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(006l.liL l9 )€l ,,rr/.frN Nryrr',aVol. 13, No. | (1999)

Thegeneric delimitation of Pithocarpahas been largely stable, with mostauthors accepting it as a separatetaxon. However, Mueller (1893) suggested reducing Pithocarpa to sectionalrank within HumeaSm., a]d.dHeine ( 1967) relegatedPithocarpa to synonymyunder Calomeria Vent. Neither of theseapproaches have been adopted by otherworkers.

Genericr€lationships

The genericrelationships of Pithocarpdiue not clear,and earlier authors appear to haveplaced unduesignificance on the lack of a pappusin the ,a featurewhich is now consideredto have evolvedindependently in a numberof generain the (Anderberg 1991). Lindley (1839) suggesteda relationship between Pilrocarpa andthe ge\!s llaaea Srn.,presumably because of the lack of a pappusin both taxa.The typeof Humeais now consideredto be synonymouswith the monotypicCalomeia, a genuswith which Pithocarpawolold, not appear to haveany closerelationship (Anderberg1991). The lack ofa pappuswas also considered to beofsignificance by Hemsley( 1905), whosuggested that P ithocarpamay be rclated.to ThiseltoniaHemsl., although recent work (Anderberg 1991,Wilson 1992a)points to HyalospermaSteetz or AcomisF. Muell. asthe closestrelatives of Thiseltonia.

Anderberg(1991) included Pithocarpa in his studyof the phylogenyof the ,and suggestedthat it was probablyclosest to taxawithin the broadlydeftned, 'lnwrencel/a complex' (genericlimits ofthe taxawithin thisgroup have since been redefined by Wilson(1992b-e)). These findingswere supported by a preliminaryphylogenetic an alysis of Pithocarpaand a selectionof six of its putativerelatives (infened from Anderberg'swork) undertakenby Lepschi(1997). In that ^nalysis,Pithocarpa appears as sister to Argentipallium niveam (Steetz) Paul G. Wilson,included by 'lawrencella Anderberg(1991) in the complex'las Helichrysum obtusifoliumF. Muell. ex Sond.l. However,as this analysis dealt with only a selectionof thepossible relatives ofPithocarpa, the'true' sistertaxon to this genusmay have been excluded.

Materialsand methods

This studyis basedon examinationof herbariumcollections from AD, CANB, MEL, NSW and PERTH,as well asselected material from CGE (photographs) andK, along with obseruationsmade on liveplantsof all recognizedtaxa. All measurementswere made from herbarium material (reconstituted wherenecessary). Conservation codes used are those adopted by theDepartment ofConservation and LandManagement (see the end of thisissue for definitionsof consewationcodes).

Taxonomictreatment

PithocarpaLindl., Edwards's Bot. Reg. Appendix vols l-23: xxiii (1839).Type: P- pulchellaLindl., lectotype,ldeLewis & Summerhayes( l95l).

Perennialherbs or subshrubs,vegetative parts covered with whitish,cobwebbed indumentum. Leavesaltemzl.s, sessile, entire, usually withered by anthesis;clustered towards the base of themain (reproductive)stems and becoming progressively more distant and smaller up thestems, grading into the outerinvolucral bracts; also densely clustered on very short,lateral vegetatiye seasonal growth lnits. Capitulaeither solitary or in opento moreor lesscompact corymbs, inserted terminally on the ...... ".'"" ElerlsnYuretsel\\ 2.... 'esJoJlu? t Iuelsa,\\-qlnosuI p€eJdseplld sJltq xeldnp el€Aclc qll^dElesd,{t t pue .(mlo :uor8erSuue,tou eql uqlr.,|\ seqJuEJq[erelel uo dre111osse 'ElntId€J 'uodo t IIe,t se 9-Z Jo squ,&ot esool uI EInlIdeJ:ecuJlns IEIXEqP eql uo uoor€u-qsrppal {J€p qll{ pe{lElu slaEJqIl?]JnloAuI stuos lstel lv I """""'quedJo esoFqur,{Jo d ls€e eBueu SuIIJ€d snotqulSelasd,(c pu€ ,{ftAo :uoISeJSuua,\\oU eql u[0l./r\ 'slnlld€t seqJuEJql€Jelel uo ,{rl?1tlos,{larer I I-z Jo squlfuo3paulJsp-lle^\ 'tJuduot sselJo elout ut elnltdec :lueruSldlJep lnoqll^\ slJeJqlDlcnlo^ul I sallalJs^ PUBsalJads ol faY

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2 Style branchesyellowish-brown to brown (in life), drying brown ... P. pulchella var. pulchella 2. Style branchesblack or very dark brownish-black,both in life andwhen dded...... P. pulchellavar. melanostigma

1. Pithocarpa corymbulosa Lindl., Edwards'sBot. Reg.Appendix vols 1-23: xxrii(1839).- Humea corymbulosa(LindL) F.Muell., VictorianNat.9: 144(1893). - Calomeriacorymbalosa(Lindl.)Heine, Adansoniaser. 2: 7 (1967). Type: SwanRiver, WesternAustraba,Toward s.n. (holo: CGE,n.v. (photo seenatPERTH)).

Illustrqtion. Blackall & Grieve (1915:849).

Perennial herb or subshrub 0.7-1 m tall; stems erect, basal portions becoming woody and glabrescentwithage, densely covered with a whitish, cobwebbedindumentum comprised ofappressed to spreadingflexuose hairs to 2.5 mm long. Leaves densely covered with a whitish, cobwebbed indumentum on the abaxial surface,somewhat sparser adaxially; lamina entire, narowly elliptic or nafiowly obovate-elliptic to very naffowly obovate, or very-narrowly ovate to linear-ovate, very naffowly elliptic to linear-elliptic or subulatefor smallestleaves, 2.5-26.5 mm long, 0.25-6.5 mm wide; base weakly amplexicaul, apex acuminate, margins weakly rec:rtrved. Synflorescenceof terminal, more or lesscompact to opencorymbs of 2- I I capitula(these sometimes loosely aggregated to form a larger corymbosepanicle), inserted terminally on the branches,or capitulararely solitary on lateral branches within the flowering region; lateral capitula within a corymb frequently lacking developed florcts. Involucre narrowly ellipsoid to more or less cylindrical when young, narowly obconic to cup-shapedor broadly campanulate(due to spreadinglaminae of the inner bracts) at anthesis,turbinate to cylindrical with a slight medial constrictionin fruit;4.5-6 mm long, 2-3.5 mm wide at anthesis.lnvoLucral bracts 3045; stereomewith a whitish, cobwebbedindumentum (dense on the outerbracts, less so on the innerbracts which alsobear some multicellular hairs to (. 0.1 mm long); outermostbracts 2-3 mmlong,0.4-0.65 mm wide,claw naffowly oblong,laminanarrowly ovate to nafiowly ovate-elliptic, scariousto chartaceous,translucent to straw-colouredwhen dryl median bractssimilar to innermostbracts butto 6 mmlongand 1.3 mm wide,with anelliptic to narrowlyelliptic lamina;innermostbracts 4.2-4.8 mm long,0.6-1mm wide,claw narrowly oblong to narrowlycuneate, lamina naffowly elliptic to oblong-elliptic, chartaceous,white in life and when dry (although bract l apicesin very youngbudsmay be flushedpink). lnvolucral receptacleQ.9-1.2 mm diameter. F/or€b 12-21. Corollqtube 2-2.6 mm long, 0.3-0.4 mm wide at the base,0.4-0.5 mm wide in the rhroat;cells t ofthe adaxialepidermis quadrate to oblong at very baseof tube, becomingnarrowly to Iinear-oblong i above thisi walls more or less straight to slightly undulatefor cells in proximal c. one-half of tube, distinctly sinuousin cells of the distal portion. Corolla lobes O.2-0.4 mm long, 0.15,0.3 mm wide; cells of the adaxial epidermis rounded-oblongto strongly rounded-quadrate(particularly at apex), walls undulateto more or less straight (sometimesdistinctly sinuousat very baseof lobes).,lrazrzas tl exserted0.1-0.35 mm beyondthe corolla mouth, attached0.35-0.5 mm up from the corolla tube base; filaments0.6-0.8 mm long, filament collar 0.2 0.25 mm long; antherthecae 0.8-1.1 mm long, 0.15-0.25 mm wide, anther tails c.0.1-0.15 mm long, anther appendageunpigmented. S/yle 1.5-1.8mrnlong, yellowish-brownto brown (inlife), drying brown, stigmaticbranches (and sometimes 2 distal portion of style) occasionallydarker; stigmaticbranches 0.6-0.7 mm long. Cypsela 1-1.2 mm R long,0.4{.45 mnwidewhen mature,glabrous;cells ofpericarp nanowly oblong, walls straight;cells of testa generally more or less quadrateto oblong (sometimesnarrowly so, oiten rounded), walls P undulateto more or lesssffaight, scatteredflat crystalsalso present,yascular strand running c. three- rt quartersthe length of the cypsela,not passingover the apex;pappus absent. (Figure 1A F) 2. v ['(6f81) lllxx :tZ I :^lo^ ddv ScU log s,prr],rpg'Ipu-Iuou ttno osolnqut.l)o:tDdrDroqttdl

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National Park,Beekeepers Track, 6 June 1993,W. Evans567 (CANB, PERTH); comer Hammond Rd and Bartram Rd, Jandakot,23 Mar. 1960,A.S. George67 | (PERTH); Nicholson Rd, c. 2.5 miles [c. 4 kml N ofForrestdale,T May 1961,A.S. George2350 (PERTH); 7.4 km ENE ofPinjar Rd on Neaves Rd,MelaleucaPark Reserve, T kmWof Bullsbrook,31Mar.1996,B.J. Lepschi &7.R. Lttlly2552 (AD, BRI,CANB,PERTH);3.5kmEofBmndHighwayonMogumberWestRd,c.36kmSSEofDandaragan, 7 Apt.1996,8.J.lapschi &T.R.lnlly2555 (PERTH);c.2l.1kmNWofBP CatabyRoadhouse on Brand Highway towardsBadgingana, 7 Apr. 1996,B.J. Lepschi&7.R. Lally2557 (AD, BRI, CANB, MEL, PERTH);4.9kmWof CockleshellGullyRdtumoffon Coorow-GreenHeadRd, c. 11.5kmENE of Green Head,7 Apr.1996,B.J.Lepschi &T.R. ktl/y2561 (A, AD, B, BRI, CANB, HO, K, L, MEL, MO, NSW, NY, US);samelocality,l luly 1996,B.J.lzpschi&T.R.lnlly2692 (AD, CANB, MEL, PERTH);Bakers Junction Nature Reserve,2.5 km NE of Bakers Junction on road to Jerramungup,2 May 1996, 'on B.J. lzpschi &T.R. ktLLy2581(PERTH);bushland remnantadjacent Kwinana Freeway ramp' at junction with South Street,Leeming, Perth,26 May 1996,B.J. Lepschi &T.R. lttlly 2602 (AD, BRl, CANB, HO, PERTH);7 miles[c. I 1.3km] from Bindoon,26 Apr. 1957, H. Leu,is32(NSW,[K n.v.]); Darling Range, 1843,L. Preiss 27 (MEL).

Distribution. This vadety occurs in the northern part of the range of P. puLchelLa,extending from Eneabbasouthwards to Jandakot(on the southem edge of Perth) and inland to near Mogumber and Clackline. An isolatedcollection (aeprchi & lnlly 2581) alsoexists from nearBakers Junction, north- eastof Albany.

Pithocarpa pulcheLlavar. pulcheLlaoverlaps with var. melanostigmain the Perth region. In this areavar. melanostigmooccurs predominantly on the Darling Scarp,with var. palchelLaon the Coastal Plain, althoughthis distribution is notentirely exclusive.To dateno €xamplesofactual sympatryhave beendocumented, b nt CranrteV 828 from Wanneroomay representa mixed popdationof P. puLchelLa var.pulchella andvar. melanostigma.Thiscollection comprisesplants with both brown (MEL sheet) and black to dark brownish-black(CANB, NSW and PERTH sheets)style-branches. (Figure 2C)

HabitaL Grows in deepsand or sandover laterite,limestone or sandstone,often with some gravelly content.Vegetation communities include low heath(often with emergentmallees), banksia woodland, eucalypt woodland (mainly Corymbia calophylla andlor Eucallprus marginata Sm.), or mixed banksia-eucalyptwoodland, the latter three generally with a shrubby understorey,

Phenology. Populationsnorth ofabout Mogumber tend to beginflowering andproducing new growth approximatelytwo months before more southerlypopulations of this taxon (i.e. south to Jandakot), althoughthere is overlapin phenologybetween no(hem andsouthem . New shootsareproduced around late May (for northem populations),and this continuesuntil approximately November (in southem populations). Flowers have been recorded between December and Apdl for northern populations,and between March andJune for southempopulations. Fruits havebeen recorded between Februaryand June.

Chromosomenumber. n = 13 recordedby Lepschi & Keighery (1999).

Conservation s/drrrs.Widespread and common, and not consideredat risk. Pithocarpa achiLleoides, here consideredsynonymous with P, palchella'var.pulchella, has beenregarded as a poorly known species,and was listed by the Departmentof Conseryationand Land Management's unpublished "Declared Rare and Priority Flora List" of 1996. 'AV)08r,Kqqsv 'peoa 'C n'V'ZL6l AEI 1; dnlureSluo sdnrn8uorodJo S:(HJ.Eg.d)'u's s^ atpuv '106I 'pu"1p11 'tunoulueerD rdv l Jo g :VI'MJS1V N1AJSA ypauluDxa suarurcadspaltalag

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'rqJsda-I tidro2oUttd Jo uorsr^er .IuouoxEt V fg Nu)trid Vol. 13, No. 1 (1999)

CANB, PERTH);Albany Comrnonage,6May 1924,C.A.Gardner 1649(PERTH);41 km W of Rocky Gully on Muir Highway,8 June1977, A .5. George14589 (PERTH); King GeorgeSound, May 1899, B.T. Goad.bys.n. (NSW, PERTH); Toodyay Road, t 20 mile peg, .r. dat., B. Grieve s.n. (PERTH); Kalamunda,19kmEofPerth, l7Apr. 1986,R.&M. Hanilton2l8 (CANB, MEL);top of Nlt Barker Hill at sceniclookout, l2 Apr. 197l, K.F. Kenneallys.n. (PERTH);junction of Albany Highwayand YellanupRoad, c. 15.5km SSEof MountBarker, I May 1996,8.J.Lepschi &T.R lzlly 2576(PERTH); 2.3km ESEof SleeRoad on VasseHighway, c. 17.5km ESEof Busselton,3 May 1996,B.J. Lepschi &7.R. Ially2588 (A, AD, BRI,CANB,HO, K, L,MEL, NSW,PERTH,US); c. l4kmSSEofCapeton roadtoDonnybrook,3May 1996, B.J. Lepschi &7.R. lnlly259l (AD, BRI, CANB, K, MEL, PERTH, US);Darlington, 23 Iune 1949,B. Roarks.n. (PERTH); Mt Clarence,Albany, Mat. 1971,J. Stqers.fl. (NSW); near45 mile pegon BrooktonHighway, 23 May 1966,GNT 191(PERTH);3l .5 km NW of Denmark,27 Apr. 1993,G.J. Wardell-Johnson3339 (PERTH),

Distribution. Disjunctly distributedin the southernpart of the rangeofP. pirlclre1la,The apparentgap in the distdbutionof this taxon may be an artefactof inadcquatecollecting, but this requires conflrmation.Occurs liom theWanneroo Midland areaon thenorthem edge of Perth,south to Augusta, Albany andCape Riche, extending inland to the southernedge of thc StirlingRange. Overlaps with vdr.puLchelLa in the Pertharea (see under that taxon for details).(Figure 2C)

Habitat. Generallyrecorded growing in sand(including sand over granite or latcritc),sandy-loan or loam(occasionally with somegravel content), but oncerecorded from clayover ironstone(Kcigr",") 13334).Frequcntly grows in winter-wetsitcs. Vegctation communities includc heath and other shrubfandformations, mallee-heath, banksia woodland and sucalyptwoodland (mai y Corvabia calophylla awJlorEucalyptus tnarginata), the latter two generally with a shrubby undcrstorey.

Phenology.Data on phenologyis lesscomplete than that available fbr var.pulcheLlcr. New growth is producedfrom approximatelySeptember to November,with flowers recordedbetween January and July,and fruits between March andJuly.

Chronosomenumbel. n = l3 recordedby Lepschi& Keighery( 1999).

Conservation slalrr. Widespfead,not regardedas rare or endangered.This t axonlasP. melqnostigm.tl wasformerly listed on theDepartment of Conservationand Land Maragement's I istof poorlyknown flora asa PriorityTwo taxon(e.g. Hoppcr e/ al. 1990),but wasdeleted in 1992following the location of additionalpopulations (R. Hearn, pers.comlrr.).

Notes.Pithocarpa meknostigrna is here reduced to vadetal rank within P. pulclrcLla. Lewis & Sumnrerhayes( 195 I ) distinguishedP. nelanosligmafrom P. achilleoitles(see above) antJ P. pulchella sers.rtr. on thebasis ofstem morphology (stens slender, ascending and 'rather flexuous'), branching 'much 'flexuous' pattern(plants branchedabove the base), involucral bract number, the natureofthe largest(= median)involucral bract margins, and the colouration of theapical portion of theinvolucral bracts(see below) and the style-branches.Of thesecharacters, only one,styic-branch colouration, is useful as a taxonomic discriminator (see under InfraspeciJicraxa, above). The charactersof stcm morphology,branching pattern and involucral bractmargin areall very subjective,and were not lbund to be taxonomicallyinformative during the courseof this study. Examinationof plantsin the field also suggestthat stemmorphology and branchingpattem arefrequently intluencedby environmental factors (seeLepschi 1997).Colouration of the apical portion of the involucral bractsappears to have been erroneouslyinterpreted by Lewis & Summerhayes( 1951).They describethe involucral bracts 'tipped 'nedium o'iP. melanostigmaas being with very dark brown', asopposed to to iight brown' for 'rrlrrtsnV .srsiqt itolourlriJ to ,{trsr:^run urtrnJ rtlloldrC .trrnplrjStsodpiqsrlqndun 'IuaSotIqd srrJo rfl.u!\isst ur pur (ara.rr:IsV :r?irpqdruD) DltDra\t\I Jo orsr^ .rL ouollt V (1661) tB lq.sdi.l ( qliid .O-L.iurpxJrlrK :rrnttn.u;\, to turruurd.O ul]rl lcnv u s:r^\) l 8j.9 dd I urd ,..uorBrdqtrad eqt to llrol:1,. japuu'I 't f SN N':t ]]iuu:rg g 3Id'dI.r:rtr li\..DN,turxt.rnt{:r1 rrru-rrsv (/86t) S N,riturt ( qu?d :luau3,3nul]npui-I pUlr uol]rr.rrsuo:).to ,trutr

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.rt)sd.-I .I D.troroqtq Jo uorsr^ar rrluouoxet V g Nuttsio vol. 13, No. I (1999)

I-epschi,B.J. & Keighery,G.J. (1999). New chromosomecounts for someWestem Austmlian Gnaphalieae (Compositae). CompositaeNer,,/sletter 33i 19-22, tewis, P. & Summerhayes,V.S. (1951). PithocarpaLil\dl. Kew Bulletin 5: 435+40. Lindley, J, (1839). A sketch of the vegetationof the Swan River Colony. Edwords's Bota ical Regirrcr APp

Marshall, J. (undated)."Wildflowers of the West Coast Hills Region." (Quality Publishing: Perth.) Mueller,F. von. (1893).Notes on the limits of the genusHumea. Viclorian Naturalist 9: 143-144. Puttock,C.F. (1994),Re-analysis of Anderberg'sGnaphalieae data matrix CompositaeNewsletter 25t l-14. Shan, F.A. (1996)."Westem Australian Namesand Their Meanings." (Universityof WestemAustralia Pressl Perth.) "Plantae Steetz,J. (1845). Compositae.lr. Lehmann,J.C.G. (ed.) Preissianae."Vol. 1. pp. 417-490. (Meissner: Hamburg,) Wheeler, J.R. (in prep.). "Flora of the South West ForestsRegion." (Departmentof Conservationand Land Management:Pe h.) Wilson, P,G. (1992a).A new speciesof AcottliJfrom the NorthernTerritory and a new combinatiooin the gedus Thiseltonia(Asteft.e ei Gnaphalieae).Nt/ltsi.a * 419483, Wilson, P.C. (1992b).The classificationof Australianspecies currendy inc]]uded in Helipterumand relatedgenera (AsteraceaerCnaphalieae). Paft 1. Nultsia 8: 379-438. Wilsoo, P.C, (1992c).The I'awrencellacomplex (Asteraceaei Gnaphalieae: Angianthinae) of Australia. Nryffia 8l 361-378. Wilson, P.c. (1992d). The classificationof Austnlian speciescunently inclrded \n Heliptetumand relatedSenera (Astemceae:Gnaphali€ae). Pafi 2. Leucochryswn,Nuytsia 8t 439446, Wilson,P,C. (1992e).The classificationof someAustralian sp€cies currently inclrded in Helipterum^rd Helichrysum (Asteraceae:Gnaphalieae). P?fl 3. Anehocarpa and Argentipallium, two new gencra from Australia lvryl.tio 8: 441-460.