080057-14.002.Pdf

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ij\dgz-t:(Zl otslinN Nq'tsia Yo|. 14, No. 1/2 (2001) partson systemcards beneath clear magic tape,field testsfor pollinator specificity, and the extensive useof 35 mm colour slidesand line drawingsof fresh flowers. We havealso benefitedfrom accessto recently acquireddata on allozyme and DNA markers.

The early history of the systematicsof south-westernAustralian orchids is reviewed here. It comprisedtwo phases:(1) the first encountersby Europeansand (2) the early colonial phasein which James Drummond was prominent. The strong reliance of taxonomists on the collections and observationsof interestedrcsident orchid enthusiastsis a recunent featureof this history,

Our taxonomic conceptsfor genera,species, subspecies and hybdds are described. New taxa are consideredto be speciesif they grow in populations(i.e. are not aberrantindividuals within normal populations),if they have featuresor a combination of featuresnot seenin any previously named species,and especially if they grow with previously named speciesand produce few or no natural hybrids. Geographicalraces with minor morphologicaldifferences are recognized as subspecies. These may hybridize and intergmdeextensively where their geographicalranges overlap. Our emphasison biological as well as morphological attdbutesin recognizingspecies and subspeciesis a significant changeftom past practice. It leads to the recognitionof more specieswithin complexespreviously treatedas single polymorphic species(e.g. Caladeniafilamentosa R. Br., C. huegelii H.G. Reichlt.)- Thereare many practical advantages toourapproach, includingimproved management forconservation, improved cultivation, and more precisecommunication and conductofpopular and scientific studies on south-westemorchids. Nevr'genera, published elsewhere,are basedon numerical analysesof morphologicaldata, aswell asthe recently completed DNA sequencestudies ofKoreset al. (20O0,2001\.

Introduction

Two centuriesago, Archibald Menzies,surgeon and naturalistaboard the ILMS. Discovery, was the first European to make collections of Western Australian orchids. The Discovery, under the commandof CaptainGeorge Vancouver, was anchoredin King GeorgeSound from 28 Septemberto 11 October1791.

"copious Menzies made a collection of ... yegetableproductions, principally the genusBantsia, which are here very numerous" from various sites onshore in the vicinity of present-dayAlbany (Maiden 1909; Heberle, undated). Three orchid specieswere subsequentlynamed from Menzies' specimens,including one bearinghis name-Leptoceras menzt€stt(R. Br.) Lindl., Caladenia flavaR. Br. andDiuris longifulla R. Br. (Brown 1810;Lindley 1830-1840). The presentseries ofpublications, basedon more than two decades'research, contdbutes towards the celebrationof the bicentenaryof Menzies' pioneeringcollections. Here,we provide a genemlintroduction to the series,and a new key to genem of south-westernAustralian orchids.

South-westemAustralia is one ofthe few regionsin the world where the abundanceand diversity of terrcstdal orchids may be breathtaking (Hoffman & Brown 1992, 1.998). In some habitats, particularly the first spring after a fire, more than 40 specieshave beenrecorded in a few hectares,in somecases so abundantthat walking without treadingon them was impossible. But such situations are exceptional. Indeed, most residentsand tourists, professionalbotanists among them, overlook orchidsand underestimate the dch diversity found in the south-west.There are several reasons for this. e roJ 966I osrJJ oes)lsluEtoq unlrPqroq eql ol lsol eJPsJelcsJeqc cruouoxel AuEr! lEql sueurlJeos 3ur,{rp puu Surssard,{q pe olsrp os sr sprqcJoruolse^\-qlnos Jo Jnopo pus .rnoloJlrslep IeJnpruls 'sueupeds eqlJo qsnl{ unrJeqJeqIsuorlue^uor qlr^{ dluo ?u1pepur sruolqordclluouoxEl eruocre,ro fypquelsqns e,req suorlelndodprqJJo ur uolleue^ Surdpnls o1 seqceo:dde^\au'{lpuoros puv

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'u^rorE ,ftotsrH I ,{Solopqro u?llelsnv uelso/A dV pu? loddoH dS Nryrsjd Vol. 14, No. 1/2 (2001) similarexample in thepea genus Splraerolobium). Additionally, observations on habitat,population vadation,flowedng phenology and pollinationare also commonlyscant or absenton herbarium specimenlabels.

Botanistsin Britainand Europe in the1800s had only a small number ofpoorly preserved herbadum specimensto work vr'ithwhen they commenced naming south-westem orchids. This hasmeant that muchofour cunentnomenclature is basedon broad species concepts develoPed by taxonomiststrying to makethe bestjudgements they could on the limited characters detectable from the available material. Drummond(1842) was the filst to alludeto theresultant frustration felt by rcsidentfield botanists who couldreadily discem distinct species that were lumped under the one name by botanistsftom afar(see openingquotation).

Themost striking example of sucha broad species concept was Bentham's (1873: 381-382) notion that all taxa in lhe Caladeniapatersonii, C. longicaudaLindl., C. dilatataR. Br. andC. huegelii complexesbelonged to theone species, C. patersonii. Bentham commented in supportof thisview lhat C. patersonii was:

"Veryvariablein thelength and proportions ofthe sepals, in theshape ofthe labellum andits fringesand calli, andin the colourof the flowers,which areusually of a pale greenishhue outside and yellowish or pink inside,the calli andend or middlelobe of thelabellum usually and sometimes the whole disk of a rich purple.The following are the principalforms this speciesassumes, rcgarded by Lindley andothe6 as distinct species,but passingtoo graduallyinto eachother to be clearlymarked out from dried specimens."

He thenwent on to recognizethe Caladenia dilatatq complex as a varietydistinct from the rest of C.patersonii.

JamesDrummond, Westem Austmlia's first residentEuropean field botanist,collected and separatelynumbered several of thenew speciesnamed in thepresent series, and even described and namedsome himself in publishedlette$ to Hooker(Drummond 1842), only to seesome of his observationsignored and specimens lumped under the broad species concepts ofthe botanists to whom theywerc shipped.

Bentham's(1873) and particularly Reichenbach's (1871) broad concepts were also recognized as unworkableby Australia'sfirst residentorchidologist, R.D. Fitzgerald. Having seen and accurately paintedfresh flowers of manytaxa in New SouthWales and some from Westetn Australia, Fitzgerald (1882)remarked rather caustically:

"Specimensthat, if submitteda few yearsago to the highestauthorities, would withoutdoubt have been considered as representing new species,are now mattenof hesitation,uncertainty, and rc-comparison, and are finally pronouncedto be varieties, very oftento thedisgust of thecollector, who hasseen them in a freshstate, and who cannotaccept the ruling that they are identical with otherspreviously named, with which he may be equallyfamiliar, and into whichhe feelsconfident they never lapse."

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Figurc I. Systcm card showing dissoctod spocimcns ol Culadcnia [ulcata

ln thisseries and a previouspaper (Hopper & Brown 2000),we concentrateon newand reinstated south-westerntaxa in Caladenia R. Br., Drakaea Lindl. and allied genera.David Jonesand Mark Clementshave describedand will be describingmany new taxa of south-westernPt€ros4)lis R. Br., Diuris Sm., CyrtostylisR.Br., Gastrotlia R. Br. and CorybasSalisbury (e.g. Jones & ClementsI987; Clements1989;Jones 1991). In SouthAustralia, Robert Bates is curently workingon Prasophyllum R. Br. andMlcroris R. Br. (e.g.Bates 1984, 1989, 1990), while work begunby the lateJoe Weber of the StateHerbarium , Adelaideon ThelymltraForstcr is now beingcontinued by Jeff Jeanes Wewould expectthe descriptionof many new taxa in thesegenera when further detailedpopulation studiesare undertaken.

It is timely to describethe new taxafor severalreasons. Many arerare, and their formal description will aid conservationagencies in identifying and managingthe few populationsknown (Hopper el cl 1990; Brown et ol. 1998). Our taxonomic researchis ongoing, but many undescribedtaxa were identified some years ago (e.g. HolTman & Brown 1984) and are well known among local orchid enthusiasts.Publication of the nameswill facilitate communicationand increaseawareness of the real wealth oforchids occurring in the south-west.And finally, the nameswill be validatedfollowing their informalmention in the revisededitions of"Orchids of South-WestAustralia"(Hoffman & Brown 1992, 19e8).

While we regardthe descriptionsin the other papersofthis seriesas a significant new contribution to knowledge of the south-westerngenem treated, it is by no meansthe last word on the subject. As our taxonomic researchis ongoing, we would welcome hearingfrom anyonewho has found orchids that do not match thoseillustrated and describedin this seriesor in Hoffman & Brown (1992, 1998).

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Finally, the currentsurge in taxonomicresearch and description ofAustralian orchidsis notwithout its critics (e.g. Heberle 1995; Dockrill 1995 - seeBower 1995 for ar independentresponse). This is not unexpeated,and indeed,is healthy for any areaof scientific research.As long as cdticism is based on carefultesting andwell documentedfalsification oftaxonomichypotheses, significantimprovements in our knowledge of Australian orchids will occur. Indeed, such an approachled Bates and Weber (Weber & Bates 1978, 1986; Bates & Weber 1990) to move from early unworkably broad species conceptsto a taxonomy much morc in tune with biological reality. There will never be a final word on matterssuch as taxonomic rank and speciesconcepts. The important advancesnevertheless will occur primarily when new contributionsto existing data aremade by carefulwell-documented study. We would cite the exciting new insights gained on the evolution and classificationof all organisms through the recent application of DNA sequenceanalysis as an especiallysignificant example of a major advancein knowledge (seebelow).

Systematichistory - the Pioneers

Given the bicentennialcelebration to which this paper is dedicated,it is fitting to review the early history ofthe systematicdescription of the south-westAustralian orchid flora, focussingon the genera treatedher€in. We havedrawn our informationfrom severalsources, the main onesbeing Diels (1906), Maiden(1908, 1909, 1921), Edckson (1969), Hall (1978,1984), Carr & Carr (1981a,b), Marchant (1982),Heberle (undated), Short (1990), Barker& Barker(1990), and tmnscripts ofJames Drummond's lettersto Hooker held at the WestemAustralian Herbarium. We havealso examinedspecimens in the majorAustralian herbariaand at Kew, the British Museumand Vienna to obtainadditional information on collectors.

At the outset, we wish to stressBarker & Barker's (1990) thesis that throughout the history summarisedbelow, right to the presentday, knowledgeof the real contribution of all involved in the collection and naming of Australian plants, orchids included, is imperfectly recorded. Proper acknowledgmentof assistanceis very much a personalmatter. Botanistsfrom all ageshave varied greatlyin their ability andwillingness to recordtheir debt to otherpeople. Along with Barker & Barker (1990),we haveattempted to recountdetails ofthe efforts ofpoorly known enthusiastsas much asthose of the well known botanistswhose contributions rightfully have been extolled in the literature.

Four main overlappingphases may be recognizedin the collectionand descdption ofsouth-western orchids by Europeansand their Australian descendants- (1) first encounters'(2) early colonial, (3) easternAustralian taxonomists,and (4) residentWestern Australian taxonomists.Here we review only the first two of thesephases, both mainly involving descriptionoforchids by Europeanbotanists, with the most notablefeature ofthe secondphase being the outstandingcontributions made by James Drummond in the 1830s.It is hoped to review subsequentphases in a future Publication.

Becauseso little hasbeen recorded, we have not attemptedto describeaboriginal krowledge, but it must have been prodigious given the 40 000+ yearsof their occupationof the south-west'and the importanceof some orchid tuberoidsas dietary items.

Firstencount€rs

Orchids may have beenseen, but were not collected,when Willem de Vlaming briefly landedon RottnestIsland and the adjacentmainland (now suburbanPerth) in December1696 and January1697 (Playford 1998). Similarly, the fragmentarycollections available from William Dampier's exploration 'suoqeroldxe 'eurllsuoJ senboellsruelog laql;o Suluq aql go esnuceqdlurrru lsa.r-qlnos eql tuo{ sprqJrolrelloc ol pelleJllsr^ s,u,rorg Jegedlelutperurut puu sJoJeqslslu8loq qsuarg "{13ur1sere1u1

'rg '4 ' y 4 reddog ('tg'a) slutnlap 2puu Ig d v "wel)'V "I'( 4reddog(rg g) zafno) DpuuDt)' tg d orloltiDl muapoloJ 1'l.IEsouof ( rg'U) suout8u stqctoJ{d se qcns 'Elle4snv uJelsa^\ uI punoJ ueeq eJurseAEq erleJlsnv uJelsse-qlrlos '(oao1J'3) rrrpelrelloc Ieueteruruo4 (g1g1) u,r,ro:gdq perueusatceds leuorllpps lure^as DluapoloJ '(9191) uBrlellsnv ualse/(\ ls.rr;eql Surpnlcur u,rorg dq peueu aret serzuel'iqdq puno5 eSroeg 3ury rxo{ papolloJ erel\ leql rededlueserd oqlJo qder8eredSuluedo eql ur peuoquerusetceds eerql eql

'(7161 lyexelg) lse^\-qtnoseql ur Suyrncco lpxelg DuoalooDro4,snue8 e1e8e.t3es s1t q1r,,rr111dsdpuecer'rg'douaa1rzJpue'(9191-Ogg1)deypurl,(q4uurcrreuo8olpolu^eleflluenbesqns sei qJrq^\ 'tg'A sotacotdaT uo\aes oluapoloJ pelxes oslE uttrotg st1,(lsool4 pne wryfqdosDQ 'suorJlw 'tg'6 "tg'a 'orpoitsDg ' 'd ' snqluotadKT wmsaldouag Jg snlrlpoug Jg d owalEdg 'sll/0sor.rfJ"Ig "tglcsrylqrop)'ouapDp)-EllBrlsnv d slt{lsotdtrC Iuolsel[ ut u,tou4,r.roue.Ieue8 '(g eqlJo z I paqsllqelseeq 191u,trorg) ,,ueuelc-uE^ eBlnsuJle aetpuEIIoHee^oN eerolc snuorpord,, 'uorlrpps Surreeuordsrq ur u1 san8ealloc puE u^\o slq Jo suorlcollor Surqrrcseppup SulIeuI ueqlIBrluelsqns eJouIJEJ sE,r sprqsro uellerlsnv urJlsel[Jo sJr]eurJts,{saql ol uorlnqrI1uoas.u,{\org

'relutuns.{Huo ',(lpelqnopun usql reqler ?uuds ur pepueluollrpedxe eql peq tuareJJlpueaq e,req p1no,,n stql ureue8 pelllero lpul-I 2211o1o'ouapop) ur sr zOsl ur pelrellor sarcedsgl oqlSoeuop (pelepuneFeqeg) ,{ueqlygo ,{1rurcr,reql ur requeceq ut SuuerroU punoJueeq ecuts er'eq selceds rno; luuotltppuuu ,{1uo 'splqcro luql lcBJaql ul pelJagar seansseuqSnoroql rreql (9191) u,t\olg ,{q poueu ,{PuenbasqnsIIe uJelse,/rr-qlnosJo selcadset pelJelloJlnq'uosBes prqcJo eql ur elEIera,A\ sonSeellot slq pue u^\olg

'o8eledrqJJVeqcJeqce1 eql ur spuelsruo 's,tep 'acuuradsg puu a,nr;ro; (yed IEuogeNpuurC e1 edu3 ,,(up-luesa.rdeq1 ut) ,{eg ,{1cn1 le SuIpue[ JeeupelJelloc puE peroldxe oslu uotltpedxe eq1 ',{renue1gl puu0I uee,trleg ,4uuq1ypunoru slelrquq Jo a8upr e uroq Eueld lcelloc pue aroldxe o1 ,{lrungoddooldue sJeqlopue pooC'u,tro.lg SuIpJoJJu 'punog 'Jole^rasuo) eql urtueds se.t' (208I .{renuut9 ot) {tuoru 11ryy reneg pueurpJoClslue oqt pue se 'pooC JeledJeuepruS aq1 e:o,r,r. ,{ued eql ur osly s>1uegqdesol rrg ;o a?euorledeql Jopunpelulodds 'u^aorg 'lors ueqog Suno{ e su,tr1stle:nleu s€ pJeoquO I08I Jequeceq 8 uo punoSe3:oa9 3ut;4 1u petoqcuercpggsaa l aql ur sJepurld.^^eqllslil{ ,{q pel EIIBJlsnyale8r,reuulncrtc o1 uotltpedxe eq1

'u.,vroJg 'sprqJro uaqod uerlersnv learl o1lsturouoxel toleru lsrlJ eqt dq dl oqs plJo,rrrclJlluelcs eql Jo uorluelu aqt 01lq8nolq oq ol aJe,truIIeJlsnV uJelselA uI sun?O pue o4rwt1aq11o acuatntco 'wS, eq1 daupfg le pepellof, oarnD 6,;o suerurcedsuro.r; sr.rzrg snue8 eql petuuu (gOri) qttuls ',{yupu.ug (E961 ,{o11opr14 suqol) snueSaqt qstlqelseol pasnse.tlt g/11yo a8efo,t s.1oo3 u1e1de3 uo s:elsrog eq1{q epuw t?tstol orlolt3ro1 J Jo uorloellocpuBlBeZ ,raN V (9//I Jats.roC? JelsJoC) oarwQatllset*sueedorng dq peureueq ol uIleJlsnVuJelsel6 ut Sutunccosptqcro;o snueSlsrg eq1

' 'vut1[ usarzuawuuacoJda7 pue ollotr8uol suryq oruapo1o31o sedf,1eql pepelloc ,{gedsq ro serzueltr punos oBJoeC3ury ls lwoc qlnoseql uo I6LI uI selzuet{ pleqlqJry ol petnqlr1l€ 'pe1q3llq8tq lBql sB^\erlsJlsnv ulelseiN rrroq sprqcJoJo uolpelloJ pepJoJallsllJ eI0 i(pearp sy

'(9661 ,{-teq8rey 'IC('rg !9161e8roegry eSprqrng:qI/6I eAIoeC)'ruey3y1,14sauof A) suocrt8ruslt1ctott1 pue sn1o1o1tp'dsqns'1pw1sn1otllrysrylqJolrAr opnJJuI lou op 669I lsn8nv ur spuelsrs1r pue f,eg 4uq5;o

'u^rorg ftotsrH I fSoloprrlrro uErtE4snv olllsar1l d V pue r.ddoH C S 10 Nulrsid Vol. 14, No. 1/2 (2001)

Juliende la Billardiere,and doctor and botanist Charles Riche with D'Entrecasteaux'sexpedition collectedplants near Esperance from 13-20December 1792, includirg the fint kangaroopaw which Labillardieresubsequently ramed Anigozanthos rulzs l-abilt. On 15 DecemberRiche became lost inlandfrom DempsterHead, and the next day had to abandonthe plant specimens he collected in his urgencyto getback to thecoast and his ship(Marchant 1982). l,ate-flowering orchids may have been amongthese, but thereis no recordto informus.

And onthewest coast, botanistsAndre Michaud, Leschenault de La Tour, and Jacques Delisse, head gardenerAnselmeRiedle, and gardener'sboysAntoine Guichenot andAntoine Sautierwith the Baudin expeditions(May-September 1801; February-March 1803), and Quoy, Gaimard and C. Gaudichaud with theFreycinet expedition (1818) all similarlydid not makeorchid collections that survived to be studiedby taxonomists.

The genus Corybaswas first describedby Salisbury(1807) on the basis of material of C. aconitiflorusSalisbury from Sydney.Another 66 yearswould elapse before the occurrence of this genusin WestemAustralia was recorded by Bentham(1873).

Anotherfamous botanical explorer, Allan Cunningham,as a memberof Phillip ParkerKing's Australiancoastal survey expeditions, appears not to haveincluded orchids among his collections madeon MiddleIsland on January16, 1818, and at King GeorgeSound during 20-31 January, 1818 andftom 24December 1821 to 5 January1822 (Cuny & Maslin1990). Again, his visits occurred well afterthe main peak of floweringfor orchids.

Similarly,the horticulturalcollector William Baxterworked in the vicinity of King George's Sound,Lucky Bay and Cape Arid in theyears 1823-1825, and in 1829(Maiden 1909). Littleisknown of hisactivities, and no orchidspecimens of hisare known. In a letterftom King GeorgeSound dated March25, 1829 to Sydney'sColonial Botanist Fraser, Baxter remarked "Specimens: I have not got any worthsending, this being the worst time in all theyear for them,but if we remainhere until thespring I will geta rarecollection." Unfortunately, Baxter sailed to Sydneyin July,and missed the spring with its flushof floweringorchids (Maiden 1909).

Thus,prior to theestablishment of theSwan River Colony in 1829,only 16 speciesof orchidshad beencollected in the south-west,all from King GeorgeSound, and the majoritylate-flowering. In addition,17 genera and some additional eastem Austnlian species to befound later in thesouth-west hadalso been named, primarily by RobertBrown (1810).

Early colonialphase

Thearrival ofJames Drummond (1784-1863) at the Swan River with CaptainStirling's colonizing partyon the Parmeliain 1829heralded a majoradvance in knowledgeof the south-west'sorchids (Erickson1969). Drummond, a Scotand keen nu$eryman, formerly Curatorofthe Botanical Gardens at Cork,Ireland, was designated as unpaid Govemment Naturalist for theSwan River Colony.

Drummondsoonbecame occupied as the Colony's official gardener, andwaspaid as such for ayear (1831-1832).Additionally, he establishedhomes and farms for his family briefly at Guildford (November1829), but then atAscot (early 1830), at the foot of Mt Eliza(1831),at HelenaValley (1831), andfinally the mainfarm 1/a"thornden atToodyay (L836).

L .. sIoAIl eseqlJo sesrnocJe,rol oqt uo .{llu.lnl€u- sro^ly us?l€) puu ?ur) eql ol uo ueql pue du€qlv uro{ JBJlou osp t .tEgI .uef 1nqpuno5 eql yo dlrurcr^ elerpeurrrraql uI epeuleJe.r suorsJnJxesrH ZI-I I rxo{ pouruuoreq eJeq,npuno5eSroag ?up1o1ees dq uo penunuoJuaql pus'lequeceq 'e8ueg 6I eql 1lun JeArUup/ns eql uo peureuraroH uofullJue 1u Suqreq eq1;o looJeql seJ€J se Jolelq eql spJeaolpelEllaued pup'quedJo pooqtnoqq8teueql ul slueq S qloq uo polcellot'relr1 ue,r,iseqt ol ocuuJtueeql le spuelsreql pele8pse^uraH,, s

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,, sueu-rnedsq1t,t Sur,tog;:e,tos,{e,tr1e ere,r outoq tleql ur selleqs eqJ aJeqls1ue3e pezruSoce.t q1r,t eps dpeare punoJslamolJplr,/$ Jo speespue sueulceds qrns eleq^\ aculv qlnos ol suorlcollocalrsuelxe o,t; lsee11z lues ,{eq11eq1 u,roul srlI u,troladu31e s:opalloc crJrluercsJoJ sleurue Jeqlopue sprrq ul{s ot peuJsal 'spesur ,{eqlpus Jo uorlcalyoce8rel u dn 1pnq,{eq1 lsaralurolos lreql lou eJe,rstueld

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:palunorer(6961) uospu3 errX sy ',{1a1uurlur 'sreef eereqgad orll Jo sleurru€pue slueydaq1 ,rou1 ol ouru, dpusJ eql dpea asaqluy

'u,raorg fuotsrH t ,{SoloprqtrrouBrtpr}snv ll-IalseA dV pue lrddoH qS Nuttsia vol. 14, No. 1/2 (2001)

Threeorchidswere subsequently named from reputedHuegel c olleclions- Elythranthera brunonis (Endl.) A.S. George(as G/ossodia brunonis,byErfi\cher 1839),Thelymitraleaosa Endl. (Endlicher 1,839)and, Caladenia huegelii H.G. Reichb.(Reichenbach 1871). The type sheetof the latter species actually has three specieson it - lypical C. huegelii (as lectotypified by Hopper and M. Clementsin Clements(1989), not by George(1971a)), C. paludosaHopper & A.P. Br. and C. ensaraNicholls (not C. longiclavataE.Colemanvar.longiclay4ldas stated by George 197 7a). Only two ofthesefive species (E. brunonis and C. paludosa) have been collectedor recordedflowering as late as late November. Hence,it seemshighly improbablethat Huegel collectedany of theseorchids himself. It is likely, in fact, that the Drummord family may well havegiven dried specimensof someor all of thesespecies to Huegel,given their enthusiasmrecounted above for the local flora of the Swan River. Huegel and JamesDrummond knew eachother well enoughfor Huegel to be included among the subscribersto Drummond's future collections of seedsin the early 1840s.

In July 1835, George Moore and Lady Stirling received letters from Captain Mangles seeking assistancein obtaining seedsand live plants from WesternAustralia. Moore was aware that James Drummond and his son Johnston,therl residentin the HelenaValley, were collecting seedsof native plantsforshipment andsale at Capetown.Moore purchased100 packets of seedsfrom the Drummonds and shippedthem immediatelyback to Mangles. Late in 1835an enthusiasticreply was received requestingmore material.

In themeantime, on September18, James Drummond independently wrote to Manglesoffering to collect seedsand specimensfor payment. A sample of seedsof severalspecies was included with Drummond'sletter. Mangles forwarded the letteron to DrJohnLindley, ProfessorofBotany ofLondon University, Secretaryof the Horticultural Society, and renownedorchidologist, who wrote back to Manglesin December1836 in favourof the idea:

"l think good Swan River specimenswould sell for two poundsper 100 papers,that is the marketprice for suchthings. Probablya dozenpurchasers at leastwould be found. But as it is not known how Drummond would preparethe specimensI recommendhim to send2 or 3 setsof 100or 150each upon trial. I will takeone set. We horticultuml people wish for ten pound worth of seedsand bulbs as a venture. Nothing but very handsomethings will do, we would ratherhave a good deal of seedof a very few sorts thana little seedofa goodmany sorts. lfthe seedsuitus, we may be very goodcustomers.,,

Manglesresponded immediately to Drummond's letterofSeptember 1835 requesting, among other things,all kinds oforchids. Thus beganDrurnmond's career as a major commercialcollector ofsouth- westernflora, with an initial focus on orchids.

In 1836he prepareda box of orchidtuberoids for shipment.They perishedon thevoyage to I.ondon, as he suspectedthey would, and insectsconsumed all the seedssent as well. Undeterred,Drummond wrote again on 23 August 1836 offering his servicesas a collector:

"If you can procureme someorders to enableme to live in the bush I will sendyou specimensand seeds,mosses and insectsetc. until you are tired of them"

It would be a yearlater in September1837 before Drummond receivedlette$ of supportfrom both Mangles and Lindley. At this stage,he was given no idea that Mangles was losing interest in his commercial collectins activitv. 's1s1ueloqqsq8ug Suour€Joloelloc e su ecueyodrurs,puourunrq 'rueqt peuurlJeuorlcefioc eqJ uonmJlqndJoJ eseql Jo eurosJo suorlducsepparcderd delpurl Suoue ereueSpue serceds,tau duurupurJ ol pelq811epsu,t oq,r'felput-I JO ol uo ureqlpepueq sefuull 'ggg1 'ralo,roH ur dpue dlqs u:n1e:eq1 uo suarulcedspayp Jo uorlJellot ISJIJs,puourruruC 3ur^recal uo

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'I

In September1838, GeorgeFletcher Moore was advisedin a letter from Mangles that the latter wishedto terminatehis seedorders. Drummondwas notso adviseduntil afterthe departurefor England of theJoshua Carrol/ on 8 December1838. On board was a major consignmentofseeds,live plants and dried specimens,consigned by Drummond at his own cost to Mangles. Also, Lady Stirling forwarded GeorgianaMolloy's collections on the sameboat.

Drummond, perplexedby the changeofinterest ofMangles, was powerlessto recall or redirecthis shipment. After arriving at Mangles' door early in 1839,it endedup in the handsof Dr Lindley, who securedthe volunteer servicesof a young GeorgeBentham to divide the collection up and distribute rt.

Lindley was pleasedto receivethe new material from Drummond and MIS Molloy, as he had yet to finalise his manuscriptdescribing plants from the Swan River Colony, and severalmore new taxa were amongthose presented to him. Drummond was not to know the fate ofthese specimensfor some time yet.

Meanwhile, in a letter dated28 May 1839 to Hooker, Drumrnond indicatedthat he had decided to make anothersubstantial collection (ten setsof 1000 species)the coming spring, in the hope that subscriberscould be found. He indicatedin the letter that he had already gatheredsome 60 species "a of Orchidaceae,and that zealousyoung Germanbotanist - a Mr Preiss,had lately arrived here".

JohannAugust Ludwig Preiss(1811-1883), was the only notableEuropean collector oforchids in the early colonial phase. His south-westemAustralian specimenswere labelled with locality information and the dateofcollection, enablingsome of his movementsto be tracedwith considerable accuracy(Marchant 1990).Preiss's orchid collectionswere written up by Endlicher(1846) in a multi- "Plantae authoredtwo-volume work Preissianae"edited by ProfessorLehmann of Hamburg. By the time this work was published,most of the speciescollected by Preisshad already beendescribed by Lindley (1840)from Drummond'scollections.

The earliestorchid collectionof Preisscited in this work is of Calad.eniaunguiculatal= Cyanicula d.eformislobtained on 19 January1839 from woodlandnear the Limekiln (vicinity of Subiaco),Perth. Preisssubsequently made orchid collectionsin Juneat Lake Monger,July at the CanningRiver, Guildford and Darling Scarp cataractson SusannahBrook, and August on Rottnest Island before venturingeastwards to York in lateAugust to earlySeptember. On retumingto Perth,on 23 September 1839,Preiss collected the type of Elythranthera brunonis in woodland on sand.He made a few more orchid collectionsnear to Perthin the ensuingmonths before venturing toAlbany, wherehis first orchid collection was of Cryptostylisovata from StrawberryHill on 9 January1840.

I-ater in 1840,Preiss collected orchids ftom Albany and the granite inselbergWillyung nearbyto the nofih. The most importantof thesespecimens was the type ofCaladenia nara, collectedfrom Mt Clarenceon 5 October 1840.Back in Perth in 1841,Preiss made a couple of orchid collectionsfrom the Darling Range, including a small variant of Cqladenia longicauda Lindley which Endlicher recognizedas forma humilior (possiblyC. longicauclasubsp. clivicolaHopper & A.p.Br.). This was an important early insight ftom a Europeantaxonomist that therewas actually infraspecificvariation in a speciesof south-westAustralian orchid which many subsequentauthors would subsumein unworkably broad conceptsof polymorphic Australian orchid species.

In June 1839, Drummond wrote a long letter describingthe plants to be seenbetween Fremantle and Perth. The letter was publishedin 1840 in Hooker's Joumal. Drummond indicatedthat between

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'!u^{org ,{rolsrH [ ,{Soloprqro u?llerlsnv lllltsey'd dV puE raddoH CS lr'a)tria Vol. 14, No. 1/2 (2001)

Cyanicula andCaladenia subg.DrakonorchisHopper & A.P. Br. (Hopper & Brown 2000) asdistinct generaftomCaladenia. By the endofthe 1839spring, however, Drummond hadceased corresponding with Mangles, and was to lose interestin making further substantialcollections of orchids.

Understandably,therc must have been a mixture of pleasureand constemationwhen Drummond "A finally receiveda copy of Lindley's Sketchof the Vegetationof the Swan River Colony", which was publishedearly in 1840and drew heavily on Drummond's lettersand his paperon the underground structuresof Swan River orchids.

Lindley, working with dried specimensand without the insightsof seeingliving plants,described some50 species,not 60-70. He also did notrecognizeCyanicula andCalqdenia subg.Drakonorchis asgenera distinct from Caladenia. He did, however,recognize most ofwhat Drummond senthim as distinct, and this must have pleasedDrummond.

Nevertheless,in a letter to Hooker dated 13 April 1842, Drummond expressedconcern at how Lindley had lumped speciesof kangaroopaw (Anigozanthos)and Drakaea Lindley that to him were perfectly distinct and good specieswhen seenas living plants (seeopening quotation). In the same letter he descdbeda specimet of Calochjlas R. Br. found nearToodyay, "the only individual I have ever found of a most curious orchideousplant".

Drummond's waning interest in orchids was evident, but he did make one further interesting discovery in the spring of 1844 while travelling north of Toodyay to the vicinity of Lake Moore. In a bdef sentence,he unmistakablydescribes his discoveryof Calad.eniad.rakeoides Hopper & A.P. Br.

"Near our first bivouac I gathereda singular Orchidaceousplant having the hinged Iower lip of Drakea but with other characterswhich will probably refer it to the same genuswith 861 of my largecollection: only threeplants could be detectedin flower and one of them I put into spirits as you directed. Our presentmode of travelling is not favourable for collecting Orchidaceaewhich require close investigation of partjcular spotsin order to detectthem ..."

Drummond's collection number861 is the type specimenof Caladeniabarbqrossa H.G. Reichb., gatherednear Toodyay in 1839.

Taxonomic concepts

Our taxonomic conceptsare describedbelow. Few authorshave provided such information in previouspublications dealingwith Australianorchids. Fitzgerald(1888) was a notableexception (see his discussionof Caladenia dilatata and C. patersonii). More recently, Clements(1989) provided a brief accountof his conceptsof taxonomic mnk.

An explicit statementof conceptsis essentialfor the scientific evaluationof a taxonomic work. Without it, future readen are unable to unde$tand accuratelythe hypothesesunderlying the names provided,and cdtical testingofsuch hypothesesis rendereddifficult. While we acceptthat an element of subjectivity is part of all scientific inquiry, the best works in the field are thosethat presentideas and data in a way that ensuresthat they can be examinedcritically and testedby other workers. '(8uoJ,tr elp seur8al 3urra1e,r,r.ro erg lnoqr sessenSlerlrur;r sprqcroer{l Jo lueurJlop eqt ot uego) reUESurlool ors .(eqtl€I{t osoluawo1lt31o s1ue1drelncrged eql yo seynqullelect?o1otq eql eJe lzq^\ dllr€xe se^lesueql JoJ lno purJ 01 pecJoJeJ€ re.,nor8eql pue re8eueuruorlp^resuoJ oql qtog asuaspeorqeqt u\ osotuaaoly' 3;o ere suaunceds1eq1 s1 smoul ra,ror8 eqt IIe Jl tclperd ',{lJelrrurs ol llnJgJrp s1s1ue1d pelu,rqyn: ro; eur8er Surrele,ra.eleudordde eql edecspueleql ur ,rol stBUpoSSol.lalen -dlpuos€es ol'secueurure lseq8tq eql uo sllos poulerp-llo^\tuo{ ?uIql,{ueeq plnoc 'pouad perdnccolzlrqeq eqt puE qluoru xrs e re,roeurl due eq plnoc uosuesSuue.^dou eql leql elef,rpur 'eJu€lsur 'saur?eJ plno^\ ernlBrolllorll osneJaq JoJ eJrJlnoqs pourocuoJroAeuuur € o1dleq lecrlrtrd optrt 'tll'suas sepr,rorde,,ueser uorlel.resuoc B uo sr DsoJuawDly'J lerp Sur,r\otr) lseoJqlnos eqlJo seunp 'uor8er pelepJlosuocuo .{Jlunoc11e;urer lseqBrq eql ul slueld8uue,troU reruurns f,pee o1 lerolsudpuu eq1 'selSoloueqd ur puelur lle,r sdorclno altuerSelourer uo slueld Suue,trogrelult,t luo4 pue secueJelol 1ect8o1oceesre,rlp;o suoqelndod sepnlc:.lr osouawol! oluapa1z1;oldacuoc puo.rq e'aldurexe rog

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sa!Jadsqnspue salJads

'u.,'1org &otsrH I l3oloprqcrouellersnv ulatsoiA dV pu" reddoH CS Nu)rsia Vol. 14, No. 1/2 (2001)

The poor predicability of such a broad speciesconcept cortrasts with that in our treatmentof the Caladenia filamentos4 complex. Land managersororchid growerswill know, forexample,ifthey have plantsofC. abbreviataHopper & A.P. Br., that they arelate-flowering denizensofsouth-coastal dune country requidng good soil moisture conditions. C. remota Hopper & A.P. Br. is a winter/spring flowering inhabitant of inland arid zone granite rocks subjectedto prolonged periods of dry soil. C. dimidia Hopper & A.P. Br. occursin the wheatbelton good soils, etc.

To highlight another significant implication of the speciesconcept adopted, a broad view of Caladenia filamentosa gives no suggestion of conservation problems, becausemembers of the complex range all acrosssouthem Australia, and someate abundantin conservationreserves. Even if rare infraspecific taxa are recognizedsuch as George's (1971a) concept of C. filamentosa vat. d.orrienii (Domin) A.S. George,they will not featurein the national list of rare and threatenedplants (kigh & Briggs 1996) becauseit includesonly species.

In contrast,our treatmentsegregates outftomthe Calodeniafr.lamentoso R. Br. complex several common well-reservedspecies but also a few that arerare, su ch as C. dorrieniiDomin andC. elegans Hopper&A.P. Br. ( Hopperetal.1.990;Brown etcl. 1998).Ifourconcepts are followed, these threatened plantswill receivethe attentionthey deservefrom authorssuch as Green (1985) and Leigh & Briggs (1996) who ignore infraspecific taxa.

Ofcourse, we do not advocatethe elevationofall infraspecifictaxa to specificrank for this reason. Such an approachleads to unacceptablynarrow speciesand an often unworkable taxonomy (see below). But where taxa meet our biological criteria regarding strong reproductive isolation in sympatry, and are morphologically distinct, speciesrank is consideredmost apPropriate.

Perhapsthe legacy of greatestconcem regardingpast conceptsof polymorphic Australian orchid speciesis the diminished value of much of the information in the popular and scientific literature. Almostwithout exception,previous authors, despite makingmeticulous observations overconsiderable periods, have used names like Cqladenia filamentosa ot C. patersonii without citing voucher specimensnor even giving preciselocations from where their material came.

This seriousoversight recurs in popularbooksand field guides(e.g. Pelloe 1930; Blackall & Grieve 1954; Erickson 1965; Nicholls 1969; George & Foote (undated);Cady & Rotherham 1970; Clyne 1970; Pocock 1972; Hoffman & Brown 1984; Woolcock & Woolcock 1984; Jones 1988; Bennett 1988;Bates & Weber 1990). Perhapseven more unexpectedly,voucher specimensare rarely cited in scientific studiesof south-westernorchids dealing with mycorrhizal fungi (Warcup 1971; Ramsay et al. 1986),pollinators(Sargent 1907; Sto]utami-:,e 1974,7975, 1981, 1983; Bates1979; Peakall 1984, 1988, 1989; Peakall & James 1989b), chromosomenumbers (Peakall & James1989a),or hybrids (Hopper1973, 1979; Heberle 1982).

Without voucher specimens,it is often difficult or impossible to relate much valuable and expensively-acquireddatato the species segregated out from speciescomplexesby previoustaxonomists or ourselves.Conflicting resultsobtained by researchersworking on the sametopic may be due to the fact that they are working on different specieswithin a complex. One only has to read the various attemptsof authorsto describebroadly-conceived species s\ch as Caladeniafilamentosa sens.lat, to appreciatethis problem.

In many ways, the developmentofconcepts regarding these orchid speciescomplexes parallel that describedby Hillis e/ aL (1983) for the American LeopardFrogs (theRana pipiens complex). These -:

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'(6961) s1ueue13fq peleco,rpesu sercedsueql reqlel sercedsqnsse 'Jleqluollcrluor pal€eJllsoqole?zllC Urrlun,pu? Ipugouou JnoelUopun suollsreprsuoc JBInurS 'sercedsse uuql raqler sarcodsqns se exel asaqlprsAaJ ol rsetu€tsurncrcq"ns ul'elsrrdo:dduerorusruees 'dupe,ro;o 1y dluregec:eluer8 qll^\ pezJuSoJereq usc exel or{l euoz eql ruor; pe,touar suorlelndod reqlo ul pueqeql ul sJolt\ouqse{ qlr^\ ue^e'qloqJo suawwa '3 nopnoc8uo| 'S esrrduoc,{eq1;re:ns 'suollElndod 'd11zcr3o1oqfuour ueeqlou eABqo,{\ ,{ueuruI deFa,roso?uur rleql eroq,{laper8Jelur pue 'I repu.qs ,{re,r eJB BXBIaseql luql ,rroqs solpnts JnO sercedslcurlsrp se seuol O trl tuelJ hl 'aldupxa (ulttoq) suawwa J pue lpur"I opnprt8uol oluapop) pepruBer(6961) slueuelC JoC

'soseJsuos ul sldacuot sorceds,trorruu ,fte,r Surlecorrpe ',trerl dq furouoxul Jleql Jo sJesnJoJ surelqoJd,fiessaceuun eleerc (uu puv rno ur ?ururellsuoc 'sprqrro ool sl qJeoJddesrqJ {uer lse^\ol peJJeJerdeq1 se sercadsSurJnoAeJ uerlerlsnvJo slueulueJl rroql ur uxul cJJJcedse{uJ,{\oJ trre^ pe4uSocar(9661 'I66I) souol puu (686I) slueuel] ',{lSuDserelul '&oSatec '€xel eler.ldorddeeql ss selcodsqnsesn peolsur pue clJrcadsBqur.rog ,{1erre,t 3o quer eql Sursnpepro,re e,req e,r'serceds se pazruSoce.Ltrou uxel,{ueru JoJ serlelJEAJo esn lsBd eqlJo esnBceg s I 'pe,r1o,turs1ue1d 'uorleArllnc orll Jo dpnls pue uorle^resuoc eql ecuulpe flqereprsuoc IIr,^i.leql suonecrldrrrrpcpourd I lueclJluAIso^eq spJqrJo eseql roJ sldeJuoc crurouoxBl rno leql Jeprsuoro1t\'xoldruoc 3o.r3 prudoal eq1;o 'Brlu:]snV uoDnloseJleurJ eq] qlr,r sV u:elso,r-qlnosu! sex?1duocraqlo lere,resro; peleedareq ppoc xeyduroc'rg 'A osoluawol{ ouapoloC aqt u sldecuocserceds ro; eloq€ pelr€lepsuorlenlls ar{J

'serceds 'slueurredxe (crdfldlod) eruuseql Sursne:oar,.{eq1 1eq1 Surgode: 1nq :ra$ roJ selJads I luaJeJJrpEursn s:oq eoselSursoddo dq elqelunoJcesen {ro^\ raqrealo slynser3urprUuoJ eql Jo qf,nry a '(peztuSocer rrrouere 97) s?oq predoelgoserceds lcu4srp snoJeunu peepu erel\ a:ar{lleql uollesltueJ l 'salnqu ',ftolrpn€ 'lsrr8otoqdroru aql 01 Eurp€al p e,rrpnporde: puB lecruoqoorq a,rqcurls-rppeq 'uo11zs1puqdq 'o.r?^\ ex€l esaqJ ou ro eplq q1r,,rdrledurds ut perrnJcoleql exel ,{upruIJBJ ur ereqllsql peqsllqelses0/6Ipuus0g6IoqtuIUo.tlplalJellsuolur're^e^\oH's0g6Ipuus0t6IeqlulslslSolooz alerqegaaSuoulu ,{cuepuecseol asor ldecuoc sercedscrdd1f,1od eql sBrnoJ ol pernpeJsu^,\ Jeqrunu '6961 eq1 fq sarpnlspcrSoloqfuoru leuo4ua,ruoc uor; xelduroceql ur peqrJJsapuaoq puq saroedsZI 'suorgess? 'slururue q?noq11y snoruoruuJepue sqnsa:Suprlguoc qtr,r u?lJo eseqllnoqt uelllJA ere^\ slededJrJrluonssnoJounN'slsrAolorsdqdlecrSolooz;osuorleJeueSroJloolprepuelseqleJe,rsleturue

ruaorg diolsrH I trSolopqcrouerl?rtsnv uralso/d dV puDleddoH CS NuytsiaVol. 14, No. 1/2 (2001)

Hybrids

Some south-westernorchid hybrids are spectacular,readily identified and often encountered. Othersare rare, and warrant specialprotection. Many found by orchid enthusiastscause considerable identification problems because,hitherto, they have largely been ignored in published works on WesternAustmlian orchids (other than by Hopper 1979; Heberle 1982).

For thesereasons, hybrids are recognizedand named formally by us if they occur as sporadic individuals among mixed populations of likely parental taxa, and where their flowers are clearly "x" intermediatebetween those oftheir presumedparents. The presenceofan precedinga species'name denotesits presumedhybrid status.

We havenot formally namedhybridswherc their identification is rendereddifficult by the presence of backcrossesin many populations,so that a completetransition in form occurs from one spgciesto the other.In suchsituations, reference to hybrids is prefenedby the conventionalformula ofthe names of the parentaltaxa linked by an "x"

Gen€ra

Since the early 1980s, we have been examining generic relationships in Australian orchids. Initially, new genera were recognized on the basis of detailed numerical taxonomic studies of Australasianand SouthAmerican terrestrialgenera (Hopper, unpubl. ms). This work establishedthe presenceof some segregategenera within Caladenia sens lat., and also identified the generic distinctnessofwhat is now known as-QrorclrisD.L. Jones& M.A. Clementsfrom typicalLyperanthus.

We informally publishedour conclusionson genericmatte$ in colour field guidesto enablewide scrutiny of our conceptsbefore formal publication(Hoffman & Brown 1992, 1998). We arguedfor a reducedcircumscriptior' of Caladenia,with the reinstatementof Leptoceras(R. Br.) Lindley and the erectionofthree new genera,Cyanicula,PraecoxanthusHopper & A,P,Br. andDrakonorchis. YJe also prcposed lhat Lyperanthus nigricans R. Br. and Z. forrestii F. Muell. be transfered to the genus Burnettia Lindl., blutsubsequently (in ms.) proposedto erecta new genus. This latter conclusionwas supportedand overtakenby Jones& Clements' (1994) descriptionof Pyrorchis.

Recent DNA sequencestudies of the Orchidaceae,including the Diurideae and the Caladenia alliance in particular (Kores er al. 2000, 2001), haveaffirrned our proposalsexcept for Drakonorchis which appeanto be deeplyembedded w ithin Caladenia(Figure 2). Elsewhere(Hopper & Brown 2000) we formally describedCyanicula andPraecomnthus, and presenteda new subgenericclassification of CaladeniaR. Br. in which five subgenerawere recognized,three previously regardedas sections of the genusby Bentham(1873) and two newly namedas C. slbg. Drakonorchrs Hopper & A.P. Br. and C. s\bg. ElevataeHopper & A.P. Br. A key to all generacurrently recognizedin south-western Australia is siven below.

Key to genera of south-westAustralian orchidaceae

1. Subterraneanhe$s, saprophytic.Inflorescence a many-flowered capitulum...... Rhizanthella 1 : Terrestrialherbs with leavesand flowers abovegrcund, nrely saprophytic. Inflorescencea raceme ruynpuelS.rouale^elJ lou'poNnJu,rop ol lBtuozlloqsl€lad :tt se.racoldel .''eJlluesurSJEru'snoJerrJnu rllEJ'paqolqrunlleqsl ueerSse,real:91 ellaJoda.I elerrqurrlsurireru 'rIIBc8uD{Jel 'F^o flesre^suL,rlun11eqe1 seduls pet qlln uaar8se,real 91 J?lnpuelS,{llBnsn 'el€AEIc 'lceJe sleled 'tl 'uuoJrpasul euruel uPr{tJeuoqs qrnu ,^^up lou unlleqe.I :I I eaelnJlds sr?^\ogeJorulotqll^\'dqsogsed€Js:ZI EUEJIBJSJBd qtnod z ! Sururro; s8ur,t pe8JBlue-qcnuq1t,tr pue Surpuacsop{yenb1lqo uurnloC qcnol uo Sunrdsfle^rlce'pa8ulq ^\elc Iunlleqe-I :€I lou S "" BaerIBJ(I peqcnod1ou s8ur,tr relncunu mouuu qlrl\ pue spJ?/^dn ! Sur,rrncurunlo3 'elqour ,{le,rrssedpue peSurq,trep unllequ-I 'gI ( ',{rr,n' .71 (sre,nogg 1-7 rrsaor(JT ldecxe) sra,,rrog(g)1 qt1,tr sedecg 5 uuruel se q13ue1rues J /dlllJ:uloJtlJesurunlloqc-I .II I 3uo1qoo1 a1e,ro ,{11unsn 'snorqe13 ,lute^es ol ,{tu111osse,reel :4 sluapeluJ (4urd sarurleruos)a^oqe ueql reqlo,{llunsn sra,rog ilJor peSrelue luseq 5 ue qlr^\ srr€tl iecrunl pete,{e1,na; s ut poseruo,{11ut1red sproreqn; :g s " "" BJaqtuBfCX lser eql ueqlre8rel qcnu puu 1ucgpur1,{c,{11crq1 rred ( 'srrud leseqeql 71 e ur r11e3 eldrndqsrlurd sledas pup sl€led :0I """",{1lecrdu e1nr1uu,{3 euruel eql Jo esuqeql tuolJ azrsur Surseercep,{11unpu.r8 t 'snoreunu rlluC (e1rq,trro,trol1e,{ r(1orer) enlq sledespuu sleled .0I euruey3uo1e snore(unu r11e3 l1npsledes pue syeled ;o ece;rnsJeddn :6 Breqlu?rql,{19 "" . '.'...lesEq 'pe8relue'7 .6 : 111e3 dssop sludaspuu s1e1edyo ece;rnsraddn ro eld:nd 'en1qfllensn sre.tog :llec lurd IBseq l pe?re1ua3ur1ce1 srpq :ucrunl pere,{ep11nruE ul pesocuesproJeqnJ g 'etnsJ[l ',fuzlrlos ruaurl ,{llensn se^€e.I / polcuJluoJlou'eldturs erurTlod utunloc o1 pesserdde ,{lasop 1ou ueql 'pulurpl ql8uel eulesJr uBqlteuoqs qcnru ,{ypnsn,ne1c unlleqe.I :g 'pelecue^rp 'utunloJ snllqJoFg salJrpnpcolur pelcprluoc lsotule prurllod o1pessetdde ,{lesolc 'eurrupJ se r{l8ualetues'c,lnplc unllaqc-I .9 reqluu se qt8uel eruus r eu8r1g 3ur11nqe1ou s?ur,truunloc uo (lueserd;r) salcuny oo{ Iedasl?slop puz slelad :g "" s!;,{lsoJald JoqlueJo qlSuel ecr. l ueq} e.rorueru8pg raqluego luor; ur 3ut11nqes8u1,,r urunloc uo selcrJnv 'urunloc .S Sursolcuepooq e uuoJ o1pelrun ledes IusJoppue sltlod peJnpeJJo luesq8 unrprcsr^ .t :BrpouruBlseer; Surlcuy'JI€l luru S ueql ra3uol ,(llensnuunloC (sryrycottg ydecxa) ',{llenuu€ luesqe alorled :edEJseql ?urpuelqns paJ?[deJse^ee-I :g s!1.{tsotd,&C " " 3uo1urc g1-7 eloqad 'lEruueJad :edeJs?uue,rog ol luacefpeproraqnl uor; Sursue se^?e-I .g pedeqs-1eq1ou s.ra,trolC sptoJeqnl tuels-lool qll^\ tueld :Z 'setuozqr .Z BlpoJlsBC pedeqs-lloqsla,i(old qlr.{\lupld

'u./!\org &otsrH I ,{Soloprqrio uerletsnv Ulrlse/v\ dV pu? teddoH (S Nurtsiavot. 14, No. 1/2 (2001)

16. Dorsal sepalhooded over column 17. Flowerssessile, solitary...... Corybas 17: Flowers bome on a scape,few to many 18. kaf fleshy, ovate. Labellum lamina with prominent fleshy longitudinalridges and sparsesmall hemispherical cal1i...... pyrorchis 18: l,eaf thin, linear. Labellum lamina lacking ridges and calli denselyclustered, incudiform, pyramidal, cylindrical to irregularly oblong and serrateon top or hemispherical...... -..... Lyperanthus 16: Dorsal sepalheld erect splaying away from column 19. Scapesubtended by a fully developedbasal leaf, many-flowered. Labellumglabrous...... Cyrtostylis l9: Scapesubtended by a reducedbracflike basalleaf, l-flowered. Labellumwith calli...-...... praecoxanthus 4: Column absentor shofier than 5 mm, with prominent ftee or united (mitral) staminodia;viscidium present 20. l,eavesterete or tubular 21. Labellumon lowerside of flower,below dorsal sepal...... Microtis 21: Labellumon upperside offlower, abovedorsal sepal 22. l*af well developedbeyond point of departurefrom scape. Labellumimmobile. Columnwith no basalfoot. Antherheld vertically and attacheddorsally at the base,with a poorly developedrostrum ...... Prasophyllum 22: I*af reduced,and bract-like beyondpoint of departurefrom scape. Labellum highly mobile. Column with a prominent curved basal foot. Anther held obliquely erect or horizontal and attacheddorsally at the apex,with a stronglydeveloped rostrum...... Genoplesium 20: Leaves channelled,distichous or flat 23. Lsaves 2 to many. Labellum prominently trilobed. Anther and stigma E sessileon ovaryroof; staminodiafree, not cucullate..,,., ...... Diuris B 23: Leaf solitary. Labellum not prominently trilobed. Anther and stigma on a short column; staminodiaunited into a cucullate mitra or free 24. Labellum petalJike, lacking appendages Thelymitra 24: I-abellum clawed, with basal calli or denselyhirsute 25. Labellum samesize as petalsand sepals,glabrous except for c basalca11i...... Epiblema 25: Labellumlonger than petals and sepals,densely hirsute...... Calochilus

Acknowledgements

The intensivetaxonomic researchon which this paper is basedspans two decades,and aroseout of a pilot mapping project on south-westemorchids conductedin collaborationwith membersof the WestemAustralianNative Orchid Study and ConservationGroup. We aregmtefulto many colleagues for assistancein this work, including Garry Brockman,Eric Chapman,Mark Clements,Chris French, -''T

tqued :luaoo8eull^lpu?'I pulr uortD^rosuoJJo tucuu8doc) f,ruo8v sorle^resuo3 '! olntsN u?lprtsnv'lrun sercedspereSu€pug oql ol uoder I?ulJ 6I oN polord dsg VJNV splqrro uEIIeIsnV 'soleoJ 'S 'srre}sre3 S1 ru.lseiA p.uelserqlpue eru o^!JJo .{SoloJauoqelndod pue scrouof uollE^r.suoJ(n66I) fO d' 'gzl el I DdDd snoautnarsiNuo\olrossv otno! pun orol! snouaSlpul?rlelsnv uols?a lr qlnosuro{ (eeecsplq6) 4 a otpotsop pttE )g i sutotSotlq)')g Eol apoto} ul sxel ,r\oN (toot) ,ffC'rj*C 'Llt 6tl .\9)9 ouallanw 'rrlpqsnv 'sotp/A 't 'uBI) qlnos ^\eN prrr Euoprn tuog (.eeruplqcro) ry oluapDlo)Jo salcods^\oN (886I) ,.'4C 'I6I rE '(oroc?plqrro) 'uB:) -Eg1 :(d9 ouananl/,lErtEr]sny orrr^ ruo4 solcods^\au B DlosDtoxDluapDl') (986I) ,^Ac ('troup,{S:ssardcruepecv) 9II 6/ dd.,ul?rtsnv ul stueldpu? oldood,, (spa)IaDS.llE)t fC'r?J.rl o-rorprsllrqe'IJo suoltcaltoc oqJ aJrrpauueqr V 1qI86I) tC'rrult t{CS'ue:) (,{eup,{S:ssord rluropDcv) .. ele4snv ur slueldpue atdoad,, (spr) (et86t) },iCS'ueJ T'I('ue3 'l'trpeJ tIeupIS:peau l\.v ?t HV)..inoloJ u! splq.ro o^lr?Nue!le!snv,, (0/6t) XS'uEqroqlout '06 tL :09 nu ^nv j,t]',A4Io iDpos pha 'puBlst aWlo lounof Et]|P"\snvuletsot Soyrg 1:16 Jo eunq pue tlou oqJ 1816I) sv'e8rooc ?i vV'o8prqrng ( uopuo'I :rot,iED ,, S08t-208t sruue 'u^\org senbunlelurld seretcPrertrsueqrqxo uelu?rc-u?A eelnsul to oelpu?lloHo?^oN oBrolJ snurorpord,, (0lgt) f ( ouroJ :tururodDuet\lpue-I pu? uonPNosuoJ isucc-uosuJoql"dV'u^\org Jo tu.rttlled.c) .. erotc pcucrsorqJs.?lt!4snv uratsetA,,(8661) cN lueqrreN d' J .leloorg ( eunoqtat{:ssord qqut) z lo^,,srd,tlt3ngot eprn5pt"rc,, (ooot) V'C'Srurrl) t'Hrr{ '68t I :(l)8 ,rs/{r/y' urn{snv uletse,^dul (oeec?urr\t- uepv]!^{aoaadsl^a1 saees g) solrodspol e psu'nunp g'oopuo sadqorhgJo uorsr^errtluouoxnt y (t66t) C'S'reddoH? HIII'reloorg '(osurruoppleo Ee-91 tlt Kuotoglo lt lnof uollt1ttsllv :opo.pp!q.ro)sluolSollqJ Jo serrodso^rtdocop lllenxos ur uortelosra^lt.npordar p.lelpaur-rol8ulllod Jo uoltlrJlsuouoc(966t) JJ lo^\og 'LgEjgt:tL uoryotproaqJ ,{ruouoxul^{ou oql Jo erucJoput (S66!) IC'rr^{og 8CC-t€f :0t uDrynpro )4J (:n:ocprql6) o0ulluapplp)aqutqns ag ur sprqrroa^rtderap trll?nxas Jo,{uiouoxvl aqt u! sroleurllodJo asneqJ (266t) lJ !0,{og .egz_gLz tn anuD.ltaH 1o11o4DNsalrll qtnos {lloxnlg / aN aW worl suoqnqlrrL/oJerlerlsnv ur (epe.pprq.ro)uouaS petelar pue lleny{ C sn1.11lcott!uy(726) d d ( spuElpoN:ssord PrlPrlsnv urotse,{Jo {tlsra^!un)I uxd 4 sro^{ouptr^\uellerlsnv urotseiA,{\ou) ot ^roH, (t961) tg'o^euCt At ltplrElg 'I) CpoJqsv:oo d' o^.ou g6E-Lgz:g IoA .!slsuelt?lsnv erolJ,, .u/ enaplqrroxxc ropro (€/8t) D'uDqtucg ')tr?d (quod :preog+e6 s8ury)q ?ltE4snvuratsey' s8ur) Jo stuEldpuelqsng et{L, (8!,6t) t{g'lleuueg '( 'Dq.,^A apralopvluud t,^oD) rllpl]snv qlnos Jo sprq.ro (066I) zf ? U'sol?g tS-6V :tt u.,rrpt rtuDtog JTQLVV )qt Jo tDulnot 'uxpl ^{euo^{t Jo uorldDrsepaqt qlra erlerlsnvuretso,r^ ul (aca.eplqr.ro)storrrl,{ snur8a$ uo satoN(066!) !'soleg 'sctB{ nn-Zn t() toulnor Dtt tsnv VnoSlo tarros plttito a^tDN rg d snltqotl7 snua8aqJ (686I) U '68-Sn :L uaplDD r1llDtog aPlDt?pV a1flo lolunol A8o'otquo setoupup uorsr^arrlurouoxel e :(eraceprqr:6)tg \ sltorrtt4tsnur8 eql 1t86t) U'srlpg '0I-6 :(It)e tr!)nor Dlo4snv qtnos 'sot8g Io {Dlros plqxto adr,/rNsrol8ul od tue slt pxp Dt,uqw{ Dtp)odaj Ot up.d- sprqrroJo uortrur od (6/6t) U 'poo^lSurloJ ( aunoqtet\t :,{leoos,{uBtogtrl)euatsds uElerlsnv) 98-rg dd.,ErsElerlsnvur ,(uEloarrlEuets,{S Jo ']loqs 'slsunllncuoq &olsrH,,(.pe) S d :r1 ?lou uerprtsnvoql Jo uollntuouncop,{lrEo .ql ur sr.qlo puesroroldxo 'srolrellor 'ro{rDg 'r.U?g Jo uoltluSorarpu? eloi .ql :pe{ooFe^osuoltnqlrtuor prru?toA (066I) U IA T I^l U

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'I 'u,{ora tz fuolsrH {Soloprqrrouerlelsnv urotsa/\A dV pu? raddoH CS NuytsrcVol. 14. No. l/2 (2001)

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BrrEllsnvJo sprqcro,,(spo) gJ'rrnt{ ? :uI (696l HA ,{l '9IZ - 'slloqrN IIZ 199 tsllontoN uoirotxtA L ert?llsnv jo eEoreprq O oql ol suo[]ppv (056I) HlA 'OLZ-LqZ 'irg 199lslJortllDN uolroplA sal.eds raqlo ro solou,{rpuns pue soDaue^ ^\!u errl{l oslE U - 'H 'sfloqrrN ouapoloX sana8 aql Jo solc.ds ^\ou V llt srllrtsnv urolse^,\Jo eBareprq O oU ol suolllppv (6t61) nA '8tl-9€t - '!sltoqrrN :t9 ltlltun|DN unuolrlt t erlPllsnv urrlsa^{ Jo epeipprqcro og ol suolllppv (/t6t) HnA ( uopuo'I i)toorqtpl^ ,{) ., serurSroqv oql Jo oSBnBu?.I:ql Jo ,ftElnq"ro^ .^ltdgcsed V osl? pue eller1snv urrlsa1t ur replas dt:tA uE Jo eJr.I InJlua^g sreo1 uoJ Jo ftErq' (t881) CC'eroo4 n6n 4Bt:t Dtst[nNat aqrlrd souaseqlJo uorsr^ad y n- (eElresorurN),?rrrysnuoSaq]ur salpnrs(9/6t).U.B,uqsElt '(sus^ qtnos r'rul {lolcos,{u?togrlleLrr.tstrS uerl?rFnv) '9€t-Igl dd..?ls?prlsnvur duslogcrl?uols,{S 'IVf Jo &olsrH,,Ipe) Sd']loqsrrl ssr.rd Jo sollllEol SulpellocuertErlsnv uatsay'A ar{f (0661)CN lunqrnt{ (qu.d :sloog looluvl .,9z8l-e09t ErlPnsnvuralso^{ qtnos ur oseq -I f rlSolsrlspue,{uolor lsuod u puno3o1 slduapu pu0 suorteroldxe qruerCIo,(pnts E olerlsnvoruttC,, lzg6t) luEr{rret{ '69t (. llst :gE salD[4qlnos ^(aNIo 'slsturlog .H &aDoSto{ou aWlo s8ulpaarordpuD tDurnof lu.uiolddnspuoces usrtertsnvJo sproceg(1761) f,uoprsj^l 9Z-9 :Z .stsrrr?rog &aDoStuotstH l\laoN uolj $nV urarsarhaW lo trrrnol uprrulsnvurolsei^ Jo spjocau(606t) H.I,ueprDl,{ 'EE-BZ 'Zn sato[ qtnos ,]/.aNlo KJalxoSloKo\ aqt lo sSulpaDotd 'urprpt{ puo loutfiot'E\n\s..v urltsrlA Jo urolJoql qll,1(lpep e^eqoq.r stslu?logutrnlrltsnv lo .prorcU lgOet) H f ,cu"o.t Cspuelp.N:ssord eltellsnv urrtse1l\3o {lrsre,rrun) € IoA .. BllBqsnvJo slueldsnoro^turrJ,, (SOOI)V 'lo^ 'au^ro-l ( spuelpeN:sscrd prlerlsnv ruelsc/A Jo {trsro^run) Z <,Erlrrlsny Jo slutld snoro^rulDJ,,(6961) V ,a!,ro.l CspuetpaN:ssard rrtt4sny urclsc1{Jo Ilrs:ro^run) I lo1 .,'BnrxlsnyJo slu?ld snoro^rurDC,,(/861) V 'rcls!i0d ',{etpur.l Iecluelogoql Jo seunto^EZ lsrlJ .ql ot xtpueddv truotoJre^ru ue^\So$ Jo uollDlaSo^eqt Jo qrtots V (0t81) I ,,ialpu'-l ( uopuol :s{e,{3prg)q slurrldsno.replqrro Jo sJreds pu! prauao eqJ,, fut{t8t) f '0zl-l:uortPcrlqnd 'st0eld ,H ,q8le-l IErrodSertruos eJllpl1l\ pur surd |?uolleNuellrrtsnv unrtel}snvp?uetl]orqJ (966t) d.f's33u8 t f 'sserdu! 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'u^\org -C.S &otsrH I ,{Sotoprqrrorerllrtsnv ruolsoid d V pu" rrddoH NuytsiaVol. 14, No. 1/2 (2001)

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"The " Salisbury,R. (1807). Paradisusl-oninensis (William Hooker: London ) h' Sargent,O.H. (190?).Pollination of Caladeniabarbarcssae. Journal of the WesternAustralian Natural History Societ, 0

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