Nomenclaturalnotes and new taxa in the Conostylis aculeatagroup ()

By StephenD. Hopper+

Abstract On the basisof detailedfield strLdies of lhe Co osrilisaculeata R.Br. gro.up andan examinationof specimensai themajor Australian berbaria, (i) C. robustt! Dielsis reinstatedas a speciesdistinct from C. acaleara,(ii) confusionconc.erning the ideniityol C. stylidioidesF. Muell. is clarified:this speciesis redescribed a1ldC. prolifercBenth. is r'einslatedas a distinctspecies; and (.iit\C. aculeata ssp.hrevifotu ssp. nov. and C. puLrcif{orasp. nov. are described;both are reslrictedendemics of the westerncoastal region of the SouihWest Botanical Provinceof w.A. A keyto the 7 speciesnow recognized in lhe C. acule.taEtoup is provided.

Introduction The Conostylisq(uleatq R.Bt. group consists of a complex of sevenspecies of perennialherbs whicb are restrictedto the westerncoastal plain and nearby plateau regions of south-westernAustralia. The group forms part of the sectior Cor?orryl/.'(Bentham 1873; Geerinck 1969), and is characterizedby leaves with marginal spines or setae,numerous ovules borne all over the placentalsurface, and a basicchromosome number of x : 8. The of this speciescomplex has been notably problematical in previonssystematic studies of the . Severalauthors have commented on difficultiesin delimiting taxa (Bentham 1873; Ewart 1906; Domin l9l2; Green 1960). Indeed,herbarium studies leave the impressionofa polymorphic but intergradingassemblage ofpopulations wjth few fornrsthat are consistently distinct throughout their geographicalrange. Elsewhere,the present author has shown that both hybridization and ecologically-correlatedintraspecific variation occur in speciesofthe C. aculeqta group near Dawesville on the Swan Coastal Plain (Hopper 1977). It was there denonstrated that while the identification of specimensfrom this region could be difficult in the herbarium, critical lield observatiorlsenabled the satisfactoryplacement of individuals into morphologically definable species and/or their hybrids. This work suggestedthat a wider field study of popu- lation variation and ecologywould facilitatea sound taxonomicunderstanding of the C. aculeqtq group as a whole. The present publication seeks to formalize necessarychanges in the tax- onomy of the group arisirrgout of detailed field surveysconducted in 1975 and 1976and a critical examinationof herbariumspecimens lodged at , UWA, AD, MEL, NSW, CANB, CBG, NE and BRl. This researchhas revealed (i) evidence in favour of the reinstateruent of C. robusta Diels as a good species,(ii) past confusion concerningthe application of the name C. stvliclioidesF. Muell.. and (iii) the existenceof two undescribed taxa While it is felt that present information allows for a reasonably sound assessmentof the speciessituation in lhe C. aculeatagroup, a number of problems regarding the status of infraspecific taxa remain unresolved, par-

* Botany Department,University of WesternAustralia, Nedlands, W.A. 6009 and WesternAustralian Herbarium, Department of Agriculture,Ceor'ge Street, Sou{h Perth, w.A.615l. 254 s,,u?qluegr'ql u.,noqse^Eq'n"rn uoro',J.i"oeqr ur,{llerredso'sarpnlspleg,, r"ql 'llenl^I ') 8uD?ts g sapTolpylrQsJ ropun {Luouou,{s ot qrueg DSow p.t pLte 'q]rreg 'sllltsouo) Dta/jp.rd , pecnper (0961) uearD Jo uorsl^er srq uI 'q1uag 'llonl{ 'C eragllo.rd'J pu€ saprolpll,{lssllf,lsouoJ Jo d]puapl aql

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Figure 1. Geographical distribution ot Cotlosi,lls robusta (.blackcircles) and C. aculeata (open circles)in the Mingenew SharkBay areaand resultsof a discriminantfunction analysis of l0 fforal neasurementstaken on samplesof the two speciesfrom near Kalbarri. Camera lucida drawings of representativehalf-flowers are illustrated. Measurementslaken on half- flowers: l, perianth leDgth;2, style length; 3, ovary to baseof highestaniher; 4, ovary to base of lowestanther; 5, ovary to baseof lowestfilament; 6, perianth lobe length; 7, anthe. length; 8, perianth width acrosslobe bases;9, perianth width at the ovary; 10, perianth lobe width at the base. 256 L9Z ')lc€Il ^Jepunoq Suole peor uEur ru{ : (,{eg uor}els eleursJ ur{ 0Z '.Igbtt A 'V 'Z16I Jo S 'a,908.92 'S,29}reqs) Jo S nZ + atuoaDS roqualdes€ "€II "92 + V IA puelslSoueH 'druEctno,teg plereH reatJ:O69 puD DuplDD'T€6I 'euefv lJr( '926I llupDlg 'uolduequoN lsn8nv 8Z Jo N wl St :086I )aupnt y J requerdos 9Z Jo iA sefur 9 :ZEO'Iua^o) U '016I g ruIg.t'!e e)]letn.B9gl 'g '/11'Ot6l r6qu0ldes '€A\ollnllsn8nv 9Z'uotplErooJo 's 'It6I.6qur0ldesllDpDlS 'sureld IZ I pu€ eun^ uaat\]rq'.IIL lloltrDlg "41 Iz E.l.\ellnw:z/89 proag 'S 'f '€16I roqopo '.uorlets esnoH uoslqcrnt :,,C),,,D$tauDg '896I fet 'snorrnJ IZ '(HJUAd 1 t 6l tII Jo IANN srlrlu 9.€ ur pesnoq JIE)pautuoxa suaunads p)paps 'sur?ld 1eUro sodolsllrquo ,{ll?nsn'spues,ro11a,( ro peJ ,(lp ur qnrJs ooll?ur-rlrrrf ur ^llu?uruoperd sJnJco sorcodsaql 'pu?lsJ 3o ?H pue ^"9 01 uotplereD luo{ trs"oc eql UDI rrrqlr^\ '"rle.rlsny)rro )r"rls Jo 06 uJelsed\ Jo lse^\-qlnos eq1 (7 arn8rg) : wrlqoLl puD uotnql.ltsre '9I 'uees : u.taqwnu atuosowultC lou sPaas seln,\o snoJounu qlr,{ ro^ o lle peJe^oxDu r/r/ : s.raqlueeql uer{l raq8rq ,{llq8rls Jo rllr,r lo^el + Eru8nseql'8uo[ urru 8 g a/&r :8uol ruur g,Z I sJeqlu€'8uolulru Z I sluelu?lg 'elBu3srq f,11q8rlssriar.riaTs i,uro1le,{ uoplof octJ.rnsJ3uuI eql 'eqnl eql 'rrqtr^\ 'aprstno ot l€nbo + pu? 8uo[ Illru 9 t soqol eql snoJqelS osolu.ruol qluDuad:8uol lxur 6',ro1ef ueplo8 ea ol! se^eel egl ueql re8uol qrnur ^llEnsn '8uo[ ruc g edecs pepr^rpun eldurrs r uo euJtc etBlrtrdBc e a?uaJ 97 '(uc 'reeurl -satolluJ aprx ruur usrll ssal A. ,{11ensn)3uo1 uc ol dn 'alerolqns I 7-9. 1 'luelrnbeA 'snoqcrlsrp ot IEU esr,rJaqlo'esEq eql lE el"JrldnpuoJ 'eAE '3uno,{ qlr,t\ luecserqelS 3ultuoceq seruqeruos ueq,r esolueuol-^oJ8 '9,/a11. sad,oaT 3uo1 urJ 0Z ol dn suolols qtr^\ porlcu?rq ,4lsnoJ3JrloJd ') ( ueoslou :osr :]al/\l '.opti pp!{no'3irp' 'ruosrqcrn141srultunu €rur3r^ uI:uotto]n adtJ 'ut8"rg 'll3nl (y7 ern8rg) (gfgt) ft :8 tr C saprorpll,{lssrr,{tsouo}

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sr pu" 'seqolqlu'ued ,rolle,{ uaploa 'auno,{ u.U^ .r..r.rq[?t*r.rl)J#ti:iln sa^Beloso]u3ruol ,{[esuep'etrq,{ sBq JerJ}oer{J (8:u) proldrp sl pu" 'ec?Ins 'uoeJ8 rouul aq1 uo pornoloJ-ruEarcare qcrq,\\ soqol qluaued'se^eel l?U .4llensn s"g euo ]srxe op "xel lcuDsrp ,{lleluosolllo.lqr pue ,{1pcr?o1oqd:oru o,{1 tsql sel€Jrpur uor8eJ srqt trnoq8noJql suorlepdod;o ,(pnls " 'Je^andoH 'uorlElndod lecqrrc euo oql Jo slueld lueleJrp Suoue pue sl"npr^rpur elSurs urqlr,ra 3ur,(re.t seurleruos 'sJelceJeqccrurouox€l elq?rloJun e.r" e.rnpnJls scueJsejou 'odeqs -ur pu? qfuol Ja^\oU J€e[ ur secuaregrpeseqt t?q1 uon?^rosqo (096I) s.ueeJg luJUuoJ uorAaJ,4eg{Jerts-uo}plBJeD eql ur sfe^Jns plau u.ro,{W August 1975,S. D. Hopper 556,22 5 km E of Binnu on road to Balla and Yuna, 22 August 1975,S. D. Hopper 561;6 9 km E of Yuna, 22 Aueust 1975,S. D. Hopper 559|.28km ESE of Binnu, 22 August 1975,S. D. Hopper 56O'.- 1 km S of GreenoughRiver floodway on Yuna-Tenindewaroad, 22 August 1975,S. ,. Hoppet 558:,4 km NE of Binnu, 22 August 1975,S.D.Hoppet557;9 2kmwof Binnualong West Road, 19August I975, S. D. Hopper 555: Bowes.3l October 1903.A. Morrisotr. Conostylis stylidioides has affinities with both C. candicqns and C. prolilbra- It bears a very close resemblance to diploid hybrids of these species (e.g. S. D. Hopper 138, PERTH) and is most probably a stabilized allotetraploid hybrid oerlvauve. C. stflidioitles can be distinguished from C. cqndicansby its sbort leaves lessthan 5 cm long and I mm wide, while it differs most noticeablyfrom C. prolifera it its tornentose leaves and its golden yellow perianth lobes (the latter character is only discernible in fresh material). C. stylidioides has been found growing sympatrically with C. prolifera at The Loop (Kalbarri National Park) and near Yuna. No hybrids were observedin thesemixed populations.

Nerv Taxa Two new taxa belonging to the C. oculeatq group have been encountered. Both are restricted endemics of the western coastal reqion of the South West BotanicalProvince. All rneasurementsgiven in the descriptionwere taken from Herbarium specimens. Unlessstated to the contrary, all collectionscited are housedat the WesternAustralian Herbarium (PERTH).

Conostylispauciflora Hopper sp. nov- (Figures2,3 and 4) Ab C. aculeataR.Br. spinisfoliorum flexilibus,membranaceis, dense tomentosis, 2 n1nllongis; foliis < 2 mm latis,< 25 cm longis;et floribusper inflorescentiampaucis, difiert. Differs from C. aculeataR.Br. in the flexible, membranousand densely tomentoseleaf spinesless than 2 mm long, in the leavesless than 2 mm wide and 25 cm long, and in the few flowersper inflorescence. Type: Hacienda,20 km S of Mandurah on the Old Coast Road, on the eastern slopesofthe SpearwoodDunes overlooking Harvey Estuary,in Jarrah-Marri woodland, in sandy soil, 1l August 19'76,S. D. Hopper l3l (holo: PERTH, iso: PERTH). f1er6, proliferously branched with stolons up to 15 cm long. Zecres green,equitant at the base, otherwiseflat, ensiform, maximum length l5 cm on average (up to 25 cm); marginal ryrnes flexible, membranous and densely tomentose, rarely more than I mm long. Ittflorescencea capitulate cyme, usually of lessthan 10 flowers,on a simplescape (8) l6 (30) cm high, equal to or fongerthan the leaves. F/ouers yellow, 8 l5 mm long' peria th tomentose outside,slightly hairy within, the lobes4 7 mm long and usuallyexceeding the tLrbe,stqnens uniseriate,filaments 0 5 1.0 mm long, anthers2 3 mm long; style 5-12 mm long, the stigma level with the anthers; placenta covercd all over with nunerous ovules. Seeli not seen. Distribution and habitat: (Figure 2). The south-westof ; Yalgorup-Dawesvillearea ofthe Swan CoastalPlain within 10 km ofthe coast. The speciesis known from only a few localitieson sandyhillslopes of the Spear- wood Dune System,occurring in mixed Jarrah-Marri-Banksiawoodland. Specine exatni ed. Between Lake Presron and Lake CIifIon, Yalgorup National Park, 20 October 1912,S. Pa tt 1361. 2511 692 're^\ou 'r 'g ', Jteq p :JEelSuno{ :Jeelornleu :eurdslEurSreul olers atlles eql tE ore exel eerql aqt Jo sue8ro alqEr€dluoJ lerreleLur{sa4 Jo s8ur^\?rp€prf,nl €rooeJ '^or|ds o.toL[DnDd (J) pue '^ou dss o.tob[!^q dssDtDa]nto (g) 'llenW 'J laptoryq$ ) 'suorlnqrrlsrpJ -Z sultrouo) (.v) Jo ,{SoloqdroruJEel pue sre^\ouJleq IeJrqdErSoaD eln8rC

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'a^U€^r.rep puq,{q pezrTrqels" lceJ ur sr serceds,rau eqt leqt (//61 raddog) oror{^\ -osle pesodoJd sr ! pue'D.tog[nnod 3 o1 ecuelqueseJ SuoJls " rBeq suoJlpuDJ '3 'dss ;o ed,{1ocel?lseoa eql p\te DJoaF)D oqalnco p ;o sprrq,{q euro5 'o,roglicnod 3 JoJ snlels cunods lu?Jre,r 01 laoJI ]uetJgns eJe socuoJeJrp?seqJ '(t eJnArC) EoJB ellr^se,'("( eql ur ,{qJ"eu SurJJncao Dtoaryco'C s1ue1dop ueql sedeJspu? sa^?alre oqs seq otolftcnod sttltsouoS 'acuecseJogurJo peJe,roU-^\eJpu" eeles leurEleu snou?lquou s1ruor.;1redy FigtJf€3. Conottylis pauciflora sp. no't.trolot}'I)e. Both specimensfrom the samecollection- HA 6 and HA 8 are field numbers. r9z 'se^?el Jolorls slr ,{q pue (sorcedsqnsJsqto eql JoJ rur.u q1r,t uosued 'sJJ^roU I I 6 -ruoJ ur 8uo[ ruru S./ EurBeJa^E) ]rllerus s]r ^q osrql uo4 peqsrn8url -srp ^lrp"eJ sr .tI oJDalnJD3 jo serctdsqns Jeqto uro.g e8ue-r lecrqderSoe8 lcuDsrp " serdnJcolnq ol porlle ,{lesolJsr o,togfiotq'dss opalnco srltlsouo3 :(v.Arn)e

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'ocuacseJogur paJe,{\oU-^\0Jeql ol sleJel laqlrde cgrcodseqa uorrerndodDloaN)D.rpunruo",oof,o.lflffl1,f{.,""{Ti"t",l;1i,:#:;}:;!: solqueser l.r?d I?uorlBN daqcuBl eql ur patcelloc (991211 3u1ta167.y -r14 pue plo lLlll'U) unuBqJeH suopJED cru€log ?JJaqueC eq1 ur ueurrceds '(9919 e'oslv y 3a) uor8ar srr{tur papelloc ueer]e,rbq spriq,{q ')-o,tog:rJnrd2clog'g 'J DloalncD SurlqueseJ suerurceds aJurs ,uollossng sE JeJ s" 'C pJalr]\trlos olob[pnod Jo e?ueJu,rou{ eql puelxs f"ru {Jo^\ pleg JaqUnC .Dtoaln)p., ploy[lrnDd.C - ,tq sfeg te^\ eql pu? ^q pJJno^EJ sJoolslllLl_AJpJr{l uJe,idleqoJnlsroru Iros ur sl"rparxJalur tr?lrq?q pordncco 'sorJods ? ,pu? _l"lueJ€d Jr{} ol a rt?ler f1111ra; ueiiod ur uoncnpel lq8rTs " pe.{?ldslp spuqfq (rra ,o/ reddoHJ ,{1r1e:o1auo punoJ uaaq 'dss tE eLeLIDioalnJD otoalnco C puz DJog[lJnDd.J uea^\lJq sprJq^q leJnlBN C. poucif loro C. qculeq t(t - hillrlopes A estuqrine f lqts V

;40 VV V v

v. v AA

AA ^a o ^ iu' t A- . a^- A. 20 .fs"' a aaa .l o'aa

10 20 30 40 50 60 Leof length (cm) Figure4. Maxinum leaflenlath and sten heightof a populationsample ol C.pauciflora sp. nov. and two of C. eculeatassp. aculeat1 (Spearwood Dune hillslopes and HarveyEstuary ffats)from the Dawesvilleregion of the SwanCoastal Plain. Data fron Hopper(1977).

Coliectiorrs of small-flowered, short-leaved of C. aculeata have beer obtained north of the Murchison River (e.g.W of Nerren Nerren, 20 October 1914, J. S. Beard 1l0l), but these difler from C. aculeqta ssp. breviflora rn having unusuallylong marginal spinesand narrower leaves. Their statusmust renain obscureuntil detailedfield studieshave beencarried out. Hybrids of C. qculeata ssp. breviflora and C. cqndicansare known from CocklsshellGully and the Hill River bridge on the Jurien-Cervantes road (Figure 5). At both localities the new taxon occupies lowJying alluv.ial flats wlrile C. c.tnclicansoccLlrs on adjacent drier hill slopes. Hybrids are confinedto narrow zonesof overlap wherethe two speciesgrow intermixed. Specimenscollected l3 km S ofthe Hill River on the JurienBay-Cervantes road (S. D. Hopper ll5) and l8 km SSW ofThree Springs(S. D. Hopper 221) have features intermediate between C. aculeata ssp. bretiflora and C. aculeqta ssp.aculeata, suggesting that thesetwo taxa hybridizewhere their distributions overlap. The subspecificepithet refers to the characteristicshort flowers of the new taxon. 262 t9z

'peoj sslue,{ra] . ,{eg uolrnl 3ql uo oSprrq ro^lx aLIturo]J o.rog:!,1a.tq'dssntrzln)D pue puq,{q D 'suDrlpuDl') nrH J (.!q3 ot I3l) Jo sra.,!\otlJlEq pur se^lal-Luolog ro^rX lll{ aqt rmu ,{E,,!lqirH plr€rg uo qalrp eplspEo.rr? ur 8ur^\or8 ^ou dss D.to{te.tq dss otD,IitcDJ dol t ernSrC Acknowledgements I wish to thank the curators and staff of the following herbaria for allow- ing me to examine their collections: PERTH, UWA, AD, MEL, NSW, CANB, CBG, NE and BRI. Mr. A. S. George provided the Latin diagnosesfor the paper. Constructive comments were also made by Mr. P. G. Wilson and Dr. J. W. Green. I am grateful to students in the Botany 300 course of 1976 (University of Western Australia) for assistancein measuring flowers for the discriminant function analysis. Research for the study was supported by Grant No. 74/692 from the Australian Biological ResourcesStudy Interim Council and by a University of Western Australia Postgraduate Travel Grant. The project was undertaken while the author was in receipt of a Comnonwealth Postgraduate Research Awald and later while in the temporary employ of the Western Australian Herbarium.

References BENTHAM,c. (1873). 'Flora Australiensis'Vol. 6 (Reeve,London). 'Multivariate BLAcKrrH,R. E., and R. A. REYMENT(1971). Morphometrics'(Academic Prcss.London). DoMrN.K. (1912). Additionsto the flora of westernand No h-westernAustralia. J. Linn.Soc.. Bot. 41: 254 283. EWART.A. J. (1906). Contributionsto the (No. 2). Proc.Roy. Soc. vic. 19(N.S.): 33-45. FlsHER,R. A. (1936). The use of nultiple measurementsin taxonomicproblems. Ann. Eugen.,7: 179-188. cEERrNcK,D. (1969). Le genreConostllis R.Bt. (Haemodoraceaed'Australie). Bull. Jard. Bot.Nat. Belg., 39: 167-177. GREEN,J. w. (1960). The genusCotosryltt R.Br. II. Taxonomy. Proc.Linn. soc.N.s.w. ffit 334-373. HoppER.S. D. (1971. Variationand naturalhybridization in the Condrttlttaculeata R.Br. spediesgroup near Dawesville, Western Australia. Aust.J. Bot. 25: 395-411

Publicationdate of Volume2 Number3

The publicationdate of Volume2 Number 3 was4 April 1977.

WIILIAM C. BROWN, GoY.rnnent Plinter, W€st.h Au5trali6