www.vireya.net total knownspeciesoccurringonitsslopes. carries which and years the million of number 1 disproportionate than more much not of rapidly very giant' arisen having , for rule' the 'proves which cou of is Kinabalu Mt. building. mountain rapid and instability i with Guinea New of rocks bare jagged the with relatively contrasts peaks Borneo fewer, in form has land the Thus island mountains. declining the predominantly secondly inp genetic but broad fairly past; a geological meant has which and origin of centre where mainland Asian south-east the to accessible ra trs y w smwa opposi somewhat explain two by maybe terms broad situation This known. Vireya largest number of species of species of number largest section Vireyaasawhole. Abstract. p hs nta suy ae ih n niain rm h Sbh Na the from invitation an with came study initial this up opportunit marvellous several A made. that were observations and material collections this up working on them presented problems taxonomic only was It expedition. 'Mulu' Departm Forestry the of Par National Mulu Gunong the to author paper the by visit a this from came in to referred rhododendrons Bornean on observations first The Never 167. about has which Guinea t rich less much is species, Rhododendron 46 about with Borneo, taxonomic subsectional the of re-asse made Some preferences. habitat and species the of strategies possible hybrids, of observations field with Sabah and Sarawak eut ae rsne o sever of presented are Results Vireya RhododendronsinBorneo by G.C.Argent any island with about one sixth of sixth one about with island any INTRODUCTION ona rooedos ih about with rhododendrons Bornean n o te oa Gorpia So Geographical Royal the on ent selves and on a return visit m visit return a on and selves ruig wt implications with groupings hls Bre cnan te se the contains Borneo theless g atr: isl Bre is Borneo firstly factors: ng l eet olcig expeditions collecting recent al rse the exception the rse the 300 or so so or 300 the k as a guest a as k s 'young a as inl Parks tional ore careful ore have their have y to follow follow to y ts tectonic tectonic ts pollination set is ssment relatively relatively f green of a New han t n the in ut af the half l and old quite a o the for ciety's d in ed cond to www.vireya.net been solvedbythisstudybutpr probl taxonomic Some wild. the in observation casual to obvious w plants these of aspects several of study the enabled has This as successfully collected species the of many grow to Edinburgh facilitie excellent the with well which visits short several on observations and specimens gather to assistance field generous very with backed was This 1988). al,. et (Argent of rhododendrons the of account an to expanded subsequently was Kinaba of rhododendrons the of handbook a produce to Department ovary shape and flower colour, which all show quite remarkable remarkable quite show all which colour, flower and shape ovary shape, leaf the lobes, corolla the of overlap va of degree the are Other 1982). Ar (Argent, (sensu javanicum R. of populations status Bornean the in characters subspecific to reduced been has mo f and ovary glabrous of basis R. the on brookeanum R. from differs fact In Borneo. in places two least at in discovered been cha indumentum this in variation showing populations whilst G., teysmann var. javanicum R. in occurring hairs with down, breaks Bornean hairs simple of absence or Presence javanicum. R. in found size of dimensions of the most covers undoubtedly and species of group this variable of collections highly is shape Leaf process. submitted been have flowers the which to pressure of degree the lar a to flowers dried the of dimensions the and shrinks tissue corolla because herbarium the in with deal to character difficult very a is size and shape leaf ovary, the on hairs absence or Sulawesi presence including brookeanum, R. from javanicum R. Bali, Java, t although Sumatra, separate to used are characters of number Borneo A island. the encircles Philippines) in Malaysia, occur (Peninsula not does distribution javanicum R. (1966) Sleumer f Hook. lowii R. and Sleumer retivenium R. Ridley, moultonii R. brookeanum R. particularly Benn., (Bl.) javanicum R. to related a problematic very been have that species of group first the Of TAXONOMIC PROBLEMS poie b te oa Botani Royal the by provided s obably asmanynewoneshavear flower size, but none are cl are none but size, flower ge extent reflect reflect extent ge re those closely those re ear cut. Flower cut. ear leaf venation, leaf Low ex Lind., ex Low . According to According . variations that or this reason this or hich were not were hich the range of range the on the ovary the on living plants. living in the drying the in Gre in Garden c ii (Miq.) K. & K. (Miq.) ii isen. atr have racter gent, 1982) gent, loi only ultonii combined combined m have ems of simple simple of lu which lu Sabah and riable he www.vireya.net are often correlated in local in correlated often are ebru. hs a a isac w instance an was This herbarium. c good sufficiently a as accepted wh questionable, slightly already th in indumentum Ovary identity. flower scented its and indumentum is lowii depen retivenium R. but ecology montane its of because R. isolated group, problematic this in species two last the Of o many on obtained is information more populations itatleastgivesapra until but problem the of t of nature resolution satisfactory totally island a not is This restricted. are they the which to ground of because isolated absolutely more specie montane much the for than being, time the for least units at taxonomic taken, the of view broader much a reason t this for are and there lowland, mainly isfo the in intermediates species potential of group this of ecology form. leaf different totally but shape flower same the exhibits fro plant a but lobes corolla overlapping and quilted elegantly from Argent (Ridley) moultonii dist the Thus Borneo. of whole the ey ag etn, ih h scent the with extent, large very aspe general ecology, with well correlates It leaves. expanding of aspect the but bud) the in leaves the of folding (the pyxis edges. upturned or straight with expand which rel leaves have other which the all unlike manner revolute a in leaves new the of i character This species. maintainable highly and distinct a as redefined has and reliable very proved has which character new okn fr ebru seies ih ny aue evs present leaves mature inci represent may which forms corolla two least occurring at and colours species only widespread another with is Lindley longiflorum specimens R. herbarium not of for limitation the have working does it although herbarium the in even species alway will It growth. of stage right the at material cultivated populations but which do not corr not do which but populations G. Api in Sarawak has short sulc short has Sarawak in Api G. rests between the different si different the between rests haracter, even in thelivings even haracter, ctical interimsolution. is group, for the reasons state reasons the for group, is lt h peec o set cou scent of presence the ilst d lwr, n i vr es t easy very is and flowers, ed ee utvto o te plants the of cultivation here ntv fr o R brookeanum R. of form inctive fr h mitnne f sep a of maintenance the for s tate, letalonethe s distinguish this this distinguish s tes of collection of tes s the expansion the s Given that the the very young very the elate well over well elate ate leaves and leaves ate pient evolution pient Kalimantan m d above, was above, d almost totally totally almost R. retivenium retivenium R. ded on ovary on ded td species ated t n, o a to and, ct wih are which s rvdd a provided This is not in several several in d o be not ld a been has se in see o herefore e high he subsp. the f arate . www.vireya.net enabled himtoseparatethesespec ut aue rfso Sleumer's Professor salute must flowe the which collections that with ho sympathize so wild the occasionally in sheet collected have a or who Those on erect. down stood apparently down hung slapped flowers been the had that praetervisum wilted badly so been pressed obviously were longiflorum R. of Specimens lost. often of posture the amazingly quite and reported, accurately rarely ecological notes were too imprec too were notes ecological t in However, flowers. hanging vertically displays praetervisum f its holds longiflorum R. whereas and colours flower different zone altitudinal different in grow They confused. been ever had wa field, the in studied were species these when surprising was res long for had specimens name whose under longiflorum R. from separate neatly herbarium the in indum which pistil the discovered in characters Sleumer 1973). (Sleumer, species w the Sleumer described Professor by christened this with synonymy to reduced hav Becc. subcordatum R. and Sleumer praetervisum R. mention to needs more investigation. Whils investigation. more needs of hybridization between R. longi R. between hybridization of eas now is species the Borneo (Argent, species this with synonymy to reduced been have Becc. subcordatum R. and Sleumer commutatum R. pollinators. different favouring rsioim tp wr goig n bnac coe y n i lo it and by close abundance in growing were Stapf crassifolium stenoph R. cases these all in 1981; and 1980 in locations other were flowers varying widely rather but morphology leaf similar colle isolated Subsequent Edinburgh. RBG the at grown been have of area Cave Mesilau the from collected first spe were description rare a be to considered on Parai Marai from described was nervulosum R. Argent. ex (Sleumer) R. problems and Stapf interesting ex f. raised Hook. stenophyllum has R. Sleumer, nervulosum which species of group Another Kinabalu. Specimens agreeing with agreeing Specimens Kinabalu. ily identified, although there is some evidence some is there although identified, ily t dealing with R. longiflorum it longiflorum R. with dealing t florum and R. brookeanum in Sa in brookeanum R. and florum ise to be of use, flower colour flower use, of be to ise euiul ciia detective critical beautifully r smtms es hn perfe than less sometimes are ies fromherbariummateriala is perhaps best perhaps is lowers erect, R. erect, lowers the flowers was flowers the d this species species this d collected from from collected was lost and lost was e herbarium he iaau and Kinabalu lone. yllum and R. and yllum s; they have they s; 1982) and in and 1982) ok which work, rawak which rawak after having after the original the s that they they that s ided. What What ided. ee, will wever, t u one but ct ctions with ctions kd very oked uberans e he hen been e entum s R. is cies R. rs www.vireya.net strongly imprintitsprogenywith alw and species several with hybridize easily rather to s appears R. nervulosum. R. 'typical' from hybrids these separate to be occu it if it with hybridize must ste R. of those to flowers identical almost has It common. very this of variety a as mountains, of number a on occurs exuberans R. 1963). (Sleumer, described formerly was and nervulosum, R. t into intrudes also exuberans R. enigma. this solve to done be stenophy R. with crassifolium R. of crossing artificial the and o work field Further evidence. in nervulosum x R. of population hybridized been has but existed t or undetected, yet as remains but examined sites the by close o part on occurs stenophyllum R. either that speculate must one of popul theory the maintain to and species the a like more much behaving Lotong G. on hybrid, a was nervulosum R. that clearly onan Ti psd problem: a this posed from This recorded been mountain. has or seen was stenophyllum R. no but present population rather than isolated i isolated than rather population Lo 'specie G. this Here in Sabah. southern studied and collected was nervulosum R. however, 1982, species two these between hybrid a was nervulosum R. if as much ra f he ajcn mutis i mountains adjacent three of area rat a in occurs that malayanum R. of form montane small-leaved, spe full a than rather microspecies a be perhaps should Sleumer classifi hierarchical neat making to seems variation but taxa subspecific seven recognizes (1966) Sleumer Seram. and Sulawesi to on and Borneo Java, Sumatra, to Malaysia thr down Thailand from distributed is It taxonomically. species un very are species the is itself the Jack malayanum of populations local while scale, broad encoun is variability great group this In are Sleumer. with deal Malayovireya to wish I that species problem of group final The most widespread and probably the m the probably and widespread most rs in close proximity. The pro The proximity. close in rs ndividuals; there was plenty o plenty was there ndividuals; its distinctivenarrowleaves s' was to be found in some num some in found be to was s' ain mosbe R micromal R. impossible. cation Bre. . aaau itself malayanum R. Borneo. n u o eitne n lae onl leaves and existence of out hra al h sgs elsewhere signs the all whereas . blem would then would blem f R. crassifolium R. f llum remains to remains llum ough Peninsula ough nophyllum and nophyllum he problem of problem he ays seems to seems ays be reticulate, be her restricted her hybrid origin hybrid ee o the on tered G Lotong G. n n subsect. in tenophyllum but is never is but f G. Lotong G. f cies. It is a is It cies. a i once it hat ost difficult ost to was ation can look bers as a as bers fr. R. iform. . Early in Early . ayanum species og in tong were the y www.vireya.net far the highest peak. The vegetation in the upper montane zones montane upper the in vegetation The peak. highest the far in mountain Bornean exceptional the in interesting is It parents. both exist intermediates of v populations great with numbers cases substantial two last these In Sleumer). Malay other the of most not species. if many, up swallowing of danger in lump 'a by concept its broaden to tendency any although species coriifolium Sleumer) and R. abiet R. and Sleumer) coriifolium Hoo no ex Low most rugosum R. Kinabalu, and f. on Hook. ex found Low buxifolium R. be between to swarms hybrid clearer much the ref already Lotong G. on nervulosum x R. of case the are Borneo exceptio Notable species. the of purity relative maintain which enable may work careful Further in itslowlandkerangas(heathforest)formwhichhasenormous str looks but inflorescences) different rather has (but montane ran its of end western the in micromalayanum R. to similar very rsmby hr ae selective are there Presumably appear tobetransient,anddonotusuallybackcrossgivehy larg in plants occasional just as occur usually they but Borneo par its of that between morphology hybr the and vicinity immediate carefu the to obvious very are, b can they wild the in but herbarium Vir the in detect to difficult Hyb Borneo. any like area an of proximity the that within least at group experiments species other hybridization any with crossing of capable appears rhododendron artificial numerous from W made. be can phenomenon this on observations general some and hybridization and hybrids to made been have references Several that Kinabalu, as has already been mentioned, is mentioned, been already has as Kinabalu, that osre. h prns usually parents The observer. l ehnss prtn aant suc against operating mechanisms d ln i gnrly lal in clearly generally is plant id ifolium Sleumer and R. buxifolium (x sheilae sheilae (x buxifolium R. and Sleumer ifolium rain niaig akrsig rbby to probably backcrossing indicating ariation R. malayanum to be split into s into split be to malayanum R. HYBRIDS ents. Hybrids are in fact qui fact in are Hybrids ents. that it is actively growing actively is it that e, and generally and e, brid swarms. ikingly different ikingly - it is also by e populations, e leaves. te common in common te ge where it is it where ge emdae in termediate is r very are rids erred to, and to, erred ns to this in er' would be would er' everal more more everal rw n the in grow is still in a in still is within the within wild the in hybrids h ovireya e know e . . (x f. k. tably eya www.vireya.net rbby olnt te og white long, the ani pollinate of probably groups different quite two attracting apparently flowers a would it but explanation complicated a seem may for This but insects. nectar the for primarily not , by sought then are climate, unpleasant an be can what from corolla the in shelter a scent the by attracted are insects that speculate to tempting t cannot and smell of sense no have visited frequently blacke mountain the are by especially flowers Its nectar. copious produces and stro a with scented is it Furthermore, landing-platform. insect colo broad, with tube corolla medium ( red bright Although categories. pollination standard tpinm ese e Pan n R saeln Semr u there but Sleumer suaveolens R. and Prain ex Hemsley stapfianum rhododendrons b handsome from most Apart buxifolium, Kinabalu's interest. R. of also tube. is species, the flowered red around another disposed regularly or ventrally grouped other All flower. the of mouth with Argent which yongii flowers red R. dark handsome of that is discovery notable most the perhaps paper St Since scent. strong and parts of numbers increased corollas, absent although Guinea, New for (1982) Royen van or (1976) Stevens by described been have which types fo found be can Parallels rhododendrons. Bornean of case the in specific groups of pollinators is pollinators of groups specific Pollination syndromes or the gener the or syndromes Pollination ar bare large the and cap ice an of disappearance recent the to due stabilization and colonization of process oeta fr rdcn nw pce t fl te aum f lar a of vacuum the fill environment. to species new producing for potential o state a in still are species altitude high these that appears dist artificially that way same the in hybrids of establishment POLLINATION SYNDROMES red flowered Bornean species hav species Bornean flowered red something which we have few o few have we which something spreading corolla lobes which p which lobes corolla spreading eas of the granite dome appear dome granite the of eas ye (Chlorocharis emiliae Sharp emiliae (Chlorocharis ye have the anthers dorsally arran dorsally anthers the have al associations between flowe between associations al tblr lwrd pce su species flowered tubular , herefore be attracted by the by attracted be herefore r bttps ih thick with bat-types are i de nt ety al no the into fall neatly not does it , urbed areas do. It do. areas urbed f flux with a high a with flux f ng sweet smell smell sweet ng and the flowers the and bservations on bservations nd nectar, find find nectar, nd h sheltering the rovide a good a rovide evens' (1976) (1976) evens' e them either them e r, t a a has it ur), r most of the of most r as Moths mals. to favour the favour to ig n of one eing e), but they they but e), r types and types r scent. It is It scent. y birds, by ged in the the in ged ey open gely ccount for for ccount h s R. as ch & Kores & fleshy , are no www.vireya.net oh rbby iie b iscs Sl pliain ol appear although wehavenodirectevidenceforthis. would pollination Self insects. by visited probably both brookeanum R. as such species th in common is colour flower yellow Guinea, New in As Sleumer. and R Sleumer polyanthemum R. crassifolium, R. as such anthers, the Sabah: hav in those favouring particularly observa made Several been have Borneo. rhododendrons for visiting butterflies this confirm to observations direct uscin Frhr ok is a work tubercles Further epidermal subsection. on sit not do they that except Guinea New Phaeovi subsect. of those to similar hairs stellate or dendroid for proposed is Dendr 1984), al., ser. et (Argent grouping, Phillipps & new Lamb Argent, the but groups main four into analysis Sle followed broadly has type scale into divisions The species. many on information new given have microscope electron scanning studie and prominence due given been have hairs) (multicellular salver-sh epi The shape. corolla using on without defined uniformly be could based the all that F., so abandoned been Copel. has corollas, (hypocrateriform) Solenovireya Subsect. island. the relat of taken is cognizance little very as particularly (1966) tha tha on advance real no perhaps stressed is which arrangement provisional be must it rhododendrons Bornean the regrouping In exerted onthischaractermaking be may which pressure selective strong the given questionable also is level at shape flower of use the and division arbitrary an lea than the more ser. of size into the on purely (Euvireya) Sleumer Javanica Vireya ser. and subsect. Sleumer of divisions The species. stenophyllum R. putting of anomaly atten drew (1978) Woods natural. means no by are (1966) account in found as rhododendrons Bornean of groupings subsectional The TAXONOMIC GROUPINGWITHI edd o sals te relationshi the establish to needed ing bright red or orange flower orange or red bright ing it insufficientlyconservative and R. nieuwenhuisii J. J. Smith J. J. nieuwenhuisii R. and with three totally unrelated totally three with N SECTIONVIREYACOPELANDf. . s with prominent with s osis outside ionships reya Sleumer of Sleumer reya umer's (1966) (1966) umer's dermal scales dermal New Guinea Guinea New pce with species subsectional o e rare, be to t of Sleumer of t subsections . fallacinum between p Sleumer's wt the with s tion to the the to tion which are which lowland e e i no is ves s in that that in s olepidon Buxifolia insects in of tions i i a is it t f the of aped www.vireya.net ar o nt O te ona seis xmnd al hs i Sle in those all examined, species Pseudov and Malayovireya Bornean subsections the Of not. or with fringed hairs are bracts) sterile or (perulae scales bud floral the whether is first The useful. particularly be to thought are which observed characters new cultivation, of number large a observing While subsec of subsects VireyaandDendrolepidon. species The Phaeovireya. scale to according distributed been have f. Copel. Solenovireya and Dendrolepidon in species like centres with broad peripheral broad with centres like point- small have scales The Dendrolepidon. and Vireya series in as division, st the of examination close Even proposed, which isagainsubdividedintotwo. are groupings major two species Bornean the for Thus de of base the at cataphylls shoots. subulate having most at or plants) manner naked a in developing young shoots the absent, virtually are scales bud Pseudovireya and Malayovireya r prominent exhibits generally which Vireya in developed well are These scales. bud deve and buds of vegetative lack or development, general the is character this with 'scales with fringed often are they (although not do buxifolium Solenovirey and Vireya subsections in those all whilst hairs of nat more a in species the grouping Section Vireya

Pseudovireya Subsection Vireya have been constantly sought that might help with help might that sought constantly been have

esting buds with sheathing bud sc bud sheathing with buds esting

ructure of the epidermal scales epidermal the of ructure flanges and it is really a m a really is it and flanges

the Bornean species over several years in years several over species Bornean the rl aho. w characters Two fashion. ural wih s o rr aogt tropical amongst rare not is (which ireya (Clarke) Sleumer have Sleumer (Clarke) ireya

Dendrolepidon Malayovireya Pseudovireya Vireya Series

atter of degree of atter a except for R. for except a

iue simple minute type between type ales, whilst in whilst ales, supports this supports ) Correlated '). omn, of lopment, ae been have a fringe veloping ah of each umer's t. www.vireya.net suoiea n bod n Malayovir narro being in this - broad flange and marginal the Pseudovireya of development of degree dif again and cushion central broad a have cases both in scales the Malayovireya and Pseudovireya series In Dendrolepidon. ser. in as stellate / dendroid or Vireya ser. in as lobed are flanges these whether New Guinea.Vaduz.(Reprint fromV van Royen,P.&Kores, P.(1982).TheEricaceaeofthehighmou National ParksPublication. ------, ------&Collenette,I.S.(1988).Rhododendron Rhododendrons fromSabah(Borneo).N ------, Lamb,A.&Phillipps,( Ericaceae. J.Linn.Soc.Bot.85:2-17. Argent, G.C.(1982).Newtaxaandco na reasonably and workable both classification canbeachieved. a species, more on observations the in Vireya within groupings t on light more shed may chemical hoped is It section. unhappily rather resides Pse it ser. where to transferred been has species this reasons two these central large have they small, very although that clearly shows examination Close hairs. simple short in covered partially face having of rhododendrons) Bornean for least (at feature unusual relevanc What scales. the in has Borneo outside have groupings structure of similarity underlying between difference the sca exaggerates of numbers the of development hm o e aty bt o fully not (but partly be to them shows scales bud floral the of Examination lobes. corolla broad with flowers ho at appears superficially it where (Vireya) Euvireya subsect. Sleum by placed is It buxifolium. R. is scheme this of neatness spec The tentative. remain they 1984). Newtaxaandcombinati future and, together with furthe with together and, future REFERENCES figd ih ipe ar and hairs simple with fringed ) ies showing greatest potential to upset the the upset to potential greatest showing ies not yet been studied and even and studied been yet not ih h sal lwrd speci flowered small the with an Royen,P.TheAlpine Flora les between these last two groups groups two last these between les hm u tee s neverthele is there but them otes RBGEdinb.42:113-120. mbinations andcomments y. h ata dfeec i difference actual The eya. s ofSabah.Sabah me with itslarge me ons inVireya r morphological r cushions. For cushions. of the scales the of o ae the have to ntains of fer as to the to as fer r 16) in (1966) er ofNew for Borneo for h abaxial the e various he s f that of es individual on udovireya these e

s an ss in w tural n www.vireya.net Sleumer, H.(1963).FloraeMale Guinea 3:1485-1911). 37:157-159. Woods, P.J.B.(1978).TwoBorneanRhododendrons.NotesRBGEd and theirsignificance.J.L species, especiallyinthePapuasian types inRhododendronsectionVireya, Stevens, P.F.(1976).Thealtit Malesia (Ericaceae).Blumea21:357-376. ------(1974).NewspeciesandnoteworthyrecordsofRhododendr (ser. 1)6:474-668.Groningen. ------(1966).Rhododendron.InvanSteenis,C.G.J.(ed.), 114. notes towardstheknowledgeoft Reproduced fromNotesRBGEdinb.43(1985) with thekindpermissiono Royal BotanicGardenEdinburgh inn. Soc.Bot.72:1-33. udinal andgeographicaldistrib sianae PrecursoresXXXV.Supplem he Ericaceae in Malaysia. Blumea 12:89- he EricaceaeinMalaysia.Blumea

f theTrusteesof ution offlower FloraMalesiana on in entary inb.