(r--lorth Borneo)
Rana df!corala
Y!IX')UARU,
1890
Cd/amaria /aff!rali.. M()( '()I '/\RD, I B90
Pd/lll'OKonia CPphd...
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(r--lorth Borneo)
Rana df!corala
Y!IX')UARU,
1890
Cd/amaria /aff!rali.. M()( '()I '/\RD, I B90
Pd/lll'OKonia CPphdllllr/ MOC 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 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
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 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 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
:~,-.'~ ~ ·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; 14>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 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'ia gnusis A slllall low land Iree frog wilh a robust body and slender limbs; snout wilh a distincl, conica l projection: outer two fi ngers part iall y we bbed. ol her. unwebbed; wavy skin !lap along: arm and 4'" fin gcr and another along. tarsus; narrow !lap on heel and crenulatc dC l1na l projection on ve n\. Desc ription \t1 ales up lO 37 111 III , lema les up 10 50 Illlll ; TLlSVL 0.45-0.57: head longer than broad: canthus ro ' (rali s ro unded; l ore~ obl ique: Iympanulll disli m;l, half the eye di ame ter: vOlllerine teeth pn:sent: linger well-
Fig, 20-1 RIIU('(}llhorrr\' appel/dim/am, from Mu lu NP,
. arawa "- .
204
Rhaco ph ori da.c
E':J ::;
ti
-,
;;...."--~_-"'"
Fig. 205 Male of R/J(fCOfl/IOr/lS a/IIl('/u/iclI/IIIII' from D aIl lIm Va lley. Snbah.
Fig. 206 Femaic of Rh{/("opllOrtI,1 (/I'P('I/{!iclI/all/l' from Dall UIlI Valley, Sabah.
in fln t nrcns o f small . inte rmittent strea ms ( Ill : Ii . bing I( 1- 1.5). 2( 1- 1.5/1). 3( Ii i ).·H I ): toe webb in' 2h). 2-5 m above the wa tcr slIr li1Lc. 1(0-0.5), 2( 0-0.5/0).3(0;0). 4(0-0.5/0-0.5). 5(0): an ova l inne r metatarsa l tuberck. no ou ter one; sk in abovt: with irregu lar low ridges: sup ra tympanic fo ld Ca ll: ( 122°(,) J\ series of soil. clid.ing. pulsed notes: note du ratio n 200-300 IllS: eaeh note increases frolll be tween eye and ax ill a. For th. 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;)"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 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.
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 t ( M.\ I I 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 &
)'''.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".
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
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 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! ~
'":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/' \. 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 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,,,,), ,,.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 ICe I largely on Sill a II ills 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 R1'1.'
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, 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 1l11a 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 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.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 \ 11
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, 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 Xl)a . I'll>, I')')I~ . 1.1. 1') '-\
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 777 ) IIO/JidtJphu l'lt,\ IIItJetlJlllJ'clJ III RIHHI 1') I"' . ?7h. t\.1 \N IHI ' 1\) ' , 27; 1\1 I I "'Il , 1 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 /'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 appearallca 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 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"
1.1r:"\ ,.~ ,IIld 1.1, 1 IIIti "., ('11\1\, BIH II Nt 01 I< I X '7 l) , 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" predI).1 is di~trihutcd frolll thc i'vtday I'c-
ninsl1ial/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"& 1 1~\ 196): 1·1'/; I') X~ 27, ~I " "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. KII1ia ~ 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,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 '1,, \ l·!. /'1< /,, \ \1 \1 ~ " , , 1()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 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 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 copulali1
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
:II''': Il'rrl' ~ lrlal I hl'~ ked 111:11111) llil li/ar.!\. rrogs. alld '1I 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 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 (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,,
, 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 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 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'
~· 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
s.. I" '" I 'N'I I ~ O H!JII"""/'/'" d')'!'.IIII,I:".I , 1\1\I
I-. 'It ,
I '),)-1 11
~ .j 7 ,
I 1
11 ,\,' 19--'l)
~
UIUl"d"l ,hl\(1111H Jr!~II . \1 "1111\ & 1)1\/11< 11)~ 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 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" 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 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. pi ' I ""' tH' toc .. til) : 'vlall\()()f ( "" Iah . 1IcJlt:a 1 IJIIIJ ~c / "' '' / fl \uafll' .
I IH Ni l \1 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';rgllllllelraflle";11 1131111-.1 R, I X5 ')) ('ull;"I,It;,\ ;111(',\ (;11111;\ 1111'1""'\ ; I Hi li ~I ". I X5 9)
IIItl
II I I I '" H
I
:-.i""
20 I
I) Ill' In r:l li( ~:
S lllta n ):'_
\ \ \·,,1 I lUl IH'll ( 'O IlIIl/I/,1\ /l1\I ' )!,t I/O 1 /.'/role/('II(t/)
I \\\ ...1 IN "'001
'\ f{)\\I " ' .... '.
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: 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 11l'Ilto// Cltr/ IIS ( KHl( .JI, B \)(J..I ·)( & Sllt NI 2(0 1). Ma, . total Ic ngth 170 CI11 . Cll rt ll.\
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I. XO ."76
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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 ~calecondar~ 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 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
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 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. 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. ) 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-rkall: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 ' ( '/lI1 Ihc ,,1;1"':: gC IlII~ '111'"1/)( hi/m . ;:"ul .I L11111 Sll,·. 10')' 275 -;!1)') f) ", I. 11 1)1) 5 ): i\lllphibi , lll~ ,lilt! reptdl's rl'l'onk-d al nalu ,\pOl , " 10" lallt! ll1l'l.:rll\.'; I'11 IIl1e~1 III Hnlll
Bihli ()graphy
(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-."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'UI,/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!(Iphi.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- 11I 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"(>0.1 . \ '1NI I', !'vi (1resd':I1.9(J): 1-10. \\ II I II, I\. .IU I . IIl)XX) . SlIa".:s ali VltIS . .I .. 112): 21)-3N . "" WON' ,. 1\..:"'1. & J\ I 11'1" (1'ds. )( I,t/I/('I/'"
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