,{lpoueue8eq ol pareplsuoJaq selJedsel8urs slt eauls,ttrolrsJo uolsnlsxs Jtll lqllll''wnsfuqrtla H 'palsleJ JO uo[Jes e se (L98I) umqlueg ^q pastuSocerst,tr xelduoc eql oq ol ponplsuoJ uoeq eaeqxelduroc nJJa:uaJn\D1aql :;.pepqcur sJoq3rr leql sercodsaq1;o 1e:o euos eulll Suol? Jod

'pepr,ro:d aru qrclep e,rrlducsap;euq dtuo palu8lls€^ul llxul aql Jo euros 'uoncallqnd Jol 'uortuoderp-ro^e puB orurl e^es ol JapJout pult 6uItuoJtllJoJslql Jo ,\\eI^ uI ullsrlsnY pesluSossl Jo erol{ oql Jo 8€ ournlo^ ur aBaflllolsv ,(llulrJ aql Jo lueuleoll e loJ uolpjEdeld ul Jo poqu3sep Euleq eru BreueEeleSe$os Jo laqunu tr lueulsor1 slql ol lol.ld uollerederd uI sl elprlsny ulxeldurotwnnldqag-unsf,L{r.tlrHaqlJo uollcJl}sslllJ e^lsueqalduoJ J^tlDJoqclloJV

'dnot? otzlnA4 aqt ol '.(pnls peleler ,(lqrssoduoxq Jrd,{louorulJullsrp ll sBpeluol oq ,p4l29 snueSoqJ loqljnJ psrrnbal 'zleels ''Jg';.g 'I8eBH 'I 'pelou aq 'r{Jrq,,'rxelduroJ p aJuahsry aql pue Druaollzs snuwullozo 'y 'd,le yqrapuv DtluDapDtg L\ unlDqdaJos[]?J uI ureql psculd pue ansfr4Jtlrg uI popnlJul 'eeslpqdrug uasq ,41snor,re:dpeq tsql selJedsuelprlsnV eql po^ourorslltl e11 aqrll oqtrpJ,\1er^Jl scq (I66I) Eroqropuv ,{puerell '(9361 l8oeH €s) snue8aql ulqll,r{ esoql ol ucql:eqler ueua6.nqlo u1 selJedsol ^lluanbe{ srselcods eseqlJo soIIuIJJ? lsesolJ oql ltll pull selcedsJosSf,lquessepluJlUD ue '(I86I sr etp'rlsnv ul poqlrcsun3r3 s3 snu33eql ;ql pesruSocu:aqunJ e^eq feql lung T prulluH '.LL6I'lD Ja ralll!]|ixtetr^l3'e) snue6l?qlgo en1ueado.rngpuc uBcuJVqlnos oql qll,^\suaue8uoJlou ateuns(ttp1ag ur pa ld sor€ds uallf,Jlsnv eql lf,ql pesluSo3sJo^Bq slslusloq Sread,4ueur :Qg

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(266r)t/€-r9€:(€)8 en nN Nuytsia vol.8, No. 3 (1992)

distinctdue to its flattenedachenes; he noted,however, that this specieswas otherwise similar to speciesn Helichrysumsect. Inrerencel/a. Mueller (1889) recognised the relationship between the onespecies ofsriaerra andthe species included by Benthamrnsect. Lowrencel/a for helisted them togetheras speciesof Helichrysum,as didBlack (1929, 1957). Earlier Black (1915) had suggesred thatif schoeniais to be recognisedas a genusdistinct from H elichrysumthen H. ayersiiF. Muell. mustbe includedin it. Haegi(1986) placed sciaenia cassiniana(as Helichrysum cassinianum), Helichrysumayersii, H. davenportiiF.Muell., and 11. sdmr/erlrle F. Muell. in sect. ktwrencellabrt notedthat this sectionshould have generic rank since the includedspecies were very distinctfrom othermembers of Helichrysum.In a recentpaper Anderberg (1991) indicated the closeaffinity betweenLawrencella and and recognised that further work wasrequired before the status 'l,swrencella' of relatedtaxa could be clarified. rn the complexhe placedHelichrysum ayersii, H. davenportii,H. lindleyi Eichler (= Lawrencellarosea Lindley), H. (with its close 'xanthochrysum' flifolium relativeshere referred to as the group),t1. spiceriF. Muell. andF1. obtusifolium F. Muell. & Sonder.I concurwith his suggestionsexcept that I excludethe last two specieswhich I considerto genericallydistinct.

Tribal classiffcation

The speciesincluded in theLawrencella complex have traditionally been placed in the tribe Inuleae.work on the tribal classificationby Anderberg(1989) has indicated that lhis tribeshould be dividedinto three,the Inuleae, the , and the plucheae. The Gnaphalieae, as defined by Anderberg,comprises the taxa that were includedin the two subtribesGnaphaliinae and AthrixiinaebyMerxmnllet et ol.(1977).In arecenf comprehensivetreatmentofthe tribeGnaphalieae Anderberg(1991) has rccognised a numberof subtribesincluding the subtribe Angianthinae Benth. in which Lewrencellaand, its relativesare placed. Anderbergexpressed uncertainty as to the systematicposition of Bellidabut, asindicated above, I considerthat it shouldbe includedin the Angianthinaeand that it is closelyrelated to Lawre cella.

Taxonomy

The Lawrencellacomplex

Manyofthe characters that havebeen examinedandused todelimit thegenerahave been discussed elsewhere(wilson 1989).There are many morphological and anatomical characters associated with theplants that are of valuein determiningrelationships, however, not all of thesecharacters can be so describedthat they arereadily comprehended. Thus the textureof the corolla.the shaDeof its constituentcells andthe thickeningof thecell walls arechrracters that are of valuein suggesting affinity,though sometimes of timiteduse in formaldescriptions becauseoftheircryptic natur;. I hav; expandedbelow on someof thechancters that I considerto be importantwhen assessing affinities.

A numberof charactersthat arc found in theachene are useful in sering eitherto distinguishthe I'awrencellacomplex from othertaxa that have been inclu ded in Helichrysum,or to distinguishthe putativegeneraor infrageneric taxa within the group. Those characters that are used in thispaperare asfollows:

,L pue,srcur,"nru,o.",,"uno,o.ounoff,""'iJ:il",1.iil i:t#'itX:l#fi:;::JTit.'#:ir;i:": rltr,rslseJluoJsrql'd4s1ro13u1puelxepusllsfllnJsc^lnolseqll,1plollap xade oql'3uol oJe suuu aql xaldwocollaruatnoToql q puers rqncs?^orllJoluolxepmessou{ortl ot$ pue'xodee4;o,{lrso11tdcd ;oaarEappueedcqsoql'suuueqlJosseu:1crqlpuc43ua1eqleJeof,ucJqlu8rsJg'(1e:nEtg)a1&5

'peuslcrql llq.&auos s[e^\ IIst oql qlt,]t\IUJIJ eJe slrel sql putr plslp eql s3 'DqiuDaflDtg our?seW oJee8epusddt aq1;o slac pru]xord at$ (s awalrozg pue'wryoqdacosfuqJ '3'e) ereuaSraqto eseqt u:wns {-tI r! aH ur pepnlru,{lpuonlpEll ueaqe^cq tcql ereusSreqlo eql qlt^\ sls[JluosprrB'/s,(elpluJ1laq]]4?poqaqpunoJl?ql ol rsFurs sr uoucn s srqJ JclloJpuo.{eq Sulpuelxo 'prolelnbe pur snoluaruclrJpuc ]oe,r ,{ro^ slcl ]oqlue eql puu Suolqo-^\orJcust llllloJ eql pus 'sllt.r 'sJlD.^\ lcrus s11ocpurBruur eql pue leuurql qlr./i\Suolqo slle3 lclsp eql {tlq1 qlr.u p:sle1tnbaI 'els^o-pllolq s11ecpurrxo.rd eq1 ro elcp:oc ,{1e1ncrst aSupueddueq1 xeldLuo: n ll)rur.t$D7 a\l uI

'seururJJr Surssessuuaq,,n anp^ Jo €Js luql sleljelcqc JcqllnJ leJJosllet luanlusuoJ 3ql lnq polou 'slleJ 'edcqs ueeq.{puenbarj srq e6?pueddureqlue oql Jo odBqseql tuenlnsuooslr Jo Surue{Jlql pue 'lueruaEus:r?eql prm'oSepueddtreqlue eqlSo ernlxel pw eduqsoql :slnl eqlJo SuItlJucJqpue e:nlxel 'qtSual '(I aql :Jullot oql lo qtSuel pw eduqseql s.Itrroqlult eql ul ecu?uJluSlsJO am8ld) soLltuv

'seqol oql olu! Surssedlourxq eerql lset oqt Jo spultlls:tlnJsl^ qlt,tr snorqelSlsotx1c pue IeJupull.{3 'snuwoqJozo 'wnlotldacosttt[S -,\^orj?usr elloroJ 3q1dnol8 wnllottsruqo w stl.lrlPLI etll ur pup. 'D\tuoaJrDtguI 'peddn-putrl8put alcueslq eft eqnl elloloJ uo sJIDq6ql.11q,,,r 's:qo1 aqlSo x:du:ql ol rlscal spu[rls J?lnrse,,reql'lqSmns410o.lql eqlJo puBlcJelcllnbe + eJBseqol eqlJo sIullapldeJauul oql 'qull Jo slloJ eql elelnueduleJpuB eqnl pf,upull.{J qlI.4^rf,lnSel fl nllojo:) eql xaldwot ollacuauto1 'eJn]clnasu^ eq1 uI acuccqru8rs oueucS reql JoJ pessossceq lsnu lf,ql stelJelt]qJ II? eJe aql 'stu:eprde Jo luelxe eql pu?'lsoJrlt pu" seqol eqlJo eplsul eqlJo .{ltsollrdcdeql rouut cqlJo sllec eql ;o fultrelcrql puc eduqseql'urnluaurnpul aql'saqol puD oqnl eqlJo elnlxJl pull edcqsJqJ, rilr.roJ

'(Z-? seJnFrC)Suolqo- oqs ole (a\t eJeqll'Dplllzg pur lutodua^Dp H JoJ 'suISJsur ldacxe uxel qe ur prelelnba 1aJI] deql xeldwot o11atuatanz7eql uI 5l?8n.uoi ql!,& Suolqo 'su18rcur - oqs eJR Ke\t wnstJqJnndT ptl/.JDlztlo{I ur e11q,t lqSleJ]Sqtll\ eJcnbslsotup elu .{oql alMDalJDtgur'rvaurl eJ?slleceql /,t s lanpqdacosfurq3usnql 8toue8uaa.aaleq puc sdno:8-setceds ues,nlcq eduqs ur dru,r duur 1nq pa1t1e:fyesoyc Sutaq se pastuSo:el ,\\ou eJc leql sepeds ur oduqs orrreseql a^Bq IBreueAur ,4eqlleql u,roqs suq wns,QtlJlpH pur wn.raldqaHu peprlxut ,{lsnot.terd seroedssnoJeurnu ur [lsal eqt Jo :edq relno cql Jo sller eql Jo uolleulurxa stwDptda :Dlsal

'dnor? ,runsfuqJoqlurx eql Jo sotcadsu pxt'DaDtarssDJDtuaoLlJS 'ustaKo 'g ur eprs Jaqlo crll o1 xedu eql ro,to ssssrd I a[tll\\'Dasot DllzJuaJtADTpuE'Il\todua^Dp wnsfuqcpall 'oplltag ul xed? oluols B ur selculurel 1xelduoc D aJuall DT eql ol peJ-nJeJ ojaq 8xgl asoql ol pJeSoJqllid 'xsdu eql Sutqcuere:o;eq sasusclt dno:8 uryo1aunsttt[c4ap eq ur pue "sse'I sfl{rr1,qr o1da7'wnpqdatosttqJ"lpuoll'?tztro1]1r ur ollq.{\ :eplsreqlo eql ol xad?eql re^o sessedyl oqlunal ntgr uI pessaqlJo xedr e1ue1sgoqs eql ur sal?uluJel ,{luelslsuot pue4s JtrlnJSBA 'uoslud 'g 'pesluEocel eql (qz66l 1ne4unstqtotnaT) dnota suoctqp wnraidlpH a\), ur snrlJ ,(pua:e: st eeurqluer8uVutrller1snvaqlJo eJeuo8 Jo suollcos,(ueur ut luulsuoceq ol sJeedd?JelscJuqs 'peas '(urnlnctde srqJ eql punon enurluoc feur ro ellrels uoqs ? Jo truoJ eql ur sI xads aql eteqa sasecesoql ur,{ln1nc[red) paeseq;o xed?oqlte oteuluJel Feu Jo'Jedordpoes eql;o xsde aql eroJaq '(1-7 alrurur:e1,teruleq1put4s rulncse,r.{ru11lose seq s.{n.^41[ l]lsal cq1 sunBtg) puDrtsnlnJSDr': D$aJ

'uoslr^|c xttdtuD D arlta/$t1 lntd Nuyrsiavol.8, No.3 (1992)

Pappzs (Figures2-7). In the Lawrencellacomplex the bristlesare linear, firm, prominently barbellate,without spreading basal cilia, they arepersistent or shedentire. InBellida, H.lindleyi utd 'Helichrysum' H. davenportiithepxppus lengthens during anthesis. In othermembersof thebris]des arefiliform or, if broadenedat base, eventudly break just abovethe base to leaveapersistent corona on theachene,

Acheneaper (Figtes 2-7).The apex ofthe achene forms a shallowrim aroundthe base oftheprppus In Helichrysumcassinianum utd in H. .zlerrii, in theother species of theZawr encella complex the acheneapex narrows and passes smoothly into thepappus base.

Achenialhairs (Figures2-7). Normal duplexhairs, when present, are thick-walled;two-celled papillosehairs are absent. The pericarpis velutinouswith minuteacicular unicellular papillae in ,It. lindleyi,nd H. dave portii, in tt leastsome of theachenes - \n H. davenportiiacicular multicellularhairs are sometimes also present. The achenesofH. cassinianum,H. ayersii,and the 'Xant hoc hrysum' group lack these papillae or h{rirsbut berr scatteredmuliicellular gland{ipped hairs in additionto normalduplex hairs.

Vasculatureofthe acheneand seed (Figures 2-7). Theposifion ofthe twovascular shands in the pericarpin relationto thesolititry vascular strund in theteslr andto theposition of thecotyledons of theembryo was suggested by Shortel a/. (1989)to be of gene c signillcance.

(A-D) Figure L Anther apicula aDd stylc apices (E-rl of bwrencello dayenportii (A &E), Bettidd eraninea @ & tl, Ozothonvws IepidophJll4r (C & c), and Crryroctphalun senicatwn (D &tI).

L '(X) dnor6 '(Q) ,uns{tlror.ltuDx,aqt pue'(V) nsnto'g'(J) unuDlulssoTH lllodua^Dp H'(I) t[alpall unstrllJlpH '(g,)Dplllzg ul unlnltdecpue euaqce eql Jo sJeIJDJBqJautos;o uosrnduoJ I slqcJ

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'pol?Ao3xe 'wnuDtutssD)'H lou sl es8qal{l dnoJS,Ld,s&qcoqlaox zql nl pue'lrJatD H ur 'H'Dpluag elrqA\estrq .rolloq lueururorde seq eua\xe ew lyrodua^Dp H puz'tblpull |ui.'Je^ent.ot\ 'wntpododto3 :xelduiroxD aJuat,$r7 oql Jo setJeds e ullusJglu8Fullo lpuls sl urnpododlut eq1

'psceld ,(lprpeur ere ,{eql dnor8 ,wnsC lJoqtuDXatltu1pue'gsnfu p'wnaDlulssDr H ';l'alrq/r\\u.todua^Dp'Hptm'tKalputlwnsAqctlap 'npylag ut (stopelhoc ol pm8er qlt.\\) pereld ,t1prele1eru asal pue fuecued aql Jo spuB{s oql

998 xalduo. ollarua/,tttl'nosly'A'O In?d 366 NuyrsiaVol. 8, No. 3 (1992)

Chemicaldata

Few Australianspecies of llelichrysamhave bcen chemically investigated and the taxonomic significanceofthe results ofthosethat haveis notvery informative. However, in apaper by Jakupovic el4l. (1989)theresults ofan analysisof I'lelichrysumdavenpo ii, H. lindleyi,andBellida graminea werepresentedalong with analysesofH. ambiguum'fucz-(= ),H. bilo bum $t aket- (= ozothamnus),and. H. leucopsideumDC.From this investigation some tentative conclusions were drawn.

rn Ilelichrysumdaveflporliiwas found isokaurenic acid anda thiophenederivative numbered 37. rn H.lindleyi thethiophenesnumbered 37 and38 andsome triterpenes were lou nd, while in B elLida gramineawercfornd thiopheneacetylenes and the desoxy derivative ofisokaurenic acid numbered 37aand the thiophene numbered 38. Thesecompounds were not foundin fte threeother sDecies of Australian'H eI i c hry sum'.

'' The chemicalanalyses therefore suggests that lhe three speciesare more closelv 'Heliclrrysrm' relatedto eachother than they are to theremaining speciesihis lendssupportrothe 'Lawrencel/a' opinion, basedon morphologicalconsidentions, th'J't the complex is a natural assemblaseof taxa.

Mycorrhizaldata

It hasbeen demonstrated by warcup (1990)that Australian species in thc tribe Inuleaecan be almostequally divided into those that form only vesicular-arbuscularmycorrhiza (VAM species)and thosethat form both ectomycorrhizaand vesicular-arbuscularmycorrhiza (Ecto species).The 'Helichrysum' Australiantaxa placed in can alsobe separatedinto Ecto and VAM species.The presenccofboth vAM andEctospecies in thesame is unusualsince normally genera have been foundto containspecies that are either all ectomyconhizalor all non-ectomycorrhiz:rl. 'Helichrysum' when the speciesare segregated into thosegenerarecognisedbyAnderberg (1991) andby myself the situationchanges, for thcn eachof the segregategenera contains, as far as is known, only ectomycorrhizalor only non-cctomycorrhizalspecics.

Warcup(1990) examined the mycoffhiza of thefollowing species in thela,t r encellacomplex: Helichrysumayersii, H. cassinianum,H.lindleyi, H. davenportii,H. subulifolium,and Bellida graminea,hefotnd themall to benon-ectomycorrhizal. This is alsothe situ ation in Rhodqnthes.l. (Wilson 1992a)and Helichrysumsrb5. Ozothomnus(Ozothamnus R.Br.); it contrastswirh the situation in the Helichrysum bracteatum grotp (Bracteantha),the H. opiculqtum Erolp (Chrysocephalum),and in theH. leucopsirleumgtoup all of whichare ectomycorrhizal.

Genericcircumscription

on the basisof the charactersthrt are foundin the achene(see above) a closerelationship is suggesfedbetween H. cassinianum,II. aycrsii,and the species of lhc,Xanthochrysrm'group on the onehand, and bctween Bellida,11. lindleyi tnd,H. d.aveflporrl/on theother. This correlates with the natureof thepappus which in H. lindleyi,H. davery)ortii,xnd Bel/rda elongates dudng anthesis to extendbeyond the corolla while in theother genera it remainsrnore or lessequal to the corolla. qll^r elqrdrls 'olsn.?[3passoJdu]oJ euorlJv xodtr ol Surpuolxe'lnols puells JelnJse^ 'oluuunoe ol elnc? 'prollep xadu e1.(15 re11ocEurpoacxo pue aurS ,(1eura4xaslltrl :Suolqo-,1aoueuJelloc 'polle,,t,t-urql :lI€urs slle3 ltrur8reur pue Suolqo sJlocIrlsrp puc Jcrpeur'peJ1l;,/rr-1crq1puep:e1rlnba 'aluproJ-,roJJf,u 'p:alclnba 'xede + slla3 purxord eScpueddc:sraqluy 1 sru:ep1de:euur ;o s11er ol Eulpuetxasputrrls rclnJse^ 'eprslno osold ,{lesrf,ds'urqtr,r osollrdDd'S seqol :snorqBlS'pedeqs lau8q qurrl :asolrd'ltrJupurl,(cflUoqs oqnt :elloJoJ olclu Jouul'pnxoslq:e1no'crqdlou.rourlcu 'snoJeltlnu s1s:olg 'sno:qe18 'l?Jueqdsrulal{ alcrldeceg dq snora3rld c sc SurnuuuoJ e^Jou pnuec'xede o16urqJBorauoarols u,4\orq,roJJlutr qlrm ourlefq'al8lros-gslJeJq :lultrpeJlot|'elcurqJnl 'lBurtrrlet ',4rslllos 'ruro;1lg'IDsoq 'sruols alJnlo^ul r1n1rdr3 se^De.I Jo oseq lD rtlr^ etrq^\ JoJ ldocxo snoJqclS'osrq tuo{ SursuBsedtrJs sseucel pJo^es qlr,r q8rq urc a1 ol qjeq lJeJeJunuuv

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'dnoJ? '11'rysnto 'H 'H ,ransfurlJoqtrr;6 sq13oserceds pse'wnuDtulsslr (Zpue'lurodaa^Dp pue ttalpu! wnstrqJrtaH( 1;o lsrsuoterauoS o,nl osoq; snddtdpue oueqJ? eql ur peNosqo srolaerDqr uo pesequreue8o.rrlolurtrxslaqlSurpr^puelrpr/r8rJouorsnt3xaeqlqllld'ftalputt'Holsrn]fla uoqtJeqtoqcue ot palBlor,{lesolJerouI am tutoduampp puenptlla g pql stsaaSnsr1sa1 s1r 3o s11ac 'rerre,,no11'snueE puuapldaeq1 go ,t8o1oqfuoureq1 crd.,{1ououB sa lr esruSoJe:o1 Surnurluoc uru 1 puelruusry.{lpcrSoloqdroureqolsmoddeopttlag'peyasasdeJtrxnar$JO'(dnofi,ansttq]oqtuDx, aqtur esoqlruo{ uede),{1rs:e,rrp pcrSoloqfuour rraql pue parlo,rur sercads,ro; eq13o esnrcsq dpred 'ssessu ol lFrgJrp slxeldwot ollacuatur1aqlu! posruSof,eroq plnoqs lcql c:euoS3orequrnu eq1

xatdu'lo2 o axual,$t l'uostttd D In€d NuytsiaVol. 8, No.3 (1992)

Fi9]uje2. Bellida grdnntel. A acbcDcwith pappusand corolla. B 1.S. achcne(senid'asBnatic). C achcnialsinsle celled prp;llaeanddDplexhAirs.D- portiorofpappus brisiles. E-seed(i.e.resra). F- epidennalcells ofrcsra. (v.s. - vascularsirand, p - pericarp,t - tesra,e - enrbryo.) From rnrSr Pdft 793/86. excenfic hollow bxse,moderately to sparselyhirsute with duplexhairs and densely and minutely velutinouswith acute single-celled papillae (in smooth achenes thesc absent), mot myxogenic, at apex expandedinto two cup-shapedextensions that bearthe pappus;pericarp deeply and transversely rugose(or occasionallysmooth), thick and cfftilaginous, a vascularstrand in eachmargin ofachcne. Seedcompressedturbinate, free from pcdcarp;testa thin, vascular strandextending to apexin medial positionand at right-anglesto vascularstrands ofpericarp. Pappus at anthesisabouthalfas long as thecorolla, in fruit almosttwice as long; bristles filiform, prominently dentate, rcddish towards apex, unitedinto groupsthat are firmly fixed to eachof the extensionsof the achene,persistcnt.

A monotyplcgenus.

Bellidagraminea Ewart, Proc. Roy. Soc. Victoria 19:35(1907).71pe: Jibberding,Wcstern Australia, 1905,M. Kochs.n. (iso: PERTH). (Figures 1 & 2)

Distribution. Foundonly in southernWestern Australia.

Moore(1917) indicated a relationshipof Bellidato H elichrysumdavenportii and treated them as beingcongeneric, while Anderberg(1989) has suggested that Bellido mxy\ot evenbe a memberof theGnaphalieae. I considerthat the similarityof floral andfruit morphologybetweet Bellida and, Lawrencellaindicate that its positionhere is correct. The similrrity is strongestto Lowrencella davenportiiwhich, if a hue reflectionof phylogeny,indicates that lanrencella as heatedhere is paraphyletic. '9769aBnag -a'elset - 'prrclls - -{ gytuo:g to,ftqu: I'dieruad d l?lnrs?^ s^) PlsalJo sllar lEuusptdo - 'ouaqce letsel ar)paas A allsuq soddt?dJoxedv C (auaqruJoxed?rEau ttlorJlqBF pu?arlu3c Jo orulns tUorJPltrdPdrJaD - - 't auaqcarrto.U saruoqrpl C (clletus8plpttuas) amqre S I B snddedqlr^\ amqre V ,rlodudtt)P Dllatuar^ry 3in3'C

'r;lcr1snVol JlurJpuJserJeds o^iI

esrq sprr^\ol sdnor8 ur polrun 'alquep ,{llueuruord 'el?uunte -luoul'unJ sellsuq lrnJJ ur SurusqtEual]nq srsoqluel3 elloroJ uBql rouol.ls'{uld lo sselJnolot 'luelsrsrod snddu4 'd4 pouolleu eql ur SurlsuruJel 'suopel.{loJ ol etuoroJer qr,lt peceld,{1p.Ie1t1 pueJls JelnJstr^'snoutrlquou l]lsel :djDruod ruo4 se{'drl aluols pauall?u Ioqs e ql$ ruroJrsnJ -,,tro:rcupaas:snEq xeldnp33ps q11.'tr pgdstq,(losnds.Io seruoqal! pollscl]lntu etf,utunsEpouallBUqlL\\ 'eso8n::o snorqecsro oellded pelleJ-I euJoJrJ?elnurur qlr,r snouDnla^Jo snoJqulS qlootus'1crql prrupreq clrerued'd.rucuadaql3o uorsuelxe.$olloq c SutuloJ palc^t3xe esuq:Jlqdlotu.{todseuaqJV 'JulloJ 'dg o1 Supuslxo'pu?-qs JBInJsc^'ploltep xodc al,ils SulpeaJxasnoluoulltJll slcl roqlue :Euolqo-,rouuurclloJ :llcursslloJ lcurBl3u'pelp.r-urql'Suolqo sllat plsrp'pelp,t-1ctq1'p:elcFnbe sllacpurxo:d:a1u,,ro-pt;orqoAepueddcreqluy'dqolSurpuolxospuers:t;lnJse,t'urqlq.,tesolltdcd'9 saqol :snt.r lqAI?Jlsqtr^\ttrorqlJo slle3 :snolnprdslqJl]lnpuelS'3^oq? atulnucdues,{l^\ouDu',\\oleq relnqnl 'clqfuotuoullJu elloJo] 'e[?u ]euul'lenxesrq retno'snore|Ilnu slejolC snorq?lS'qloouls '1urd'cudlllo 'pedsqs uorqsnr elcrldeceg Durluel :euoerels ltg 8uo1qoue ql!,a\eullefq'Suolqo 'Jeln8urxrl-^roJJcu 'snoecwDqJ 'Suolqo-.,Y|oljuu ,{\cl3 :$Jelq lsourJeuur:ulql ouroe.rels:olclllc ol JelnSueul-,rolJcuslJcJqelBrpeurolur pueJolnO 'olf,r.las-9-tslJl]Iq:[EJueqdst uJq olJnlo^uI luclpl]J 'pseq-qns 'sedaosssel1uel Jo seqJu?Jq ol pururol 11n1rde3'peue11eg:o elorol 'eteuJollu ro elsoddo ro eurlntr3salea'I e.{oq?peqrucJq'lf,ere'e18urs ualg 'asoltd:o snolrueqndre1npue13'sqreqpnuuy

dalpurl '(Lg}l)tlgtt' ''q1uag DasotDUaraatMDI :adq 1t4sny g ( Aa1pu!1)ollatuautDT lxes wnstq)!pH

'(6t8I)tZ'loJ U uc^\S'3:r1q:t:15'^rlpur-l ?llaru)r,raB'I

xa]('ltlloaqtaru"l^\t t'vostt/rA O Ined Nuytsia Vol. 8, No. 3 (1992)

Lawrencelladavenportii (F.Muell.)Paul G. Wilson,comb. nov. (Figures1 & 3)

Helichrysumdavenporlii F.Muell., Fmgm. 3:32(1862). - Helichrysum lawrencella var. davenportii (F.MueU.)Benth., Fl. Austral.3:616( 1867). - I'lelichrysumrosdrm (Lindley) Dru ce var-davenportii (F.MueU.)Domin, Vestn. Kral. CeskeSpolecn. Nauk. Tr. M Lt.-Pir.2:120(1923).Type citation: 'In Australiacentrali ad flumcn Neales.J.Macd. Stuart.' Lectot!pe (here chosen): Neales River (MEL 579934',isolecto: MEL s.n.).

Bellida major S.Moore,J. Bot.55:100(1917).Type citation:'Western Australia, Mulline; J.E.C.Maryon,1916.' Type n.v.

Distribution. Foundin WestemAustralia, Nofihern Territory, and South Australia.

Lawrencellarosea Llndl(y, loc.cit. (Figures1 & 4)

Helichrysumlawrelctll.lBenth.,Fl. Austral.3:616(1867) llom.illeg. - Il elic hry sum ro s eum (Lindley) Druce,Bot. Soc. Exch. Club Brit. Isles 4:626(1917); Donin, Vestn. Kral. Ceske Spolecn. Nauk. Tr. Mat.-Prir.2:120(1923) comb.illeg. aoz (Hook.)Baillon (1886).- tlelichrysumlindleyj H. Eichler, Taxon12:295(1963). Tlpe: VasseRiver,on the South WcstcoasrofNew Hollan d,1839, Mrs Molloy (holo:CGE photo seen).

Distribution. SouthernWestem Australia.

FisuJe4. IawrenceUa ln /.];. A - acbenewilh p,tppus.B - L.S. acbene.C - T.S. achene(semidirgmnutic). D - achetriatduptex hair. E - Apex of pappusbi slle. F - seed(i.e. testa). C - epidemralcell! of resra. (v.s. - vascutar strdrd, p pericar?,t - testa, e - enbryo.) From P .G. Wilsol 12353. '!p!^ 'f'ld 'lJ 'ulllnuuruntrAl uouadq ',(i'lorag 'qH) aBroeg lrsru apun 'rd urlsny'JlsrpurqsH, :uo4c1rcod.{; '(906I)619:9t'lsfS'qrquf'tog's1erq ra8roaS slla1opo4

'(-tdl,\l ''podxa :otoq)€/8I s.assog:rdf,J ',essoC '(t/8I 'ruBrrg''11onyq 'g :o?3tosrluoru trrunr^ uL :uoqlllrJodfl )19I:g ttsnto wnstuqcttag (sr {{r0tg't I sorn8rC)'(9I6I)0t8:6Euremsnv'StoS ,{oU'rordT,'su0{ ( onl^t'C)rsra,{e eluaoqrs

dnorSqnsBrueoqJS

'cllsJ,tsnVol Jrrrepuasermds eArC

'(seueqJe aluolsaql uo luolsrsDd)snonprJop ro lualsrsJad'asaqle Euu uoqs B ul pallunlnq ee{'.rollef Jo ssolnoloc'o1e1uep 'xadu spm.,rlo1 ruroJrIrJpu?e$rqteraeurl-^{oruu sallsuqsnddsd 'uoursodprpeu eql uI pees SuqrJIf,ue,{lelelduoJJo 'snooJeuoJ 'fuerued €/Zpuels JDtnJsr^ ,{[urql?lsel ot lus-rsqpeIo uro4 eoq poos:pele^ecxe lou 'rulnuur es?q:lu?cgruEls ur urnrpodofuuJ:uoltrsod le peu urpw sactrJIssrop pw prlue^JoeIuoJ ur spur4srqncse,r ',{:edBd ro 'snoaculsnrJpur urrll'snour8EluluJ fuDJued:snddcdgo escq Surpuno::ns pur posrersou-rDolllos urSreur l?Jrde !uoserd osl? srl?q poddp-pue18 goqs's:ruq pr6ut?q,retuos qlr,r 'plo^oqo-possoJduoJ elnsJrqdlosuep ol ,(lesJeds ol oleJoleueqcy dq ol Surpuetxepuc lnols pue:1s ntnxs?^ 'esoJlrded'elnJc 'prollep xedu eIFl5 Jt]yloceq1 Surpaecxe puc euU ,{leuroIlxo slrrl :Suolqo 'pe1p,,w-urq1 -,roIJtrurullo. :luslelrnbspue lptus qlec purSretu puu3uo1qo s11e3 lclsrp puB lurpetu 'pelp.,t|-{JIr.0 'olDproJ pw Ierelellnba+ slleJleurryord a8Bpueddu:sreqluy dr1o1 Eurpuolxa spue4s 'peqol rBIn3se^'urqlr,lne$lntrIloJ roeso11tdcd saqol:snolrueqnd-re1npuel3 flesruds -E'qu1etcurqrnl 'oleu 'Jrqdroruourlotr -aorr?u qlt,lt pcupuq,(c-,trorruuellolo3 Jouuroql pue J?nxesrqrelno cql 'esolldud 'xe,tuoc '(alrq,r slerolC Ju1npur13:osno:qe18 elrcldeceg ro) ,rotta,{ro 4urd'cpd11euuru.rel 'Euolqo 'ouoeJe$ :ouoerelsEuolqo w qlr,4\snolll]os .,!\cla:luasqtr ro luesa:dslcuq rauul leu uoeI8 J31n6ueu1cqlr,tr fssolS 'alc^o-snouDJs fl3BJq Jalno :slJsrq Jelno oqlJo euooJols eql uo snolruaqndro snorqclSosr^ueqlo 'alerIrJ ,{lloo,,r1 sou-rlletuos 'eleues- g J slJs.rq:lou Jo tuurpcJ 'lr3upurl,{o-,rorlcu ol pc1:eqdsrueqe:cnlo^ul ruroJrqu,&oJro .&8lllosBIntrdBJ 'elerol ro Suolqo'el?uJolla lo alrsoddo sa,rm1'sno1n:aqndJslnputrlS ro'olnsJrq ,{luoqs Jo'rs.^oJ ,{ltoo,ll e Surcnpo:drunlnordu snoluaurulrJ '8uno,{ 'lBql 'sqreq palm3 B lBoq uaq,r srpq oleuosruntJoqs qlr^\ snollolJrq tJeto lonuuy

'Ilent{ d r',ruls,srsourrl '(ES8l)SIt:SZmeuurl' uoSodotatJ'.(ueso\3ent4)ad&opa7 llentri Csrsdorrrdr laH'txesaoSodoratd

f,t tg.'y wn11auat wnrafi !p H '(ZgBD\EZtt 'log 'I 'let| 'l3es :adtJ JsrII prEC ^re) s.Je{ooH t/ owadsotuag u raidlpH

'zxnlwn!p!!l!t '(I "zornlunstuqcoqtuoy unstuqtoqtuoy:ad{1 S8I)66I:I/tZno3sol l setsllurn'leN'dru1':ogllng

'[zl$1S( qJIpn?C)DuDtutssDTS =f 4eots,Dllolllsoddo ,raraqrs :(uesoqcereq) ad(rcpaT '(Zggl)gl.I:g 'ldtsrH 'uollrcq (zleets) otuaoqrs't oswnsf-tqrlpH - (ES8I)SIt:SZeeeuul.I "[onhl'{ (zleelS)DruaoLlrs 'pas uoSodotat4

'(st8I)08t:I 'ssrord'ld"urqo'I ul asetseluaoqJS

x.tdurox D aruahrj'uostrnd O Ined Nuylsiavol.8, No.3 (1992)

Distribation. Foundin southernWestern Australia, Northem Territory, and South Australia.

A speciesreadily distinguished from Podolepi.r, in whichgenus it wasplaced by bothDiels (1905) andDavis (1957), by its triangularstyle apex and its turbinaterugose achenes with couse hairs.

Schoeniacassiniana (Gaudich.) Steetz in Lehm.,Pl.Preiss. 1:48I(I845). - Helichrysumcassinianum Gaudich.in Freyc.,Voy. Uranie 466 t.87(1830). - cassimanarz (Gaudich.) DC., Prod. 6:216(1838).- Pteropog,oncassinianus (Gaudich.) F. Muell.,Linnaea 25:415(1853). Lectotype (herc chosen):Baie des Chiens-Marins,C.Gaudtchaud (P photo seen,isolecto G-DC photo seen). (Figures1 & 6)

Schoeniaoppositfolra Steetz in Lehm.,Pl. Preiss.1:480(1845). - P teropogon oppositlfulius (Steetz) 'Speciminaintegraleg. F.Muell.,Linnaea25:415( 1853). Type citation: cl.Roo in Australasiaaustrali- occidentaliinter flumen Swan-river et sinum regis Georgii III. (V.s.in herb.aulico Vindobonnensil)', nonvidi.

Distribution. Foundin WestemAustralia. Northeru Territorv. and South Austmlil.

Figure 5. Sc/roenraayarsii. A - achenewilh pappus. B - L.S. - acbene.C - T.S. acheoeGemidiagr.rnatic). D - achenialduplex hair. E - apex ofpappus brisile. F seed(i.c.lesta). C - epidemralcells oftesta. (v.s. - vascDlarstmDd,p - pericarp,i- resla, e - embryo.) Fro l/. Ds,urz 6588.

,L 'f1gg4 :osr)pp;aqa1g ,174'gg6t 'tdos 'oupueN 'SaU 'cro 'lsrlsnv 'ld'unug 'reupruC :rd,tJ wou (IE6l)EgI J tunvulqrnlopnasd 'ros '[.Eqr{ wnstulcllaH-'(106I ,iq/'{)€Z: I'oN isIII lcN'trlsnv'/r\ l wuDutqntwnstuq4lag

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1\ l. \

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xad\uo. D aJaar oT'uos[/\A'D FPd 374 Nuytsiavol. 8, No. 3 (t992)

(

Fienrc1 . Schoeniartlifolia slbsp. lubulilolia, A - acbenewilh pappus.B - T.S. achene(semidiagramatic). C - achenialduptex hair. D - apex ofpappus bristle. E- seed(i.e.testa). F- epidermalcells of testa. (v.s. - vascularstrand, p - pericarp,r - tesra, e - embryo.) FromA. Moriro4 30.ix.1904.

Key to subspecies

1. Involucreturbinate to cylindrical;ray 3-6 mm long 2. Plantsingle-stemmed; involucre turbinate...... -...... snbsp.filifolia 2. Plantmulti-stemmed; involucre cylindrical ...... s\bsp.arenicola 1. Involucrehemispherical; ray 7-10 mm long...... stbsp.subulifulia subsp.filifolium

Helipterumtenellum A. Gray,l.c.

Helichrysum turbinatumW.FiAg.,l.c. - Helichrysumpseudoturbinatum C. Gardner,l.c.,nom. illeg.

Distributionand habitdt. inlrnd south-westWestem Australiafrom Mullewa to Kalsoorlie and Lake Barker in saline areas. 'roqlo qJca olur ep?.t8sarcadsqns o,,Y\leql leql lsoSSnssuorlJeltoc sruos lnq'srllq esuapejour pw Je8uol qll,t\ souoqJeJoBJel pue clnlrdcc:a8re1qcnu; 1l:l.t*r1m o11ol11{'dsqnsuro{ sre;Jrpscrcodsqns stq11o lutrru,'r 1tctd,{1aq1

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'lcls 'uosll7t6'9 ou l3'quoJ 1ne4( 11ontr'^\trC) srtoJrlnqns dsqns

'(3uol ururt'0'Jsrr?q qlr,r pur pro^oqolou) srnq xeldnp(3uo1 turu Z'0 f,) oqsqlr,r seu:qf,€ '(IuJuaqdsItuaq olerel3ur^eq ur pu? ]o olsulqrnllou) orJnlo^urIuJUpull^J u 8ul^?rlur'(saqcucrq roltrrupro^os qlr.^\tou) pauruels-e18ursSureq ur ,rolly''dsqns uo{ sJoJJrpsoraodsqns srll

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'uolplcjec se qlnosJeJ sepunoJ eruo se,rlr teql sele3rpur'l33roosr ltup.(trp]ol aqlJr'.{eguorduuLlJ uro{ uortcalloJoqJ 'cllD.llsnVulelse,t\Jo ueJl] uo^leuJl|3 Jqt urslIqpucs ol peurJUoJ ,{ll uJp r^A Jo qoqpaDuounqu$tO

'(HIdAd) 'uo^ruureJ uastNJ.'W6I reqot.O/'ldoS :(9rS0II '688I ',(eg 1fl,N) tsat.to!stw uordu?rlJ :VITVUISOV NdAJSa,t uaassuoura n plnotppv

'3uo1 urur7 g c'1c1tlsrreq xoldnp :etarelsaueqJvuuSZxS J euru?l ele^o,rollo,( qll,r e slJejqlsourJeuur :q8rq uru 1'c'pcupur1.(c 'squ-r,{roc aronlolul uadotro a3uaJsologul q8rq urcgg ol'a^oql EurqJuoJqqraq lcore lonuuv

'(sNvJ'Ius :osr:HJugd '6S6I '1dag 'uo,rnuru3 'rlllt;:tsnv :oloq) 2099 aSptqng J N y Jo A'S sopurg urelsai&;sn d,(J 'sr3uo1 urur7'g c srssercsnqrcrldnp 'urur sqrd'serolunruoqJv g'Zx g', srelnlsue^o srururel truluur eolf,clq :unlp uur 1'c urnecerpurt,{c 'ttsoqtrl,{roJ unJJnlo^ul exel crluof,seJouq clle urf,0t pe Esourcrurscq Drdnsclcojo eqjoH

'a.ou 'uosy16 'dsqns dsqns 91nr4 elorlua.re

'epeu ueoqe^eq suollaelloJ .reJ 0 ,{luoecurs cxul cgrcadsu:;ur 're^e,rorl'serradsqns raqunJoqucsop ol alqruosearluase.td p lou srlr o,tr1:aq1o oql Jo uolleSoJ8as eql qlr,r ua^a, Eoloqd:oruur alqeue,rluq.reuos surcurer zl,olly' dsqnsFuouoxul e^oqe eql uI

',JWl llDSc Jo sJJoqsJql SuolDs{JoJ Jo s:Jr^JJ ur, pelielloJ se,r ed,{l sll 'seuaqJ? snolnprdsrq dlesrcds ,{ro^ 'rplnSu?Jprnb '^\ouuu pue'slceJq 'uorlJoJloJ IeJJnlolurJouur oql ol oeurtu[lelq^r (3uo1uurg c) l.roqs8ur.r.uq ur sre;yp,{lqrssod leql ruo:;u,trou1,t1uo pue eJn1rrurtur srqlrq,,['waourq-mt p 1oed,41eq1,{qpalueserdu luurrvl eqls?ereq,r 3uo1uru.r 9-7 enutuel FeJ,ro edpue seueqJu pro^oqo scq urrolly''dsqns Jo luurn,rpcrd,Q aq1

xe4lljor D "ruatuq'uosrr?l CrnPd 316 NuytsiaVol.8, No.3 (1992)

Schoeniamacivorii (F. Muell.) Paul G. Wilson,comb nov.

Helichrysummacivdrii F. Muell., S. Sci.Rec.3:99(1883). Iype; GascoyneRiver,1882,J. Forrest (iso: MEL two sheets,PERTH).

Disffibutionand hqDitat.Gascoyne River areaon sandor loam.

Schoeniaramosissima (F. Muell.)Paul G. Wilson,comb. nov.

PteropogonramosissrzasF. Muell., Linnaea25:412( 1853) . - Helichrysumsemrlertile F.Muell., Rep. Pl. BabbageExped.14(1859) - Hetichrysumramosissimum (F.Muell.) Druce, Bot. Exch. Club Brit. 'In Isles4:626(1917) nom. illeg. non Helichrysum ramosissimum Hook.(1848). Type citation: planitiebusarenoso-argillaceis inter montes Flinders-range etsinum Spencers-gulph etiam non procol a rupe Cudnaka.'Lectotype (here chosen): In planitiebusvalibusque sterilibus propc Cudnaka et Wrtlendunga,Oct. [18]51, Dr Mlueller] (MEL 604822).Poralectolypes: Cldnaka, Nov. Holl. austr. inter.,F. Mueller(MEL); Intcr montisFlinder's & SinumSpenceri, Oct. lI8l5l, F.Mueller(MEL).

Helichrysum semifertile vx. xonthoglossumF. Mucll., Rep. Pl. BabbageExped. 14(1859). Type 'Wonnomulla, citation: Elizabeth Creek'. Zype: Smith's Walcrhole, aaoa. (holo: MEL).

Distibution. Found in Northem Territory, Queensland,New South Walcs and South Australia.

This specieshxs a variant with white and a vadant wilh yellow bract ray laminae. All of the syntypes of Pteropogon ramosissimushave white ]rminae and evidently for this rcason Mueller 'Babbage' describedas a vrricly a yellow-rayed collected on the expedition.

Thepresumedholofypcof H. semifertiLev ar.mnt hoBlo s sum is a small specimenmounted in a blue packet that is labell cd.'Pteropogonramosissimus JJ flavisima, Smith's Waterhole'; it beilrs no other information. Smith's watcrholc is one of the localities visited by B. H. Babbageon his expedition 'Report' in 1858. The locality is rlso rcfclTcd to in lhe by the Aboriginal nameWina-Wirralu which is statedto be near Elizabeth Crcck, the place name mentionedby Mueller in his protologue to vff. xonlhoglossum.Since the specimenagrees with Mueller'sdescription, and is theonly collectionin herbMEL of thaf speciesfrom the'Babbage'expedilion, I rm confidentthat it is the type.

Acknowledgements

I am gmteful to Ame Anderbcrg,Laurie Haegi and Jim ArmstroDg,who mademany consfucdve comments on an early manuscript, and to Ma-rgaretMenadue who provided the illustrations. The figures of some of the transverscsections of acheneswere basedon slides kindly provided by Phil Sho. '9ZS-Ll9t8Z 'to ,&lsruaqror,{qd .sapedsunrat(luaH ,rxtlerlsnv tuorJ stoantllsuorraqto ps! saootrElauadl4lnbsas (686I) p 3'olavz .9tt-6tt:8 ?r.t{nN kns[r\rorn /1 Z wd (e?oq?qdeuC:asarera!sv)utuatdlpH ]clpapqtul fnuaDm salredsuellerDnv Jo uollPrlJlssElsaql (qZ66I) O d'uosl/A '8[]-6Lt:8 plsl,(nN I !?d :(a"all"qd€u9 :3"atsersv) 'uoslllA aeue3 palylar puy wru4dtaH u papnpu! Iluaunr serreds uPllPrlsnv eql Jo uorlsrgrss?I3aql (ez66l) C d 'Iot-9/:r els nN (eerllEqdDug :a€lnq iae.ieJa]sv) DwlzdsololH souaS3ql Jo trotsr^arV (686I) O d'uosllA\ 'dn 'Z8I-6LI:Z eua anhl (aeecarersV)epatnut u?rlersnvJo sronPlross?l?z!quoc{u eqJ (066I) H f "/'tt '6r-a9:z euellenhl '(es!lsoduo3) luoslrr' "S amFul eqrto sraquaur uellBrtsnvJo ,(uoruu! lriuJ aql uo saloN (6861) f'uo|?N ? g.)l d'rroqs (eunoqlaw'raluudluauua^oD),,sluEHuelFrlsnvJosDsu3lrnEluals,{Spuores,,'(688I) C'rall3nhl 'ssau ( uopuo'I crueperv) ., aEllsodtuo:)Jo ,0lsluraql pue ,{Solor8aql, (spe) .I -aeolnut 'sulo1' 'rattnurxral rerunJ g ?oluoqreH gI'poo^\ftH'HA,/ ^\e!^arctreuats,{s (Z16I) Hlatsseod?p d H l '$g-Eng8z ' ' x{sfrr|aqro qd sabads perEpJpu"' unsKrqr4aH oe!PrlsnV urorJslu.nlgsuor roqp put s.milq!C (686D lrra l'r'^odnryi 'z[z-Igl:Zg r'ros 'umt - 'I 'y'l 'pr"llttH ]og f 2?mpqdeurJ eqlsodruo:l q srdacuocaueueB auos (tg6l) B t}|n8 a O ( aprulapv 'relugd 't 'ralltvluttrlrqnaH '1 'r8aeH aurua^oo) upA ..'e!l?rsnv qlnos Jo erotJ,, Gpe) u"{aoJ U H ?9doss.t d I .4 (986I) (?prslapv'reluud rueuru3^o9) Z upg,, ?lterlsnv qlnosJo ErolC,, U96I) ( apr?lapv'raluud lueuura^oo),,'!rl"llsnv qrnos Jo erotJ,,'(626I) 'Z'8 €Z8r6fsrrerlsnv qlnos ros ,tou '.ord 'ltetg ? suerJ ,{uDro8(l} e llertsnv {nosJo suolSal ulalse^\-quou aql olq uolllpadxeu? oo setouc'Jtl[rlr S 1SI 6I) ,{ I ' ',,slsuanPlNnv 'tuE$uag ( uopuo'I o) T, 3^aaU lla^o', € 1o^ "rou.. 098I) o '96I-I:'0I E petog ?redo (emrglatsv) aueneqdeuo5qr4 aql Jo ,{ua8ol,{qdpue truroooY"J (166I) V V'Siaqrapuv '96ZZ- LnZi Lg'!og'l verpEueC (asotsarsv) aertnq aquraqlJo uoltsrlJlss?lr3-rpue.{ua8ol,{qd (686D V V'&a+"puv

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