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080057-08.022.Pdf 'l]loua8 IIts seqsorJeds puouppe eseqlJo eJnlBI3ueuoupuc uorlducsurncllc1ca:roo aqJ .rau eql Jo reqloro auoot poncJsrffJlaq IIr,r r.atrs{-tqJlldH]Ulpsarld,{llu rnr sarJodsrequnJ leql olquqoJd sr 1r ,{11cn1ue,rg'annidtpH Jo wns{-ttl)llaLlJeq1e ol Fur8uoleqse petuer1ueeq pcq rlauoS,\\ou o,rl eqlur ps^[o^ur serJeds eq] lueutssesse s.SJeqJepuv ol Joud xolduor,saprotfuoJs,DlpJuaLADT 'serceds 'wnt 'puoaas aql ur Sraqrepuyfq peeeldele,\\ qJrq,r ;o rno; xrs sutt;tuoJ oduuaSlv 'dlg/{unpLtdaros[/qJ xql ur ulrq.(q peJeld euo pue o8ulquesst] ,atltuppoqlJ, eql ur SJeqlepuv ^q poJuldserJeds euo s apnlxtrr'DdlDJowauy lsJrJ oqI ?JeuaS.rouo,rl Surqucscptuc l radudsrql u1 ',{p$s roqunJpopeou puc tueulte.l1srq ur paquosunJ.rrJsn cncl.{qd/ilod ore,r [elpurl aUtuDpoLlYpuc AxIpurIDlpruafiD1 Eroue8eql teqtpelcJrpur ]eq nJ"1, do'SJeqJepuV 'tlrouo8 lJUUSrpJo requnu ? ur poccldeq plnoqsueql ur popnlcur,{lsnor,re:d solJods usrlc4snv 'Jsues eql13q1 pu? lrlf,Isnv ur rncJolou op lJr]lsJql ur'nrJUJSssJqt lcr{l peluJlsuoruop uoeq seq 'I861 l! ereq/$'(I66ISroqrepuv'686I uoslrld lllng ? pltrlllrH)sndudluacar pre,res ul possnrsrp uooqsuq urleJlsnv u\ re\\\\I uns[,!.lru a H puB')AwntatdtlaHtrIouaA oqt Jo uorlduJsuncnf,eqJ uorlrnpoJlul 'tg)d snuwDqtozOol pelulet flesolJ lsoul ^lluopr^e st lr :I z3JnI sapoqdal snuwDUtozolsaporqdai Vpue'['llen] gr-/arrdr wnst.rUulaHl ttaJtds V't4eelswna^nt 'l'ltqv-IwuDqpap 'lrepuos urualduzH\ una^ru V unstry)lpHl wwDqp2p V unualsi opuDlq unstrqJlpHf anaDDlstlopaDlq V :sarx.ds:eqloe.tr1 sapn1cur puc lropuoStu rllal snlqo uns{-11.!r!pH] 'C 'dleld uoslrld Insd (lopuos) wntolswqo unupduuaStv uo poscq sr snuo8 puoJos aql anpy dacos tttq 3pnu uosIr ld g Inrd('Jq)wns{rtlJoJnaTerznaBeql ol paptet,(losololsoru Inuerudds sr lr:[uoslrld'g lntrdallpxDswnnidtlaH] sUuDrDsV ptrc'l'letIN'Jwnrydappod wnstqJlpHl tunrprdalopod y :sorccdsJoqlo o,rl sopnlourpue eJlu:1snvuJelseld uoJJ DloJnp) ndtD)ouauv sal3ads,lreu ? uo poseq sr snue8 1s:r; aqJ eeurqlurr8uv :ororluqdru5 eqt ur pequJsap enwryllorluuaBtgpueodtotowauytnue8l[auo^\I'(266I)09V-LVr:()grrst.{n61 'ulltrlsnv tuo{ 'rar r:eue8 ,,neuo,tr1 ! Dduu a3 tV pw od.tu owauv E lr[.d.(ecerIeqdcuC:euecDJa$v)utnsKrqJtpHpuv wnraldtlaH ur papnlcur ,tpuel]nc sercedsusrlellsnV auos lo uortDJrJrsselJollJ Clnnd'uoslrld lJBrlsqv zsl9 R'tvllsnvurals:lA oruoJ tol xoBod 'lu.ua3eu"t1,l 'unrrqraH po€.IpuB t|or]p^rosuo] Jo tsaluued.e u"rprrsnVtUalseA uosul( I Ined BlleJlsnY ruo{ BJaua8A\au oa\l tMn\pdrluaSrv pua DdlnJouauv C lJsd :(aEallEqdEu3 :eBaJuJalsy)unstq4laH puB wnrafinag ur pepnl)ur ,{1lua.r,rnrsalJeds ue!luJlsnv aruosJo uolluJglss€lc aql (266r)09',at' :(g)gprsfnN NuyrsiaVol. 8, No.3 (1992) to be elucidrted,a taskthat will entail fuflherresearch and will involve the studyof materialin Australianand overseas herbaria; for thesereasons only thosespecies are treated whose names are requiredin regionalfloras and whose taxonomy and nomenclature is clear. Methods Dissectionsof achenes,florets, and involucralbracts of represenhtiyeAustralian species of 'Helipterum','Helichrysum', rnd of relatedgenera have been mounted in Hoyer'ssolution (King & Robinson1970) and examined under the stereo and high-powered microscope. Achenes have also beenembedded in Spurr'slow-viscosity embedding media and sections stained in ToluidineBlue. In additionmicroscope slides were studied of transverseseciions of achenes,some of which were preparedby P.S.Short, K.E. Wilsonand J. Nailonfor theirpaper ( 1989)on acheneanatomy while otherswere prepared for me by my colleaguesR. Cranfieldand C. Parker. Morphological studies The morphological charactersused in discriminating the generaand specieshave been described (or referencesgiven to their description) in a number of earlier papers(e.g. Wilson 1989, 1992b): supplementarynotes are here provided on some of thcsechffacters. Achenial hqirs. The natureof the hairs or papillae on the ircheneswas studicd. Various typesof hairs and papillaefound in the Gnaphalieaehave been described by Anderberg (1991) and their taxonomic 'Helichrysum 'papilla' significance assessed. In the elatum' grotp an additional type has been observed,in this two cells ofthe pericar:pepidermis are lateralty insteadof verticalty placed;they rre transprrentand often myxogenic. Papillaeof this type arerefcrred to as2-celled feneshae* . In some speciesof this group lhc fenestraeapprrently consist of only one longitudinally elongatedcell, this cell is usually not myxogenic and is not raised above the surfaceof the achene. In Anemocarpalow, rounded,2-celled papillae arepresentwith the lower cell slightly exceeding the upper. This papilla type, which was describedby Anderberg (1991),is comparableto that found in the Waitzia grotp of genera. In Argentipallium short narow-obovoid duplex hairs are present;these are similff to the more elongatedduplex hair type that is found in OzothamnLrs. Achenial neryation The vasculatureof the achenecan be determinedby mounting it in Hoyer's solution. Where the testais free from the pericrrp, or where it can be readily sepffated from it, the vasculatureis preferably examinedin an achenein which the seed(i.e. testa,endosperm, and embryo) hasbeen removed. In all Australian membersof the Gnaphalieaewhere this organ hasbeen studied, with the exceptionof Myriocephalusguzrilae F. Mucll. (seeS hort et al. 1989),there are two yascular strandsin the pedcarp. In the testathere is always one strandand this passesfor a varying degree longitudinally arcund the seed;it is bestobserved by studyinga whole mountofthe testafrom which the embryo and endosperm(which is usually I free from the testa) hayo beeo removed. * 'papilla' Thi s ty pe is alsofoDtrdi\ Bracteartha as is Dotedby E.E. CeorgeadouTvrtkovic Sahin (1982) who refen ro rhem as '2-celled idioblasts'. It is anlicipatedtlntthese willt'e morefully described,and tbeir taxononricsignilicaDce assesscd by ber in a forthcoming paper. 'lou op snuaDqlozo snucSJcllurts ^lprrSoloqd:ou aql ur serJedsalrq,r 'cznlJrocfurol:a lddJo1Qun4pduuaSry qtoq) ropuos wnuDDlsi.topu'lq H puu lllqcl wuDqpap wnsf,rqrlpH '(1661) Sreqropuy ,{q por0ld str,\\ wnrydappod H srrue?qxrq^l' wwnpqdarosf,.q) Jo sanads qlr,r slsclluoc osIDlI uzrquo3,{ruotJe wJolleql DtzltDAlprn ans{-tuJoJnaTEJeueS pclcler .{leso13,tlo^rlund ew ur sorcadsqlr^\ slsr?rluo3 'czrquoc,{uro13e srql urJoJol lou punoJsu,t (Dd.tDJowauv)'\entv'lwnpdalopod unst4JlpH 'palsol ueeqoAf,q Ipnls luesJd oql ur pe^lo^ul sor3sds,reJ 'serreds t dluo ,{leluunIoJUll yuzrq::oJrruolae-uou,(luo Jo lezrq:-Ioc,turolce,{luo ull?luo, ol punoJ uooq reJ os o^uq erouoSoleSorSes eseql JIos,{u dq pue (I66I) Sraqrepuy fq pesnrSocorerouoS eql 'JaAe,\,\oH'lczrqrros,{rrolco-uou olur paleSsr8esJrB sor3eds 3ql u0rl,r Ip ro IEzrquoJ^ruolsoIle mqlro .{1pnsneru snuo8elrsoduoJ tJo sarJods'p.€ue3 ur'olurs Jcnsnunsc.r snll lcrll lno pelulod dnJrui6 'ezrr$JoJ.{rrrolJ3 rurolwns tJq Jlp H to unDfitl2 H tatllraurpspnlcur .{lsnor,rsrdueoq pe q ltql soneds uf,rl?.rlsnveqlJoJIBqlnoqu,{luolnql(066I)dncJci&,{qu,roqsusoqsDq]I'suollDlJossDlDztqtroJf,Il .snuwDl.llozo ur punoJl?ql ol JclIUls sr uot1cnllssrqJ peue{Jrql pu? elu^rlJ aulloq.ffq llurulel orll q1^1 spJu^\dnetu11:qrcq aru,{aq1 '3SCq 'lualsrsJed 'palrun e^Oqtr{CeJq lou op puB 3JD ,{l oqs ,{JeA3Je sensuq aql an! oduuaS-tvuI 'pJUJ)JLIt tOUInq lUJUrUOrd pu€ pelseSuoJaelleqnq purrursl oql qlr.arspre,,$dn olclloqreq olc sellsuq eqJ. wnsttqJornal Io ptrc dnor? awDla ansLqJtpH etll Jo selJodsur selsuq.ql op snJouueu eu?s eql ur eslq eqt e^oqr] r-rturI J {cerq ,{lFrfllua,redeql ]nq luatsrsredpull eer; eru sellsuq snddcdeql ndJD)owauv u1 snddDJ 'lou op sleqlo elrq,&sauoarels elutsouoJ o,\trqol ll]eddBsarceds suros'(I66I SJoqrepuV)poqucsunJJrJ,{lluolncse'snuu,tDt!1ozOsnrrefloqluI'louopdnorStunulawnstulttlap eql lo pue Ddn)owrlly Jo serJodsscoJoq,{\ sourooJels eleJlseueJ e^uq 1rtlrllDd\uaSlV s\uefl e\l Jo 'aJnpa3oJd 'xelduroc sermdseqJ srql ol olqcuorucoslB aJc xoldruor srql Jo sorJodsueqc.qsny eq1 unst-ltp!|2 H - wnlloqdoug egl u !ile'le8 eleJJsrpolur sar3adsJo uorlBferSaseqt ur tsrssBot (I66I) Sreq:epuy,{qpue (186I ) nJng puuprulllrg (q pesnueeq srq:al JuqosllJ'suor8eranbrdopausprql 'slore eql uee,rleq luaseJd eru',ee.qseuaJ,se ol paroJor oroq luocnlsucrl peua{Jrqlun leql pue erlue lou sr euoe.rolscql uoua8 suos ul lcqt lno pelurod eH (016I) ,tlnro .{q peqrrcsep,t11ucgur 'etljoe'lels 'slJcrq lsrrJ sB/( oql leJJnlo^uroql Jo uouod psDq snour8Elrusceq L sDDJqplJnlo^ul 'rg')I 'pacetd, snuwD\JozO|resB oql osp sr qrrq^r llerpeuroJe spu0Jls Jslnrs'?^ eql lanqpd4uaStg u1 'drror8 ru lDp wnstt{JtaH eql Jo serJedsur peJeld ,tlle:elrl osp eru daql 'uoslrld 'g 'tpuoi& InBd ('lC) uzs,tr{JoJ[r7 pue ,lzt.trr4{se^lleleJ lsasolc e^Uqnd slr ur aru feql se suopal,{1oceq1 ol uonBlerur peJtrld.{ll8laltrt orf, spuBISrcln3se^ lBruoqreoql Dd.tmowauv nI 'reded slql ur pessnasrpsdnoJS-serseds lo 'oueqcD l]JaueEeqlJo nsJo serJedse^rlrtuasoJdeJ ur pe^Jesqo ueaq seq uonlsodnoqJ erI Surlcesse,(q '(tueredsuul 'eueqJB ro dltuorJqJnsoJe €seqlJr) slunou eloq,\\pell]el3 Sururur[xe,{q eql Jo suoriJos esJe^sulrJlSururrulrxa,tq pourur.rclcp eq,{twprn ('ltt do lon lJoqS)eJupJrJruBrs tuouoSJo srodjqure aql Jo suopel.{lo3cql ol e^rlBlsr fuBJued pw elsel 5ql Jo spu0lls JelnJsl]^eql Jo uorlrsod aq1 tuhtpdltD3tV pue rtl tDrokly uo\t'M C lnEd 450 NuyrsiaVol. 8. No. 3 t1992) Genericstudies In a recentpaper Anderberg (1991) recognised the genera C hrysocephalum Walp., Bractednthq Anderb.,Ozothamrus R.Br., and LawrencellaLindley serru /aro as segregatesof (Australian) Helichrysum.I havereclassified th e taxl.in lheLqwrencella complex (Wilson 1992a) and most of theAustnlian
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