllla.:) Sauna. Ikrlll1 . 1911) 13- 1(>. \'Ok l'. 11." & IU 1'1, ,11( (11)9~) ' hog ,Ibuntla ncc all)l1g , lrC:II11S III 1I0n1C,lII lon;,ls <. \lIl"'I"\ . Bllll., 'II \) 07'I-MG , W \(,11 H, J. (I X\O). Nalllrlll'hl'~ S),lelll d"l \Illpillblc'n, mll \\lr;III!!.:hl'lIdcr t 'lasslliclIhlll d.:r Si iu~dlllo,;rl' und \ i\gd IJ to. (inlt:I''l:hc nlle:llh.tlldIIUlg). \I I l'i-l PI' \\ \II . I 11'12,) \ halld-li"l or Ihl: Sll
I'"''
(,32 '\\' \I I " I. A R II Sh<) . I he \ ,I;day ,\rdllpe:lagn LOIIdOli (M:le Itll,lIl & C" ). W 'II " II. V & /\ .M II \I " ' 111)%): Oil Ih e Idc:nllly and 'lalli' of' \/I/lli/n ,, 'J/lIII//' 1/11 1'1 "1('. I ){(,2 (Scrpclllc~" ( 'nlublldac:) 11 ;lllladry,ttl. 1Jura,. 21 . 1,-1 X \.\ \II \1 II. V & R. t;1 " 1111 H (I ,-)9X): VI 'CITa I ,lIlaIOm) "I' 111.: 101101 'sian 'na~c' gl:IIU' \·('/If/I'/I/c!im/. inclmhllg a '!a,lIsll, anal~"" allli alillealion II> a nCII 1:111111)' (:-'I:'l1c:nles: :\c'lIl>pllllllld",') I\ll1plllhla-R.:pllha. L.:idcn 19: JX'i-1111 \.\ \ \II~ 'I I I ~, H (I tJOO): I til' l\..:nl1ll1" d..:d ialtulIg Om", L. ,\hll ller I\lus . II I:I~ . \i'II~..:rJ.. .. i>resd':I1.9(J): 1-10. \\ II I II, I\. .IU I . IIl)XX) . SlIa".:s <.rlhc ()lIenl ' :r ehc.:" 11,1.
-116
I\lalahar, 1' lOtlll,1 (R"hcl1 I' Kn ..');.:r PlIhl ('0) 1-\11 I IX.I pp \\ 1\( \ 11'111. II (1<)(, 7 )' I "Ic dt.:1 l"/CIlIe'II .!\lllphiblCII IIlId R"plilicn : ,\ g: III1HI:r ' n .IS TIC'IT<.'lc'h, Ikrl ill. :-:1>. XII I 1'2 7 PI' \\'I I(N IIL I. (1i\L)2) . elkl ':1111.' 1..1<.'111<.' ('nlle'cll"" \(111 Rcpltll en lind Halr,Khlc'1ll \ on l\Jias. .I:illlc.,llCr. Ahh 11:11111"\\ VCT \1a!:!dl'bllr~ : 2-1X-:~:'-I . II X')(,). /-\\l'lkl Ikill; l!! lUI' I krp"'II>ln);lc dCI IlIdo:tusllall~d)(; 11 R"~IOII Verh. 1.\\01. HoI. (io,;, WI"1I.4(,. h-:'
11'!lO). \ IlCl 1I0001Il' IId.:r ,,~IICIlC Rep"II':1I d.:~ lIalurhi~ IllIISI:II':1I 1\111'~11Ills III 11:llnhlllg.. II I IlkchSl.'1I .Ih. Ilal11h. \\i,s. An .. \.. '2 7. :! I'killcli . I (, *\\'11111111 \I). J. II X'I3) '1hc' .:"plnralion 01' K III,II>al". Nonh Homc" Londol1 (,IIIIICI 8:. .1,,0,;"'011). 1.\ 7 pp. *\\ 1111 \lORI. I.e 119X-I J' ('I'lll'"'a l ralll jlllC,1 Ill' llil: Far 1-:1'1 I. olldoll (Oxl'mllJllI\ . Pres'). "Ihl ) (1')X7 ): 1!logc",lglaplll..:al cH,lull\l\l of lhl' M:llay .!\rchlp<.'l:igll (hlmd (CIlllglc'_ !!.:salllllldl auf ('illl:l Re iSt.: UIII dl~ Erd ..'. \'''n 1)1 1'..1 ,1' I\kycl1. sl.:henk ,\bhalldlul1g. -,\l11phll1";11 V.:rh . Kals. I.cop. o,;al'lli. A"ad. Nalluf.• Iialle 17( I,. 1)0;5 2(,X. \VII II \ IS. K ,L. & \' \-\ "I "" (I 'IX'!). - 'Ina"cs ol'llic world "nl 1. Sy"oP'" of ""I"': g"'I1<.'lic nal11e,. I\lalabar. 1' 1011(1:1 (R,>lx:n I t-..nl'ger). 1111 - 23·1 pp. Will I. S. 119.1(,) I{C\ ISlIlIl dt.:1 Untc:rg:llllll1g N"III iI/I"Onl\ 111111. Rallk, 1\111'.. Sl\Ig.lporc, I '2 ' 1:'7-21 7 \.\"" (,. i\ (1')1)-1). P"plilalhlli t.:cnlo/!) pi aillphihians III dilll:rl'l1l alllindl.'s Ill' K illahal" Park . :-'al>ali VltIS . .I .. 112): 21)-3N . "" WON' ,. 1\..:"'1. & J\ I 11'1" (1'ds. )( I<)'J(,): t-.. III:lhal" - SlIIlIIIllll)j IIl1rn,'ll. t\ rc\ \Sed and .:\palllkd hhllllll 1\.01:1 KlIlJhalli (S.lhah Soc. "" S:lhah Pario.s) ''''Ialrc' ,,>bra ( \ellel IIl1jel) 'I'':CI':, I:olllpk\ ill slIulh ...."1 '\ '13: rcll,lbrll1) and randolll r<.'s:lInpling. 111('1. 1. I Inn ')oc, l nlld<>n . .II,: .19 1--10'1. Y ' \lI,. n .-T. ( I I)l) II ' Ph) IngclIl:"" .,ysl"IIl:Jlics of Ihe 11I1iI/II/1I glnlll' 1>1' railid Irllgs 01' SOlllhca,lt.:nI ,\'>Ia and Ihe (,rt.:akr StilltJa I,lallds. I:iddiana /:(101 .. ( 'hiL'agll, IlC:W SCI'. (..I : Iii I -121'1' '\ "" ,. II S & I. R \~II \ (Il)X 7): ('onqll",l or Ihe land ' Di 11:1:1 d.:\doplllCIlIIll a I\I:1ia) '1:111 'JIII/al/III\ Tr<.'e I·rng . Nalurc' I\labY":lII:I, KlI;11:! Lurnpur, U( -I): -1-7 '\ ()K~I. ( ' I). (19X3) ' :-'ur\,I\'al nrCll1hl)~lS :lmllan'at: "l'lhc Img 1'"Ill'ec!"lt '. , " '1/''1111/1 'IiiI' III Malay,ia. .I 11<.:'11.. 171-'): '2.1 ,~-2-11. III' ,II H. I. 8;, W. 131>1/\11 (1')\)61 ' l.bc:r das Ikulc:"pC:"lnllll \'on IO/'{lIlIl.' cllIIlI"1I11I ISI III 1-1 ,11 . I x.~I)) . S~I:lIl1alldra, Rhclnhach. I:!IJ)· 20.1-21 II.
1',,"
Indc.\
Index B ahho//I, I"'I'/ohm, frlllln 'II
""ICI//" , ""lfllI/lI
, INtlh,'" holio"/I'I/.\ ha/IiI//el I'll III
hlihot/l'Jrlllll hllhod4 ' lrllJII . (;Oll,~" \ IO\tlJlltl
a/lilli,',
) ("'11,1:,1'/"'011111 ha/ioi/""'/11n
f)I'(/('" 21~
, lltO"/III/11 {,( 111t/"lillell/II ,
) I )c1l"1''''"I''' I I .wlt/"IIII<'alw
lIlhll/II,I'l '1/\ , /."1 '"11'''1,111,1 IIlln'di, ,\i,h<'lIiJ1l1'J/ 1"111,1
,·1-1 2)U
)
also In/{'rllll/111 IlIl' l 'I,1
) 'l / el';,'/lIp'1I1 '"
, tlll o/"I" 11111"""1'01(111111"
, 1111"/"1" ", /I 'HI 'IIIIUI/111111 ~ /111/(1 ('(/I 'i/I'IIIIIIIIIIIIII , 11110/"1" jCl'h,," , ) ai,,, :\/t 'I';,I/O,l:(,/11 :1 :\/CT/\lt)S.!..c 'nrs H 1II/I ,lre'{/lh
. IIII()/o/,,' J... III0hohll'lI\'j, . -t . rmolu/}\ (W,'/III()( '11£'1111 ,\ ~
,\/('ri\I(}~(,"\
\ "il1(/h{lIIl(,J/\i,~
'\1('1'1' I01!..! '11\ ', O'phllfJt'I1l'I1lIS
<11111 "'''lwlwlIllI , :\ " '1'''/ " ,!.!I '1/1 ' I
1-19
30:" 107
. tJIII '/If('\1I1<1 \111"(/1\0"-<'1/\/\.
,; . 1111/ 1/1/('\111(/ ,\'orlll\'tJ( '( '/I\('
, IIIII,fri,H '. '/lIs IJeI'<'
R/w , "1'''01'1/1 20 I
111111",/"111 , /I"'glI d<'llllrol,llll(l J 10 , '" ,1'1111;11 61
,
17
glll/1( '1 6~
.lOt)
""1I0,!OIl
J 12 1 1"
IU"Indea lIigrU'c,/),\'
Ho;,!!."
'/('l/(lmllllll"
) /I(lIga '/('l/dml,/II/" "1111('( /"1/1
XO IJ I hUI'II,'''I/,I;,\', 1'''U,III/,hl'l I ' 2-17
11(I/,f,oll/('lI, 1,<'1,1"/,11/'\'11" />"/,/Wl'li.';S , .\Ii< mill '/a
,(,f,
hfll 'I/{ '('II.\iv ,
,)'/oli(": /..,O J('l
bon/t.'clls/\,.
Inllli'I"t ' \ltn/\
,1 7)
hurlll'll.\ls. (,'O}lO( '( 'I,IIIIIII \
2·11
Hot/UHI)\ grolllil/('II'
) 1,.1II[( '!l ',\IWU,' /JO/lt. 'UJ/'llll/ \'({IUlIII
"""Tillgll , 1<"'11<1 210; I HI'(I{'hYIllc/l' , ' '27 1 "rl.'/I('/ls/( 'III1 , l~l 'lh()lJ
/1/'/lIIc/lfJ{'l' llI
227
ri., 10 lelia
~2X
l
frlllli/II 'III (,7
iXtJ
I il/"l, ,,1,,/('1
IJnll" I"" d" !.'l/llh"III, '11I11 I.;I/(//>,t/I/('I/'"
/IIIII:,/'/'glllI 711 1"lIll 'l olIIlI 7~
/J,.Oll lltou·/a
'/}/II/t/i/<'I 7-1 . ln WJIIllI lee/oIH/s .
Hlljil ---7
also
I}I\UIIIII ,\/'1111111/1 "
fO Wl/lid g,/lihei
-4 ... 1:-;\)
,'flllI/lI;"'i,,' 125
/llgn S!. lIll1r1\ .
dilt'l:t!('1/.1
71>
11l.\/lI\/ItT
77
HII/" "'1,, '1 ) ai,,, HII/" 1II\I"'I"'/' /JII/n ltil'{}/'{'(I/IIS , > /Jill" ,!/lell!., 'II '
~26
, IIIII/II/iolil 11lI'"/a ,
7"
, 11"""lIlI/I ,\ (J/,/llIlti
IJu/o
U/~/iJ hi/Jun.-alll\" rliI '( 'I't!( '}I\
AI'/I'I:I'J,:ot/o ll 27() l 'il/l//11I1I 270
HII/o /''1'1''1'11,'
/JII/i, /1I"g/l/"IIS ,
"1//", IJrucll1 'I/,,'I" ,1 2T!. "I'm'"i , I ,"II/ll/,i/i/\' <)X III"OI )OI,/ti,\ 1>0I'HI.'('I/.\I\·,
I 'III It/IIIII \
HII/n
hi/Jornllll\. hll1fJI'OIIlI\
"I'IJ"//lj,('/i/a/II" HIli" "f'''UI'II,\' 2().j
tlltHIIIIIIIIII ,
3X(,
"I'"' 272 ,11111111;11 IlIliglll"(I
/1I",I!.lIn'
()rllll/a
( '''/''II/''/'/{/
hO"'1('( 'I1 ,\'\', S/('gu IlO/II,
//(11 '111'1111.1 ,'lOll
. III \'(mlll mlliltla
III( "I", ;
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111(,
l/l'goll"n ' l
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1IllIllIdilli"
/>,,111('1/1 i,l , "'''/''1'''1'1/111,1 120 1/(/"",1/.11,1, 1" '1,/olm" II"",, ,'I
1>"",,,1/1 i,l ,
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I iX
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sare /HO c ( '11\'( '
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('/1I/(1Il'1I",
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I
ab(l ( ', 1'11I'/,,('/d1l.l
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laUlt''''''
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nll,-iu ~72
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'27
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-l I X
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( \ rfur/ul'/l'hl,\' 1'(JlI,\oltl';'lI/\
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7
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2"\7
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/ ),-c./( n 0"\( 1/1"11\ 0"\( 111"11"
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Index
1)1'(1/ t) l/11111(I'W/tl\·( 'ltllll\ .
1)/'{/(()
'l'III"I"t '/{I\( It/Ill'
/Ol/gihlll'hll
f)ro ('(1 ." IJllnl'/em" ('(lntll/II.\
100 1I1lIl'IlIlIa 1I/'I1,IIIIlI 11/'0,1I11lI
-4
I')"{/I'IIIII,I
dlllil('III'I\ , Rh II/,"I,h 1)1'/1 I ' 20<)
l:'Iul,I",
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grahll\' I~ I I
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-4
t.'/'I/',' Il'Inl/at'lIi(l
) :\/(lII( Ol'll hil'lIgl//a le/rall/('IIIU
nl//" Ih"I"'.\s/. -4 ,lful /illl'll il/I, ·,IIIIU/" Ih, '/wIII
Jr.')
(;oIlYfJ.\o/IIl/
11X
1110
gUJlHllgellsl ', t t'IJ/ohrllc/llulII
111111111('/1/, , II/SOIlIiI 1>7
hilll:s.:.U/ll
JX(l
itw'/'/II(llli, RI/{{c rlI,itllrll' 21 ·1
1/(,lIIillu"~r/IlS
-4
Ihlu,\"hc'\ X2
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''''/IIip''ylllJ(lu ('~r/ll,'
1·11
'/l/ldl/( '/rllI.I
,()() 1,/l1l1l1la
2h2
fI,
( 'hlll','I'I1I1I
1.1 7
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,.J1I
IWllelmll/oltl ,\.)o
I I/)
Rana
1111" ('/,,11"11/1/111/
3(,0
III'UI,I , f'lIlIl
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1I..t"
gll/'llOIlI , 11('//Iidlll '/I 'itl,1 21>-1
I " '/'llde,111I hor/>"lIllo
gll/lIli, /(//((('''I,ItU/,1I1 212
IItI"retl/"
~SX
/l1/1It/0/((
mrHlal'l 'llIl\'
260
(;eo( 'IJI\ 'c!a 'I J(,II .t!.h 'f I \th 'l/g/e Ji (1,\ U
Ihl'll, 'o
) l'ulllwel"" '1 11'11, (l1/1I " 'al 1<'1' 1111""'111" I>lIl'hllllll 'lI , NIIIIII (/1111('11/1111"
,,,,,It 11//("0
I !t 'I(//'(/1I11 11111/'1111')/(/ 2S()
(,('IiA" 2(,()
'1" II
1, '/11 '01111"/111 (NIIIIII)
( ,halto/IUIU
) /((///(/ 1"1
(;
(;,·/iYrII
" "11111,/11 '1/1!(/"cI I ,It I I' 11 '/1111
1111 ill U(, /~\'dl'l/hl{/h(·"
IlIligill<'(/ , 111.11111111 I,.J III"
}
1/('(1,1(''''.1''' 3<)7 ,I/!IIIIISlI .llJ7 11111/1, I?II/III 1111
£lol,h" 333
1'011,,111 111 '1011""", -4 /" '0" '1111 '1 In 'IIIII/IS , I h
ro IIl/lIIIo/((
I (;"/"
It,/I/II
/I'" 'in'l''' hU"II'IISI\, /JllI/gunl,1 " X.J ,,/ill ('II 10
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2h-1
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21;;
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2 ()~
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,1
1;'II"rilllll~ lillll'I'II1I1II', /ir,lI'()
'V I',II,witll III/illS
)
/I('/i('()p,\II1I1('.\ /"{linll
F
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t)J
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<'1 '('/'i·II/. OII,!!.I"lol/ 3.Jl)
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1 ' .J
2·11 gn~It,()"" , ( 'lI/WI/"nO 3 1'1 glld1c·;. III,ol1ll1 6." grllllrlll , CiulIl/( '('pllll/ll.l
I !tc,·lm /111 '111,
!JIIIII(I(h"/'I.1
'I X
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Rill/III h(lIlT/I/!!}1 "1'('1'('111 /I/(itTrI,I'('eli,l , Rllo('ol,II",.,,1 21 ()
lilSI 'IUIIIS, IIlIlIgII 1'/1 " F('j('I'I'UI)'U I J.J
11'1
1I\:<'I '('I,h"III/II
grll"lIll ,I~ 1'/, ( '11111111111'111
II
1/t'U ,\('/JIl ',\ \'/11110'(/
({'lrafU ellia . :\/lHic oro
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J3 7
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".!!." ,
( 'I r/orillcll Itll
)
(;'111.\'01'" iI' ,17
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1;'11111('1 '," hill ..,.,
,tI", (;"/1/11 "'1,111111,,
/)(JI'I/('II'I'
I-.'/(/III"'"'IT/hm)"{/, -4 al,o 1:'1111,1", 1",,"IJlll'o/(/ F:llIl,h(' ('n 1111'111 '111' alsll F.III/,h" II." ()lilll 'II/(/
l:'Ial ,III,1 !!.rah()II ',I~ 1'1,
, (;lmnn '/J/wlll' hfWIlf.'Il.\I'
"(WI/C('II.'"
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JJ4
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el,hll/II.\
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etlll ar""/{/<" ;\Ieg()l,hn ',1 10'i UlIl'h ,' J.12
1~'I11I ',\' .'IIIJlO\t/
) also
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dllllleri!Ji, j lllWIIII 2lJ.J
11I1\'(1lill(,II/(I
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Gil/we('p//(/Ills 2·10
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I)/,(/(o ,'u lal/ \'
tlrill~l , L('/'/(iloim
nr\"(I/,/1I.1
• f)r", (I ('(lntll/II.\
(,'1J".~yl(J.'OIl'fI J.3~ f,,,ltod"lnllll /llIlwd"lnl//l
111'1111'0/1(/
""" 'i
1l1'i(//'(/II(/
!t"/II{f.\O ,
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"(/1.///1(//11
/r'''li • 11111111 /11,111"1(1
-7 /(11111/
111 'ft/nllll' IIIIt 'Hllh,t ,U /'t/It/\,ulle ' I1\I\
) ,tI'I'
",,",,'/1/\1
~ 1IIIIIItIll"{ '/r ','
l'O/CH 'OIl{'II,i,
Illlart/llu
IIIt 'ohartl'lI\"
) Rlllla
111'1'''0,.,('1/''''
419
Indn
~ NUllo \;,.:lIcll(l
,i!!Ju l/a
111'I0/(lIltl
~7'
11I( ' III\ '(.' IIItII1\1
1/\ /onJlltl JIII 'ohtlni'II\I,\ Ill 'p,i('{ll",," 2-15 AIiU.dud"t'JI\/,\ 24'"
1 '0 170
IlIl( 'nlh" ('i1{,IIII"ril/
Icll(}l'lIlil/(I/II.I, ,\I(IIlroll I."flid(//"/( '~r/lls
~h(,
':(,(1
/'lIJIUIU'n,I ,\
1-1-1
/( 'I""'/lIII\ . lilJlllflJl, '(I,'.I'
" 'flll,YOI,ili, 2, I 'I 1II1J1lf1thro/t.\ 21 t) Idlfl/du)/I\ --) j\j 'w/uruhdlolt IJI'/(,/'n'H\fl, \/11/1111'0 27(1 11I,t: e rIlIHI I iX hall/r'U\I\ I ~X 11I'...!..('II . {l"rlot/Oj ' (1 '/1/\ .2 ,,(, IH~("'1. elllllllllfis
t"I'IJlbrtl{'liellll XX
/1/<'1 ",1
11 '1"01>)'(1<,11 .. /10 1II101'( 'lgl
hulu,
halll\ /(JIJ ,t:, /(
halll.\
--7 /"'(1/(/ '01\ lid/lifo/"
rtf)
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,SftuJrO/x
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1.1'1'loi>f'III'/iilllll hl/h" 'II""
---7
I.("/I'o/woc '1tl/II1l .'~I'ill'il('
) al:-:o /'( '/1'01111, ,, 1II'IInd
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1<'1'10/'1'0('/111/111 ""' ,\I,lti
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I ''Illoh",/( /,illlil 11111111111111111 --. :11,,, 1l'{!llIhl'lll'lIilllll 1I/(1
l .e"lo/lIlax (17 fllt/nl/'
.I ,le l /hllllnl 11(/\100
' /'/IiIII/flll\ (11'110/0
.J(//~("II"(/ II1'.!,n/ah,." ,
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./"1,,11111"0 (Jj"I/alll
I/)/WIIIU{/\ "'-Jlllla
-,
11111,"1('11, I/oigl/ 312 il /\/H'IIl'J'
i)c"-
dnll,!!1 IOn
mil'''''' \ 102 "il'II/' 10:1 1."11/01010\' gJ'fIl'I//\ 01'(/.\
Nil /II 77
/Hl"II'
1.('!'Iump/I;' .\-1 -1 IIlhlllll'''IIS \·1,1
/.(,/ IIIII'ol,Isi,
12J
Aol,,/,II"" IItI\ /,/( ·w'O,\'IJ,f.!lIIl1 hll/W'fI 'J~
rut/uf/lln 11/1,~ ji/t'lIrn'lJgmd /d('Urr"'i,1!.Jf/(I ,~t 1\0 A,dl/('/lI;,\ , Aa{,'I,/in 1111,'
1.''fIlIll'lIlll1i,1 ,,/1·,,111\,, '11,1 I /. '11"11 111 I'S!,/I', l'oil 'I'('''I/I< ',1 I ()-I li1JlJlfl(·hol'i." / t '/( '1'\'01'1'(/ 1.15 h"rI/"1.'1/.1/1
-'
/£'{U 'O.t:,(/S/( 'I: ( ',dIlI Hllnll
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) /(01/(1 IlIe/JIII,\lI
ril""(l/l i~(""'·,'·
j}/tllro.\lig",o
139
lille/II 1-11 /'l/hllI 1-12
12~
l'ul[,
l-l-l
"'I'fII 'IIIIII ,),~ 5
150 j, illl/hi l ilit 'II \/,\, ,\iJ/rl'llf III/~ "l,/rl'\' 2X(1 1,,,1'1/1 '<1,1/111, ,\I"g"I,lin' ,' 110 /'1Ih111 , 11111111//1( '('/, " I ~ 2 ).,lIfi1lil1ll1j'/lSI\, :, 1"I;\'/fJ,1!.l'I/\ 'S
/wlu \"lI J/c "n\';\
1..t ~
l. imlloJ/{·/ 'tf'\' /('!JtJl';1I0
I iOI,('llil h(//;odt'lrlll
) /_iJJ/110Il i 'C'lt'S
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/f'I'(W;III1\ ha/iud('il'lil1l
1.(II/(){/('II"lflll
IlJllgin'f/.\ , r'hilllllfllS 1~6
/u II !..!, It !JglIll .
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ai.
L"/)("I"II:I'/I £, XO
III/IIII"l/Pili" I I I) hl/iI/C/l.I/' 12() ,J/c ' IfI'(i.\'{i,t:,IiIU
\/( ')!/I/J/II"I '\ /'(dll('l/\I,\
t:m, 'di" , --. :1 1"(1 1., '/,10111/(11 IIl'/Il'lII --. :i1", Lqlflll.l'(I, 1//1/11/ III>/lli/(i :
'-',/IIII/>/'(/C";11111
/(/lo/)UIO/(I/II ,\
,\/II11,ni, IleI/lifor
IIflltilur
11if/1/.1 1'1'1('I's/
l)~
IIInH/fIlUIIII
J ~I "\
tI"'I'tlS'l. ( '
l!lt'''II1t.\
'-, '/'Io/a llll' (/I'
"Is" I ''1l1o/'I'u,.he/l"
haltft'II.H,\
IXf,
1\11111\ III/u{1ll1mo",,\ ,
-
Ll'{I/ohmdlilllll ') I o/1/'olli ')1 gllllliJlgc' I} \'(' 91
,\-12
1I1/('" liIlO/;'
> ; i1 ~"
1.t'l'I"hnlf'lic'!11I hoIIlCII,I;I
/"/ (',."1/1,,,, , 3 11
/J/otO( ('(lilli'
X')
h.df("IIS/\'
1)(11"'(1 'lO
. 1J/\fJlIJU
70
I .upltn-II" hUI"/I('II""
(I{ 'It(',,, grolll"l i '11.\ ,
)
,\u/tu/Ji furl ;". I JlhTlltllll\'
10<'111, (It ('Id"-\,I!.a
I. II IIIl'rof"l'i" .; 7~
,lI so
Inlllen ' \IJn/\ /uII'j';OJ'fflll
11I('luo,\lI. /1<1 11<1
hIJllhn('(JIc/"u.
1-l2
1'7
I.ye""",,
16<;
( '"/d f1lllrJtI
J? 1
J-l5
,ub( ill( 'ln' s('olci /.1< 'lIdll/l IIlholllll/ll'
.'\..+6
) 1.<'I'llIml,ltis IIlh"liIS"1I.\
Indt: x
L" ,I !,I','OIlIll
~1I/(/hll!t/('IISI1
1," 0 I ~2 ll/ill"I!('{/c!i 153
S"II<'lIl1l11lill'/1II.1 a//I'(''/i ) Hio/'tt hour/Jlgil
Il/ll""ll(ll('lIIil
~llIal"rll" 'II",' . -> .\il/i"IIOIIII!!I,III/\
~1I111"(/III"IISil
al", 1!t ,1'I '/lIg('l1l I
;\I" I'I."/(Ig/'III" /,Tholl
l .Y}!,(I\(J OIlI 1I1 {,IIB 't.'lIhu , ,/.
L.\ -g.o."H lIlI ulin / ('('I1 111
("'/'/1111)( '11 4..'111 i,\
/4tllHl)ru/cIJJ\ 1I{( ' lIl1 '(' nhlf/\1
f)tl.~i(/ nIh lIn.'lI
-t
I Xg"\l1 1I1lI 1(,lIill('/1I1I11I
~ al ... o ."phc '/IfJJllrJI"/JIIt"
1/IIIlllllla
.\, "" '1/11/1/111'1 ,1111,1 ~ 11I(/"alll<'II,1I 1
' p.
12 7
,l/it 'l'l\alll' "(//11,'1/.1/1.
I'I'IIIwlal'lIIlI.l : S"II(,IIIIII/OI,{I/illl IIIldll .\I/lIalllaills , I , I,!!, (I,II!lII
127
:I1elil/J/II),lI elll/
) ,'>iJ/II ' III1I1UI/I,IIIIS Il'III1I1 ' lIllI.\
L1 J!,U,\OJIIU \ 'Urieglllllli i.
1.1'1111111/1"'"'/111
I IX I-I X
cr, IIl'f,IIIIII'II"/lII'
lII ellilull'el,/ia/II' IIId,,1I0l '<'I,/ialll' I X,!;""'"I" al/ln/i 4 1~\)!.(lw)lu(/ /l()l\rlllgi
~/lIlIhllfl/('III/'
cf.
) )'ihl 11111'/11 I
/. ,('(}(IIIII 1II<,lallli(l'p lwllll
IUo/,
1.12
f''''I'I,I!,t'I/(/
('Tllld{/( I\!t" hallll'II'I '
1 IIlgI '1't1l11l hll/tIl'IIIIS
'\/;L'I'o /ty ll/ 110 htlm. ,,'III/' 1.11 Pc '/o/ ,hn lI( ' x)
111/.\'('1"(1,
I X7
mi"""lgl , 1'''i/ilIIIIII
.1I11l "'11.1 I"", rIllgl
M ;\10 hll icl
) ,l/a"1I1 II
III1I)WI'e/III,I. (Jd~"
MI/IJllyl/ '27 S illd"lm 'II "' / 27 (,
tl. l 0 1l11lll )),1/11 (' 1 lill.
27h
III lilli/a " IIIIa 1'11.1/,1 27X I'Ilg;I"rrt
Nllif!/{ hll" rtll,!!,il
J:!2 260
11100'( ',\(0, ( 'tI /tlN/onu
1111'11/1//1
IIl l11lt}(-//rtJIII, "III" I "IJilll
0I01llUIIUIII. 1~ ('{}lo"rllChll"l1
27'1
,(IIII/( 'I' II;;
2 J l) 9.'
27(,
1II1I11t11l1'/'I11111, '\I(/hll\'(/
M al>II' a 1II111'//,(//,;lIa'a, -, ,I/ahll' a 1II""I>I" 'lIla
11I1I11;1'llIillIllIlII.l , ,~i"I(,IIIJ/I/(/l'f,/tIl.1
,H lIITIII';,' llwdoll J 17
III IIrIldl' lI II,', RIIII""tlf ,/t /1
1'11"'/11111<'1111' .,.+7 IIIlICH)\t
1I1111iflllll
(·/is . Nluu 'olJluWII,\
111<1('1'111;" l 'o /II'l't/IIIL'I IIll1Cit/alus,
('\ 'c'l'c 'lIl
(;,,1/1 I'll 25')
2 10
1\
1')('
'·('''o,\/,h" ,\ Xl
III(/ni/
lVl/;1I 3X7
,170
~xx
.\I//JIlI/nUIlI
11/<110(,(111/111'. 11I" .,.,1lI111 1
.,.+.1
'\IIjo Jlu/u
mateo/"II ,
17'
Valli frI/JlU/ttIlI S_
1 ;" 111( '1"" .\111"1(\
/II11I),:'lIrilfllll .\, (IfHlI 'O/J/t"
,1/ali(1I1'I/ ill'''''IIIII/'I'.
J.'" 7
Valli
,( 'aI/lilli/III 1I11"llIlIa!t, III<'f'al'l;
,\lul/ ( 'u,.u 1I1/( ',\111/0/i\ nigl'o ltlt'Jli{llll\
) (
all,u/III/\
III/,'III/wlt, r//('f'(/III
(1
,1/('gaf"I'III"''' .
Alihlll'I I,IIIil IW,IIII.I
) ,Vel/II ,\lIl11u/rUlla
112 1IIIIa/(I/; SIIIIII'OI,I 17 1 1111 1 /1/a , 1/('J!,III,iln 'l 1
/tn 1(/1;0.:(/
, f(/IIII>""f '/II' (IIn',II II ·!!./I,1 • .'11111 ,111".1111<1 Ilrl1 'l/nll/\
'\Olrn JIIt/I ' II/tllIl ,\
\ 'j' lIoclrful"lI.\ 11/(/(11/(/111\
'\'a/I'ir JIIltl'/It/CIl.\"
) /(/111"""/,11/1 111111'11.1"11.1'/1
;1I, :!!."I,hITI
NUln \ ,\'UI"l I1 '( /( '('//\I,\ .
~
V", '"1'111'1 lit' (//111111/1/1/1 .
1111
IOI ,hnJIt ' {'1','I'('1I1
\'( 't
lo/ ,hn lit ' gllt 'IJlllt'ri
'\<'1 1(lI,fIlTIIt' 1II1 ~. '1'1I
-, /.( '1)/0/>/'(/' ///11111 a"""III
,1/('g"I""" ', I' a"h"III ,1/('.'0'11/'''1'1 .1 g/'(II tll,1 ,1/<'glll,III"''' 11<1'\01''''11.
, " Iso -+ ,Ii,,,
\('IlIhlll 1II/lIh"I',!!,1
I "f'lnl" '1I1 fllllill 11111111
IIt("O/'Ul"lc. 'lI.\t \_ NUflu
,I/"g"/,/IIY,' 11/1)/11;('11 /11 , > alstl 1/''gIl/''" '1 I ~ofl(lI '(/\"1/ :\/t'gtl/ ,ltn ',\ IIIUllll nJla ill/S tOll ---7 '\/j :e..O{'/II.,.,\ 111/\'11/11
abo ;\I('g"f'''"rs ~ohal (/I"i, )
al'(1 tk'!!'!'I,III"s
Jl/j '/tlIIOI)o,t!.fJ ll lli,t:,riOI'P<'lIdh·/flalll\. !)ru e I}
al"u ",.. Ioldlli/lt ' 1111\,'1'0
1'' ''''/,/IITIlt' 1111 I('/'(/ 1,"II/(I/"-
/'ltllIl ' lIl/1
16:'
I ,,,II/}J"iJ/cpi\ 27. I )ro{'u IIH ' /dllfl/NJgolI 2.17
"i.~l'icll)I't'lIdl{ 1t!0(1I.\ ,
nfl/go "' I ~ 11 H!,l'Igll/ol'i.\ , ( 'ol/l/JI,rOU\
2J7
'r.:i
II I !!./'fIplIllIIlI III I
,
2.' 0 1<11(/, (11,1,,"-'11 ~ I " Tn)/~lcI"/~hnrll'
\"or/wu h"j ·t'Urlt \"ol'h('o 1110((11101'(11
l..t t)
)
~ / )c. ,du,\(thc. ''\ IIIINII/tlllI\
1II,':.!.rt('('/'\ ,
~()h(/1'al'llI l
JII('llIlIl t('('I ,/wfll ,I' 1I1(,/
1-1 7
111c ' /n\I 'lIll1l1'·J.
"\ _III WJIIllI /lI lt C,IlIt 'lI
,ai,,)
f."I'lollI/a, IIrtlllll
\(/1'(/1\ (/( "('11.\( '
~ / )n/o\lIhc ' \ (' I 't'U'1I1
\t '(
\ {'( IO{,},n 11(' "UI( II/"Id
;I/".!'''f,''n ,1 II(/SlIllI ,
11l1{ JhJ(',\I IIO
\'(,I1(Jrh lol,/tI\ (' lllll,1!.ullgt'rtl\
:,V tl/n\ Irltlllgllflgl'ru
lOt)
(1IJ/(l/"fljJtdClm11\
;\illtl \1I11101l '( lIla
23(1
112
M (' /,;" I/I)I <'II.\' "
\'(I/o ,\III/IlI/nlllll
.lI'\l
'\,/lI'il //(/, 111'0111
,l h gol'llI:r.' · IO .'i hll/II(,III;,1 10(1 • ,d.,-,mllll
:. al,()
/1'l/JlIt/il lll ,\ \ , If m in/e'I'i\
'\lIll'iI
/1 '11", 257 Illl1l1rIlI, f "'I)/o/lIllI\' 102 IIWI,I IIII , ( 'rrtO"".
1I1rlril1lllS , /)V(I(
2X7
,(d
IIIH
II a It.\ , l'on'cJ\
ht "
(e"'''
) li -u{nt/oIJhnrll\ h. '( curll
~)5
42 1
Index
I> I lIIilli \
:V"c,;x lIllI" 171> II/cll/\ 17h
IU~tTI
o
I X-I 1 'h IXh
/Ollglll'll.\
()CI' id" ~.rgll
.~
I )1'1/("0
OhS('tll"ll,\ 1111.\( IIn(\ ,
/'/11/,,"/11.1 (111/WIIIIIIII 11I/lllIiI/II11I1
. ' \ '{'r( '111
lII;nh, 'I').!,i , "ili/Olllill /'l'lenl
J50
PIIl/III1I/{\ !'e rer\i
273
/)(I.\llI
I )I"{H
r)
1{1I111111/t'/o\'c'jali{\
(hwIlI,rl ',l..!(/ h(l/II('II~i,
23')
) (l¢ 'c'lllu:\-gu
IUUlco/)horlis LT( 'n' fI,
)
2·17
honlt '('II.';S
24X
('('/,h,,11I1I1
()JliSf"II'rllfJi,~ \52 /l'lli( 'U,"" 3).1
/'lto,II'/'/II"',I' lIig,ri/lI/ms, --> " Iso ""li\'O/II".,."
( ) rc 'ohrlfrucIJII\ h"/lIt'II.\I ,\
-4- ()({'Idn = \'.1.!.(/ hrl/twn'l;s
11/'1111/", , 11)"Ui//lI//\ 11(, (w,liIlIO('/H'HII\, r'\ {{,/,i\/rJj.!.t'li I ,
152
oli/O/,IIII,\ , 1'''/' '1 " ,,''1/(,,1 1'17
166 I"' /liS , 1,('}liti/a/1l \ I (l ~ I'i( III,' , :\'I '('Ii'''/1/1 I 7()
1';ClllroW , Rallll
--> (,('H,} 11I(Jllllrl'}lIt'
1'/111'-'/11,11'1,11 111111/(/1','/111'
~ 71 (,0111 '0"" 011/(1
"It',IS
"('1'/'''''''"
,(h ''';''iJ~l'gli h,,!J"'/I.\'i,
1)IIITIlt!g/'J\IIIS /'ol1l('/I\i,1
11/(//1,1 I,i('/II," /)(,11111''''111,111 ,1 J,~O
(J1'"phi" ,17,
(1\ I'g /mlll,'
--> 'vn 'ln"III\' I'i('//I'
Il/uJ{II/III\ ,\/"t Itltlll1\
t>lIIIXOP 111:1' I 2-1 7
2511
ll\yn.'/)ha/lIl11 ,
ei'ollo/JUllt 1t1111\'
jJ/IIlallllt.\ mloh. '/:':!.1
HI{I ('n l,\('e /i\
()I)IIII, 'S ,,/1>"/11.1< '11,1
,/1<)-'(,111
I'" IllS,
/'}II/III1II/,'
abo
)
"01lIc '11S;S
0,,";,\'11111'/1.\ 1."11 /",II/k,,/,n
R/frll 'n!Jltllrtl'
1I11gllIt ro,\'lJ'l \
J5_
",TII'hm!t.1 1('1'/,'1>1'11 /'1 !ollgdNlrlw,
also RiI"n'l,iI"I 'II'
['III/lIl1ll1 ,' I>III1W 'It/.III1.1', -?
1" I
IJllljJttro't( '(' JI\
oli\ 'on '(/
""i/III1I", ,,"ri/
,-IX 1...1')
",10/i//I'1I1111
-? I'hi/'IIIIII'
IlI(I'lIlIlillll1
3"0
1II 'IO/IIII'iI/lI.l , O/ig"c/"I/
Oligodoll
1\)1
.I'111,,'}'i
155
157
/de1'l.1
I x()
1,,''',/,\'1
I ~
}'''/II,'I/SI,I
I X7
IIIj"h<'I p,1
1X
"hO'fllI
/)/0'-\ ",\'011111 ,
J]l)
72
111/!lIrrl\/I,I!.lIId, ""/"/'/11" '1111,1
--> ()' 'I'ic/t} - I'g (l /11('1 il
PII~I'f1, ·"atl',"
p
I~"
I 'J I
1')4 1'l6 Oli/ol,IIII,1 1')7 1('I/COI/II 'SIIl,1
jJtrlll \ 'UJlCllSI\ , LIIHllo/H'I'le,
1..+5
I'tlmil/ll 1'"nllll" I, ( '''1'1 ' '''111'/' '11
/Jllrdll/ls, ['(In'I1,,
.12(,
111111('11/,11111'//\ 21 (. .15':;;
!life/lUlls 15':;; HU'( 'r I S IIwi ;\ 1',1/'1''',
111111'1'1111,1
PO/"I'ei/llies 1I/'/WIIl/WII/IIIUS, --> Rh,,,',,/d,,)!',,, 11/'1" '1IIIiI 'II/, I IIIS J'oIYfJ,.'do/c,\ ,igllaltf\ .
,llIlenltll//.\ IClt "'IS
I <'1'II'''J'{f(,/H'II.1
/ 't/\S('1'I10 /}rtISlllrl
Ihuclulla !wt/,\iJlCl
IJrllSlJld
X2
fllilt lI/all/\
1'1'111'/1/(1 I'I'lIS illll ,
1'11/01'1'11/"11/11.' Jot ~
( 'iln's"l"k" 127 P"/".-i/('/o's hi/J/'fJIII ) al~ll
"""I)/1II1l1
) !JllflgaJ'It.'
'/1/\'1'1(1111,\'
,-,' i'l/dOI'llI"lio/l J 'i ~ r~I '(I\'
X':;;
/''''''llllIl 'II(' "rel 'IJi< ' I , 1'd1lll'c /gOIlIll • ',/I}WIIllIl
/-'. 'ml"fI 1111111/11111.1 IJ['I(,/"II , }'III/IIIJlII.I'
Phillll/IIIS 17X
17() IXI
al,,) P"/"I,hrl'll(' lIIi,'(,1'lI --> ['iI","!,II"I'" ('(,/,/",111111
) (1':/II'J'O IIIlIldow
//1.1.,.",1
1.12
~60
"~)'d/ll ~II(J1I
2(.7
1(,X "'0/011/" 12(.
rill/Cop/W/'II.\' I"t/, '11)'0,
1 Xl)
Iw/ngt!/ltl, ,\ '/II"'Ohl 'ltI
tl1I1W1l11l1l1
--> /J,I"/IIIII"'~I'I/I1.\1('1
p"/l ',' rllll'lI/IIS
""//l.Il'i ,15 ~ J 5')
"t'I"JlhrY/lt! .~4
Ol1'Ot'III1,\
~O.'l
357
p,\('I1(/ohoo/c/,\ciala .
/Ie//(/.I',
noglllllll
J77
))(,
I',\(/Jllllloj>ili, /l/tln ' nt/I ' lIla ,
Ih/"'IJi,< ·, ,,/111 IIdlll/.l , --> , l nlO/ll11 /itiigill"u
IIIISI ' /'ll
, 1/I11<'1ld/"
1~.\(lIIIII1IJl(r"I.l,"ll·,\
'''''/o,lIih es XI <".,n'l1l
HOllO sigllalll
II 1'tlmh/"he,1 3-11)
p/'{lI'/nJIIIIl/i"
<)()
~
)
JlfJ/wiUrllJlI \lIha/II , TnIlW1'('\"lJnl\"
1'"i1 't'I'II/CII/II.\', /'slllll/lIlJd\'l/(/ ,'!cS 111I1,/JII/'(I,\( '('I/I , ()hgodu/I
I~rtl/llll
,X')
I>I'('il(,II.,lellli 3X')
151
357
Illtil:x
YJO
r(,/ICII/II/I/\
2 10
('\ 'Crt'''' 1I1C1('HI\'( ('/",
gal/I/i ~ 12
{'r/I/{III ( 11/'1/1 ' "rei/,'II"/('I11/
h"I'''',\ \(l1II 2 1-1
Q
I/I,I!,I'IIIICI/II/(/II/\ 215
I'"n/ali, :!Ih
239
'1Ii1//{///('/II\"IfI/II' /Ullg/"UI''''', /)r(/("
lI
1
R
) 1<111110/,11(1/1/,
Ollgllft,."" ,,.;,, I SH
Rallli
RI/{ll ()/,II"I'II" h/llIll(II/0//II ,
I "l) 160
dlll/Cli/W/<1
(,/'1'/ltro"11 //I/\'II
1
I ()J
IIl t u'rO\ (c/r,
I (l'~
IIle (Jhllrlc ' Il.\' ,\
'i.~IIII/1I
->
H/llill
"IIII/il
/ /1//111'11( ' , /,'\ AIIII/"
limll(JchurI' ) also /.illlllllll, 'c'/n
/(01/(/ hll'lllI
halt
->
1, /111/11/11( ', /("
HUI/,'
//I},\('I ,
/1,/1111
/1111/(1
"lIhllI ,
) /
HUIIU iJlIJlliI('!JU/'IS,
Nlto(
)
"'(lcnitlon
-=)
-,
l'IIIIUJlJ( '( '/" .\ 1f.'/)u";11/{\
/ill( /11
-. / ,///I//(lIIC'c'/n 1,"1(/1'<111<'/1.\;,' 1/111//(1//(', /' "
11<11'11'/" \II
~
RUllO \ig lJuhl ,i.oJ/III/0.
Rona
III"el'ilillgH/\
NWIll '1'/1I/,,11('(/'/i
---,
(//,\'0
SUhOlllf\,
S'uttHI/\ IlIhcnllll.!.!JIIS
I ('lnc/ntiIlClyll/.\'
r('/ic/lIIIIIIS, /'I/h o/I
\1 '/II/<'/"'''cI,
\ a/slI
JlllIn/( /CII."S
2()"
11l1/'/I/( t,'''10 ,\'(///( '1'/ , Pill/illi/II' 1')1
~07
\t trO\! 'fU', ' IlSt ',
S i hYII IJphi"
~Hl4
~2)
) \I(I/m nl 1II1t11l/LI" 'lu/a
3o.l(,
,;(,5
nw/ullfJccplllllil\' OIe/,IIIOe -('/,11,,/11.\
-> /(/1111>'/"1,1/1,\ c Itn\l/lX",
alsll /(//iIhel,,/,III,1 111111'1/'/"11'/\ I
,\"" 1111/'"'' );('///1//<1/11,'
:Jh"
Hl"i
\ ''''1 '11111,111'
/1/('10110, "/Ij,(//II" ///('/(111111'1 '/"111"" (Irn '\(lIgo\
,"g/lil/(I,
/(lIlI,t I h(,
I'JK
<111.1'11111'''\/1'1 ,' 2111 "I'lwlleli< 11/"/11\ ~1I 1
1·"/II, ' I1.II,\'
\,d1'a/or \(!/\UIOI ; 'arooll \
\ (,lI/CI , I.H 'Oc/OI1 \lIh('llh ' /II\
Jh ~
177
.\i,/J('1I01llorIJhlls )Xt)
S(/lWII\ 1I11i/',/a,\nllIU\ .
I
RIt {tcopllll f'II ,\'
279
\chll'gc·/r \('h/" :~t'11. ( 'a/{I/llorla 'c /lIl1i'/lI , ('"Iall/(/I'II/ J2 ,\
..,(,(,
.I')!)
RIIlI'" /oplti,' :1(,1 clll ysar}.!,fJS 162 ({J/I\/';I'III,III1\
ring" lX I :!7X mf/II,\ nt///I\', (1 '1i1U/IOI,/i" ,X II
"iI,,,III , 1i'1I1/('/'(' ,III'1/' 1"'//('/111'//111
NUllO pi(,(II"o(O
J'\ fcri\/ugelll" hllu,halut'I1,\'I\ nlllllll('II,\"\ ,
"1-.17
s
~1Ih1/1
, ,1/,'/'/\10);<'/11"
('1','1,'111
1flllllll '(I
I'I(gi/, 'I'II , \/(/h1l1'l/
) Ralld .\I,t !,Jlrllll
illllla IJlI/(/l '/II/"/Is/\
> Rhc/( '/lI"w/,II,'
hUll
mdi",
LiIlIIlOIl(,(,(4.'\
Nwltl
26X
Rill{lll 2X I
/1I11'/"llll('ll\'I\
NOlla lI}hO/('/iI
, /'11"1",'/11/' " ""/"1"'11\
ho:..,· uoll
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lnde
.~mphicsma flol 'ifron Tropi 10,,011/.' fila IJatu . tmochrophi.f lII(Jell/mus. -+ also Amplliesmo ,wro...a ell. Tropidonor . aral'tl eLf. -+ Amphie ilia Mlrall'aeen Trvpldo"ot arawa emis. -+ Amphi ma S/Ullwon'll. c
alfrf!d; 2 3 .ranola 'fflJU 2l!:1 ilJahalurn.'
tellui
lid
290 TropidophoTIIS ~
'Plte.nolllorphIL
ubunu.\·
bee-car;;
Iso phenonwrplru:.
Tropidonotllf IriDngli/igeru.r. IriDlI$:lI lil!f'rUS TTOpiliophorltS 2Q 1 ~!C
also phl'nomorpltll,{
lilli/I;. quaI/utili"
typu typu, Ht'tnfp/ry/lod tylU.f 265 unicolor, Xl!nop Iti
ILbdJll: llAr eo/ei, L,I'codon ~
V III/rll" u
soll'olor 'alva lor --2 \ n ebruli. ~ rrebroli , Olfgodon 352
W 11"0
I rl. Tropidolo
'Whi/
we.niuro gmbtlM',fk)'/, E/oplll:' i l l ntllu. ph/momorphu 290 I trotu nio. ofliophi bMrgala ~ th('pI1.' i. olliophl5 IIlIC ritJ(lIi, 1J:\fi
( 'III
ntJri ' miotympanllln, Gonoct!phalu bornt'nsi t an ,"/lgen/J , IU)Chrophis ill Triglypllf>don ja,.fpid~lm. 8Qiga ja.rp: doo Tri",t're!lurln>' ill
mol"/llm; 7'rimereSllnl! ,t umorronllJ n/olt:o/ml. Trlmerc urn rna/calmi. -t al Trimeresllrus popt'ion/fll snOOh; TriOll)':C SlIhplunw . -+ DogfJlliu ubplona IA~1ff1l
378 Rhahdophi.v cbrysorg"
7'mpiJnrll)r" con.w/cilla/II.,. -+ con picil/a/IIS
424
I'llipsi/I'r W Xenochrophls 370
macula/us 370 Irioll ~ulig, n.r.t ill /!lJodOlt purpura. ..ns.
unic'olor
. aba"i
Rhabdophi,~
1.5.3
(,M/Aphis 369
~
TTlmereSllrlLv pllfII'cellS. -t n '(mert'J>UYlLI bortlei'Il,ris Trimen.'.fl/Tll.I Ju.marranlJ.<. -t 31. 7'rimeresllnlS
379
//IUS
headi. Mt'ri fogen)o'
.. etllIpel'
malcolmi l1b jJQP.fifJnlm abah; i l l Trim T'i!.fllMl.f ehaspni. --+ Omphis Ita en; TTlmt'reslln poprorom. Trim re.fllnlS pope/orum
Hogler; 9 TropiJoNQlm chry.fargO.1.
ill
du",erilii ~
"llfOlltlll
umatrtmo, aJa IIndona. #Claphr)'"cllo L!1.
Tr 'Pill
_\93. l!H
irgulafa. Caloma1';a 32 Api rygodcm l1!I
'lIhp/ana, /)QgfJllio 122 II/uelui . Calam(1J'io 3_4
n....n.ds
TmpidophorllJ b«caril
1)phlops sp. ~ I}picu, . Opi l"fllrT/pi., 3 3
hQTlIf! 'm'; ~ 3D6. lellia sa/,'U/or. . ' ronJJ.I , a!\'alor 'ulvaror aliczAoia ~ whalr a StooC'"',.iill :!n2 bonU!e'l~ is 3til
hQ
'Uri! ill
TropidQphofUJ mocql,ardi. luberillng/lis. '{ollrois 112
p;nosa, fieQsenJ}'. 3 7 .lpiflUlifer. Ansonia N IIIItTO/; ~ IOIopo/m illS l:::!l nalalOr ill rubt'f'i!inKUi., 12 II!NonolU!t .l6b
lIochrop"is
m
ill..
Ol/gOMn
N,l
Z looe)' {u ("Us.
Po
~
Ill: ('U,
purplJr(1~
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