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(roo?)t69-tt9:(€)€r,rr[,Ar 572 Nuttsia Yol. 13, No, 3 (2001)

Thomasiainvolucrata was described correctly as six-merous by Turczaninov(1852), however, the usuallyhexamerous feature of Lysiosepalumwas notconsistently recognized in followingtreatments. Mueller's(1859) generic description stated that L barryanumwas five-merous,rarely having six stamensand ,and Bentham (1863) elroneously described L rugosumas five-merous. Type specimensandothercollections ofthese original species ofly siosepalumarc,however,all six-merous. Confusionwas still apparentwhen Moore (1921) described a third new speciesThomasir hexandra Moore,the epithet referring to thesix-merous flowers. When he later examined Lysiosepalumheldat BM andAustralian herbaria (unnamed), he foundthem to behexamerous and subsequently transferred the taxon to trysiosepalum (Moore in Sargent1923). This species,although currently not recognized (Hnatiuk1990), having previously been reduced to synonymyof L. involunotum(Blackall & Grieve 1956),is reinstatedhere as a distinctspecies. Of thetwo additionalspecies described in thisrevision, oneis 6-merousand the other is 4- or 5-merous.

"Flora As therehas beenno comprehensiveteatment of Lysiosepalumsince Australiensis" (Bentham1863), this paperpresents aformal revisionofthe as part ofarevision ofLasiopetaleae "Flora in preparationfor of Australia"Volume 7. In addition,a lack of publishedinformation on anatomy,cytology and phylogeny of thegenus is redressed.

Methods

This revisionis basedprimarily upon morphologicalevidence obtained from examinationof vegetative,and fresh and rehydrated flowering material from collections,including type specimens' from thefollowing national and intemational herbaria; AD, BM, BRI, CANB, CBG,DNA, K, MEL, NSW,P, PERTH,W. All definedtaxa have been examined in thefield.

Seed-coatmorphology was examined on seeds from the authors' fieldcollections, orthose ofKings ParkBotanic Garden (KPBG). Examination ofpollen exotesta ofl,lsio sepaluminvolucratun(KPBG 196/86,identity verified from seedlings)was facilitatedby an EnvironmentalScanning Electron Microscope(Danilatsos 1993). Seedlingswere grown of L. hexandrum(C.F. Wilkins 737), L. involucratum(KPBG 196/86) and L rugosum(C.F,WilkinsT32). Seeds were pre-treated by being coveredwith boiling water and soaked ovemight. Seeds were germinated in petridishes on filter paper noistenedwith 17oPrevicur fungicide and those with emergentradicles were transferred to potsof pottingmix in a glasshouse.

Anatomyand cytology.Anatomical comparisonof fresh material was facilitated by fixing in glutaraldehyde,embedding in GMA resin,sectioning by microtomeand staining with ToluidineBlue pH 4.4 (Feder& O'Brien 1968).To observechromosome number, fresh anthers were fixed in 3:1 absoluteethanol: glacial acetic acid for 24 hours,rinsed in 707oethanol and stainedin alcoholic hyrlrochloric acid carmine for two days. Anthers were squashedin 45Vo glac\al acetic acid and chromosomesfrommeiotic pollen mother cells were micrographed using aZeiss Axiophotmicroscope and6 ASA Imagelinkfilm.

Cladisticanalysis. A dataset offive identifiedspecies of I-ysiosepalum, two o\tgrouptaxa and 25 morphologicalcharacters was establishedin MacClade 3.05 (Maddison &Maddison 1992). Thomqsia angustifuliaSteud. and angustifolia (Turcz.) Druce were chosen as outgroup taxa as a cladisticanalysis of all speciesof (unpublished data) shows to be monophyleticand Guichenotia and, Thonasia to be closelyrelated with similarstipules, and anthers dehiscencingby apicalpores. The outgroup species chosen retain the plesiomorphic features ofthese '(0 .97 1'0 t) lq8leueql erorr = I :g-l = O:elncol rad sap,ro go requnp '0 .tZ (0 I 11uorsry o699 < esBqpesnJ Suol = I :uolsnJ% 0S> ?suqpesnJ Uoqs = 0 :srsq elDgqJo esBg '(0 I'0 I ) lueserd= I :luesqe= 0 :srrEq,&E^o enryg eZ '(0'l '0 .ZZ 1)rutu I ueqlerour = Z :uru I upqlssel = I :luesqE= 0 :aqnlpurursls .IZ (0 I'0 1)uItII S'ffi 0 = I :ruurE'0-0 = 0 :qlpl.alueu?lld '0 (0'0 I)pe^ooJ8flSuoJls=Zie^ooJ8^\olleqs=l:pesru,(lel?lduoc=0:seqolreqtueJouorsnl;oeer8aq.67 .6I (0 0'o I) eseqreqlu? ot erodurorJ lllds pueerod = l :,(tuoeJod = 0 :ecuersrqepreqluv .(0.0,0.1) tuesqe= t:luaseld=0:slBFd Sl . (0'l'0 I),{ioueupuB pBorq SupeuIelp = I :sr1tpr,,rrpnbe = g :seqolx,{FJ (0.I ,0.t)00t-t6=Z:08-u=l:0/-I9=0:qr8uelx,(pcp1o1goo6€sBseqotx,{lBJ.9l (0 0'00 I)sllecZI-Z=l:92,c-Zl=On3eynsprxBqeeloelrprquo4EqetellelsursrulBJorequnN.gl '(0'l 'O'I . ) urnfueuofasuep = Z:srEqesuep = l :,(lrsuapunlpe(u = 0 :eceJlnsprxeqB eloelcBrq uo sJrEq elEIIelS (00'S0)tueserd=I:luesqB=0:eloelcEJqJoeceJnslprxBqEuospuelSetE^ep'pellBls'€l '(0r '0r) .Zl asotueuol= € :esuep= Z l,,(llsueprunlpau = I :peteD?Js= 0 :ecBJJnsplxEpE eloelJEJq uo sllpqelEllels (O'l'Ot)sselrol:92?lE^o=[:olorutol:€Jrldrrle.{l^\oueuroelE^o,{l^\o!?u=0;odBqseloelcprg.ll '(0 1'0'l) espql? pesnJ,{Foqs= I luesqu= 0 tuorsnJeloep?lg'01 '(0 'O 1 l ) pnqernpu Butsolcue= y :pnqernlpu tursolcue lou = 0 :pnqur seloelceJg 6 '(e 'S .g 0€ 0 O)Suol urru 6'0-S.0 = Z :3uolurtll t 0-1.0= I tuesqB= 0 :lecpeduo srrBqlolnpu?lo '(0 '0 ., 1 1)snoreul-g = I :snoreur-t-t= 0 :re^{olC '(199 0'ft8 0).reeurr= t icudlle ,{l^\olJEu= Z :elB^o,([^lo.u€u = I :elB^o,(lpEolq= O :edEqs]Eel llnpv 9 (g'0'9 0),(^s^r/pedsut= I :tq8rE.Ils= 0 :ur8reuJealtlnpv E '(0'l'0 1)alnlo,re,r = Z ipe^rncer= I :lpu = 0 :urSreugeel t1npy 7 (0 0'0'l) elepror= I iel€nuellE= 0 :eseqJEel llnpv € '8 'Z (r99 0 0) ruur0-9 I = t :ruurg l-O € = Z tuur l.€-S i = t :u{u 9.t-9 = O:WAuel elotedJEel lFpV .l 10.1,0.1)lcera= I :pexeuaJtolBluozlroq = O:uouBluel.toJeal lFpV 'retcoJBrls qcEeJoJ sle{cuJq ul pelEclpurar€ ,eeJ] lse oqsalSuls eql uro{ secrpuruonueler pu? fruelsrsuoJ patepJounoJe^r lsej aql puu peJepJo eJe^\ ZZ, OZ,gl,VI,ZI,g 't 'Z sJel J€qcelaspp141 'exet dno:3tnoo/r\l plrE wn\odasols(I a/.J'ltoJ peJoJs sJetc?JEqJ .l alquJ

'wn1odasotstT 'olsowotll puu ogouar1rm7 ur snoSoloruoqAureq sE pa?er1are serod punural aql ,,(pnls srql ul luaurdoleaap,{lrzeuleleuldnserAureqJeqlu?eqlpe^\oqs,urlTodasotsKT o1Aflo1oqd:ou:eq1ue ur.rupurrs srtlcr\$'olsoworlJur(996IIeeH)fueuldolaaapreqlueSo,{pn1sy.,{leszqalDur8uo,(sru,{oqtlnq 'uousodlmrdueqlurEuraqseuorsr,rarsrqlupeqursepe&seJodraqlue6l:alcereqcrog.sr.ur:aprdaaqt spocuoc,{lrsueprmq aJoq^r osoluauot pu? 'olqrsr^ Eurumurol sruueprde aq1 qtrrrr paddey:a,ro ,{18uons 'f1erete1 ,pet€J?das eJ?srruq orl uoq^iesuap Euqcnollsnf aresJ.pq eql uaq^\,{lrsuep unrpatu lle^{ ,elplnpun eJEsJlpq eql uoq^\peJa$pJs se pougepsl ,(llsuepruq oJeq^\tl Jelc?reqc lo ,(^u^rEureq sE paugeperu sur8reru pedsuc ereq,r 9 :elceruqcroy ldacxe ,{roteuuldxe-;1as e.ru suoqrugep JalceruqC '(Z elqeJ)peluesord sl xlneura?p SutFseraq; .cqdrour,(1odse papoc se,n uox?l aql ,uox€l .peJepJoun e ulll/r\ luasajd ss,!\apls Jalc?l?q3euo ueql sJoluelar&\,\ ual eJe^\Jepur€rxaj arD alrq.{\ 'parapJo s?pelea.! ara,[ odeqsJo ezlsul suorsse:Eo:dpcl8o1 peluese:der qrq,,rr srelcenqc elElsrlFI '(I I elq?I) pepnpu eJe^\sJatJsJeqr etzlsplnur a,rqelrpnb pu? a^n€Uu€nbqloq puu ,fteurg c:eue8

, .V.t tumlodasols(1snue8 uelpllsnv ruEso { oql Jo uorsnel r(louoxsl V tddeqJ pu? sup{lr^{ .C.J 574 Nu)tria Vol. 13, No. 3 (2001)

Table 2. Datamatdx for a cladisticanaly sis of Lysiosepalumand two outgroupsp ecies in Guichenotia '?'. andThomasiq.Missing data are coded as

I 11111 11112 22222 Taxon 12345 67890 t2345 67890 12345

Guichenotio 01010 I1000 00000 00000 000?0 an|ustifoliL 22 3 lll

Thomasia 02000 21000 00100 00102 000?0 angustifulia I 3 1

Lysiosepalum 03011 t2ttl 10101 2r0r2 l0l0l abollotum ll ll I

L arofiaticum 001ll 00211 10110 2to02 00100 I

L hexandrum 13020 322t1 12t20 21002 lllll I 2

L involucratum 13020 32ttl t2t20 2r002 lllll 3 I 2

L rugosum 0l0lr l20lI 10010 2tot2 00101 II t2 I ll

Phylogeneticrelationships were determined using the program PAUP 3' 1 1 (Swofford1993), The outgroupmethod (Watrous & Wheeler1981) was used to polarizecharacters. Character states were equallyweighted for abranch and bound search to look for equallyparsimonious trees. Bremersuppon (Bremer1994) and bootstrap frequencies (Felsenstein 1985) were used to estimatestability of each node.The total support index Ti (Bremer1994) was determined by summarizingall Bremersupport valuesfor nodesin thecladogram and then dividing the sum by theobtained tree length. A bootstrap analysis(branch and bound) was performed with 1000replicates.

Resultsand discussion

Examinationofherbarium specimens and fresh material revealed five discretetax ainLysiosepalum. Theseare each sufficiently distinct to be recognizedas species.

Inflorescencestructure. Lysiosepalumlnflorcscences have been incorrectly describedas axillary (Mueller1859) and as racemes (Mueller 1859, Bentham 1863, Moore 1921). Actually they have - opposed,monochasial cymes with 2-12 flowersfrequendy oriented towards one side ofthe inflorescence. This study confirms the inflorescenceform of Lysiosepalumis ^s describedfrom one species L. involucratum,byBayer & Kubitzki( 1996).The flowering shoots are sympodia with repeating units of two or threevegetative nodes and a terminalinflorescence that is overtoppedby the axillary product ofthe distalleafwhich leads to a leaf-opposedinflorescence (Figure 1). Thisstudy also confirms that Lysiosepalumhasthe basic repeating bicolor unit (Bayel1 999 , afterTheobroma bicolor where it was irst observed)which is a synapomorphyfor thecore (now circumscribed as , sensl Judd& Manchester1997). Thebicol.or rnit is determinateand bears three bracts, one ofwhich is sterile 'e^Jnoer 'JelleursauroJeq '{eqt d1a'tlsser8or4 eqlJo rluoJ srnletlt eql sl qtq^\ sulEJ€upue J€euI[ eJour '3uolqo 'epou ellq^\ 'se^ealllnpE [?ql Je8relserull rno; ot dn puepg arese^?el qlue^eso] qlxs eql trlo{ .e4luo ,l?upu? els^o eJe sa^?al o^rJ lsJg e{L pueel?ulelp eJe salceds pe^lesqo Jo se^€31luenDasqns 'snoJqBIS'arpue 'cpd11e 'ueer8 'el?loqed puJ rr.rg uorleul,tteS-rege tuipuedxe ,(pq311s puz l€U 'l€ealda e.ruleri1 suopal,booqlr.t uogzuruuaSre1&ocoreuuqd .toqs s?u\paes wnpdasots[7

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'luouac?ldslpluecsalnscar se pmSer l{sllqn) pu8ref?g qtq^\ ^ ?lsp erou pa?nlls 'ocuocseJouu om sre^\ouluanbesqns puE puoces eql Jo lcelq Eupuelqnsoql eql Jo Jo^aoulsuluxel '(I or0sr ll sel€rlpulel?ls (966I) Drsllqn) T' re,{89tlcq/'\ ernEtd)lcElq Eupuelqns ou sEqJa/t\ou '(666I lsrJ aql'slc?rqelFels aerql;o xflecrda ue a^8q sra.{rou wryodaso1st7u1 re'{ug)slcurq ellrols aeJtp;oxfpclde uuol peJnpeJeq u?t lI sJe^\ouel3uls ul Jo'sauJ,bFlalel puolqnso^\] lo$o eqlPUE

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wnpalasoF(Tsnua8 ueq?jNnv uetsorrdeql Jo uolsl^ercrurouox?l v'llldd?ql'VI pue surxllrlAJJ 576 Nu)trid Vol. 13, No. 3 (2001)

leaf and stipule. In addition, seedlingleaves in the glasshouseand in the field, have larger, more scattered,stellate hairs than thosepresent on leavesof matureplants. Stipulesare absenton the first two leavesand develop at the third leaf node in all speciesexamined and at that stageseedlings are 1.5-3 cm high.

Andtomy. Transyersesections of leavesof l,. hexandrum (K.A. Shepherd68, J.A. Wege264) and L. abollatum(C.F. Mlklns 1213),show them to be doniventral. In the uniseriateepidermis there are single cells or groupsofcells (Figure 2A) that are larger than the remaining epidermalcells and stain reddish-purple. These are consideredto be mucilagecells, as Toluidine Blue stainspolycarboxylic acids,including alginic and pectic acid, reddish-purple(Feder & O'Brien 1968). Mucilage cells and canalsare reporled as characteristicof Malvaceaes./. (Metcalfe & Chalk 1950)). The palisadelayer is biseriateor with a poorly organizedthird layer and the spongymesophyll cells are loosely packed with obviousair spaces.A bundlesheath around secondary veins is uniseriate(Figure 28) andconsists ofmoderately thick-wal1ed,achlorophyllous parenchyma. Primaryveins aresurrounded by two layers ofthick-walled parenchymacells and a crescentof fibresis presentdirectly below the phloem. Species examinedare hypostomatic with guardcells that havemucilage inclusions. Crystal drusesare present ascell inclusionsin the palisadeand spongy mesophyll layers (Figure 2,A). Clustercrystalsofcalcium oxalatehave becn rcportcd as a featureof Malvaceaes./. (Metcalfe & Chalk 1950).

-"819{6Pfu,& gc

Figure 2. A - tranverse section ol le f of Lysiosepalun abollatum (vo[cher C.F. Wilkins J213). Key to labelling: ab abaxial palisade; c cuticle; cr crystal; e epidermis; gc guard cell; gh glandular hair; mc mucilage cell; scale bar - 0.1 mm. B - secondary vein of leaf of Lysiosepalun hemru)rum (voucher K.A. Shepheftl 68) bs bundle sheath; e epidermis; h hypodefmis: ph phloem: x xylem. 'estrqdololpuores 'c9olaLLI3ul^\oqs - g 'clloloru u'llod rrnSr{ 'eseqdPlerutsrg 8u{Moqs - v 0l = u qlt^\ ut1lonnp^ut wnqdasols!'1 Jo sll') rrqlotu €

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G. angustifolia

T. angnstfolia

L, atuhaticum 8632

8 100 2 L abollatum NodeI 58 231716 ll t09 8 Node3

196

L, rugosum 138 2

5 L hexandrum 99 Shortesttree & steps Node4 No.of treesfound I ct 0.906 24t4t24 HI0.5l6 Rr0.800 L, involucratum

Figure 4. One tree found by a blanch and bound sealch of five lrsiot?parrtn and two outgloup tax.. Numbersbelow the line aie chamcterschanging unambiguouslyon the branch. Numb€rs above lhe line in italics are Brcmer support values and in bold are bootstrapfrcquencies to estimaie stability of lhe nodes

Lysiosepalumaromaticrm is sistertaxon to theremainder of thegenus. It retainsthe plesiomorphic featureofa 5-partitecalyx. Autapomorphiesarelong leafpetioles up to 7 mrnlong,consistently cordate leafbased,broadly ovateto ovateleaf shapeand long glandularhairs on the pedicels. The otherfour speciesare delimited at nodetwo by flowers being 6-merous,broad filaments 0.,f{.5 mm wide, and more than eight ovules. L rugoswn andL. abollatum are sistertaxa to the remaining two species L hemndrum andL involucratumandtnited at nodethrce by having anthersthat split longitudinally from theterminal pore at maturityrather than only poricidal dehiscence.They alsohave mature ,{1pprc11nro1e llr4r 'relnqnl 'Ktp DwEUS ':epunb pseq eql uo sJleqelgllols ollrl^\ cllssost{lt^\ ro 'sJruq 'pellao-mo1ro snorqulS'uuo;1yg'eyduna1fig snlncoy:ed911lceJe saln^O attllels s:reqeydurs 'elnuru'e1rq,trqlr,tr,41arerro snorqel3,{1ureu selnJolJoeJEJns Jeuur'eseq Suolqo Suol ol;oqs e ruJoJol 'elrq,,r'eluuq 'elrsses 'elrxe posnJses€q ller qlr,^\srruq etullals Jo SurJe^ocesuep € qlr,r eJ[Jrnsrelno uoqrlueceld 'J?lnJol-E'pro^o 'olrssas,ftu^o :etuuuoJ Agenlel sladto3 (g oJnBrC)olBlncuer ?urxo 'aurJlno 'peuelleu 'etrq,r 'eleJodlocut ur cqdllle tuq,retrtos ua11o4 Aynlcut 1r slqs purpnlr8uol se 'soJod 'e1e,ro popuelxoeuoJoq (seneds o.tr1ur) qrrq,tr Ieuluualqns ,{l,norreuruoJg e3uecslrlap esJo.rlur 'peller-Z '(ol€upeuua^) qlr.^\'el€^oFl.^\or?u e.€Jrnsraqlu€ ol pesnJluaruu[JJo xedtrqtr^\ pexuuluo^ 'seseqJeqluuuea,^\leq sduS,tro::uu qlr,r 'eqn1uuuo3 ol xedepe,uncurlu Surqrnol flpJJr?l 'lJorl^\auo ur 'peueDEU sraqruv tt\oJmurc pEoJq uoruod ee4 :eqnl lEuruels u olur pesnl fp:oqs Jo eeq sruawDlrl 'sno1e1ade1ue 'olnuru 'ae.rJ 'gFg sueuels :Juln8usul Jo ?qnt lpurruBtsJo xed€uo ro tuesqusepouluels 'alppJocqo 'pa: 'tquozuorl :peJ tq8uq Jo poJ {JEp wnlJaoJpuv Jo el€^oqo tq8r:q ro paj I.mp Jo 'ee{ 'elrsses'alrl-el€Js 'luaso.rd 'aseq lJeJa Jo luosqes/r1ad pauelcrql uo.rJ eqol eql otur Surpuelxe 'sJr?q squ lueururord g-€ pue esEqpeue{J[ll qlr,r\ a3e]ns [Erx?qEeql aldurs pur? Lrn]rpruu l'0 ol s:lpq elullets peuJu-g-Z alrq,r ouu Jo runlueunpur ssusppue xedeo1 asu q peuelcrqt urorl Surpuelxe 'Suolqo 'alc^o sure,rpesserdurr pur esuqpeuolrrqt posrslqlr,{ eJEJnsIerxupe eW Jo Jrtdrtle,{l,roJ.rBu '{urd-qsr^nuur 'proptad ,{l,rorvu saqol:sur8J3u pexeuar qlr,r Jo l€U reqlre oJuleql soqol Je^\oueu puE seqol lug apr,r,rSuqeurellE pur? eqnl oqs ,{:aa.u q1r,n'el€loJ 'luolsrsJcdr:trlrJ sno]Jru-gJo -t 'spJe,ru,rop sr tf]Il.lt wnJuDworo I ldeJx" snoJeru-g ?urcu1uauolg x,{yec3ur1rnr1 Sursolcua-e: 'pnq ,41ptgcd's:a,tog uro4 pe^JntaJ'uru I-€ 0 es€ql€ pesnJ eJnlEueql esolf,ue,{letelduor ol paruds 'elE^o 'x.(luc ,4lpnbe ot ats^o ,{lppojq luelsrsJad .troloqf pcaJrp'aetqlspDrq xtlz;rrk xedutu :epr,r,r 'paJ 'e1nce ':eeurl 'Jo^\oU ,{lleur8reruuego s;laxlpad xadr :o alulnqlrds luelsruod lucrdeSutpualqns 'Je,\\oU o,^\tpur? sra,^\ou alurporurolur Burpuelqns euo leuruuel Surpualqnseuou sT:rr.rgTpct alcunped :uosEesSuue,rog snor,rerduror; tuelsrsled s,4r,4\luuels ecueJseJouurpue alcunpcd oql'sje,roU 'sJe^\oU 'pasoddo 7 1 o1dn q1ra,reru,(c paqruerq u ,{1ewr pnxesrq 8-Z ;o eur,(clersuqcouour ctr;3uo1e -JEeluaJuaJSaJo L[ul quprurlueururo:d qlr,r oJeIns Istxeqe:pesserdrut,{13uo.l1s sura,t rofunr fluo ueq,tr ur3;uruernue qlr,nroso8nr ,(nurEJ Jo 'pssserdrur,413uo4s surel rouru pua Jofcurueq,n ur3:eur padsrrc qtr^\ eso8nr ,{lSuoJtsJeqlre'qr:plul passerdur fldaep u puu uorleue^Jounu otr?lncllclrllr,r ecegns 'olrnuctle lErxspB:oluuurd sa^ual eJnleurpue eleurpdruurd se,rualelrua,lnf;o uorluu?A Jo alEp]oc 'Suolqo-Jeaur 'ale^o 'etE^o'ele^o 'uea:3-prur espq l ro ,{l,rorEu ,{lp€ojq ro uea:3-,{a:3sa2r1g poqs.ro Buo1sa1o4a4ql,tro:3 3uno,( uo pepmorc'pexagaj:o SurpueJds'lcala 'aldturs'elEuJotle ra,1r2? esntqo 'arnlxel xedupuc peunJeJ ,{13uor1sro ,{pq8rlsurBrrur snoaceuoce Sur,ruqur alrl--}rol :$3ual Jrel lo qlual auo IlolBrurxorddE'ruroJruer ro alEAorxroJ eJnl€rx 'elr,ro-,(lpuo:q epulq alruc^n i 'cnbrlqo esEq 'etB]rdqs 'tuetsrsJad'(wnlDtJnlo,lt1l :o elrsses ?) ruolse uo luosard,{lqeuea ro lueserds,{um1u saJndlt5 'osolueluol 'lJere'eteJel luaJsejqslS suals oJnluu saqtuntq 3unotr sJluqJl?lnpurlt Suol Jo uoqs qtr,,!\paxrtu ueuo'sezrs pexruJo sJrEqetBllals alrsses Jo/pue pa{pls Jo tUnluctunpurqtr,\\ urnrJao.rpuE 'sqr;?s 'pounuals-rllnhtr pue sleledldecxe sareJlnsluuId'tuesqe roluese:dluecs fuedded lcrdruoo

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Figure 5. Reticulirtc cxotcsta of white, prolatc pollcn of Lysbsepalum invoLucratum (voucher L. involrcr.rllrn, Kings Park Bolanic Cirrdcn sccd 196/1986, Magenta Rd, S of Ncwdegale). dehiscentcapsulc, outer surlace with persistent,bdttle, stellatehairs. Seeddark brown, ellipsoid, smooth with scaltcred,simple and sessilestellate hairs, aril cream, cap-like, with two longer lobes extending up thc hilar surface to half the seedlength, testabitegmic, endospermpresent. Embr)o straight.

Distribution. Lysiosepalumis widespreadin the wheatbeltfrom Yuna (north-eastof Geraldton) to Ravensthorpe(Figure 6). Plantsoccur in open woodlandor shrubland,often at treebases, in lateritic gravel,yellow-brown sand, sandy clay or sandyloam.

Nores.Chapman ( l99l ) obscrvedthatBentham ( 1863)misspeltthenamelyslosepalumas Lyriosepalum "Flora in the index to Australiensis",but had the conect spelling in the text, and Schumannmisspelt the name as Lysiopcldrln (Schumann1895).

Lysiosepalunris most similar to Lasiopctaleaegetera Gay, lttsiopetaLum Sm, and Guichenotia Gay i11having teminal, poricidal antherdehiscence rather than dehiscenceby extrorse slitsas in the threeremaininggeneraofthetibe,HannqfordiaF. Muell., Gay andSeringia Gay. Lysiosepalmr is, however, unique in Lasiopetaleaein having features that Mueller (1859) considereddistinctive when he describedthe genus. Oneofthese is threeepicalyx bracts that are shortly fuscd at the baseand equally spacedto completelyenclose the maturebud. Other Lasiopetaleaehave one epicalyx brrct (l,asiopetalum), no epicalyx (Keraudrenia and ), or three unilateral epicalyxbracts (6al chenotia,,l,Qsiopetalum,Thomasia)thatrf ovate ratherthan filiform, cover only the cxposedside of the bud ratherthan completelyenclosing the mature bud. The second distinctiyefeature ofthe genusis the almostfree calyx lobes,compared with other Lasiopetaleaewhich have longer calyx tubes. Mueller (1859) also describedrJsiosepalum anlhersas being distinct from thoseof LasiopetalLrmin not splitting longitudinally from the terminal pores,however, mature anthers of L. rugosum anLIthc new speciesZ, eboLlatumdo sharethis character. 'uur 'aplnqlzds.lo qtr,t pe8uggsulEreur I-€'0 x 0I-g Suoyqo-reeu-llstrr.tg3uo1urur gg-g1alcunped :peja./i\ou-(g)gro -t'3uol ulru 061tr acuacsatoylulsJPq peJe ms qlIA'eso8nrf13uo4s eceyns :urnlueumpqesuap ? qtr1'\oc?Fns I€IxuqE :sJluq allssas Eu/Fepun ou lnq spuBISele^€lc I?rxEpu 'esnlqo allq,r pu? srr?qel?lals pe{sls t{ll^r secqJnsqtoq 'paNncer ,{leturopotupuz pedsucutSreur xode'eppJo'-qnsasuq'(qI{\oJEEuno,{uluurtxg€,)ultu€-Zx9I-0I'elu^o,4l,l[outu'snorolocslp 'uaert '(qltro:8 prurapelq :Euol ruur I-g 0 eloped:tupeelds ro laeresaADrI 3uno,{uI utrr 9 I xt ?) urur€'I-90x g'I-9'0'ollssesluasard dlpnsn saprdg5 Suolururl'9 e spuel3e1z,re1c alq,t 'allssasqns '3lua}uE] puu ru?ryuIuI 0'l-sz 0 sJIEqoleuels ro ollssas8ul{lJe^o tlll,it\ellqa suuEpuu "njutp Suolurul9 {lBls peJE qll,\\ urut E Z-l sJI€qelsllals pauIJ?-Z I-? parallPcsJo lu$uaulnpul 0rZ'0 'slrErq 'selollsd :sJI?qopuels e8rel uror; ecuereadde,(1oo.,n ol1qm qur (slecrpedpu? salcunped 'uI 'se1ndr1s'saqcue.rq Euno[) satuaJserouulpue qltro.r8 Sunoi(:luesqE luats S I x S I ol qmryS

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stalkedstellate hairs. Pedicels5-I2mmlong, Epicalyxbracts shordy fused at baseto 0.3 mrn, ovate, 6.5-9 x 3-6.5 mm, apexacuminate, margins fringed with stalked,stellate hairs; adaxialsurface red, with scatteredsessile, flne, white, stellateand simple hairs and claYate glands c. 0.1 mm long; abaxial surfacegreen base and purple apexwith mediumdensity, long'stalked stellaie hahs to 3 mm diam.' which protrudefrom the margins,overlying white clavateglands c. 0.1 rnn long' Flowers 6-merous' Calyxlobes ovate, elliptic or oblong,6.5-11.5 x 3.2-3.5mm, aPex acute; abaxial surface with sessile andstalked, white stellatehairs c. 0.4 mm diam.and scattered short, clavate glands c. 0. 1 mm long, ribs with additional scattered,stalked, white tan-centred,stellate hairs to 1 mm diam. Petdls usually present,usually horizontal, rarely erect, flat, darkred, 0.8-1.5 x 0.8-1.3mm. Androeciumdarkrcd', staminodesabsent. Filaments ftee ot shortlyfused into stamind tube0.2-{.4 mm long; free portion

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Figure 6. Distdbutionof Ltsiosepalurnin westemAustralia. A - whole genus,B-L obollatuq E, L aronnticuma and L involucratum O. C - L hexadr m D - L rugosum 'LEE6 - qpauuaN C)I tuo{ u^relc (tuur I = rsq I T C'g) IPsIo I .^leF - uo puelS lelxeperewue - J Iqed - g lruru Sz = lDq) ren'oB ( (u(u 90 = Jeq)eloelrerq Jo 5rBlns FlxeqP - - arn8r[ pu? 4?q aFIIsls - J {urur 9 Z = Igq) soce}ns Jsel g (ul. I = rsq) lJre\ V wnfi oqo uqodasolsK1 L "T#;,#:e ffi W,,

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0001 (PERTH)l Wongan Hills, l5 Sep. 1983,KF. Kenneally 8830 (PERTH); Wongan Hills, 3 Sep. 1984,K.F. Kenneally 9331 (PERTH).

Dtstribution. Restrictedto one locality in the Wongan Hilts range of hills, north of the township of Wongan Hills, WesternAustralia. (Figure 6B)

Hcbitar. This speciesoccurs in orange-brown,sandy clay with laterite,in open mallee woodland (to c. 5 m) at baseof Eucalyptusitrees.

Phenology. Flowering time is August to October. Fruit not obseNed.

Consenation status. CALM ConservationCodes for Westem Austalian Flora: Priority One. There havebeen four collectionsin the sameapproximate area, however, it is not known ifthey arefrom the samepopulation. Currently known by the authorsfrom one or two small populations on a private reserve,bordered by cleared farmland. Further survey urgently required with probable need fbr recognition as Dcclared Rare Flora.

"a Etymology. Abolla (Latin) translatesas robe of thick woollen stuff wom by soldiers and philosophers",and alludes in this speciesto its woolly appearancederived from the loose covering of long-stalked,large, stellatehairs.

A.lffnities.This speciesis similar to L rz gosum inhavingalong inflorescencewith up to eight flowers andleaves with rugosesurfaces and recurved margins. It differs in having shortpetioles 0.5-1 mm long, rather than l-3 mm long, epicalyx bracts with an acuminaterather than acute apex and a woolly appearancefrom long-stalked,large, stellatehairsrather than aclose tomentumof sessile,small, stellate hairs.

2. Lysiosepalum aromaticum C.F. Wilkins, sp. nov.

Speciesfortiter amaroodora, foliis late ovatisvel ovatis petiolis longis, stipulis petiolatis,flodbus 4 vel 5 meris, antherisrutilis a congeneribusdiversa.

I)pusr StateForest 52, Highbury block, EastemBoundary, Westem Australia,23 November 1998, G. Warren 23, C. Taylor, P. Rose. (iolo: PERTH 05315700;iso; K).

Shrub to75 x 100 cm with strong aromaticscent; all plant surfacesexcept adaxialepicalyx bract surface,calyx, androeciumand gynoecium with dense,multicelled, long-stalked,red-tipped clavate glandsto 0.7 mm long, overlying dense,sessile, 10-12-armed, white stellatehairs to 0.5 mm diam.; young growth with tau hairs. Stipalesalways present,stipitate, 0.5-l mm long, blade 2-6 x 2-7 mm (seedlingyoung growth bladec.10 x l3 mm, petiolec.4 mm long). lzaves reflexed Petioles4-7 mm long (c. 15 nrm in seedlingyoung growth). Blades grey-gteen,scarcely discolorous, ovate to broadly oyate,16-22x l0-12 mm (seedlingyoung growth c. 50 x 30 mrn);blade strongly rugose,base cordate apex acute or rounded, margln crisped and slightly rccuwed Inflorescence 25-50 mrn long, 2-4(5)-flowered;peduncle9-20mmlong. Bracrsspathulate, 3-7.5 x 0.8-1.3 mm;adaxial surfacewith medium densityclavate glands; abaxial surface with densestellate hairs and glands. Pedicels54 mm long. Epicalyx bracl.rvery shortly fusedat basefor 0.3-{.5 mm, broadly ovate,6.5-8.5 x 4.5-6.5 mm, apexacute;adaxial surface with mediumdensity,sessile, white, fine stellatehairsc. 0.2 mrn diam. and simple hairs. Flowers 4- or 5-merous. Calyx lobes elliptic, 5.8-6 5 x 2-3.5 mm, apex acute;abaxial surfaceribs and blade with scattered,sessile, white, 2-5-celled stellatehairs to 0 5 mm diam. Pelals 'alJunpad '3uo1uur eql s?urnlueunpur euI?seqt qlr,tr g 11s1actpa4 ?uoJrttut9'g ol spuplSpeJeuu3s 'uIIu 'atelnqteds prr" sJr€qelEllals esuep ol unrpau qlr.r serElns qtoq g I*9 0 x 1-7 strD.ra sJ'-tp 'ellsses',4llsuap '8uol ruu ? 0 , srnq et€llatselrqa umrpeu 8ur.{pa^o unu , 0 ol spuelSet€^?lc peJ 'pelJets-8uol '8uoy '3uol qlr./t\ urur gg-91 elcunpad:pua,tog{8)9(g) uru 0 7-gg acuacsato[u1 3uo1 ', 'sJpq 'uEt 'Sul,|oqs Iuur I 0 spuelSpaJa eJsqlr,r ua$o etpllets esuepqlr,tr qupru fluo qlr,n ,{1pnsn 'r "ruerp ecuJlnsprx?qs :3uol ruu 1 g spuuy8elu,re1r pe{I€ts-uoqs parellurs r0r.r uolJo luru t 0 , 'eug 'elrsses',4lrsuap 'qloorns sJrer{elellels elH,.'\ unrpour qlr,r Jo eso8ru ,4lluluJe3uJ]ns l€rx€p€ 'oJnua 'alEnueDE :pa^rnaal ,{l3uoJ)s Jo J€ln8efi ^[lr{8rlsJeqlle ur8J?ru'esnlqo xede eseq'urur Z-g I 'cqdqye 'snoJoloJsrp'uear3-far8 'Buoy x 11-g ,{1,tro.ueu:o Suoyqo-reeuq sapop ruru S l-9 0 'lxeJe 'wrw 'alsses 'lueseJd '8uoy sapuad sau)zl t-I x I-S 0 sfe,trp salzd4g ruur g g r spuel8 'pelpls tpeJel?cs ' etE EIc peJ qll^\ pexru flur€ru ru€rpruul t 0-2 0 sJreqel"Ilets paJluac-uBlqlr,r ',Jo allq,r lo u€l'peuue-?Z runluauol e rpr saqcue:qSunod:luesqe luacs'ruc 06x 06otq lUS

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FiEuJe8. Llsiosepalumaromaticum. A-habit, B - juvenile l€af (bar = l cm). C- flower (bar= 2.5 mm). D-petal. E - antherabaxial view. F- slaminode.G - antheradaxial view. H - pistil (D-H bar = 1mm). I ovary hair (bar = 0.05 mm). J - glandularhair and stellatehair on abaxialsurface of bracteole(btu = 0 2 mm) K fruit with sepalsremoved. L - fruit within sepalsand bracleoles(K & L bar = 8 mm). Drawn trom G. warren 23 (A J) and CF. lvrlkinr 1418(K,L). 'Lgl su!111i1JJ pue8q pD4tu\s x LUorJufAric lluu l0 -c -g = = req) rpq fue^o - D (ulur 90 = wql ? d mqd unrceorpu?Jo 53sJlnsplx?pE (lxul z I?q c T a) - D^\oU-( (nru 1.0= wq) alool.srqJo oopInsFI\?qv uo sirlq olgllalspuv rPlnpueld J so.qrns qloous qll^\ so^Pel - arnt?lrl pue oJqrns esoBnrrltr^\ Jeel ollue^nl - B (urc I = ,!q eF.s) llqPq Y wtttPuDxaq uqDdasolsl'I 6 ern8lq

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Selectedspecimens excrmined. : I l5 milepegon GreatEastem Hwy, l3 Sep. 1962,T.E.H. Aplin 1986(AD); S of Narembeen,8 Aug.1.963, A.M. Ashby 162 (AD); 2 km by roadE ofPederah,17 Oct. 1917,W.H. Barker2467 (AD);5.5 km E of Tammin,1l Sep.1976,R. Coveny8318 & B. Habersley(PERTH);26 km dueSW ofBodallin, 16Sep. 1982, R.J. Crqnrteld 2349 (PERTH), HoldenRd, Shire of LakeGrace, 24 Oct. 1986, E.J. Crotford 5346 (PERTH);6 km W ofBunacoppin, 23 Aug. 1995,R. Dqvies85 (PERTH);Kukerin, Oct. 1913,M. Koch2185 (NSW, MEL, PERTH); 25 km NE of Dumbleyung,20 Sep. 1964,R.H. Kuchel 2007 (AD); Tammin,Sep.-Nov. 1909, J.H. Maidens.n. (NSW); 4 miles[6 km] W of Nyabing,14 Sep. 1961, R.D. Ro),ce 6704, (PERTH); 9.8km E of Benderingon ReserveRd, 8 Sep.1994, KA. ShepherdKS 57, J.A. Chappill,J.A. Wege (PERTH,UWA);500mN ofJilikin Flat Rd on Bueffner Rd, 1l Oct.1996,J.A.Wege;.2.(PERTH,IJWA); 50kmNNWof Menedin,3l Oct.197 4,D.J.E. Whibley 4746 (AD, PERTH);Konongoningnew school site,Williams-Wongan Hills road,5 Sep.1.992,C.F.WilkinsCW 167-168 (PERTH, UWA);Boolanelling NatureReserve, Pannell Williams Rd, 9.5 km N of Quairading-Bilbarrinroad, 25 Nov. 1994, C.F.Wilkins CW 736(PERTH, UWA);Tarin Rock,9 Nov. 1968,J.W.Wrigley s.n. (MEL).

Distribution. This speciesranges from Konongorring to Nyabing, in the south-west of Westem Australia. (Figure 6C)

Habitat. This speciesis found asunder storey it Eucalyptusopen woodland,or rarely heath,in yellow or brown clayey sand,usually with laterite.

Phenology. Flowering time is August to October. Fruiting October to December.

Conservation statuo'.This speciesis well collected and mainly recordedas abundantin the areaof collection. No special statusis required.

Etymology. Naned due to the hexandrousflowers.

ATflnlties.This specieshas the distinct specificfeatures described by Mooreof long red glandularhairs on the epicalyx, epicalyx that are greenishrather than orange{an in colour and mainly narower than in L. involucralum, aDdpetals that are usually absentand if presentthen fewer than six per flower whereasL, involucratlnr usuallyhassixpetals, Similaritiesto L, involucratumareleaves with smooth or slightly rugose surfacesand strongly recurvedleaf margins.

Notes. Two specimensfrom Chiddarcoopingand Narembeen ale anomalousas the long red glandsare rareor absentrather than foming a densecovering asin other specimensexamined. Thesehave been determinedas L. hexendrum as they areapetalous, with greenish-tanabaxial epicalyx bract surfaces ratherthan tan and without the dense,white tomentumfound on the epicalyx bract adaxial surfaceof L. involucratun.

4. Lysiosepalum involucratum (Turcz.) Druce (Druce 1917:634). - ThomasioinvolucrataT\tcz. (Turczaninov 1852: 143). Type citation: N. Holl. Drum. V. n. 255. Tlpe.' south-westof Western Australla,Drummorulcoll.5,n.255 (holor KW,?.v. photograph PERTH 05070341;lsor BM, K, MEL 724048.PERTH 01625608.W).

Lysiosepalumbarryarr.rmMuell. (Mueller 1859:143). Typecitation: In planitiebusad flumen Phillips Mx. ?ype.'Phillips River, Westem Australia,Maxwell (holo: MEL724049)

Illustration. Grieve (1998: 653). 'Frenuef ol Jeqruo^op Sugtnr; ':aqure,to111o1 .(1n1sl eruq 8:.rr::,t,ol1 KSopuaqJ

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Figtne10, LjsiosepaLunl itl,,olucratun A-habit (scalebar (bar)= I cm). B-mature leaves(bar=2mm). C-stellate hairson outersurface of bracteole.D - stellarchair on innersurface of bracteole(C & D bar = 0.I mm). E - flower (bar = 2 mm). F - stellatehairs on outersurface of calyx. C - simplehairs on innersurface of calyx. H - petal(bar =0.5 mm). I - adaxialsurface of androecium.J-pistil (I&Jbar= l mm). K-ovaryhair (bar=0.1mm). Drawn from K. Shepherd5l and CF. wilki s \lll.

Chromosome number. n =10 (Figure 3).

Consen)ation status. This species has been poorly collected and mainly recorded as occasional or 1-3plants, rarely frequent in thearea. Conservation status deserves review after sepamtion ofnumerous L. hexandrumfrom L. involucrqturlcollections.

Etymology.An involucreis alarge bractor whorl ofbracts;the epithet involucratum presumably refers to the whorl of largeepicalyx bracts covering the buds in this genus.

Affinities.This speciesis similarto L. hexandrumin havingentire, rather than wrinkled leafmargins but differsin havingthe pedicels and epicalyx with an indumentumof longred-stalked stellate hairs insteadof longred glandularhairs over short white stellate hairs, by havingthe epicalyx orange tan insteadof greenishtan andpetals that are rarely absent rather than rarely present. (C9 ern8Id) EllErlsnv 'Sutuueuru€I u:etsa,ld I ol (uolpleJoCJo lsee-quou)lle,^\Itod ruorJspuelxe solsods s\ll Ltounqy$!Q

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Fignre11 . Lysiosepalunrugosum. A - habit(scale bar = I cm). B - leaf adaxialand abaxial surfaces (bar = 2.5 mm). C - bracteoleabaxial surface stellate hairs (bar = 0.25mm). D - flower (bar= 2.5 mm). E-hairs on adaxialsurface of calyx (bar = 0.6 mm). F - petal. G - antherwilh staminodeand staminal tube, adaxial view. H - anlherwilh free filaments,adaxial view. I-pistil (F-I bar= I mm). J - brittle hair of ovary (bar = 0.1 mm). K - seedwith aril (bar - 0.J mm). Drawn lrom C.F. Wilkin! 216.

Habitat. This species occurs in red-brown clay or loam with laterite, as an understorey shrub in open shrubland or open woodland.

Phenology.Flowering time is Julyto October.Fruiting pedod is Octoberto February. 7-

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Bayer, C. & Kubitzki. K. (1996). Inllorescence morphology ol some Auslralian Lasiopetaleae(.S13rouliacea.). Telopea '721 '72'7 6: . Bayer, C., Fay, M. F., de Bruijn, A. Y., Savolainen,V., Mol1on, C. M., KubilzK, K., Alverson, W. T. & Chase, M. W. (1999). Support for an expandedfamily concepl of Malvaceaewilhin a recircumscribedorder Malvales: a combincd analysis of plastid atpB and rbcL DNA sequences. Dotanical Jaurndl of the Linnean Society 129: 261 295. Bayer. C. (1999). The b'color unit - homology and translormationofan infloresccnceslructurc uniquc to corc Malrdles. Plant S)'stenntics atul Evolutiotl 214: 187-198. "Flora Berrham, G. (1863). Australicnsis." Vol. l. (L. lteeve & Co: London.) Bremer, K. (1994). Bfunch support and tree stability. Cladistics l0: 295-304. "Conposilion Brown, R.W. (195,1)Leprin! 1979. of Scientific Words." (SmithsonianInstitution Pfess: Washington DC.) "Seeds Corner,E.J.H. (1976). of Dicotyledons." (CaDbridgeUniveLsity Prcss: Cambridge.) Danilafsos, C.D. (1993). Introduction to the ESEM iDstrumenl. Micrrrcolr) Research and Techniq a 25.35+361. Druce, G.C. (1917). Nomcnclalorial noles: chiefly African rnd Australian. The Bot nicrl Sociely Exchange Club of rhe British Isles Rcport for 1916. Suppl.2: 601-653. "Encyclopaedia Ellior,W.R. & JonesD.L. (1990). of AustmlirnPlants." Vol. 5. (LothianPublishing Co. Ply. Ltd. Perth) Fcdcr, N. & O'Bden, T.P. (1968). Plant microtcchnique:some prirciples and new rnethods. Anrcricdn Journal of Botan! 55, 123-142. "ChromosoDre Federov.A. (1969). Numbers oi Flowcring Plaols." (Nauk: Lcningrad.) Fclscnstcin,J. (1985). Conlidencclimils on phylogcniesran approachusing the bootstrap. Etol tion 39:'783J91. Cay, J. (1821). Monographic dcs cinq genres de plantes,Lasiop6lal6es. lvtunoires de Museum d'Histoire NatureLle7: 431-468. "How Cricvc, B.J. (1998). to Know WcslcrnAuslmlirn Wildflowers"Pa 2. 2nd edn. (Universityol'Western Australia Prcss. Perth.) Heel, W.A. van (1966). Morphology of the androccium in Malvales. Dlunrca l3t 17'7-394. "Census Hnatiuk, R.J. (1990). of Australian Vascuhr ." (Australian Government Printing Service: Canberra.) Judd, W.S. & Manchcstcr, S.R. (1997). Circumscription of Mrlvalcs as dctcrmincd by a prcliminary cladistic analysis of morphological, anatomical, prlyonological, And chcmical chaft\cters. Btiltonia 491 384405. Maddison,w.P. & Maddison,D.R. (1992). Maccladcr Analysisol phylogcnyand charactcrevolulion. Version 3.05. (Sinaucr Associalcs:Sundcrland.) "Anatony Mcrcalfe,C.R. and Chalk, L. (1950). oi thc Dicolyledons." Vol. I. (ClarendonPress: Oxford.) Moore, S. lc (1921). Alabastm divcrsa. - Part XXXIV. The Jouflnl of Botany 59:243-251. "Fragmcnta Mucllcr, F.J.H. voo (1859). PhytogrrphiaeAustraliac." Vol. I, pp. 142-143. (Cove|llmcnt Printcrl Mclbourne.) Sargenl, O.ll. (1923). Notes on wcst Australian planls. zre JounaL af Bohnf 6lt 285187. "Die Schumann,K. (1895). Slerculiaceae.hr EnSlcr, H.G.A. & Prantl, K.A.E. Natitichen Pflanzenfanilien." Vol. 3.6, pp. 67-99. (Verlag von Wilhclm Engelmann:Leipzig). "Phylosenelic Swofford, D.L. (1993). analysis using parsimony". Version 3.1.1. (Illinois Natural History Societyl Champaign.) Turczaninov, P.K.N.S. (1852). Dccas seplima Senerum adhuc non dcscriptorum adjcctis dcscriplionibus nonnullarum spccicrum. Bllll?rir de La SociAG Imptriale des NaturaListesde Moscou 25(2). 138-18). watrous, L. & whcclcr, Q.D. (1981). Thc outgroup comparison method of characler ^nalysis. SJstematic ZooLogy 30: l-11. Wilkins. C. F. (1999). Keraudrenh eraslia a\d Keraudrenia kdutona (.M^lvaceau ) new species from the Kimberley r€gion of weslcm Australia. Nr)/rid 13.233-242. Wilkins, C. F. (2001). Taxonomic revision ol HannafatuJia(Lasiopetaleae: Slerculiaceae (Malvaceae r.l.)). Arrtrdlidn SJstenatic Bolany l4(1)i in press.