www.vireya.net these factorswouldseemtohav clim quaternary and geography present of light the in discussed Distribu Malesia. of rest the of that to compared when Celebes, geographic Preliminary f Rhododendron species. Papuasian the of richness the Malesian emphasise analyses West in variation floral of characteristics main Papuasi striking most the of two of absence complete almost and colour, flower in variation infraspecific notable types, corolla lobed deeply of preponderance moths birds, be to seem Papuasia in Rhododendron of pollinators a Malesia, throughout markedly, less although occur, also which con flower and this colour flower of changes The group. the of evolution and studies, phylogenetic for unsatisfactory are group t of Most chromosomes. apparently vegetativ of characters in uniformity considerable by underlain Rhododendro Great of features. vegetative and indumentum, floral, in primarily species Malesian between variation The ABSTRACT. by P.F.Stevens Towards AnUnderstandingOfTheirEvolution Malesian VireyaRhododendrons- lns I i lrey geographica largely is It plants. fascinating a is Vireya sect. Rhododendron subgen. Rhododendron u lc ohr utclua his Semr 16) te corolla the 1966), tails long (Sleumer, have seeds the hairs 1976), (Stevens, markings multicellular zygomorphic other lack but to tax All 4000m. over to belonging sea-level from growing 1), (Table occ region Vireya species sect. of six species c.297 the only of 287 whereas are subgenera, there Malesia dis is In area separately.) Celebes-Moluccas The Islands. Solomon the including t islands the and Guinea New includes Papuasia and Philippines; Born Islands, Sunda Lesser the to Malaya from area the Malesia, Islan Solomon the and Guinea, New Philippines, the fro to area Peninsula the includes hab Malesia paper, subalpine this and (In Malesia. montane tropical in mainlan diversification Asia great southeast the very on showing growing species few a only having e hadadifferenteffectinthe he subsections and series curr series and subsections he INTRODUCTION l sprtd rm h rs of rest the from separated lly two halvesofMalesia. diversity in these characters is characters these in diversity type with altitude in Papuasia in altitude with type atic changes in the area; both area; the in changes atic aaoy fut se, and seed, fruit, anatomy, e in n dvriiain are diversification and tion e ulnd Te or main four The outlined. re , butterflies, and bats. The bats. and butterflies, , tan dsuso o the on discussion strains ently delimited within the within delimited ently n lwr ye, r the are types, flower an lora, and also that of the of that also and lora, sc. iea is Vireya sect. n a have scales, have a te Malay the m h genus, the eo, and the o the east, the o t either at r n that in ur group of group s West ds; tt in itats cussed lacks other , yet d, www.vireya.net group may also be distinctive be also may group pigment Corolla mesophyll. the in embedded are bundles vascular structur its of details & 1951: Keenan Hayes, 1888; (Breitfeld, multilayered and mucilaginous least pointedthere),theepidermis/hypod at are they (or end Total Euvireya series Solenovireya Albovireya Malayovireya Phaeovireya Siphonovireya Pseudovireya Subsection e the in although stands outisourignoranceoft highlighted, be to problems evolutionary some Malesia. montane in changes vegetational and geological recent di and distributions considering and diversity, high especially emphasi Malesia, throughout species of distribution the section discuss the dive in types the flower on of significance observations ecological summarize possible shall I species, its delimit th subdivide to used characters the of discussion brief a After o similarity the against out stands characters. group the within divers Thus characters developed. well not are hybridization interspecific featur vegetative and floral in

Saxifragoidea

Linnaeoidea Stenophylla Buxifolia Taxifolia Javanica Citrina

Table 1.DistributionofRhododendr

Malesia

West 15 11 98 40 16 1 1 1 8 5

Moluccas Celebes- Celebes- 34 15 4 1 2 4 4 4

ae o wl udrto) ad h smaller the and understood), well not are e

(Spethmann, 1980). Despite the gr the Despite 1980). (Spethmann, he evolutionofthegroup. Papuasia s ht sal ics, genetic discuss, shall I that es 163 32 23 12 26 39 14 3 1 6 7

Mainland SE Asian 8 2 6 on sect.VireyainMalesia.

Australia 1 1

Total 296 87 43 12 38 14 15 41 32 1 4 1 1 7

ermis complexis

est against versity scin and section e I hl then shall I . zing areas of areas zing ares to barriers t o some of ity eat diversity eat This allows This st and rsity Cowan, d what nd s in the the in s other f www.vireya.net levels inthesection. taxonomic all at uncertainty underlying this to due partly is changes) patristic evoluti the discuss to order In R.javanicum =WestMalesia/Celebes R.zoelleri =Papuasia /Moluccas R.zollingeri =Celebes /WestMalesia R.malayanum =Celebes /WestMalesia R.quadrasianum =Celebes /WestMalesia R.lindaueanum =Papuasia /Celebes Notes. hs caatr. t a b diffi be can It characters. proble these are there yet taxa, these separate differences character subsect delimit shape; and size leaf and type, to corolla type, scale were series Sleumer by used characters of sets three The caut subsequent My group. re the within phylogenetic about uncertainty the convey to also and group, variat important taxonomically the of idea an give to both not, b s and Vireya, sect. subdivide look to used been have that characters shall I evolution. understanding to view a with designed classi its understanding; an such from far Howe are we Vireya, sect. taxa. the of relationship the understand to begin then can one hypothesis, an such With characters. derived unique, of possession their by recognized ideally lineages various the with phylogeny, its of hypothesis is, includingallandonly known sub monophyletic largely and coherent seem all podium, little a incised deeply its with Kores), fide - hooglandii R. excluding leuc R. (including Phaeovireya subsect. and scales, of covering yellio sizes R. two (excluding of Albovireya scales dark dense, its with Malayovireya, subsect. Albovire (atypical yelliotii R. to (Pseudovireya) saruwagedicum recently (1982) Royen van & Kores portions. central the of size n or Siphonovireya or Pseudovireya subsects to belong specimens t has then one and 1978), Stevens, 1973: 1966, (Sleumer, scales TAXONOMIC BACKGROUND on of a group satisfactorily, on satisfactorily, group a of on descendants ofanancestral sp i ad . auclcru) wt is dense its with lagunculicarpum), R. and tii ut o eaae nie rm s from entire separate to cult ion in discussing evolution discussing in ion e should have an have should e scales, borne on borne scales, basically, single basically, fication was not was fication ecies. ome that have that ome ya). However, ya). (cladistic and (cladistic ihl incised lightly o wti the within ion sections, that sections, eue R. reduced ify t the at riefly s ih all with ms ggs but ogigas; o decide if decide o lationships e, within ver, subsect. , ot on the on ot os and ions www.vireya.net prdcly n te gop (s Malayovireya) withoutbeingaccordedtaxonomicrecognition. groups other in leaves narrow els sporadically similarly nothing and shape, have leaf their Stenophylla from ser. apart common of species Bornean and th Guinea 1978) Woods, 1976; (Stevens, noted been has As 1982). Royen, van & Kores in composition series in changes also see 1978; Woods, 1981; fro Buxifolia ser. of separation at useful doubtfully be even may and character, key a as simply species, discriminating in help to 5 length the and recurved, f margins the linear, to orbicular be may lamina the long, very may petioles - considerable is shape and size leaf in Variation ahr hn caatr ht nec that character the a up than dividing for rather used character key a is shape and size Leaf Siphonovireya and subsection inparticularismonophyletic. Solenovireya flowe other are subsects so distinctive, distinguishes that type tubu narrowly long, the although However, characters. taxonomic furni and below, discussed be will as rhododendrons, Vireya the gre show characters associated and colour, size, shape, Corolla aeu suy f vn hs 'lsi' hrces il doubtless import will taxonomic of information characters 'classic' these even of study Careful e bqios n h gop other group, the in ubiquitous be comple hypodermis / epidermis distinctive the Although used. be rget eoe uig n fw s few a in fusing before fragment fr trace one be to section thee Philips and Philipson variation. nodal Even 1888). close Breitfeld, (cf. and interest of quadrasianum be may R. zoelleri) in open are they the (e.g. of bundles shape midrib the petiole, and stem in sclereids of distribution m ser. Javanica (Sleumer, 1973; Stevens,1976, Javanica (Sleumer,1973; ser. m om a single gap, although the tr the although gap, single a om when used at higher levels it u it levels higher at used when tps ad t s nla wheth unclear is it and types, r 150mm. Although there is much in much is there Although 150mm. ance, but there are other charac other are there but ance, n 16) osdrd h basic the considered (1968) on bet Pedvry, Phaeovirey Pseudovireya, ubsects saiy rns eae species related brings essarily eis Tre ude wr fou were bundles Three pecies. set o te ntm, such anatomy, the of aspects ace might briefly might ace structure shows structure sually functions functions sually be absent, or absent, be sh very useful very sh lat to strongly to lat r h former the er at variation in variation at ters that may may that ters eil and petiole yield more more yield tutr in structure x seems to seems x ht a in as that, the leaf to to leaf the together. together. variation, a flower lar d n the in nd crop up i R. in d as the the as New e and a in e is www.vireya.net Brighton, 1972). grou the in uniform is number chromosome that suggests evidence trees almost are others whilst form and small are some these, (of branches erect have species va also gravity to stems the of reaction The horizontally. less he usually are leaves seedling although 1985), (Rouse, seedling leaves erect has saxifragoides Rhododendron held; are they and how flush, the along disposed are leaves in the how variation flush, per much is there is growth there pattern, basic Philips this 1981; within (Stevens, However, inflorescence the below leaves of axils replac by overtopped are and stem, Inflorescences way. similar a in grow all rhododendrons Vireya bey R. of petiole the in bundles ha I culminicola. R. and leucogigas, R. commonae, R. of petiole sfl n itnusig between distinguishing in useful may size Fruit similar. superficially be to tend they rho although Vireya of seeds the of study detailed little been has There stro are leaves the and innovation growth pseudoverticillate. each of beginning the herbarium in however, stem, the r lack to and r less or more scattered buds being leaves expanded entirely, leaves terminal vegetative naked completely have to at Edinbu at himantodes leaves R. of plants Young reduced flush. the of inconspicuous beginning and few having Pseudovireya and e some are there that subsects in species some with bud, vegetative the in characters suggests also work Argent's shrubs. small unor generally are be stomata Foliar should observations these genus, the of classification major In intranervatum. R. of that in blade. t to parallel orientated are quadrasianum R. and taxoides R. as ) and others have spreading branc spreading have others and ) view of the importance of nodal of importance the of view erinckianum and R. citrinum, an citrinum, R. and erinckianum specimens there are a few reduc few a are there specimens ettd bt n narrower-leav in but ientated, aa eg Semr 17) Pre 1973). Sleumer, (e.g. taxa mn sot aiig rm bud from arising shoots ement anatomy in the ries, thus some thus ries, oeie be sometimes d five bundles five d egularly along egularly Malayovireya Malayovireya he axis of the the of axis he ve seen three seen ve d aa such taxa ed hes and form and hes terminate the terminate dodendrons, ed leaves at leaves ed rgh seemed rgh hummocks, extended. d oe or more ld much how p (Jones & (Jones p n 1985). on, even as a as even s in the the in s liminary educed xciting the ngly www.vireya.net itntv got fr. o nld te et aein Euvireya Malesian question critical the approach West We relationships. about the ignorance our of expression reasonable include To wo group serie this in citrinum) R. excluding (possibly rhododendrons form. monotypic growth the in distinctive placed a with rhododendron Euvireya saxifragoides, Papuasian a just is Saxifragoidea, R. Thus coherence. have that subgroups into divided be cannot stage this sin at which a form well, as Siphonovireya subsect. of However, species Papuasian discussion. biogeographic and evolutionary ensuing the and abov Phaeovireya, delimited as (all Malayovireya, Albovireya subsects paper, this of purposes the For oor b 20m Sees 17,i. A, u tee s defini a is there f but 2A), other 1976,fig. with (Stevens, 2000m species by colours than numerous more flowers proportionally becoming greenish or orange yellow, flowers, red-coloured have these of 24 3000m; above or at only grow spec colours other of flowers nine only 3000m; above grow species red-flowered 50 nearly elevations. higher at commoner being corollas red with species Papu in especially and rhododendrons, Malesian in colour flower between correlation clear a was There yellow-orange. basically basic red, basically into colours flower divided (1976) Stevens questions. l us tells taxa supraspecific separate recognized currently the fl these of distribution The evolve? types flower these of some her point the to particularly and Also, relationship? Papuasian monophyletic, Solenovireya subsect. Euvire the in species other to than zoelleri) R. (e.g. Papuasia compar to phyletically related closely more javanicum) R. (e.g. funnelif large, with species the are instance, For Papuasia. and Malesia West in species between FLOWER TYPESINVIREYARHODODENDRONS much above this altitude. About 2 About altitude. this above much f h ntr ad ubr f the of number and nature the of orm, orange-yellow corollas in West Malesia Malesia West in corollas orange-yellow orm, e) are tentatively accepted as accepted tentatively are e) with a comparable direct We direct comparable a with Rdfoee seis r cl are species Red-flowered . ya group; and is and group; ya e, how often did often how e, ittle about such such about ittle able species in species able 8 species grow species 8 useful units in units useful ally white, and white, ally ower types in types ower uld also be a be also uld the rest have rest the phylogenetic relationships st Malesian- st In Papuasia In liue and altitude particularly e ek of peak te ol then could sa with asia, l group gle oe of some l the all e with ies lower early s www.vireya.net n diin atog tee s gr a is there although addition, In (Stevens, 1976,fig.3B-D). d be not would 3000m above abundance of peak secondary any that means below) (see 3000m above habitats t non-volcanic of althou absence for area, relative Borneo-Sumatra holds the correlation for and general region same Celebes-Moluccas The 3000m. above abundance and has corolla lobes about half the length of the corolla; it corolla; the of length the half about lobes corolla has and fu less or more also is type flower yellow-orange commonest The but appearstobeabsentfromWestMalesia. Celebes-Moluc the from three and Papuasia in species 11 perhaps repr is type flower This 5-merous. than more themselves flowers the and thick rather be to tends corolla the lobes; long has and scented, and co other The Malesia. of rest the seven and region, Celebes-Moluccas the in six, perhaps or ty four, this of flowers have Guinea New in species 30 About scented. a thickness unexceptional of walls has corolla the tube; narrow type flower common very Another pe altitudes. is this be; may sm second in occurs frequently it because the that suggests comm.) (pers. red-colou Kores by shown corr correlation particularly general the not than is other altitude, type flower first The Malesia. Western t throughout species some in and Celebes the in species several t flower This corolla. the of side one on placed tube, be may stamens the of zygomorphy any if little show that flowers similar red, with species c.36 flowers. are scentless zygomorphic, there below), see - exception single a but (with alone Papuasia in and Papuasia, types In altitudinally. or geographically flower such region, whole hr ae n diinl .0 pce with species c.40 additional an are There mmon white flower type is larg is type flower white mmon all leaved species that tend tend that species leaved all eat variety of flower types types flower of variety eat r nt itiue a rno e random at distributed not are s ht ih wie ool an corolla white a with that is may or may not be nd the flower is flower the nd to grow at high throughout the the throughout e, funneliform, funneliform, e, lhuh the although ype occurs in occurs ype nnel-shaped e flowers. red ltd with elated r ih in eight or sne by esented cas region, cas e et of rest he e a do as pe, a long, a d tubular, tubular, etected h the gh rhaps ither he www.vireya.net uua foes n wie long-t funnel-shaped white, and have flowers species tubular of majority wi species flowers; coloured variously long-lobed, the campanulate, Malesia, Western In ty this of flowers flowered versionsofitarerare. have Papuasia from species 10 About scented. nectar source for such birds, but details of the general behavi general the of details but birds, such for source nectar ver a form may generally Ericaceae The Guinea. New in altitudes nic nectar-eating the that shown (1981 Beehler there. occurring often so conditions cloudy cool, to become will insects many whereas temperatures, all at active 300 h at above predominate to pollinators occur Oreornis) ten the of theory general the with agrees and This 1976). (Stevens, Myzomela Melidictes, (Ptiloprora, repres which of Meliphagidae, the pollinat by bird-pollinated probably are particular types flower tubular red with common types flower Papu particular from coming of mostly association and tentative albeit evidence, is There altitudes (Stevens,1976). are notfo tubes corolla long particularly and flowers red with expect be might that those than altitud to type flower given a of restriction particular any be n does there Again, 1982). Royen, van & Kores see - for allowed the at variable notably is there additio In more. or way half to lobed are species the half over th in 43% three- is fully but figure region, this Moluccas corolla; whole the of th length one the lobes more) with flowers 19 have Stevens, species Papuasian also of (se 35% Perhaps emphasized be should species Malesian West coroll long relatively The 1982). Argent, see yongii, (R. Sabah zygomorphic red, with species lwr oor n ausa r Papuasian in colour flower ooedos s ehp getr tha greater perhaps is hododendrons igh altitudes (Cruden, 1972); bi 1972); (Cruden, altitudes igh d rm h foe clu, altho colour, flower the from ed ures n h Ws Malaysian West the in quarters e n id pootoal incr proportionally birds in he lwr hs eety en describ been recently has flowers nrseii lvl (infraspec level infraspecific blr lwr ae nomn a uncommon; are flowers ubular naie o fu nectar-eatin four of entatives und atthehighest nl oe other zones inal a lobes of these of lobes a our and feeding and our n, flower colour flower n, fc aito in variation ific rds will remain will rds dency of such of dency ae a high at eases ors. The two two The ors. ) has recently has ) th red, short- red, th members of members ein and region, g species ugh Celebes- e y important y ot seem to to seem ot pd n the in rpid genera g earlier n d from ed sa of asia, e red pe; single single r (or ird 76). 0m to www.vireya.net (Kores, pers.comm.). pol probably is bats by pollinated probably is type type flower white funneliform the and sphingids, flower white tubular long The below). see also 1981; (Bierzychudek, butterflies for combination colour attractive an is red by surrounded yellow glabriflorum); R. also (see lobes red and tube yellow arethoseofabird-pollin R. christi of syndrome floral the of interestin is case latter This 1976). (Stevens, christi R. e.g. utrle hv be osre t vst fw pce wt larg with flower species funnel-shaped broadly few yellow, a visit to observed been have Butterflies asynchronous. flowering the unless sources food additional for Meliphagideae pers.comm.); Argent, 1982; Kores, 1982; Royen, van & Kores 1980; Smith, 1978; (Kores, flushes in rhododendr some least At 1980). Snow, & Snow (e.g. ornithophily obvious the have not do increasingly that flowers many becoming on feed birds is clear, As sparse. are Meliphagidae high-altitude the of Yeo, 1973). ( areas small five to restricted tube the of nectar the base to access himantodes R. like species in and pouched, the but concealed, well less apparently nectar area the from species of flowers open more The kinabaluensis). brook subsp. javanicum R. (e.g. combination yellow / red a with v butterflies notes w (1985) Argent 1976). absent (Stevens, trends clear almost is evi evidence available pollinators; to types flower assign circumstantial to attempting even Malesia West In pollinator. parti one of preserve exclusive the as considered be not should types floral knowledge, of level current the with especially addition, In (1982). Kores in photographs excellent the of several in shown clearly commo quite be may robbing nectar syndrome, floral the Whatever cf. the aptly-named 'revolver' aptly-named the cf. , n as sm red-flower some also and s, hs a ices te ed of need the increase may this g; although aspects although g; ated flower,ithasa flowers: Proctor & Proctor flowers: dence shows no shows dence stn flowers isiting cular class of class cular & van Royen van & characters of characters seems to be to seems van Royen & Royen van d species, ed au var. eanum lse are flushes , orange- e, ae the makes ons flower flower ons iae by linated n. This is This n. s often is the hen www.vireya.net meitl aprn t ayoy h hs tae ee a ite w little a those fewbeatenpathsthatexistinMalesia(cf.Stevens,1981 even strayed has who anybody to apparent immediately Rhododendro as genus well-collected apparently and conspicuous, composition during the Quater the during composition dis abou pseudopro of about talking imperfection talking the when avoid addition, and Hence, try to vandeursenii). has (R. one distributions, Guinea New Papua of appare Cen the in endemics' 'point the few very the of one in of loss the and result which saruwagedicum) R. (1982) of (that pattern distributional Royen disjunct a of disappearance van & Kores by proposed evolutionar and vitis-idaea R. of circumscription in prob changes the by shown aptly geographic chance affect concepts species representing How events. unimportant perhaps hybrids, almost are few occasional a unity; phylogenetic necessary no taxonomic with major species morpho are the species the of that remember to some has one of above mentioned unreliability the to addition In Lflr 18) Te reln tdy s p o 00 eeain i elevation 4000m to up is today tree-line The 1981). (Loffler, th in early only current those to close elevations reached have However 1981). Raven, makes the & today that chain in (Axelrod mountainous more B.P. or years 1500m million to 15 up Miocene, elevations with emergent become have to Guinea, New of part middle the forms now which belt, mobile The hist differing them, toalargeextent,independently. shal the I region, of Celebes-Moluccas the and Because Malesia, West germane. Papuasia, also is history and Tertiary Malesia, in Rhododendron of evolution the and distribution great very the that realize we subsec and species of distribution the comprehend better can We GENERAL DISTRIBUTIONOF nary are central to an understandi an to central are nary hne i bt vgttoa lim vegetational both in changes tributional data even in a so so a in even data tributional MALESIA RHODODENDRON SECT.VIREYAIN up the spine of New Guinea s Guinea New of spine the up ). and R. yelliotii yelliotii R. and tral Highlands tral ng of both the both of ng e Pleistocene e well-known, is believed is tions when tions oe late some blems. In In blems. places, n discuss l wl be will n groups re of ories certainly eems to eems ay from from ay es is lems t and its logical the , ily nt t www.vireya.net Eastern Highlands of Papua New Guinea and other blocks, and so and blocks, other and Guinea New Papua of Highlands Eastern joins almost which Mountains), (Snow Range Maoke The confluent. wo which of vegetation 'subalpine' period glacial the Vogelkop, hs mi bok icue h mount the include blocks main These 00 attd hv seis of species have altitude 2000m o blocks main the of Many above. discussed that as such history vegetational a with expected be might this below); see (but high be to seem Wilhelm, Mt. Giluwe, Mt. and the of flora ericaceous the of study their in noted (1970) Vink Hope & Walker and (1979) Flenley interestingly, Equally Pleistocene. the of have much may through tree-line common lowered a Such 1975). Peterson, & Hope 1981; Walk 1979; Flenley, & Walker (e.g. 2300m about to down was line years 10-11000 some (until glaciation last the during However, lower somewhat generally although This clearly has important implic important has clearly This western thanineasternNewGuinea. veget that note (1979) Flenley & because o now is it than drier the of part Also, grassland. alpine of kilometres square 55000 v of lowering the of effect One and ofcommunityconstancyovertime. affil ecological of degrees varying are there community; stable s be cannot and years thousand few last the over element only developed of synthesis post-Glacial a represents forest subalpine roecn got - hs oe & Hope too thus - were growth conditions arborescent which in places obviously were there island. area, the of lengths great for continuous peaks was isolated land such on grow; perforcedly and 3100m above only growing now su species which in type the of periods glacial the in have vegetation probably would that 2000m above land of area vast a was f the exposure of the broad Arafura shelf;Walker f theexposureofbroadArafura and the Kubor Range, local endemis local Range, Kubor the and gttoa lmt cn e readil be can limits egetational hddnrn Tbe ) restricte 2) (Table Rhododendron ational limits may have been sl been have may limits ational ations for distributional stud distributional for ations eesn 17) siae that estimate (1975) Peterson epcal o ioae mount isolated on especially , 18) ugse ta te curr the that suggested (1981) is f h nrhr pr of part northern the of ains ies. As Kalman & Kalman As ies. of Rhododendron of ightly higher in higher ightly se. There seen. y oa Peaks Doma uld be almost be uld chain may be may chain ago) the tree- the ago) iation of taxa taxa of iation f land above land f ihn this Within m does not does m aid to be a be to aid er & Hope, & er ta has that s d to them. them. to d hr was there akr & Walker harsh for for harsh forms an forms with the with pported been could ains. the ent www.vireya.net of Rhododendronknownfromthere. at about 2000m elevation, some is v is some elevation, 2000m about at sma very a has Britain New blocks. altitude high main other the are block, Suckling-Dayman the especially and block, Finisterre very a has mountain this south; block especially is now rather effectively isolated, since Mt. Mt. since high northerly most the is isolated, which effectively rather now is especially block Th vegetation. elevation 1200k high of block largest the over far by forms altitude 2000m above land of chain continuous almost mollianum, R.papuanum,phaeochristu 1. TheVogelkop.R.brachypodariu mountain itself is fairly isolated, andisformedofserpentine mountain itselfisfairlyisolated, 4. Mt.Doormantop.R.extrorsum (alowaltitudespecies). 3. WandammenPeninsula.R.asperrimum 2. Fak-Fak.R.vinkii(a altitudes). undercollected. several speciesofthesmall- 5. MaokeRange(SnowMountains). T endemics isexceptionalforasinglemountain.

Some endemicspeciesofNewGuinearhododendrons. lowaltitudespecies). leaved ser.Linnaeoidea,but thew , R. lamii, R. simulans, R.subu , R.lamii,simulans, m, R.erosipetalum,hatamen here areabout20species res mountain of this block, is rel is block, this of mountain interesting vegetation. Both th Both vegetation. interesting m, R.porphyranthes(mos Table 2. olcanic, some not; there are there not; some olcanic,

rocks.Thenumberofapparent hole rangeismuch losum, R. taxoides. The R.taxoides. losum, se, R.hirtolepidotum, se,

tricted to it, including toit,including tricted tly speciesofmoderate atively far to the the to far atively e Owen Stanley Owen e

e Saruwaged- e ll area of land of area ll isolated from isolated Amungwiwa, five species five

long, m www.vireya.net 6. CycloopMountains.R.bryophilu n R vlouu ae atclry epeig Rooedo atr Rhododendron perplexing. particularly are villosulum R. and hooglandii, R. of those like Distributions 3. Table in given is these of selection a patterns; distribution anomalous produced has extinction d long-distance that suggest which distributions some are There report being maius further south. R. only Block, Stanley Owen the in area Road Amungwi Mt. the in found all are emphasized, be should it taxa, revers is relationship the christi R. and maius, R. pleianthum, the on mainly occur alticola with whilst west, the to stations individual with but Stanleys, R. and the rubellum on R. and Stanleys; block Owen Suckling-Dayman the on occurs carringtoniae R. Thus, qualify probably will known better are they when distributions altit lower at growing species one than more on geographica that seems occur it Although atropurpureum R. and hooglandii R. like species and fragmented, much are Guinea New Papua of Highlands Western in blocks altitude high addition, in 3000m; below slightly only generally gaultheriifo are var. gaultheriifolium 3000m above growing wo R. However, blocks. altitude high major single to restricted today Rhododendron of Species 9. Thesouth-easternmountains occur onlyontheOwenStanleys. giulianettii,R.detznerianum, 8. OwenStanleys.R.comptum, Guinea andisalsonon-volcanic. 7. Mt.Wilhelm.Noendemics,althoughWilhelmisthehighes cyclopense. Anisolatedareaof None ofthesespeciesgrowsa 11. Saruwaged-Finisterrerange.R var. wrightianumisawidespreadtaxon. goodenoughii isamemberoftaxonomic 10. D'Entrecasteauxarchipelago.R.goodenoughii,wrightianu in thearea. t thehighestelevationsonMt.Su . R.archboldianum,armitii,R not very high hills; again serpentine occurshere. not veryhighhills;againserpentine . hellwigii, R. pachycarpon, R. rhodostomum. . hellwigii,R.pachycarpon, m, R.dianthosmum,laetum, udes (Table 2), a more sophisticated analysis of analysis sophisticated more a 2), (Table udes lium are spread rather more w more rather spread are lium ally difficultgroup(cf l isolation has affected the affected has isolation l . Hunt,1981);R.wrightianum andR.spondylophyllum . rhodoleucum,R.truncicola. ckling, thehighestmountain t mountaininPapuaNew m var.insularum.R. R.wrightianumvar. R. atropurpureum, R. R. womersleyi, R. womersleyi, R. the Eastern and Eastern the ed - these latter these - ed mersleyi and R. and mersleyi spread of some of spread idely and grow and idely this statement. this opurpureum

wa-Bulldog sesl or ispersal ed from from ed Owen block.

www.vireya.net egahc ags ad pit endem 'point' and ranges, geographic coherent fairly have species most however, speaking, Broadly 1975). Smith, scarc the th east with the to dispersal connected to barrier the be or habitats appropriate might Giluwe and Hagen Mts. of east from absence its and range, its of part eastern the in species b a is saxifragoides R. thus examined; are compared species the altitudinal and habitats, ranges, total the when modified be to dist they that those of mountain single the by affected similar was occurrin species numerous were there but mountains, the the mountainsbetween speaking, two broadly between that, flora found ericaceous (1970) Vink & Kalkman th on genera) other altitude of in mountains isinexplicable. species 3900m (and almost rhododendrons reaches in and difference Wilhelm Mt. to adjacent is Mt Kerigomna. Mt. on latter the and Wilhelm Mt. on grows former kno th is on occur hooglandii R. it and it both which although 1974); (Stevens, in areas two the in differentiation local shows R.villosulum R.yelliotii R.vitis-idaea R.womersleyi R.atropurpureum R.hooglandii R.saxifragoides Owen Stanleys;Smith,1975). there highest is plants alpine herbaceous of endemism 2); Table Ra Maoke the on occur locality single a from only known species

Distribution ofsomeNewGu

Giluwe + + - - - - -

Kerigomna + + - - - - -

Wilhelm + + + + + - -

on selectedmountains.

Kubors + + + + - - -

studied. To a certain extent certain a To studied. Table 3. Amungwiwa inea speciesofRhododendron + + + - - - -

c ae o ntby common. notably not are ics

Saruwageds + + + - - - -

Irian Jaya Stanleys Owen C.Highlands & Mostly eastern Central Highlands Total range Central Highlands west To Elsewhere

mountains to the to mountains at may exist (cf. (cf. exist may at , this finding has finding this ,

preferences of preferences e Kubors, the Kubors, e nge (but see (but nge n o occur to wn (and on the on (and . Kerigomna . og-dwelling g on only a only on g Several two e t of ity t of ity the ; ance www.vireya.net konori, and R. macgregoriae all macgregoriae R. and konori, R. fr zoelleri, R. only nummatum, R. lindaueanum, absent R. like being species few A this, Vogelkop. to close comes beyerinckianum R. and throughout habitats available many in altitudes high at occurs culm Rhododendron species. Papuasian widespread few a are There ee e ae o itnus bten h ClbsMlca ae ad 'the and ar Celebes-Moluccas The rest'. area Celebes-Moluccas the between distinguish to have we Here pattern ofdistribution. interglacial the during confluent ex that vegetation of areas highland those for tendency less be Thus Sumatra. in 350m of i depression a than indicates (1981) less Whitmore considerably be to believed is Malesia West in line volcanic depressio the addition, non In elevation. only 3000m above the substantially are Celebes the in mountains and Sumatra, Mt. altitude; great no reaching ar peaks highest non-volcani are the Peninsula Malay the of of highlands The many volcanoes. and region, whole the in altitude, i.e Rhododendron, of species subalpine for suitable land little 100 above land 3; & Wa see - extent wide of 2 however, (Figs disjunct more much and less much is is m is area land situation total the Although the area Celebes-Moluccas the and Malesia West In Arch Louisiade and Bismarck the in it and Doormantop, Mt. of east grows superbum R. altitudes. low of length the extend quite not do hri) nvrhls spot a ary ih lr o sm 34 some of flora t on occur these of (most endemic are 28 which of Rhododendron, rich fairly a supports nevertheless therein), periods glacial the during drier small formofR.konorihasbeen (e.g. Whitmore, 1975; Walker, Walker, 1975; Whitmore, (e.g. ea, although small,andinpartp ea, have wide distributions, althou distributions, wide have . o srrsnl, e e a see we surprisingly, Not s. uch greater, the area above 20 above area the greater, uch collected veryrecentlyfromM the island; these species als species these island; the lker, 1982, fig. 30.1). There i There 30.1). fig. 1982, lker, iaau n aa, t Lsr in Losir Mt. Sabah, in Kinabalu plgs nt ta wa i a is what that (note ipelagos s also relatively relatively also s . above 3000m above . o grow at quite at grow o gh the first two first the gh isima Island). 1982, and refs and 1982, erhaps notably 00m elevation 00m ist to become to ist ey different very tree- the of n e Guinea, New Papuasia; n occurs also he island of island he prnl a pparently c, although c, pce of species there will there northern different. active e om the the om peaks m is, 0m inicola www.vireya.net Guinea, New and Borneo, Sumatra, l at island, large a not is Sulawesi Although itself). Sulawesi there; the first two have a basically north-western Malesian distribution, the the distribution, Malesian north-western basically a have two first Malesian the there; West zo the R. However, malayanum, R. area. quadrasianum, yel Celebes-Moluccas R. the in to Papuasian occur both close lindaueanum, R. is and zoelleri lagunculicarpon R. and (Papuasia), R. 1976). to closer (Stevens, be to Papuasia tending perhaps position, intermediate an suggests types considera Overall species. Papuasian of number a in occurs that the with impositum) R. perhaps ar there type, flower zygomorphic distinct the have area the in species no Although Papuasia. and Wes between intermediate clearly is region Celebes-Moluccas The 1981;Dransfield,1981). fauna asawhole(e.g.Whitmore, rela lack the with contrasts Rhododendron the Interestingly, up. broken highly is this and altitude, Fig.2. Land above 2000m altitude on Sulawesi(shadedblack). Fig.2. Landabove2000maltitude Bar represents300km. large broadly funnel-shaped whit funnel-shaped broadly large e species (R. lomphoense, R. buruense, buruense, R. lomphoense, (R. species e t a a usata ae abov area substantial a has it of diversity of the flora the of diversity of lnei ad . aaiu als javanicum R. and llingeri,

east compared to compared east and much of the of much and ie iest of diversity tive ive Papuasian ive e flower type type flower e tion of flower of tion Malesia t 2000m e species, occur o liotii R. www.vireya.net only 6%. These figures should be should figures These 6%. only i figure the whilst varieties, have Malesia West in species the notin worth perhaps is It scattered. widely and small are 2000m above currently land of areas but mountains, eroded old, of consists Borneo of range central The 1964). Steenis, (Van flora microtherm more a carried and been have once may nearby ranges lower that suggested been has w the in mountain highest the is Kinabalu Mt. six it. to some restricted only currently but it, near or Sabah, in Kinabalu, Mt. on occur species 98 these of Many Islands. Philippine the to restricted are 12 Male of rest the of whole the in species 98 some only are There Philippines). Island (Sunda distribution southern basically a have two second of that from considerably Pleistocene; ericaceouspollen differ Oligo-Miocene the that from in elements record montane The elevations. higher at flora static a suggests t it, Borneo, northwest for record pollen long a quite is there Although malayanum. R. and longiflorum R. jasminiflorum, R. javanicum, R. Rhododendron of populations ar there of Nevertheless, Malesia. age) (or isolation greater this is recordedinthelatter(Mulle e a few widespread West Malesian West widespread few a e treated with great caution, bu caution, great with treated

oo, is not one that one not is oo, r, 1972). hole area, and it and area, hole Ppai is Papuasia n the also but s, t may indicate may t g that 20% of 20% that g pce are species sia, of which of sia,

species - species in West West in h Pli- the higher www.vireya.net ipitc o el fetvl wt te rbes oe b past by posed problems change. the with group effectively the deal of to biology the simplistic of knowledge current Our taxa. between barriers ecological apparent the of some down breaking by this do may it but (Ste stress or change environmental of periods during diversity mai the in helps flexibility comparable that suggested been has i where vegetation ridge in collected been has it Jaya Irian in New Guinea R. saxifragoides is c is saxifragoides R. Guinea New alth Thus, surmise. pure is this although isolation, ecological may above discussed habit and leaf in Variation standing? long that ecology their of aspects other or pollinators and species relat the are Quaternary: and the of part least at through existing fauna, the in problems increases oscillations considerably comparable by accompanied surely habita the discuss composition as and distribution vegetation in long oscillations so general, does In hybridization undisturbed, 1966). (Sleumer, groups other with hybridizes Malayovireya subsect. distinct the even 1976); (S excepted only / perhaps 'bulldog species, problems different very apparently cross to possible Vire sect. of species Malesian between crossability to barriers a in possible interactions the co the suggest (1980) Snow & Snow and (1981) Stiles out; worked bet interaction the of details types and possible pollinators would seem strong evidence for their their for evidence strong seem h of prevention would the in importance pollinators possible and types d The diversity. that of maintenance the in involved today diversificati even in involved been have Malesia throughout species between together differences floral marked the how unclear is It Despite these uncertainties, one can reasonably suggest in the the in suggest reasonably can one uncertainties, these Despite ween the plants and their pollina their and plants the ween e-rpcl iuto. hr ar There situation. neo-tropical DISCUSSION o se t b ta feun. oee, the However, frequent. that be to seem not haracteristically a denizen of denizen a haracteristically biiain n aa rwn t growing taxa in ybridization hn pce ae osdrd as considered are species when emr 16, 93 Sees 1 Stevens, 1973; 1966, leumer,

ionships between ionships tors remain to be to remain tors t is caespitose. It caespitose. is t we see today of today see we ed above were above ed uh n Papua in ough iversity of floral of iversity members from members tussock bogs, tussock fw known few e gte. The ogether. a ad t is it and ya, es 1981), vens, Chihuahua' tnne of ntenance also help in help also n n are and on that grow mplexity of mplexity ecological ae of case s too is entities this 974, is t www.vireya.net accentuated itsspeciationrate. of some has it so ground, exposed seeds, of numbers large producing and nevertheless initially, woody grower although Vireya, sect. Rhododendron r-type characteristics. with species favour to tend would particular, in o country (1982) that than Walker saturated speciation. coupled less this, that and be Malesia, aided to flora probably Papuasian has the considered areas the of diversity altit 2000m below land by limited coloni aggressive specie many of particularly distributions The ones). a low especially islands, seem not does Rhododendron but complicate plate Australian advancing the of north the to chain five last the within been probably Papuasia, in Rhododendron of diversification the of Much 1981). icuig h wsen at f t of part western the (including outlier, an as Celebes the of part Au the when years million 20 last w the from Papuasia entered certainly almost have they Malesia, bee have rhododendrons Vireya long how uncertain is it Although funnelifo large and white, tubular zygomorph types, flower distinctive most the of Three these diverged. since characters floral and vegetative in evolution great there that particular in Malayovireya and Phaeovireya subsects. imrk rhplg, h Slmn sad, n Nw una itsel Guinea New Celebes-Molucc and the Islands, Philippines, Solomon the the Archipelago, in Bismarck islands many thus - taxa montane of distribution the to barriers as act to appear would a numerous had, has always probably and present, at has Malesia flowers necessarily representative 5-merous than more with species proba that are emphasizing three types, All shapes. leaf distinctive have as once, than of anancestralcondition. a sl-etlz (os, 1985) (Rouse, self-fertilize can e eee: aitn 17; Aud 1979; Hamilton, Celebes: he with the extremely unstable nat unstable extremely the with rm white, have almost certain almost have white, rm stralian plate, possibly including the eastern the including possibly plate, stralian d, n te xrm rle an relief extreme the and ude, and many species are noted co noted are species many and these characteristics, and th and characteristics, these ilo yas te existence (the years million ae n otc wt S Asia SE with contact in came ly evolved more evolved ly s the argument, the s , each capsule capsule each , or even colline even or s seem to be to seem s ure of the hill the of ure f n island an of reproductive reproductive est within the within est at least, has least, at two groups is may have geological d ley-Charles, c e, long red, ic l derived bly a been has reas which reas lonizers of lonizers n in West West in n are not -Malesia s the as, , have f, West f slow a t of st www.vireya.net the world(ClassAves). Auk98:681-714. Cracraft, J.(1981).Towards aphylogeneticclassificationoft Verbreitung. Bot.Jahrb. Syst.9:319-379. Gruppierungundzurgeogra in Beziehungzuihrersystematischen Axelrod, D.I.&Raven,P.H Borneo. NotesfromtheRB Ericaceae. J:Linn.Soc.Bot.85:2-17------(1985)VireyaRh t support Nevertheless, interglacials. would that Australia, and between that also and Malesia, West in especially areas, large for water by isolated been always Breitfeld, A.(1988).Deranatomi 13(suppl.): 54-58. mimicry complex.Biotropica Bierzychudek, P.(1981).Asclepias 837-861. TheHague. Guinea. InGressit,J.L.(ed.) Beehler, B.(1981).Ecologicalstru Oxford. Line. InWhitmore,T.C.(ed.),W Audley-Charles, M.G.(1981).Geologi Guinea, 918-941.TheHague. New GuineaFlora.InGressi Argent,G. C.G.(1982).Newtaxaandco met. origina rhododendrons, where area an barrier for 20millionyears,anditis cross to difficult a less or more of case this in only is it too, groups, other many of (S generally Ericaceae the of distribution the in line dividing b to seems area this Although respectively. western and eastern reach Phaeovireya and Malayovireya subsects. as groups defined Celebes the in that interesting perhaps is It gaps. the across G Edinburgh,43,1:53-61 . 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n inthetropics,554- story oftheEnga xford. ary history.In olution, with , T.C.(ed.), Sunda-Sahul 63-285. The sion of the inb. 37: