Contributions from the Biological L University (1979), 25(4):

Transcription

1 Title Amphibians frm Sabah I. Systematic tes Authr(s) MATSUI, Masafumi Citatin Cntributins frm the Bilgical L University (979), 5(4): 0-46 Issue Date URL Right Type Departmental Bulletin Paper Textversin publisher Kyt University

2 Cntr. Bil. Lab. Kyt Univ., 5(4), 979 AmphibiansfrmSabah I. Systematic and atural Histry tes* Masafumi MATsui Abstract Thirty-tw species, including tw questinable and three unidentified frms, f anuran amphibians frm Sabah, rthern Brne, are reprted with special references t mrphlgical variatin and natural histry. Twenty f the species were cllected frm Kinabalu regins and the thers frm varius lcalities in the surrunding lwlands. Tadple f MegPhr7s baluensis and unknwn tadple f Ansnia are described; variatin in Rana kuhli is analysed and striped frm is newly reprted frm Brne; Rana rugulsa is first recrded frm Sabah; distributin f Kalula Pulchra Pulchra is cnfirmed; Rana kenepaiensis is first recrded frm eastern Sabah and differentiated as a distinct species frm R. Paramacrdn; tw prbable new frms f Amlps are cmpared with knwn Brnean relatives; distributin f PlJPedates teucm]stex leucm]stax in eastern Sahah is a significant range extensin. Studies n the amphibian fauna f Brne Island have a lng histry since the end f last century. Early wrkers intensively studied herpetlgy f Mt. Kinabalu (e.g. SMiTH, l9). Mre recently, IGER (966) cmpiled a mngraph frm vast amunt f systematic and eclgical infrmatins n Brnean amphibians n the basis fhis experience frm several lng trips t the island. As a result f this valuable publicatin, mre is knwn abut the systematics and bigegraphy f the amphibian fauna f Brne than f any ther part f Sutheast Asia. As better understd the amphibian fauna is, additinal field wrk may cntinue t prduce significant infrmatins. Accrding t IGER's mngraph (p. cit.), 89 species f amphibians ccur in Brne, and even recently, additinal species is als reprted (IsKADAR, l978). Sme f them are cmmn and widespread, while thers are rare, restricted in distributin, and little is knwn n their natural histry. As t amphibians f Kinabalu regins, little bservatins have been made n their behavir, reprductive habits, and micrhabitats, inspite ftheir lng histry ffaunal survey as mentined abve (IGER, 978). Cnsequently, the cllectin and bservatin f even a small number f amphibian species in this regin can be particularly ntewrthy. Tw bilgical expeditins f Kyt University have made herpetlgical cllectins in Sabah. The first expeditin, 976, perated field bases at the Kinabalu atinal Park (KP), Telupid, and limestne caves n the East Cast, fr tw mnths * Cntributins frm "The nd Kyt University Bilgical Expeditin t Sahah",,.

3 04 Masafumi MATsui (KBAyAsHi and HTTA, 978). A small cllectin f amphibians was made by Dr. Tsuneaki KBAyAsHi, a mammalgist, at KP in August 976. The secnd Kyt University Bilgical Expeditin t Sabah, rganized by Dr. Tshitaka HiDAKA and cnducted by Dr. T. KBAyAsHi, Mr. T. HiKiDA, and myself was perated between 0 March and 4 April 979. Entmlgical staffs f the Frest Research Center, Sepilk, headed by PrÅí Ry6z Ysii, jined ur party and assisted in making herpetlgical survey at KP. We spent twelve days at KP (lcality descriptin is given in KBAyAsHi and HTTA, I978) and furteen days arund Kta Kinabalu. In this shrt trip my intensive effrts were cncentrated in studying the habits f frgs and tads in varius habitats, making temperature (air (AT), water (WT), and grund (GT)) recrdings f respective micrhabitats, recrding infrmatin n appearance in life with phtgraphs in clr, tape-recrding the mating calls f several species, and cllecting specimens fr future study. Cllecting specimens in KP area is cntrlled and mst f the specimens were presented by generus native peple. The specimens thus cllected are nw in the cllectin fbilgical Labratry, Yshida Cllege, Kyt University. 7 ils 7 f -", i.q b' r -t XM 7e s 5 ' A. lt '.. L's i. k. i' { x, tt!- --. lgå~ "s,i. q '-'-' '-'.,; Lt,:. ;; n' A t,... i'e'' / t""p i"i" c e E-G HrJte,ee- M e P K-L vo -f'j"'-'"-'' L,iv(-SL-'".s,t"'L SABAH r.-.,",k,i/s. Q -S ' s, ", R T Ve e s e Ue ''?- - `lj '.d,. ' -: S,,. - Fig.. Map fsabah, shwingmajr lcalities at which cllectins were made. A, Papar; B, Gaya Island; C, Kta Kinabalu; D, Tuaran; E, Panar Lavan; F, Pwer Statin; G, Kambarangh rad; H, KP Headquarters; I, Liwagu River; J, Silau Silau Stream; K, Bundu Tuhan; L, Kundassan; M, Mamut;, Pring Ht Spring; O, Ranau; P, Gunng Alab; Q., Telupid; R, Sepilk Frest Reserve; S, Sandakan; T, Batu Puteh; U, Madai Cave; V, Mt. Lucia. 6e 5

4 AmPhibiansfrmSabah L 05 In additin t the cllectins made by the first and secnd Kyt University expeditins, the fllwing three small cllectins frm the basis f this study; () cllectin f the Osaka Museum f atural Histry (OMH), () that presented by Prf. R. Ysii and () by Mr. T. HiKiDA. The first cljectin was made by Dr. Mitsuru HTTA, a btanist, at varius parts f Sabah, between Octber 968 and February 969. The secnd was made spntaneusly during mstly at Sepilk Frest Reserve, Sandakan, by Prf. R. Ysii and entmlgical members fthe Frest Research Center. The third cllectin was made bv Mr. T. HiKiDA at severai lcalities f Sabah frm J July t August 979. The purpse f this paper is t preliminarily discuss the findings f interest t herpetlgists that have resulted frm study f the specimens and field bservatins. Particular attentin was paid fr mrphlgical variatin, thugh sample size is t small fr analyses in many species. Analyses f stmach cntents and mating calls f several species, tgether with taxnmic cnclusins f the specimens whse identificatins are at present uncertain, will be reprted in future papers. Specimens f several frgs and tads related t the species treated in the present paper were als examined fr cmparisns. These are in OMH and Field Museum fatural Histry (FMH) cllectins. Basically three measurements were made fr transfrmed specimens: snut-vent length (SVL)-direct Iine distance frm the tip f the snut t the anterir margin f the vent; head width (HW)-the greatest width f the head; tibia length (Tibia L)- straight length fthe tibia. Other measurements were taken n sme species: internaria] distance-the distance between centers f the external naresj diameter f the tympanum-greatest diameter f the tympanic ring; partid length (PL)-greatest length (excluding cn.junct warts, if present) f the partid gland; partid width (PW)-greatest width; diameter f the disk f the third finger-greatest diameter; ft iength-distance frm the prximal edge f the inner metatarsal tubercle t the tip f the ]ngest (furth) te. Fr larvae, tw measurements were made: ttal length (Tt L)-straight line distance frm the tip f the head t the end pint f the tail fin; bdy length (BL)- direct line distance frm the tip f the head t the distal pint f bdy. Tail length (Tail L) is calculated by subtracting BL frm Tt L. Tadples are assigned t the develpmental stages f GsER (960). Ratis were rutinely calculated as the standards f bdy prprtins, but n statistical analysis was perfrmed fllwing discussins by ATcHLEy et al. (976). Almst all the specimens f transfrmed frgs and tads, lacking evident secndary sex characters, were sexed by the results fdissectin. Species Accunts Leptbrachium gracitis GtiTHER Fig. Liwagu River, KP, 500 m, larva.

5 06 Masa{inni MATsul A B Fig.. Tt Lateral (A) and drsal (B) views f tadple t' Le.t)tbrachiitm.gracii.y, St. 8, L 6 mm. BL,5 mm; Tt L 6.0 mm; Tail L.84 f BL. A larva in stage 8 was cllected in a stream with mderate current and gravel bed. It was difficult t find the tadple whse pa]e bdy clr well matched with clear pebbles n the bttm f the stream, Leptbrachium hasselti TscHuDi Figs. -4 KP Headquarters-Kambarangh rad, KP, m, 6 males + females + l subadult female + 46 larvae; Silau Silau Stream, KP, m, 5 males + female + subadult female + larvae: Liwagu River, KP, 500 m, larvae; near Pwer Statin, KP, 900 m, male + 6 larvae; Pring Ht Spring, 555 m, larva. Females cllected in August cntained small pigment free va, and thse cllected in mid March had large va (. mm). These va are grayish white in clr. Tw males and ne subadult female were fbund by day hpping n the frest flr near stream r n radside in lgged frest (GT 9.5 C). The remaining transfrmed frgs were caught at night n the bank fstreams r attracted t light fr feeding, In August, a male was bserved calling at night n the edge fa small stream and the cailing male clasped a female in captivity (Fig. ). Metamrphsing tadples (stages 40 and 4), shwing the characteristics f adult L. hasselti, were cllected in a side pl f a mdestly flwing stream in mid March. The tadples f this species were mst abundant arund Headquarters regin and were frequently fund in pls fsmall streams (WT C) r near the shre f Table, Measurernents (m mm) and ratis t SVL t' Leptbrachium ltassetti. SV'L Range MeanÅ}SD Head width Range Median Tibia length Range Median 8 Ad. -. -M , ? Ad "l.l.f ,E 'l, 9 ii r r)L "Ot

6 AmPhibiansfrm Sabah L 07 re Fig.. LePtbrachium hasselti' clasping at KP Headquarters. wide streams. They hid themselves under stnes by day and were actively feeding n substrate materials at night. Larvae with n limb buds varied in size (Tables -), and the suitability f applicatin fgser's tab]e fr this species in pre-limb bud stages is an pen questin. I tentatively include all the pre-limb bud stage tadples int stage 5. Size variability in a given stage is great even in samples frm ne pl. evertheless, the bdy size in the present series is larger than in tadples frm eastern Sabah reprted by IGER (966) (Table 4), Table. Measurements (in mm) i' larvae f LePtbrachiztrn hasselti frm Kinabalu regin. st, Bcly length Range Mean+SD Ttal length Range MeaniSD F { r). s iL6 LF a e. -S O f O. 5 s jr i- O

7 08 Masafumi MATsui Table. st Tai rati t BL f larvae Range. 6-L r Median L L f LePtbrach ium hasselti frm Kina st Range balu regln. Median L O. 99 Table 4. Cmparisn f bdy length f larvae f Leptbrachium /tasselti frm Kinabalu with thse frm eastern Sabah and Sarawak. Data n eastern Sabah and Sarawak specimens frm Ic.ER (966) and larval develpmental stages are adapted t GsER's (960) table, Stages 6- Stages -7 Range Median Range Median Kina balu 5. -S Eastern Sabah l Sarawak Als, the present series has fewer rws f labial by Inger (Table 5). These differences might partly habitats frm where samples were taken. teeth than be due t in tadples differences described ln -+ mlcr- Tab]e 5. rws Frequency distributin with respect t number f divi in larval LePtbrachium hasselti frm Kinabalu regin. ded labia tth SL - -4 Upper lip Lwer lip Jr Jr

8 .l/,...,/ An?,phibians,frm Sabah L 09.?Leptbraehium nigrps BERRy and HEDRicKs Fig. 4 Kenipir River, near Ranau, ca. 500 m, larva. BL I9.8 mms TtL 55.0 mm; Tail L.78 fbl. Labial teeth I:5-56-6:I. The identificatin fthe larva is uncertain, resting n its resemblance in clratin with the descriptin fpresumable larva fl. nigrps (IGER, 966). The larva in stage 6 was fund under a submerged stne in a mderately flwing stream. The dark markings n back clearly seen in this tadple were never fund in large series flarval L. hasselti specimens (Fig. 4). IGER's assignatin f tadples f LePtbrachium with light bdy and brad dark A tm"' i':lt/tlte,-,m,.,,..,'ti'rs/ltst,,. ':X',I B slt..' twy,ig-"an ig: #/,.',i.rr.ta"i,,m.,l,,lre.,m'.s',i,ts,i. mekv.,. tt"./: ". tt c D E-' li:w ttt' ttt'ti'ttt 'nyin t. /Sfi.'.l.',',,,"' t.t pmt. "' Fig. 4. Lateral (A. B) and drsal (C, D) views f tadples f LePtbracltium hasselti, St. 5, TI. 66,mm (A. C), and?l. ni,grps, St. 6. Tt L 55mm (B, D),

9 0 Masafumi MATstTi markings n the rt f the tail t L. nigrps has further investigatins are required t ascertain LePtbrachium. been questined (BERRy, 97), and the taxnmic status f this larval Megphrys mnticta nasuta (ScHLEGEL) Fig.5 KP Headquarters, ca. I650 m, female. SVL IO.0 mm; HW O.484 f SVL; Tibia L O.45 f SVL. The female had small white varies, apparently shwing pstbreeding cnditins. In this female the rstral appendage is almst absent while the triangular prjectins frm upper eyelids are mderately develped (width f eyelid, including prjectin, is O.6 fsvl). The length fcalcified skin n drsum equals O.56 fsvl. In these characteristics, in additin t its large bdy size, the female represents an intergrading ppulatin f m. mnticla and m. nasuta. The mst remarkable characteristics fthis specimen is the cnditin fskin flds n drsum (Fig. 5) : ne pair flngitudinal flds in this specimen are shrt and abut half length f SVL-the distinct traits f M. m. Iigayae frm Palawan. The mixed cnditin f subspecific diagnstic characters fund in this specimen supprts IGER"s taxnmic treatments n this species cmp]ex (IGER, 954, 966). Megphrgs baluensis ÅqBuLEGER) Fig.6 Kambarangh rad KP, 800 m, males + larvae. The tw adult males measure: SVL 4.0 and 45. mm; HW O.45 and O.44 f SVL; Tibia L O.48 and O.44 f SVL, respectively. These males, cllected in mid March, had well develped nuptial pad cnsisted f black clusters f fine spinules cvering the drsal and medial surfaces f the first finger Fig. 5. PrfiIe f female tvegpltrår's mniicla nasuta frm KP Headquarters, shwing absence f rstral appendage and shrt drsal skin flds.

10 Amphibians.frrm Sabah L and the drsal surface fthe secnd finger (cåí IGER, 966: 48). They were fund by day under piles f wet stnes belw a small waterfall, which is frming a small stream crssing a rad in lgged frest. Three Megphryne tadpies having funnel muths (Fig. 6) were cllected frm a quiet prtin f the small stream (WT 7.5 Cl) nly tw meters apart frm the place where adults f M. baluensis were caught. Cnsequently, the mst Iikely parent f' the tadples is M. baluensis. Tw tadples in stage 6 (with shrt limb buds) have BL 8. and 8.8 mm and Tt L 4.5 and 7.8 mm, respectively. The remaining tadple is in stage 8 and has BL A B c t.tttt t D.'".'.i :"r?.l:' b z':p.l,,,d " Fig. 6. Tadple l' Vtte.aPhrJs baiuensis, St. 6, Tt L 7.8 mm. A, drsal view; C], ventral view; D, muth ( Å~ 0), lateral view; B.

11 ! Masal"umi MA'rsui f IO. mm and Tt L f 7. mm, Althugh three larvae resemble larval A//I. mnticla nasuta described by IGER (966), they have slightly smaller bdy size; IGER gave BL values f larval M. mnticla in pre-limb bud stage (up t stage 5 fgser) mm (mean 9.9). The present tadples agree structurally with larval t4, mnticla described in VA KAMpE (9) and IGER (p. cit.), except in clratin. Cmparisns with the available M. m, ligayae tadples frm Palawan (OMH unnumbered) gave the similar results: venter f ligayae is mre lightly clred than back, yet guts are almst hidden under clred skin and thick abdminal muscles. By cntrast, venter fthe tadples in questin is mre r less transparent, with scattered pigments, and intestines are discernible externally (Fig. 6-C). BuLEGER (9) als indicated the dark belly f MegalPhrrvs mntana (:=Meg- Phrls mnticla). Since the tadples fthe genus Megphr7s are structurally very similar with each ther (see keys in PpE, 9 and Liu and Hu, 96), ntable differences in the ventral clratin between tadples in questin and larval M. mnticla is cnsidered t strengthen the assignatin f the frmer t M. baluensis. Fig. 7. Drsal view f Bapf J'uxtasper t'rm Ranau, female. SVL 80 mm.

12 AmphibiansfrmSabah L Buf bipcratus divergens PETERs Sepilk Frest Reserve, O-50 m, female. SVL 46. mm; HW O.9 f SVL; Tibia L O.46 f SVL; PL O.78 f SVL; PW O.50 f PL. The female btained in December had enlarged dark pigmented va. Buf juxtasper IGER Fig.7 Ranau, 504 m, 8juveniles + 5 males + 5 females; km SSW Mamut, 00 m, female; Mt. Lucia, Tawau, 50 m, juvenile. The presence fa single vcal slit is ne f the diagnstic characters which differentiate juxtasper frm asper (IGER, 964, 966). In my series, hwever, tw ut f five males with well develped nuptial pads have paired slit-like penings. The ther characteristics such as lng partids fthese tw are quite similar with the remaining specimens. Hence, the number fvcal slits is nt a valid criterin in differentiating juxtasper frm asper. Females cllected in March and August had cnvluted viducts but va were small and unpigmented. Onejuvenile was caught by day amng stnes in a ditch n grass field near human habitatin. All the remaining juveniles and mst f the adults were fund hpping at rainy night in the same field. One f the large females is fund by day n the frest flr f lwer limit f ak frest. In mid March, callings were heard n the bank f a wide river and sme males cllected emitted mating and release calls and tried t clasp female individuals in captivity. Althugh neither larvae nr egg masses were fund in the river, it is highly prbable that this species is a stream-breeder like the clsely related B. asper (VA KAMpE, 9; BERRy, 97). This species is very active and jumps lng distance and, when captured, it sn secrets pisn and smells badly. Table 6. Measurements (in mm) and ratis fbufj'taxtasper. SVL HW/SVL Range Mean+SD Range Median Juv. 8 Ad.? Ad Z L Tibia/SVL Partid LISVL Partid W/PL Range Median Range Median Range Median Juv. `li tl Ad \Ad tl tl

13 4 Masafumi MATsui Ansnia fuliginea (McQuARD) near Panar Lavan, KP, m, females ( specimen OMH unnumbered). ` The tw females measure: SVL 40.8 and 4.5 mm; HW O.7 and O.46 fsvl; Tibia L O.77 and O.9 f SVL, respectively. The female caught in August had large unpigmented va. Ansnia hanitschi IGER Figs. 8- Silau Silau Stream, KP, m, 9 males + 7 females + immature males + immature females. The specimens agree with the descriptin fa. hanitschi in mst respects (IGER, 960, 966), except the presence f a weak but distinct tarsal ridge in mst individuals (Fig. 9). The cnditin f the tarsal ridge in them is like sme specimens in the type series f A. minuta (FMH 774 and 7747), but nt s develped as in ther A. minuta specimens (FMH 774, 774-5) and A. albmaculata type series (FMH 8975, 8977, ). Te webbing in adult mlaes (mdally nly ne phalanx free f web n the third and fifth tes) is fund mre prnunced than IGER's statements (tw phalanges free) (Fig. 0). IGER (l966) cnsidered sex dimrphism in this character nly slight, but as shwn in Figure 0 and in Table 8, adult males have mre develped webbing than females and juveniles. Adult male has vcal sac and nuptial pad, bth are lacking in twjuveniles. Mst f the females cllected in mid March had large unpigmented va (.8-.5 mm). Immature females had small white varies. Cmpared with A. minuta and A. albmaculata, the present specimens have lnger feet. Althugh the relative ft length t SVL is nt markedly large, ft length related t tibia is larger in this species than in A. minuta and A. albmaculata (Table 9). Mst f the specimens were cllected by day n gravel banks f shaded streams f mdest currents (WT 7.5 C) in dark frest. They were fund sitting r walking slwly amng stnes (AT 0 C). One was fund at night n Ieaves f a shrub ne meter abve the grund. calling was heard in mid March. Table 7. Measurements (in mm) and ratis t SVL fansnia hanitschi. SVL Head width Tibia length Range MeanÅ}SD Range Median RangeMedian s y ? Y Ad O ? Ad Z 4 L tl

14 su s th AmPhzbiansfrmSabah I 5 mm "" "'th#$..ktwygei.it:'i' ima'mapaw.,ill//je.w /,--mmm.,//x,r, i em,,ige,agi/fi,ltxe,.ira.,i""'mi'ma:,/ma/ig,ma aglsiifsmli. ' g fiw zasu fi s ee lr trt S.n in is"s".i im is St'. I.. sm. ex va Ml #M ta an Me is llitiratztes. I\ezaM'l:m " ul S-w".".. sas l i EE.Ee S::a" za "" s ssm vae { e s t s Mfi t i.} tm l: ies ES ff ntutl an " MS.s."E zst S :/ l s ls ' lh tu me.mm -:tfian s -"".,i"imil.h:sdi i i, an" ti. sgg l tṯwt p" M {i-. :, l M te = i : ; lit l l l : i's i A - B t. ewll. esills getw x tntmu me s- tk g ee twtw an:ean t an -se : pm : r :esre m -Rmfi" s E i- ss s I Rt za E -. ee P!" "vk an ts-' = m"ll anm. es :iii" ff fi. mn.. Mtl -. za Seu Fig 8 Drsal (A, C) and ventral (B, D) views fansnza hanitschz frm Silau Silau Stream KP A, B, male, SVL 7. mm, C, D, female, SVL 7 mm :,lfi/me-,:/g.twee"ee'ge.itsift.hffs'.,.,eem/tma''iee'men asmm za. i s sus.u. ".ut exe,.".',`u -.mi:.lg.:z#,ml ' Se za m an m

15 6 Masafumi MATs(TI Fig. 9.. "arsus f'ansnia hanitschi frm Silau Silait Strearri, K-P. shwing tarsal ridg.e, Ansnia sp. Figs. - Mamut River, Mamut, 60 m, larvae. The tw tadples in stages and 4 measure: BL 9. and. mm; Tt L 4. and 9.8 mm; Tail L.60 and.66 f BL, respectively. The labial teeth are II!III in bth specimens. adult Ansnia were cllected in the immediate vicinity and pertinent identificatin must wait until mre infbrmatin is accumulated. These tadples resemble prbable larval A. minuta (IGER, 960) in having the halves f the anterir beak apart frm each ther and in having inframar,ginal papillae 'Fable 8. Variatin in extent f "reb in Ansnia hanitsc,lti. Figures indicate number f specimens, Free f phalanges rd te 4th te 5th te / /,, 4 i/,, / S Y. L)?)r: 8Ad l 9Ad. 6 +

16 AmPhibiansfrmSabah L 7 i:''n. ÅrJ Fig. 0. Planter views f three Brnean Ansnia species. A-E, A. hanitschi frm Silau Silau Stream, KP (A-C, male; D-E, female) ; F, G, A. albmaculata frm Sarawak (F, FMH 8975, male; G, FMH 9606, male); H-J, A. minuta frm Sarawak (H, FMH 7744, male; I, FMH774;J, FMH 7747). Magnificatins: A, Å~.7; B, Å~.7; C, Å~4.; D, Å~.; E, Å~.; F, Å~5.; G, Å~4.6; H, Å~4.5; I, Å~4.;J, Å~.9.

17 8 Masafumi MATsui

18 AmphibiansfrmSabah I. 9 Table 9. Cmparisn f relative ft length amng three Brnean Ansnia species. Range FtfSVL Median TibialFt Range Median A. minuta 8 Ad. Y.+9Ad A. atbmaculata 8 Ad A, hanitschi 8 y. 9 y. e Ad,? Ad, rr L L9. 4 L 8. 6 n psterir Iip (Fig. ). But the anterir beak halves are mre widely separated in the present tadples than in A. minuta. They have irregular, but assuredly mre than tw series f numerus large and small inframarginal papillae n larger length f psterir lip. Inframarginal papillae in A. minuta cnstitute tw scattered rws. Whether r nt these differences are related t larval develpment is unclear, but smewhat distant lcatins frm where A. minuta and the present tadples were btained seem t suggest that they are clsely related but nt identical. The lder tadple has well develped hindlimb with light bars drsally (Fig., B, D). Te webbing is develped and abut ne phalanx is free f web n the third and fifth tes. The presence ftarsal ridge is als suggested. Ofthe seven species f Ansnia hithert recrded frm Kinabalu regins (IGER, 978), nly A. albmacutata has been cnsidered t have tarsal ridge. In additin, A. hanitschi is prved t have ridge as discussed abve. Tadple f A. albmaculata, hwever, lacks inframarginal papillae n lwer lip (IGER, I960), different frm the larvae in questin. The knwn lwer limit f vertical distributin f A. hanitschi (75 m) is almst same height with the lcatin where the tadples in questin were cllected ( 60 m). Hwever, the highest degree f speciatin is ccurring in the genus Ansnia arund Kinabalu regins and it may be mre reasnable t cnsider the present tadples as distinct species clsely related t Sarawak A. minuta than assigning t A. hanitschi. The larvae were cllected in mid March under submerg. ed stnes in a clear stream having strng currents. KalutapulchrapulchraGRAy Fig. 4 Kta Kinabalu, 0 m, 5 males; KP Headquarters, I665 m, female. The five males were fund at night hpping n grassland in the twns f Kta Kinabalu. The single female frm Mt. Kinabalu was fund at nig. ht feeding n insects Fig.. Lateral views fhead f three Brnean Ansnia species. A-E, A. hanitschi frm Silau Silau Stream, KP (A-C, male; D, E, female); F-H, A. atbmaculata, male (F, FMH 8975; G, FMH 9608; H, FMH 9606); I-K, A. minuta (I, FMH 7744, male;,j. FMH 774; K, FMH 7747). Å~5.

19 . Ii.ti'/l 0 Masafumi MATsui A g$k ff/ il,- ',- B c D \.ll///i'/ifii l,x,k,, - -,,-.m.,:,gta,ur.. -.' m,.", /t..t s" $'l" t I-i,..-' tms. ill/,i t s'., 'X" ' t.' l,i" :I btt't' f Fig.. I.ateral (A, B), ventral (C) and drsal (D) views f tadples f Ansnia sp. frm Mamttt River. Mamut, A. C, St.. Tt I. 4.4mm; B, D, St. 4. Tt I mm. attracted t lights. Only ne authentic recrd has hithert been reprted frm Brne (IGER, 966) and the present series might belng t recently intrduced ppulatins..?kalphrynuspleurstigmapleurstigmatschudi Figs. 5-6 Silau Silau Stream, KP, 60m, male + immature female, The male and the immature female measure: SVL 4.8 and 8. mm; HW O. and O. fsvl; Tibia L O,47 and O.96 fsvl, respectively.

20 - I ' Amphibiansjlm S abah, stsp. va.zsif(`-tk}s ed XkQx,Åí.flS[e:L;K.'.e.., ged}tu.,,-- rd::a$p'p Fig.. Muth fansnia sp, frm Mamut River, Mamut. x9.6. The male has tw slit-like vcal sac penings in the flr f the muth, but it lacks nuptial pad. The yung female has prly develped varies. The presence fnuptia] pad in the adult males is stressed in PhilippinesePleurstigma and reemphasized in Brnean ppulatin (IGER, 954, I966). The absence fnuptial pad in the present male is pssibly due t later develpment f this rgan than that f vcal slits, but the specimens have sme ther discrepancies t be assigned the name f Pleursti,gma. Furth finger is shrter than in Pleurstigma (Fig. 6r"y). The tw specimens cmpletely lack inguinal cceli (Fig. 5A). Absence fcceli is fbund niy in 79i, f the Philippinese Pleurstigma examined by IGER (954), and TAyLR (I96) described the specimen lacking cceli frm Thailand. Brnean specimen has been reprted t lack ccelli (IGER, 966: 4). In additin, SVL f t,..,, ;,, '"i""- - :'X'.l' i';" l#'ia,,i lll,,, ':s: -l'. i-ii'ti' el'i Z'z l."l, l'l:kl.-l,, /t :I tittt!t: t'.ttt;.'.ittltttt'e. ' I I- l!.ge,suxde, -uamge ''i i,:',.g IIS.g':,S.li,g-/l-lii,fil,.,i"',, l i/s'i./f,i i li ; 'f,l:,i,tk,f/,: i:,il-,i,,,i,l.:e,/l li IG.lii{ ;-l'i'i,.'... i l-//f':l,}'.l " '-:';i"' i.'l'il.'l..' i/,, i: l.: :-.i,i' lli ',k ':,, --. ',,.,l,'i//rtri/;.,l/ /.i:'ttis.ttr-i'tny, i//k"gil"il,i';is i-.i,;",i i, I.i i-iii'ii l,i/ i/ii,ee ttttrenttttm/.:ttmtsehit/l Fig, 4. Drsal (A) ai)d ventral (B) views f Kalula Pulchra Puichra frm KP Headquarters, feinale. SV'I. 6.r) rnni.

21 Masafumi MATsui Table 0 Measurements (in mm) and ratis t SVL f Kalula Putchra Range SVI. Mean+SD Head width Range Median Tibia length Range Median 8 Ad Jr Ad the present male (4 8mm) is slightly ut f the range f the species ( mm) given m IGER (966). Tw Cambdgean Pleursttgma exammed (OMH-AM-57-8) had large cceli and stuter habitus and lacking small dark bltches under thigh and tibia, which are evident in ur specimens (Fig 5, B) Otherwise the present specimens have the cmmn traits with K PIeurstzgma and are tentatively identified as that species. The specimens were cllected n the frest flr near a stream (GT 0 C) The male was fund by day resting under a lg and thejuvenile was fund walking at night Chaperina fusca McQuARD Lidan River, Bundu Tuhan, 990 m, female. SVL 4 mm; HW O 7 f SVL; Tibia L O 500 f SVL tvl.. is,itwh,"'ni.'ret.mumml$'gki.,m,,,.tsee.ha:su.'de-everu'x lll'.`s:,\fi/"k'$tny,{ee.eet--,.ge,ee"""g -..:m' lza MII m i."ag-g-ee,:lkg,ut iisasitige.s::xx-su-ljg,.ii-wli--mge."g,..sgvafu ew.tk.musslj/l Fig 5 Drsal (A) and ventral (B) views f PKalPhrymtsPieurstigma frm Silau S]lau Stream, KP, male, SVL 4.8mm.

22 AmphibiansfrmSabah L A B Fig. 6. Palmar view f hand (A, Å~4.6) and planter view f ft (B, Å~.9) f?kalphrnuspteurestigma, male. The female with enlarged darkly pigmented va and n drsal spts, was cllected by day under a stne (GT 4.5 C) n the bank fa stream in mid March. Micrhyta brneensis PARKER Pring Ht Spring, 555 m, l female; Sepilk Frest Reserve, m, immature female. The adult and immature females measure: SVL. and 8. mm; HW O.4 and O.5 f SVL, Tibia L O.6 and O.758 f SVL, respectively. The adult female cllected in late March had enlarged dark pigmented va and was fund hpping by day n the frest flr. Rana blythi BuLEGER Murk, midway betw. Ranau and Telupid, ca. 500 m, subadult male; Sepilk Frest Reserve, lwer than 50 m, immature female + subadult male; Madai Cave, belw 50 m, females. The tw females measure: SVL 85.0 and 96.8 mm; HW O.48 and O.55 fsvl; Tibia L O.598 and O.570 f SVL, respectively. Tw immature males with prly develped mandibular prjectins and mderate sized heads measure: 60. and 78. mm; HW O.85 and O.7 f SVL; Tibia L O.6 and O.54 f SVL, respectively. One male was fund by day perching n rcky bank f a stream with Amlps j'erba and jumped int the stream when frightened. Tw females were cllected at night n gravel stream bank. Females cllected in mid August had large yellwish va,. mm in diameter. Rana cancrivra cancrivra GRAvEHRsT Kta Kinabalu, 0 m, metamrphsedjuveniles; Gaya Island, 5 m, males + subadult males; Papar, belw 50 m, male + 4 females + 4 subadult females +

23 4 Masafumi MATsur Table. Measurements( in mm) and ratis t SVL f Rana cancn'vra. SVL Head width Tibia length Range MeanÅ}SD Range Median Range Median Meta. Y. 8 Y. 9 y. 8 Ad. 9 Ad juveniles; Sepilk Frest Reserve, belw IO m, 8 larvae. Of the females cllected during late March t early April, specimens larger than 55 mm SVL had darkly pigmented va, mm in diameter, and thse 5-55 mm had a few small pigmented va. Males larger than 47 mm had vcal slits and nuptial pad, but ventral asperities (IGER, 954, 966) culd nt be detected, partly because f pr cnditins in preservatin. The minimum SVL values f adults here treated are almst similar t thse given in IGER (966). At Gaya Island, several cancrivra were fund with R. erlthraea near sea shre. They were fund by day in the bank hle f an artificial small pnd. R. erthraea was fund in water, but mst f cancrivra hid themselves in hles n the bank facing the water. When disturbed, theyjumped int the water but sn they climb again n the bank and remained in the hles. Several metamrphsing yung were fund in ditches in the twns f Kta Kinabalu in mid March. A series ftadples ranging frm n limb bud t undifferentiated tes was cllected frm the manglve swamp in late July. These larvae well fitted the descriptin f larval R. cancrivra by IGER (966), having teeth rw I :I-IIIII. Rana chalcnta raniceps (PETERs) Meliau River, Telupid, ca. 00m, male (OMH unnumbered); Sandakan, lwer than 50 m, metamrphsedjuvenile. The male and metamrphsed juvenile measure: SVL 8.4 and 4.0 mm; HW O.97 and O.07 f SVL; Tibia L O.58 and O.550 f SVL, respectively. The metamrphsed juvenile with a trace f tail was cllected in early August Table. Measurements(in mm) and ratis t Bdy L f larvae frana cancrivra. st. 5 5 Bdy length Ttal length Tail rati Range MeanÅ}SD RangeMeanÅ}SD Range Median O O L

24 AmPhibiansfrmSabah L 5 Table. Measurements (in mm) and ratis t SVL f Rana er]thraea. Range SVL Mean + SD Head width Range Median Tibia length Range Median Y. 8 Y.? Y. 8 Ad. 9 Ad i-. 6.7I l n a bank f a drainage ditch. I culd nt detect the presence f nuptial pad and humeral gland in the male cllected in late December. Rana erythraea (ScHLEGEL) Kta Kinabalu, 0 m, males + females + subadult female + 9 immature males + 9 immature females + 6juveniles; Gaya Island, 5m, immature males; Papar, lwer than 50 m, 8 males + females + subadult female + immature males + 0 immature females + juvenile; Ranau, 504 m, 6 males + 6 females + immature males + 5 immature females + juvenile; Batu Puteh, Kinabatangan, lwer than 50 m, female + immature female. The smallest mature size was determined as mm in male and 47 mm in female in the present cllectin. These values are almst identical with thse reprted fr Sarawak ppulatin (IGER and GREEBERG, 96). Females cllected frm mid March t early April cntained va develping in varius degree, and sme had pigmented va, ls mm in diameter. At Ranau, males were fund calling at night alng the edges fditches and paddy fields (WT 6.5 C) in mid March. This species is mst abundant arund human habitatins and was frequently fund at night in temprary pls in the city f Kta Kinabalu. In Ranau, erjthraea was hardly fund by day in the places where calling aggregatin was bserved at night. Sme individuals were fund by day resting under lgs n the cast f Papar. Drsal clratin is changeable in an individua] frg frm strng green t pinkish brwn. Rana kuhli DuMERiL and BiBR Figs. 7-8 KP Headquarters Kambarangh rad, KP, m, males + females + 7 immature males + immature females + 0 metamrphsedjuveniles + 0 larvae; Silau Silau Stream, KP, m, 5 females + 9 immature males + 7 immature females; near Pwer Statin, KP, m, male +8 immature males + 8 immature females + metamrphsed juveniles + 9 larvae; Kundassan, 65 m, immature male + female (OMH unnumbered); Bundu Tuhan, 80 m, males + female + 0 immature males + 0 immature females

25 6 Masafumi.TVI.xTsul v.ge Drsal Fig. 7, views fi'emales f' Rana kttltli frm Silau Silau Stream. KP, shwing tl]e absence (left) and presence (right) flight vertcbral stripe, xo,7: metamrphsedjuveniles + 0 larvae; Pring Ht Spring, 555 m, male + femalej Gunng Alab, 740 m, immature males + immature female; Sepilk Frest Reserve, m, l female + metamrphsing yung -- larva. IGER (966) bserved kuhli frm eastern Sabah had mre numerus warts than Table 4, Measurements (in mm) and ratis t SVL l' Rana kultli, SVL Head width Range Mean ii LSD Range Median 6 Y E Y E", 8. 5i ,48 8 Ad, Å}.8.4"-. 50.l6.? Ad L, 7, Y. 9 y. Ad. 9 Ad. Tibia length Range rvledian " ,E7. 506,l Internarial width Range Median

26 AmphibiansfrmSabah L 7 Kinabalu Pring Gunng Alab Sandakan Table 5. Frequency distributin in the cnditin f tibial warts in Rana ktthti frm rthern Brne. `+' and `-' indicate presence and absence f tibial warts, respectively. 8 Ad. 9 Ad. 8 Y. 9 y. Meta. Y. Tt Ol O 8 Sarawak specimens. In my sample, mstly frm Kinabalu regin, nly a pair frm Pring had tibia with numerus large and small warts and all the remaining had at mst nly very weak warts n the distal furth f tibia (Table 5). Thus western Sabah frgs mre resemble t Sarawak samples in this character. Mrever, ne female frm Sandakan had smth tibia cntrasting t IGER's bservatin. A y B rtcs ; c Å~Åq= /, Fig. 8. Lateral views f head f Rana ktthli, shwing the tempral stripe variatiqns. A, wide; B, narrw; C, n snipe,

27 8 Masafumi MATsui Table 6. Frequency and width f tempral stripe in Rana ktthli frm rthern Brne. `W' and `T' indicate wide and thin stripes, respectively and `A' indicates absence fstripe. Kinabalu Pring Gunng Alab Sandakan 8 Ad. 9 Ad. e y. 9 y. Meta. Y. Tt. 0 0 OIO O O A The cnditin f black blique stripe alng tympanic regin was examined n the basis flger's criteria (Fig. 8). The result is summerized in Table 6. 0fthe eight specimens frm western Sabah examined by IGER, six (750/.) had wide and tw (50/.) had narrw stripes. In ur sample frm Kinabalu regins, hwever, (80/,) and 5 (6O/,) f85 frgs had wide and narrw stripes, respectively. The number fsamples examined by Inger is t small fr precise cmparisns, but ur sample seems t mre resemb!e with specimens frm eastern Sabah in this character. IGER (p. cit.) bserved kuhli frm Sabah lacked cnfluent drsal spts. In ur sample, hwever, 7 ut f 8 frgs (00/.) had such spts (Table 7). Again in this character ur sample resembles t Sandakan frgs, but the bserved difference might have been partly biassed by subjective grading different frm IGER. The presence f blackish stripe n the anterir face f the upper arm was als examined (Table 8). Ofthe 8 specimens frm Kinabalu regin, 0 (40/.) had such stripe. This is almst the same freqeuncy bserved in eastern Sabah sample by IGER. Fllwing the analyses by IGER (p. cit.), relative size f the space between the nstrils is examined (Table 4). The calculated ratis and thse given by IGER d nt verlap in ranges: my sample includes individuals with even wider internarial space than in Sarawak ppulatin, which is reprted t have nbstrils mre widely separated than in Sabahan kuhli (IGER, 966). The reasn fr this cntradictin is unclear, Table 7. Frequency and cnditin f drsal spt in Rana k4hti frm rthern Brne. "H' and `L' indicate heavy and light drsal pigmentatin, respectively and `A' indicates absence f drsal dark spts. Kinabalu Pring Gunng Alab Sandakan 8 Ad. \ Ad. 8 Y. \ Y. Meta. Y. Tt. Ol O Ol Ol OIO LA Ol 0

28 AmPhibiansfrmSabah L 9 Table 8. Frequency distributin in the cnditin f dark stripe n upper arm in Rana kuhli frm rthern Brne. `+' and `-' indicate presence and absence f dark mark, respectively. 8 Ad. 9 Ad. 8 y. \ Y. Meta. Y. Tt. + + z Kinabalu Pring Gunng Alab 4 Sandakan e but again, the way f measurements may have been different. TAyLR (96) fund a few f kuhli frm Thailand pssessed a white mid-drsal stripe. IGER (966) mentined nthing fr this character in discussing variatin in Brnean ppulatin. I culd find 7 (0.90/.) ut f 56 kahli, mstly frm western Sabah, had light stripe mid-drsally (Fig. 7). Frgs with drsal stripe were fund mre frequently amng yung individuals (Table 9). Althugh the sample size is t small fr discussin in adults is evident, the tendency that the stripe is mre frequently fund amng smallerjuveniles than amng larger nes suggests higher mrtality in striped juveniles. Table 9 als suggests the clinal increasement fstriped individuals with altitude: striped frgs are mre frequently fund in higher elevatins. There were n tangible differences in the habitats ffrgs with and withut stripe, and bth frms were fund simultaneusly in an aggregatin f metamrphsing frglets. Cmpared with ther ranid species (e.g. R. timncharis: MRiwAKi,95; IGER, 954; KuRAMT, 968), scarcely any investigatin has s far been made n the frequency f iight stripe in R. kuhti. Future studies in the gegraphic variatin in this character must be based n detailed reginal surveys. Fr the size at sexual maturity in Brnean kuhli, IGER (966) gave the values f4.9 mm fr males and 50.7 mm fr females. In ur sample, sme males arund 45 mm in SVL had nuptial pads n the first finger, but thers lacked them. Therefre, sexual maturity is reached at slightly larger size in ur sample than in samples presented by Inger. Table 9. Frequency f mid-drsal light stripe in Rana kuhli frm Kinabalu regin. Lc. Alt. (m) 8A. 9A. 8Y 9Y Meta.Y. Tt. (O/,) Pwer Statin HQ-Kambarangh Silau Silau Bundu Tuhan Pring Oll O Ol O 5 Ol Oll 8 O7 O9 OllO 8 7 OllO O!O ! O (. ) (0. 0) (6. ) (L7) ( )

29 0 Masafumi MATsul Table 0. Relative grwth frmulas fr regressins f head widthsnut-vent length in Rana kuhli. Regressin equatin, lg y=a lg x+lg B r Meta. Y. s y lg y==. 09 lg x-o. 94 lg y== 04 lg x-o. 4 O. 975 O. 989 \ Y. 48 lg y==o 978 lg x-o. 4 O Ad. 9 Ad. 7 lg y==. 6 lg x-o. 64 lg y= OOI lg x-o. 90 O. 998 O. 99 Like previus authrs (BuLEGER, l90; Liu, 96; TAyLR, 96), I culd nt find vcal sac in males. The absence f vcal sac in R. kuhli is cntrasted with clsely related Japanese nami ei. The latter species has a pair f penings in the flr f the muth, the sitatuin first pinted ut by IGER (947) and ascertained by myselåí This insures the tw as different species. The smallest female with enlarged va measured 50.8 mm, almst the same size with that given by Inger. The enlargement fthe head in males is the result fchanges in grwth rates which ccurs with sexual maturity (Tables 0, ). Females cllected frm Headquarters in March and August had large va with pigmented hemisphere. The active breeding seasn is suppsed t last lng. Mrphlgy f tadples in ur sample well agrees with the previus descriptin f the species (IGER, 966). Arund Headquarters, tw distinct size grups f tadples and metamrphsing yungs were seen in mid March. These larvae were fund in ditches alng rads in lgged frest r in the places where small streams frm small pls (WT 6.5 C). They hid themselves by day under submerged stnes. In the shaded pls, kuhli tadples were seen with larvae f LePtbrachium hasselti. Metamrphsing frgs were seen hpping by day in ditches r near small streams. Transfrmed larger frgs were fund Table. F-values shwing the differences between samples f relative grwth measurements in Rana kuhli. Differences in slpe n upper right and differences in psitin n lwer left. Single and duble asterisks signify påqo.05 and påqo.ol, respectively. Meta. Y. 8 Y. 9 y. 8 Ad.? Ad. Meta Y. F7; O. 5 Fsis O y. F7i,.0 F,.95s F,t. tro 9 y. F,} O.04 Fsi. 704 F,i, O. 059 e Ad. F4ii 9. s** Fii *? Ad. F,} O. 84 Fi, 67. 5s8**

30 AmPhibians frm Sabah L Tab)e. Measurements (in mm) and ratis f larvae f Rana kuhli frm Kinabalu regin. st. Bdy length Range MeanÅ}SD Ttal length Range Mean'.IJ. SD Tail L/BL Range Median tt) 0. IL e t l O O B-. 8. Å} ' L9.Jr. 8 under stnes and lgs immediate vicinity fwater (WT 7.5 C) and sme were fund at night hpping n mssy banks. An ld larva and a metamrphsing frg cllected frm Sepilk in August were in stages 4 and 46 and had much smaller bdy sizes (BL 8. mm and SVL 9. mm, respectively) than Kinabalu sample (Table ). Rana kenepaiensis IGER Figs. I9-0 Sepilk Frest Reserve, lwer than 50 m, male; Tanegan besar, Kinabatangan, lwer than 00 m, female (OMH unnumbered). The male and the female measure: SVL.4 and 8.0 mmj HW O.64 and O.74 fsvl; Tibia L O.5 and O.574 fsvl, respectively. The male cllected in December has vcal sac penings n each side fthe muth. The third and fifth tes are bradly webbed t disk, but the furth webbed bradly t distal subarticular tubercle. The thrat is heavily pigmented with brwn, cntrasting t IGER's riginal descriptin (IGER, 966: ). uptial pad is absent. The female cllected in late vember cntained enlarged (. mm) va with densely pigmented hemisphere. It had less develped te webbing than in the male described abve : the third and fifth tes narrwly webbed t the first phalanx and the furth webbed bradly t the third phalanx. The cnditin is nt cnsistent with IGER's descriptin (p. cit.). The identificatin f the female is slely based n dark tympanum mask. The thrat is white mttled with brwn. The back is rather smth with feeble lateral ridges in the male, carsely shagreened with mre numerus ridges drsally in the female. The presence f vcal slits in the male and symaptric distributin with R. paramacrdn paramacrdn in Sepilk (recrd f R. P. Paramacrdn in IGER, 966 : ) suggest R. p. kenepaiensis t warrant specific recgnitin as in the case f Pllpedates leacmrstax and P. macrtis.

31 li: i t-tfit as// rme'as.. Masafumi MATs('i A B...mu ma,,va' ' 'ii" iii$åq. Z'w eweesgtw#,..ww ee d si xt". tt t!mmut lt ' '/Svae.tdi t=t t/.. ag: wa I wa.maxi k'p/ re'.ec,.- e'. I;ma i.,eei-as m,mug.ff/,el/tame/;g/mai.m...'./k'/#-iiilkee,//,//-ma,t-meff,'s,.,,,//eeua.va.,'":.,,il gil.i,i- Fig. I9. Drsal (A. C) and ventral (B, D) views f Rana kenepaiensis. A, B, male frm Sepilk Frest Reserve, SVL.4mm; C, D, female Åq{dentificatin tentative) (OMH unnumbered) frm Kinabatangan. SVI. 8 mm.

32 / t Amphib'ianslblm Sabah. ''et,-.,l t tt ttt ttt ttt te tttttt tu te. t' tttttttu.t tt ttt. tt tt tt tt ttt ttt tt t / s/ets.t.t ''t!t u/ett t.,m m.l,-, M/ t' ts Ht ' t.t. tt ---. l E' ttt t.t. ttttt t t -. s '.I', ', ' ' A B Fig. 0, Prfiles f male (A) and female (identificatin tentative) (B) Rana k'enepaiensis, Rana tuctusa (PETERs) Bundu Tuhan, l80 m, males + 6 larvae. The tw males measure: SVL 54.7 and 55. mm; HW 0.09 and O.6I f SVL; Tibia L O.547 and O.55 f SVL, respectively. The tw males had humeral glands, but lacked vcal sacs and nuptial pads, as stated by IGER (966). All the tadples except ne had dental frmula I: 4-4/I-:III. The exceptinal ne had upper lip with divided uppermst Iabial teeth rw, Advanced tadples (stage 4) had brwn limbs with light bars as in the adults. Rana mierdiscapalavanensis (BuLEGER) Fig. Bundu Tuhan, 80 m, juveniles. The three specimens, immediately after metamrphsis, measure: SVL mm (mean.7) ; HW O.77-O.407 fsvl; Tibia L O.580-O.6 f SVL. The three specimens cllected in mid March are t small fbr pertinent identificatin. [["he presence fdisks n te tips, pr develpment fwebbing, and absence f uter metatarsal tubercle, are characteristics n the basis f which identificatins were made. Table,.)vleasurenients (in mm) and ratis i' larvae f Rana luctuesa. st, Bdy length Range MeanfSD Ttal Iength Range MeanÅ}SD 'rail L/BL Range Median r L L) L , + O J: r) L L 6L4-O , I l. Jr -, cs. t) L6-L7. 4"

33 tht..tt, 4.Masac fumi MAFrsiTi "v lj t gem' $ -; eq,e" l /{"'ifl g 'gedi 'i'!"'l s," m'ii ' ' 'gk',k //e i: li,/ll'li itth/e,.s,, /ttss.l,lltr g. 'l der' twif- 'pa pt gsi,- as ede as,l ge, x th su'-,an' ll., asgexs/ :-,Mi's 'te" me xu"eeme' va" el eesesva ';' #' 'a;... tttttt.; Fig.. Drsal (A) and ventral (B) views f Rctna micrdisca Palavanen.si.s Crm Bundu r]"uhan. juvenile, SXt'I. 5 mm. These specimens, hwever, differ frm R. micrdisca described by IGER (966) in the presence fridge alng median edge fthe first te, and in the number fphalanges free fweb n the furth te being three. Tw f three specimens have dark pigmentatin n thrat regin and anterir and psterir surfaces f thigh. The remaining ne has dark back and small white spts n dark upper lip. It has indistinct tympanum and slenderer and mre numerus dark bars n hindlimb than in ther tw specimens, suggesting. its different taxnmic status frm the latter. Rana nicbariensis STLIczKA Penampang, lwer than 50 m, male. SVL 7.8 mm; HW O.96 f SVL; Tibia L O.57 f SVL. The male, with vcal sac penings and humeral glands, was cllected at a rainy night near temprary pls n g.rassland fhuman habitatin. At the same time tw immature R. rugulsa were bserved. Rana paramacrdn IGER Madai Cave, lwer than 50 m, immature fema]e. SVL 4. mm; HW O.8 f SVL; Tibia L O.56 f SVL. The specimen was cllected in mid August and had small varies with small unpigmented va. It was fund at night n the bank fa stream tgether with R. blythi. Rana rugulsa WiEGMA Figs. - Tuaran, lwer rhan 50 m, 58 males + f'emales + 4 subadult females; Penampang, lwer than 50 m, immature females.

34 ' /,re irg fi,,,... Amphibians frm Sabah L t-)' ee s l,, g,, li, g : f A B :, l ' x,, "i'i l }/ #/ g,wa tt ff pe. g fi g ttt in -- il, lfivag, wtw IE-. f g g - if: ge.#- emi.. gt.'-'g ew Fig.. Drsal (A, B) and ventral (C, D) views f Rana ru.gutsa frm Tuaran. A. C. male, SVL 85.5 mm; B. D, female. SVL 0. mm, This species, nt listed in IGER's mngraph duced frm Taiwan fr fd purpse. Taxnmic relatins f the Rana tigerzna lng been discussed (BuLEGER, 90; (966), is said t be recently intr- ' cmplex,includingthepresentfrm,have PpE, 9; TAyLR, 96J DuBis, 974),

35 6 Masafumi MATt ui i pt 'i glg E l i l v. lj', -ig.. ge,:- F, := fi- :,p. ;.i-li.\" I t.ttt..tt t.,./ t. IStmttltl.t'!t'. s.t/tt.m l;asgl/ll/l: lsl,l's/r.; i'i 'li,i: x-il,}t'ig/l.i',l;,als.,',s:- l.;l i,' '' Fig.. tt t tt tt : sllissi,'i ; illl.l, i I ' i ii,i.,il.'i-',.' ' Drsal (A)..:.trget./.,.. m, - -',l' ' '.i.l, B-.,..i.. and ventral (B) views fmale Rana ra,gulsa frm Taiwan, SVL 87mm. but n sufricient cnclusins are yet reached. The Brnean specimens examined had the characteristics f rugulsa r tigerina pantherina, having nly slight differences frm Frmsan specimens in relative tibia length (Table 4). I use the name f rugulsa rather than pantherina, since n significant difference is fund between the tw in the descriptins fthailand ppulatin (TAyLR, 96). All the Tuaran frgs had been sld alive in the city market fr fd. They were said t have been cllected in the paddy fields nearby. Twjuveniles, 4.5 and 47.8 mm in SVL, were cllected in August n grassland near human habitatin. Mst f the females btained in late March cntained enlarged pigmented va. Oeidzyga baluensis (BuLEGER) Bundu Tuhan, 80 m, fhemale + immature female. Table 4. Tuaran 8 Ad. 9 Ad. Taiwan 8 Ad, Cmparisns f bdy size and prprtins f Rana rugtttsa (l'm 'uaran and Taiwan. 58, SVL Head width Tibia length Range rvlean+sd Range Median Range Median { Å} t e r,

36 n" tt #ts/, AniPhibians,fi'm )9abah. 7 ttttt. %i,, "l'l ttt x{.ṃ, i 'tva fi'*.tx slr,. lz g ditt:-,--,st '$i :i,itk i!'t.e;,l,,,l.l,//l E, i nytj,twhk'g,,..,.m.,l/.-.'.i 'tt..,,,e,-. t su. ttt tt ttt I,., t 'i/'- ' l'/l/ : \,l Fig. 4. Ventral view f tadple f Antlps cavitympanum frni rvlamut River, Pring. St. 4, SVL. mm. The adult and immature females measure: SVL 8.9 and 5.5 mm; HW O.9 and O.400 fsvl; Tibia L O,P40 and O.50 f SVL, respectively. The bdy size in the larger female apprximates t that in females examined by IGER (966) ; sexual maturity in the females f this species is cnsidered t be attained when they reach abut 9 mm in SVL. The adult female cllected in late March had varies with a few small pigmented va and the smaller female had small unpigmented varies. t } Amlps cavitympanum BuLEGER Fig. 4 Mamut River, Pring, 550 m, larvae (OMH unnumbered). The yunger tadple in stage measures: BL I8.6 mm; Tt L 47.4 mm; Tail L.55 f BL. The lder ne in stage 4 measures: SVL. mm; Tt L 49. mm; Tibia L.6 mm. The labial teeth are III:8+8fl+ :V in bth specimens. The tw tadples have undivided upper beak, which differenciates larval cavitivmpanum frm ther Brnean AnzlPs tadples (IGER, 966). They well agree with the descriptin f cavit.ympanum in every respects. Metamrphsing lder tadple lacks uter metatarsal tubercle and its tes are fully webbed t disks. These tadples were cllected in February frm a wide river with strng currents.

37 x., :t./ ' 8 Masafinni rvlatsui,gee ewmt. x ig- "ec rt i}/ IS ll Fig. 5, Drsal (A, C) and ventral (B) views and prfiies frm Silau Silau Stream, KTP. A, B, D, immature female, SVI, 9. mm. (D. E) f tlmlps A'inabaluensis male, SVI. 57.8mmJ C. E, Amlps jerba CGtiTHER) Fig. 8 KP Headquarters, 665 m, female; Lidan River, Bundu Tuhan, 990 m, males; Murk midway betw. Ranau and Telupid, ca. 500 m, male. The three males with nuptial pads and vcal sacs measure: SVL mmi HW O.9-O,54 f SVL; Tibia L O.699-O.708 f SVL. The female measures: SVL

38 AmPhibians.flm Sabah I. 9 ",,. $i B c Fig, 6. Amlps sp. A. t'rm Liwagu River, KP. male. SVI..8mm, A, drsal view; B, ventral view; C]. prfile, 65.6 mm; HW O.4 f SVL; Tibia L O.709 f SVL. Males were cllected by day either amng pebbles and rtten twigs n rcks in a river (WT 0.6 C) r under stnes n a shaded mssy bank fthe river. Several males were seen clinging. n a rcky bank fa clear stream in jungle frest by day. The female cllected in mid March, cntaining enlarged pigmented va, was fund at ni,ght aln,g a stream in mixed ak frest. ' ' Amlps kinabaluensis IGER Figs. 5, 8 Silau Silau Stream, KP, 60 m, immature males + female; Gunn.cr Alab, 600 m, female (OMH unnumbered). The immature males measure: SVL 54.0 and 57.8 mm; HW O.74 and O.67 f SVL; Tibia L O.659 ando.678 fsvl, respectively. The tw females measure: SVL 8.6 and 9. mm; HW O.56 in each; Tibia L O.656 and O.649 fsvl, respectively. Tw immature males lacked vcal sac penings and nuptial pads, althugh the larger male is almst the same in bdy size with the smallest mature male (SVL 58. mm) described in IGER (l966). They have relatively larger tympanum (O.07-O.074

39 .',l,ig 40 Masafumi MArrst'i f SVL) than in the males f kinabaluensis described by Inger (O.058-O.065 f SVL'). Otherwise, the tw specimens agree with the descriptin fkinabaluensis. The females cllected in mid March and in mid December had large va with pigmented hemisphere, abut mm in diameter. Three frm Kinabalu were cllected at night n the bank fa shaded stream, abut ne meter abve the grund level. One female frm Mt. Alab was cllected frm radside wet place in lgged frest. Amtps sp. A Figs, 6. 8 Liwagu River, KP, 500 m, male. SVL.8 mm; HW O.40 f SVLI Tibia L O.70 f SVL. The male with a pair f vcal puches and well develped nuptial pad, has the blunt snut unlike knwn members fbrnean AmlPs (Figr, 8). It has distinct tympa- L m/m,'tsua.-... s,...,.ln Sa.l' wa ge'k, g,ect, as """"'i :sti'min' me llxts.liiw,fi,il'ik'kl::,,m ";ll/as'idi'../ew -#,ii 's :i,, ri s :tg ma gel,wm i.,,,m"ge'lt' sl "an.'vzpt. i.':i.",# "'? li Etk'k. ll'g:''/....l ug.te,-'..-ima"rm ' 'en ttttt t :.mepe..p...taewp.,,\''.,,, tlm 'eql.ta ee -//,L ' ttt t t,g #., m en..,cu:lfi! wrm me mpny eqma it/b"'tru A "' B ''' tt t..di ttt ' '.sll''di,, ' iw tl Fig. 7. AmlPs sp. B. frm Kambarangh rad, KP, immature male. SVL. mm. A. drsal view: B. ventral view : C:. prfile. g ' l' ilgsul,.' -,, m..ian

40 AmphibiansfrmSabah L 4 num differing frm A. cavitlmpanum. It resembles A. j'erba in the bdy size and in the presence futer metatarsal tubercle, but differs frm the latter in the fllwing characteristics: furth te with tw phalanges free f web, head wide (rati t SVL ut f the range cvered by A.7'erba), drsum cvered with small granules but nt shagreened, nuptial pad large, extending t disk n the first finger as a narrw fringe. Frm A. kinabaluensis, this male is easily differentiated by the furth te webbing (full webbing in kinabaluensis), presence f utermetatarsal tubercle, wider head, lnger hindlimb, and much smaller bdy size. It resembles "sp. B", described in the fllwing lines, in the pr develpment f the web, but differs frm "sp. B" in the presence f uter metatarsal tubercle. The male was fund hpping in a rainy daytime in mid March alng grassy bank f a clear stream. Amlps sp. B Figs. 7, 8 Kambarangh rad, KP, ca. 700 m, immature male + l immature female. The immature male and female measure: SVL. and 8.7 mm; HW O.80 and O.76 f SVL; Tibia L O.78 and O.679 f SVL, respectively. The tw specimens have superficial tympanum and are different frm A. cavit]mpanum. They resemble in bdy shape with A.j'erba, but the web extentin n the furth te is in the same cnditin as in "sp. A", i.e. tw phalanges are free f web in these specimens. The bdy size fthe immature male (. mm) falls in the range fmature A. j'erba males ( mm, IGER, 966 : 65), but the nuptial pad and vcal sacs are absent in this male. Further, these specimens lack uter metatarsal tubercles, sharply cntrasted with A.j'erba and A. "sp. A". Althugh A. kinabaluensis lacks uter metatarsal tubercle, it has full webbing n the furth te. The extentin f webbing is determined by the time f metamrphsis in AmlPs tadples (IGER, I966: 7), therefre the cnditins fund in spp. "A" and "B" are nt attributed t grwth vari-. atln. The tw specimens were cllected by day in mid August under stnes in radside ditches in lgged frest. AmlPs species frm extrateritrial lcalities have nt been examined s far and pertinent identificatin fthese specimens, as well as "sp. A", will be made in future. There are little pssibilities, hwever, that species "A" and "B" are parental species f either f the three Brnean frms f larval Amlps, whse parents are unknwn (larvae `CA", "B", and "D" figer, 966), since these larvae have fuy webbed tes in lder stages f develpment. Plypedates leucmystax leucmystax (BiE) Fig. 9 Papar, 0 m, male; Ranau, 504 m, males + 6 females; Marakau, near Ranau, 480 m, larvae + l egg mass; Pring Ht Spring, 555 m, female; Kundassan, 440 m, egg mass; Kambarangh rad, KP, 680 m, egg mass; Sepilk Frest Reserve, 50 m, female + 6 larvae + egg mass; Utara Rad, Sandakan, 7 males + 6 females.