Copyright (c) American Society for Taxonomists. All rights reserved. Delivered by Ingenta to IP: 192.168.39.151 on: Fri, 24 Sep 2021 18:12:04 DOI 10.1600/036364411X553270 © Copyright2011 bythe American SocietyofPlantTaxonomists Systematic Botany carpha left outbecause Dittrich (1977) , inhistreatment ofthe thistles of theCompositae” ( Heywood et al. 1977 ), they foundintheCardueae. Inthe“BiologyandChemistry ates onthecorolla lobes;”bothcharactersare similartowhat in thethickenedrathersmooth exine”and“thelackofstom- on thepollens’“…prominent complexcolumnarstructure (1973) put (1890). There itremained until Stix (1960) suggestedthat of itsowninthereestablished Arctotideae; Hoffman agreed (1873) moved placed Platycarpha ogy andspinynature ofthisspecies. Lessing (1831) described part ofthethistles,nodoubtbecausestylemorphol- (1800) described mining theproper assignmentforthisuniqueclade. Thunberg Platycarpha Robinson 2009 ). been placedinatribeofitsown,Platycarpheae( Funk and based onmolecularandmorphologicaldataithasrecently out dissent( Norlindh 1977 ; Robinson andBrettell 1973 was amemberofthetribe Arctotideae, althoughnotwith- ), but and thepappusismadeupofscales. with smallhairs.Theachenesare smooth,ribbedorrugose are saggitate),andthestyles haveasmallswellingcovered and thecorollas are deeplydivided.Theanthershavetails(or the discoidheadshavefew-to-many, pinktopurpleflorets, subterranean branches.Thereceptacles havenarrow pales, cle. Thetaxaare eitheracaulescent, withoutbranches,orwith by atightclusteringofheadsonthecrown oftherecepta- wagon wheelorare arched. The‘secondaryhead’isformed spread outaround thesecondaryheadlikespokesofa out aerialstems;theleavesusuallylieflatonground and three speciesofthePlatycarpheae are perennial herbswith- Africa;” thetribehasnotpreviously beenmonographed. All the categoryof“oddgenerathatare endemictosouthern The generaoftheCompositaetribePlatycarpheae( The complexhistoryoftheclassification Until recently there wasasinglegenus, 1 U.S. NationalHerbarium,MuseumofNaturalHistory, SmithsonianInstitution,Washington, DC20013-7012,U.S.A. the riseofGreat Escarpmentandclimatechange. November toearlyFebruary. Thebiogeographicpatternisconsistentwithoneofawidespread ancestralspeciesthatbecamefrag andflowersmostlyin August toOctober, and of Namibia,andsouthwesternBotswana;itspeakfloweringtimeisMarch-July. cies are different. has thelargest numberofuniquefeatures andisthesistertaxontoothertwospecies.Thedistributionsfloweringtim etic group onalongbranch: etic group withinthesubfamilyCichorioideaebutnofirmsister-group relationship hasbeendetermined. Thethree speciesform such asleaftype,headsize,styleandcorolla length,andpollentype.MorphologicalmoleculardatasupportPlatycarpheae formed bytightlyclustered headsonthecrown oftherhizome. Thethree speciesare easilyseparatedfrom oneanotherbasedon traditionally beenplacedinasinglegenusbutwere recently dividedintotwo( Less.and Keywords— Abstract— Platycarpha A oorp o te ml Tie ltcrha (opste ) (Compositae: Platycarpheae Tribe Small the of Monograph A shouldbeintheMutisieae. Robinson andBrettell basedontheThunberg . De Candolle (1836) and Platycarpha (2011), 36(1):pp.191–208 Platycarpha The tribePlatycarpheaehasthree species;theyare foundfrom South Africa, toNamibiaandBotswana.Thethree specieshave Platycarphella Platycarphella seaee, Acoiee, boegah osaa, Get sapet, Nmba, ple ot Africa. South pollen, Namibia, Escarpment, Great Botswana, biogeography , Arctotideae , , Cynara glomerata 2 South African NationalBiodiversityInstitute,PrivateBagX101,Pretoria 0001,South Africa intheVernonieae. Bentham andHooker Platycarphella carlinoides backintheCardueae (thistles)based from theVernonieae intoasubtribe Platycarphella carlinoides ) reflects thedifficulty ofdeter- V. A. Funk&H.Rob.)fallinto thusindicatingthatitwas isthemostwidespread ofthespeciesandgrows innorthwesternSouth Africa, thecentralhighlands . . Funk, A. V.

3 Author forcorrespondence ( [email protected] ) Platycarpha, Platycarpha Platycarpha Cmuiaig dtr Vcoi Sosa Victoria Editor: Communicating n and Platycarphella parvifolia 1 Platycarpha glomerata , andit 3 Platy- (now n aid Koekemoer Marinda and was 191 treatment ofthe Arctotideae excluded be placedbackinthe Arctotideae, and Norlindh (1977) inhis (Cardueae), rejected the analysisof thecpDNA data placed itnearpartsofthe as thesistergroup totheLiabeae- clade. However the ITSanalysesproduced results thatplacedPlatycarpheae in theanalysesofnuclear and cpDNA.Thecombined ment ofPlatycarpheaewithin thesubfamilydiffered slightly unpubl. data)withinthesubfamily Cichorioideae.Theplace- ( Funk etal.2004 ; three speciesformamonophyleticgroup onalongbranch Funk andChan2009 ; FunkandChan family Cichorioideae. that wouldlinkthissmalltribetoanyothergroups inthesub- according to Wortley etal.(2007) thepollensuggestsnothing tacle, theachenesare notdistinctlyribbedorwinged,and yellow, red ororange,theyhaveslenderpalesontherecep- there are norayflorets, theirdiscflorets are pinkorpurplenot tribes: theyhavenolatex,theinvolucralbractsare notfused, characters thatwouldplacethemineitherofthetwosub- 2007 ) accepted recent classificationsofthe family/tribe( Bremer 1994 ; Karis and Cardueae. appearance isdifferent from thosefoundinthe Arctotideae minute hairsisfoundinmembersofthePlatycarpheaebut its it isfoundinsomethistles(Cardueae). A smallswellingwith found inthelarger ofthe twosubtribes,theGorteriinae,and in othercladesthefamilyorare plesiomorphic)butitisnot other charactersthathavebeensuggestedare eithercommon hairs locatedjustbelowthebranchpointonstyle(all was basedonthepresence ofaswollenarea witharingof Bentham (1873) thetraditionaldefinitionof Arctotideae ysis ( Funk etal.2004 ; Funk andChan2009 in thetribePlatycarpheae,havebeenremoved). Beginningwith from theanal- This istrue evenafterseveralgenera,includinggeneranow Arctotideae doesnothave muchsupportforitsmonophyly. 2000 ; Leistner 2000 ; Funk etal.2004 ) haveshownthatthetribe al. 1992 ; Robinson 1992 1915a , , 1994 b ; ; Robinson andBrettell 1973 Bremer; 1994 Norlindh 1977 ; ; Karis et Herman etal. ‘unassigned tosubtribe.’ The analysisoftheDNA sequencedatashowed thatthe Data from avarietyofsources ( Cassini 1816 , 1821 ; Beauverd share more charactersandare sister-taxa; isfrom inlandeasternSouth Africa andusuallyflowersfrom Platycarphella parvifolia Platycarpha Platycarpha Platycarpha and 2

Platycarpha Platycarphella isfoundinnorthcentraltonortheast inthe Arctotideae butlisteditas and andsuggestedthatitshould ). All have‘secondaryheads’ Platycarphella Platycarpha Platycarpha glomerata es ofthethree spe- asamonophyl- amonophyl- alsolackthe characters mented by . Themost

Copyright (c) American Society for Plant Taxonomists. All rights reserved. Delivered by Ingenta to IP: 192.168.39.151 on: Fri, 24 Sep 2021 18:12:04 either isto Funk &H.Rob.,are more closelyrelated tooneanotherthan Hiern) V. A. Funk&H.Rob.and that (Funk andChan,unpubl. data)issuperimposedonthe distribution; itshows Platycarphellaparvifolia Platycarphellacarlinoides and collections atPRE. mountingsolutionortakendirectly from thespirit slides usingHoyer’s [Volume 36 examination, floralpartswere rehydrated andmounted onmicroscope were supplementedbyinformationfrom theliterature. Formicroscopic kindly sentbythoseinstitutions.Thedataderivedfrom thespecimens and informationimagesoftypespecimensfrom BM,K,andUPSwere collections atK,M,MO,S,andUS.Imagesofspecimensfrom MandS cies andtheextensivecollectionshousedatPREWIND,smaller patterns ofsouthern Africa. the distributioninlightofgeologichistoryandrainfall members anddefinethegeneraspecies.We alsoexamine len, isdifferent from theothertwospecies. SYSTEMATICstyle branchesandcorollas, thickrhizomes, andunique pol- BOTANY P. glomerata and Chanunpubl.data).Thismatchesthemorphologyinthat that thetwospeciesof ships amongthecladesofsubfamily. are beingaddedinanattempttobetterresolve the relation- them inthebootstrapanalysis.Newmoleculardataandtaxa maximum likelihoodtrees, there waslittleresolution among mothamneae, Arctotidinae, Arctotideae-Gorteriinae, Platycarpheae,Ere- subfamily (Cichorieae,Vernonieae, Liabeae, Arctotideae- Arctotideae. Although theindividualcladeswithin 192 Fig. The morphologicalstudywasbasedonnewcollectionsforallthree spe- In thistreatment wedocumentthecharactersthatunite Within thetribePlatycarpheaeDNA datashowclearly Platycarpha glomerata Platycarphella 1 1. Map ofsouthern Africa showingthedistribution of Platycarpha glomerata withitsspinyleaves,larger heads,longstyles, Heterolepis (triangle= aeil ad Methods and Materials (S.Moore) V. A. Funk&H.Rob.Thephylogeny (Oliv. &Hiern.)V. A. Funk&H.Rob.;square = isthe sistergroup oftheothertwospecies. ) were present inallparsimonyand Platycarphella, P. carlinoides Platycarpha glomerata Tub) esn (Fg 1 ; Funk 1; (Fig. Lessing (Thunb.) P. parvifolia (S.Moore) V. A. Less.; circle = (Oliv. & Platycarpha

Koekemoer 12564 in asummerrainfallarea. We collectedthisspecies( appears tohaveawinterrainfall floweringpatternbutsits rain inSeptembereitherof thesebioregions. Thespecies 2006 ). According to Rutherford etal.(2006) there isalmost no cent EasternKalahariBushveld(savanna)( Rutherford etal. falls withinthedryhighveldgrasslandandonadja- and 90%ofthemfrom August toOctober. Itsdistribution to Octoberwithoverhalfofthecollectionsfrom September South Africa andithasbeencollectedinflower from July most oftheyear. species simplygrows andflowersafterarainthroughout sible that,especiallyinthesouthernpartsofitsrange,this ( Mendelsohn etal.2002 ; Okitsu 2005 ; Kundell 2007 time andtherainsare unpredictable, especiallyinthesouth); itispos- In reality, mostofthedistributionrangeisdry ter) butexceptionsoccurespeciallyat23°latitudeandsouth. out across theyearbutmore frequently Mayto August (win- southern partoftherangemore likelytobefoundspread July (latesummerearlywinter)timeframeandthoseinthe being more consistentlyand frequently foundintheMarch- dictability ofrainfallwiththoseinthemore northernareas that there issomecorrelation withthelatitudeandpre- Examining theherbariummaterialbycollectiondateshows predominantly insummer andautumn(February to April). annual rainfallismore unpredictable; below150mmand to 350mmwhileinthesoutherndrierdistributionarea the January toMarch andthe averageannualrainfallcanbeup northern distributionarea thesummerrainsfallmainlyin winter rainfall. According to Mendelsohn etal.(2002) inthe time followsthesparserainsoflatesummerandearly from March toMay(autumn; Fig. 2 the collectionswere made from March toJulyandover60% ). Thisbroad flowering around, exceptforNovember, andDecember, but80%of atic.” Thisspecieshasbeencollectedinflowernearlyyear tics andpatternsbasedonpointmeasurements are problem- often difficult touseand estimatesofrainfallcharacteris- statistical descriptorssuchasmeanandevenmedianare Sahara Desert. According to Kundell (2007) “Conventional Within Africa theclimateissecondinaridityonlyto erratic rainfall;92%ofthelandishyper-arid orsemiarid. 2007 ). ThePlateauhashot,dryconditionswithsparseand of whichare partoftheGreat Western Escarpment( Kundell Plateau andcomprisesmountainshighlandareas, all plateau region thatisacontinuationoftheSouth African Botswana. Mostofthecollectionsare ontheNamibianinland ( Rutherford etal.2006 ; Fig. 1 ) inSouth Africa, andinnearby and southernNamibia,theKalahariDuneveldBioregion carlinoides the degree square cell. brackets thatidentifiestheNWcorneroflongitude(S)latitude(E) specimens examinedthesedegrees squares are identifiedbyanumberin tion map( Fig. 1 ) isdividedintogridsquares basedononedegree, inthe quarter degree gridsquares ( Edwards andLeistner1971 ). Thedistribu- WIND, are partofaconsortiumthathasdividedsouthern Africa into ning oftherains. rainy seasonand sothisspeciesflowersjust before thebegin- ary floodplainnearariver. Maybetheriversfloodduring Platycarphellaparvifolia Current DistributionandRainfallPatterns— The herbariawiththetwolargest holdingsofthisgenus,PREand iswidespread onthehighplateauincentral ; ; Koekemoer andFunk2866 Discussion occursinthenorthcentralpart of ) onadrysecond- Platycarphella Funk and Copyright (c) American Society for Plant Taxonomists. All rights reserved. Delivered by Ingenta to IP: 192.168.39.151 on: Fri, 24 Sep 2021 18:12:04 and westofthedistribution cle focusedontheCapeFloralRegion(CFR)whichissouth with PeterLinderandRichard Cowling).The2003Linderarti- was takenfrom thesetwosources (alsofrom emailexchanges Partridge (2000) and Linder (2003) andmuchofwhatfollows evolution ofthesethree species. seems unlikelythatthebedrock hadmuchinfluenceonthe of thesedatesare olderthantheCompositaefamilyandit younger intheKaroo Supergroup, around 200—300mya. All Rubidge 2005 ). Bedrock for Supergroups, whichdatebackto2,600mya( McCarthy and tions oftheVentersdorp, BushveldComplex,andTransvaal The Damara Supergroup andGariepComplex(850–600mya). Supergroup (300–180mya),NamaGroup (600–543mya), varies from theKalahari Group (70myatopresent), Karoo cates thatthebedrock inwhich 2009 ). ThegeologyofNamibia( Mendelsohn etal.2002 ) indi- the closestrelatives was around 45myaorless( Funk etal. of theCompositaeanditssistergroup theCalyceraceaefrom Africa— classified asthewetsummerrainfallarea-coastal. corresponds tothepeak rainfall timeofthearea sometimes of thedatesavailableare from Novemberthru February. This been collectedinflowerfrom OctobertoMaybutover80% face andinthe Albany Thicket( Rutherford etal.2006 ). Ithas eastern edgeofthesubescarpmentgrassland/savannainter- 193 Eastern Cape( Fig. 1 ). Itoccupiesanarea onthesouthernand South Africa ininlandareas ofKwaZulu-Natalandthe tions thatwere gathered duringthattimeoftheyear. FUNK AND KOEKEMOER:PLATYCARPHEAE 2011] tions are ontheedgeofrangedistributionforspecies. conditions were establishedinsouthern Africa. A majorfeature 10 mya(Miocene-Pliocene boundary)the summer-drought aridity worldwideandthere isevidencethatapproximately It seemsclearthatbythemiddle Miocenethere wasseasonal other elementsoftheflorawere present athigherelevations. Cape duringtheMiocenebut there isnowaytodetermine if sil evidencethatthere wasanowextincttropical florainthe the drycoldphaseofOligocene (33–23mya).There isfos- Eocene climateswere warmandhumidthiswasfollowed had aneffect onthemore centralandeasternhabitats.The but neverthelesseventsthataffected theCFRwouldalsohave Climate changehasbeenrecently discussedby Tyson and Geological HistoryandClimaticChangesofSouthern Platycarphaglomerata Fig. P. parvifolia 2 2. Flowering timesofthethree speciesofPlatycarpheaebasedonherbariumspecimensexamined.Theboxcontainstheperc It hasbeenestimatedthatthesplitofancestor distribution area includessomeolderforma- occursinthesoutheasternpartof Platycarpha glomerata Platycarphella carlinoides Platycarpha and Platycarphella parvifolia Platycarphella s much is grows

has2collectionsthatwere madein April bydifferent collectors.Bothcollec- P. parvifolia. and thiswould matchwiththeseparationof Restionaceae showingacommon autumn-springdistinction in sistertaxaand Linder (1995 , 2001 ) hasdocumented thisin September (spring).Differences inphenologyare oftenfound (autumn toearlywinter)and mentioned above, the Highveldandevenonto some oftheKalaharisands. As ment inNamibiabutdriftsdowntheeastslopeandon to Platycarphellacarlinoides was eitherseparatedormigratedtotheNama-karoo biome. ond oldestbiomeofthethree. Inthenorthwest inhabited awidearea onthehighveldgrasslands,sec- tor of the uplifttoexplainsuchapattern.Meanwhile,ances- phylogenetic positionof cies. Sothere are manyfactorsthatcanbeusedtoexplainthe blooming patternthatseparatesitfrom theothertwospe- corresponds tothefactthat ent, withthehighveldhavingacolder, dryerwinter. This the basalpositionofthisspecies.Theclimateisalsodiffer- the three habitats ( Cowling 2005 ) whichfurthersupports grassland ofthesubtropical thicketpatches)istheoldestof The Maputaland-Pondoland-Albanybiome(shrubland/ tral taxonwhichwouldhavebeencutoff intheHighveld. tion oftheancestor this escarpmentmayhavebeenresponsible forthesepara- tropical Lowveldarea justeastoftheCFRsouplifting species, isfoundsouthoftheGreat Escarpmentinthesub- in thewinter. is more oceanicwhilethe interiorplateauiscolderanddryer ( Linder 2003 pp.617–618).Theclimate ofthecoastalplatform succulent Karoo (leeward side)wouldhavebeenenhanced” it “…wetterwhilesummeraridityinNamaqualandandthe the rainfallpatternsineasternpartofCapemaking near thecoast.TheupliftingofGreat Escarpmentchanged effectively cuttingoff thenarrow bandoftheLowveldarea north onboththeeastandwestsidesofsouthern Africa and The mostrecent uplift(ca. fivemya)isU-shapedstretching in theNeogene,firstca.20mya,thenagainPliocene. to Gondwana.TheGreat Escarpment hasbeenelevatedtwice of southern Africa istheGreat Escarpmentwhichdatesback Speciation— Platycarphella carlinoides

Platycarphaglomerata P. carlinoides P. glomerata is found mainlyontheescarp- P. glomerata P. glomerata P. parvifolia n and flowersfrom March toJuly from therest oftheances- , thesistertoother ; Linder (2006) hasused P. parvifolia hasasummerwet flowers mainlyin P. carlinoides entage ofthecollec- couldhave P. carlinoides and

Copyright (c) American Society for Plant Taxonomists. All rights reserved. Delivered by Ingenta to IP: 192.168.39.151 on: Fri, 24 Sep 2021 18:12:04 in characters suchasthenumberofflorets perhead(especially plant withouttearinguptheherbariummaterial.Forinstance, difficult todeterminetheexactsizeornumberofparts mary headandthecorollas are usuallylavendertopink.Itis heads are discoid,havethree topossibly100florets perpri- viduals are closetogetherandtheleavesarch upward. The secondary headare flatontheground, however, someindi- the crown oftherhizome. Usuallytheleavesaswell line secondaryheadformedbytheclusteringofheads on areas withrocky andsandysoilinfullsun. discussed above.Itsmembersoftengrow inopendisturbed the distributionsofthree speciesisfoundin Fig. 1 and They are foundinlowelevations upto1,800m;amapof Mpumalanga, NorthernCape,andNorthwestProvince). South Africa (EasternCape,Free State,KwaZulu-Natal, carpheae are nativetocentral andsouthernNamibia known chromosome counts. dark. Pappusof7–12persistentwhitescales.There are no end; antherspurple;styleslavender. Achenes 3or5sided, surface ofapexeachlobe;tubessparselyhispidatdistal nearly white,varyinginsize,lobeswithglandsonabaxial bisexual; corollas purple, mauve,lilac,orpink,occasionally gins entire, apicesacuminate. Florets 3–60(possibly100), more slenderwithinnermost bractsresembling pales;mar- 6–20×1–5mm,innerbecomingsmallerand 7–40 in3–5series,lanceolatetolinear, usuallyglabrous, outer more orlessfusedintothe crown. Involucralbracts(phyllaries) ary headonthecrown of the rhizome; headsfirmlyattached, Inflorescence sessile,few-many headed,grouped inasecond- mostly glabrous, adaxailly withdense,whitetomentum. in thelowermostringandlarger; bladesabaxiallygreen and radiating from acentralcrown, in2–6ringswitholdestleaves separate from oneanother;nomilkysap.Leavesprostrate, forming matsanddensecoloniesortheindividualscanbe 9 SSEAI OAY [Volume 36 SYSTEMATIC BOTANY P be thatthelineageisoldbutspeciesare recent. and longrelative tothebranchesofthree species, itcould branch forthetribeisstrongly supportedasmonophyletic two sistertaxa because ofchangingrainfallpatterns,andevolvedintothe western populationswere isolatedfrom oneanother, perhaps Platycarphaglomerata ment separatedthesoutheasternpopulationswhichformed ancestral taxon.Themostrecent upliftoftheGreat Escarp- 194 ( collections were examinedindetailandherbarium specimens 2696 erata three collectionsinspirit,oneforeachtaxon: of theindividualsaspecies. Fortunatelywehadavailable heads andlengthofthepales cannotbedeterminedforall be countedexactly. Likewisethediameterofindividual Platycarpha glomerata latycarpheae The speciesare easilyrecognizable becauseoftheiracau- Habitat— and Distribution Perennial herbs,prostrate, acaulescent,rhizomatous, Based onthesedataonecanhypothesizeawidespread Platycarpha glomerata ( Less. 47: 24–27,2009—TYPE: ), and Koekemoer 3466 P. parvifolia Platycarphella carlinoides V. A. Funk&H.Rob.,CompositaeNewsletter aooi Treatment Taxonomic . Subsequenttothiseventthenorth and ( n and ) ), Koekemoer andFunk 2866 ) andthenumberofinvolucral bracts Platycarphella carlinoides Platycarphella parvifolia Platycarpha glomerata The three speciesofPlaty- n and Platycarpha glom- P. parvifolia ). Thesethree ( (Thunb.) Koekemoer cannot ) . The . distal ends. of thickenedcolumellaethatare prominently branchedatthe inner columellaelayerisover 2µminthicknessandconsists fine columellaeandisapproximately 1.2µminthickness; the by multilayered internaltecta.Theouterlayerconsists of tectum comprisedoftwodistinctcolumellaelayersseparated poles. Instructure thepollenisecaveate( Fig. 3 ) withaninfra- apertures are tricolporateandoccasionallysyncolpateatthe tributed overasmoothandminutelyperforatesurface.The 3µm across theirbasesandwithmore than100spinesdis- echinate withspinesapproximately 1µminheightand2.5– greater detailelsewhere ( Wortley etal.2008 and 3) ), thepollenis ( Fig. Berkheya been isolated.Thelatterhavesofaronlyreported from Vernonieae where sofarnoditerpenesorthiopheneshave however, excludesaplacementofthesetwogenerainthe may besupportedbytheco-occurrence ofditerpeneswhich, they pointedoutthattheproposed relationship to these compoundsa4-methylgroup ismissing.Furthermore, aurene derivativesandtheir glycosidesare present thoughin to in thediterpenes Cardueae. Thehighconcentration andthedegree ofvariation posed positionof lide 17.Theyconcludedthattheresults supportedthepro- ent-16 4–10, thenorkaurenes 13–15, thethiophenederivatives16, P. carlinoides Arctotideae andCardueae. Theirexaminationofaerialparts as anindicationthatthegenusisintermediatebetween some germacranolides.”Thesefindingscouldbetaken gave inadditiontothiophenacetylenes,typicalof reported that“Thechemical investigationof lated from aerialparts.In Zdero andBohlmann(1989) they were isolatedfrom roots andtwogermacranolideswere iso- contrast tothat of noides buttwospecieshavebeensampled, can spread overalarge area bymeansoftheserhizomes. ondary head.Thus,onceanindividualhasbeenestablishedit head andgrow somedistancebefore developing a newsec- other times,arhizome willformfrom belowanold secondary ing, anewcrown canformwithanothersecondaryhead. At old heads.Inaddition,onceasecondaryheadispastflower- are inflowerthere are newsmallheadsformingatthebaseof any onesecondaryheadare ofdifferent agessowhilesome in thesurfaceofcrown. Thecrown iswoody. Theheadsof crown. Theheadsare strongly attachedandevenembedded are produced atthenarrow circular perimeterofthe of thecrown orontherhizome nearthesecondary heads,the crown oftherhizome withtheroots emanatingfrom thebase growth pattern.Thesecondaryheadsare formed bythe the spiritcollections. taxon couldbefoundinthepopulationalvariation of thevariationfoundamongdifferent collections ofeach were compared tothemestimatethefinalnumbers. Most reported that The minutelyperforate exinesurfaceconsists ofdistinctbut The pollenof Pollen— Chemistry— Perhaps themostinteresting aspectofthisgenusisits Atractylis n and b -hydroxy-kaurane-19-oic acid andthegermacrano- specieswhichalsocontaingermacranolides. Platycarpha glomerata The pollenmorphologyof (tribeCardueae, subtribeCarlineae)where nork- P. carlinoides gavetheent-kaurene derivatives1,2,3aand P. glomerata Little isknownaboutthechemistryofthis Platycarpha glomerata Platycarpha Platycarphella parviflora P. carlinoides ( Fig. 3 ) isidentical. As describedin hadfivethiophenesandadiolthat betweenthe Arctotideae and . Bohlmann andZdero (1977) mayindicatearelationship Platycarphella parviflora ( Fig. 3 ) isinmarked and Platycarphella carli- P P. glomerata . . Corymbium carlinoides Berkheya , .

Copyright (c) American Society for Plant Taxonomists. All rights reserved. Delivered by Ingenta to IP: 192.168.39.151 on: Fri, 24 Sep 2021 18:12:04 with permissionfrom Funk etal.2009 ] k. lateral;l.fractured grain.Scalebars:whole grains=10um;fractured grains=1um[Figure prepared byJ.Skvarla; someof tured grain; e-f, 195 FUNK AND KOEKEMOER:PLATYCARPHEAE 2011] Fig. 3 3. Scanning electron micrographs of Platycarphella parviflora : e.polarview, f.apertural,g.lateral; h.fractured grain;i-l. Platycarpha and Platycarphella pollen.a-d Platycarpha glomerata Platycarphella carlinoides: : a.polarview, b.apertural,c.lateral,d.frac- theimagesare reproduced i.polarview, j.apertural, Copyright (c) American Society for Plant Taxonomists. All rights reserved. Delivered by Ingenta to IP: 192.168.39.151 on: Fri, 24 Sep 2021 18:12:04 Platycarpha glomerata ophate polleninotherCichorioideaegroups. Species1. grains echinateandapproaching thestructure oftheechinol- “bow tie”fusedacross the pore dividingitintotwopores; ous around eachcolpus, bow-tie-shaped, narrowest pointof nar areas, grainsare prenanthoid, withfusedmuracontinu- distinct butirregularly arranged ridgesseparatedbylacu- scales. Pollenwithminutelyperforateexinesurface rounded apex,glabrous. Pappusofnonoverlappingwhite [Volume 36 part ofstyleshaft;branchesnarrowing toaslendersomewhat branches 3.5–5.0mmlong,longerthanthepapilloseupper heads with20–60(–100)florets; corollas 20–23mmlong;style lobes curvingupwards. Inflorescences ofsecondaryheads; blades ellipticupto30cmlong,margins deeplypinnatisect, Leaves prostrate inrosettes, radiatingfrom awoolycrown, rescences, proliferating bystoutrhizomes; withoutlatex. Platycarpha SYSTEMATIC BOTANY 1. Leavesentire, dentateorrarely pinnatisectbutwithoutspines,abaxialsurfacesmooth;primaryheadssmaller, 3–4mm 1. Leavespinnatisectwith2mmspinesatapexofeachlobe(spiny),abaxialsurfacerugose; primaryheadslarge, 10–25m in two pores. Spinesandspinebasesare somewhatgreater than bowtie-shaped, atwaistfusedacross thepore dividing itinto nanthoid, withfusedmuracontinuousaround eachcolpus, in theVernonieae andGorteriinaebutthegrainsare pre- somewhat resemble thelophatepatterncommontopollen irregularly arrangedridgesseparatedbylacunarareas that 196 series; outerbracts broader atthebase,15–20 ×3–7mm,straw Involucral bracts, ca.25–40,lanceolatetoovate orlinearin4–5 fused tothecrown surfaceandembeddedwithinit( Fig. 6A ). discoid, subglobose,10–25mm indiameter, baseofhead ary head3–10cmindiameter; headsvaryinginage.Heads secondary headformedonthe crown oftherhizome, second- tomentum. Inflorescence white sessile,ca.3–20headsgrouped ina surface adaxial 6A); glabrous withage,abaxialsurfacecovered withdensewhite D,5 4B, (Figs. tomentose whenyoung,becomingadeepglossygreen and spines long mm 2 in sizetoward proximal end,toothed,spiny;apiceswith tisect, lobescurvingupwards, alternate,broad, diminishing to 2–3cmwhere itmeetsthecrown, margins deeplypinna- 2–30 ×0.4–3.5cm,inlarger leavespetiole-likebasebroadens lying flatontheground orarching upwards; bladeselliptic, Leaves prostrate, radiatingfrom silky-woolycrown, either to ca.6cmindiameter),fleshy, forming matsordensestands. Perennial, rosettiform, acaulescentplantswithsessileinflo- Perennial herbs,prostrate, acaulescent,rhizomes stout(up P 2. Larger, lowerleaves8–20cmlong,margins entire, dentateorrarely deeplyandnarrowly pinnatisect;florets 3–11 (fre 2. Larger, lowerleaves1.2–6.0cmlong,margins entire, somewhatundulatingorminutelydentate;florets 5–20;heads1–20 . . 1831. glomerata sheet no.18566,S(2);scannedimagesatUS). no. 18567,scannedimageatUS:Isotypes:UPS-herbarium December, TYPE: SOUTH AFRICA. EasternCape:ZwartkopRiver, Stobaeaglomerata apssae o vrapn,mrisetr . pappus scalesnotoverlapping,margins entire 20–100 florets; corollas 20–23mmlong;stylebranches3.5–5.0inlength,muchlongerthanpubescentupperpartofsha clsoelpig agn reual ett . scales overlapping,margins irregularly dentate 3–14 (20)florets; corollas 8–13mmlong;stylebranches1orlessinlength,shorterthanpubescentupperpartofsha parviflora heads many(uptoca.400);secondary2–10cmindiameter . ed – mi imtr . heads 1–2cmindiameter Cynara glomerata Tub) Less. (Thunb.) Less.,Linnaea6:688.1831.—TYPE: and Thunberg s.n. P . . (Thunb.)Spreng., Syst.3:394,1826.— carlinoides (Thunb.)Lessing,Linnaea6:688, Thunb.,Prodr. Pl.Cap.141,1800. (holotype:UPS-herbariumsheet . Theapertures are tricolpo- Key totheGeneraandSpeciesofPlatycarpheae Platycarpha thy inthelacunarareas. weaker) andlesscomplextectum;thisisespeciallynotewor- branches appeartobothsupportaswellformafiner(i.e. larly thickenedanddistallybranched( Fig. 3 ); however, these P it formsdense matswithprostrate leaveshowever inmore pers. comm.).It hastwogrowth forms,in someopenflatareas infrequently probably becauseof itsspinyleaves(N.Barker, common inmanyareas oftheEasternCapeitiscollected lawns (Bathurst;Koekemoer, pers.comm.)andalthoughitis town itisreported tolinethesidewalksandcreep intothe stony orsandysoils( Goldblatt andManning2000 ). Inone cially disturbed,areas andalongdirtroads, growing in previous discussionand Fig. 1 ). Itisprevalent inrural, espe- Natal andtheEasternCapeatelevationsbelow500m(see the easternpartofSouth Africa ininlandareas ofKwaZulu- above theothers,are smallerthantheolder, lowerleaves. available. Onlarger plantstheyoungerleaves,whichare just depending ontheageofplantsandamountwater conditions. Thesizeofthesecondaryheadandleavesvaries what uprightpositionespeciallywhengrowing incrowded prostrate, thisonealsohasatendencytobeforced intoasome- developed involucre. Whileallofthespeciestribeare and longstylebranches,itslarger headswithawell- cies inthetribebyitspinnatisect,spinyleaves,longcorollas gins nearlyentire, apexacuminate. Figures 4–6 . persistent, whitescalesofvaryinglengths,4–6mmlong,mar- of unequalwidths,ribbed,glabrous. Pappusof7–10linear, oblong, ca.5mm,whitewhenimmature turningdark,5sides narrowing toaslendersomewhat rounded apex. Achenes branches allthewaytoapex,style3.5–5.0mm, stiff hairsfrom baseofswollenarea alongbacksideofstyle 1 mm,swollenarea justbelow thestylebranches,veryshort 1 mmlong,filamentsca.5–6mm;styleslavender, 22–28×< purple, subequallysagitateatbase,apicalappendagesacute ( Fig. 6D ); stamensca.13mmlong,thecae4–6dark ing underwithglandsondistalendofabaxialsurfacelobe 1–1.5 mm,glabrous, lobes 3.0–3.5mmlong,recurved andcoil- long, tubes14–17×<1mm,sparselyhispid,throats 6.0–6.5× corollas purpletopink-purple, fadingwithage,20–23mm slender, ca.20–60(counted)orpossiblyupto100(estimated); for apexwhichissomewhatstiff, glabrous ( Fig. 6C ). Florets overtopping theimmature florets, ephemeral,soft,except pales. Pales(receptacular bracts) linear, ca.10–15mm×<1 slender andapexlessstiff withinnermostrow resembling with 2mmspinytip;innerbractsbecomingsmallerandmore colored, stiff, glabrous, entire, apicesacuminateandreflexed, region. There issomestructural similarityto rate withsurrounding ridgescommonlyobscuringthepore . . itiuin n Habitat— and Distribution Platycarphaglomerata carlinoides ...... inthatthelower(inner)columellaeare simi- . 2 . canbedistinguishedfrom otherspe- . Platycarphaglomerata m indiameterwith in diameterwith quently 3–4); ; secondary Platycarphella carlinoides Platycarphella parvifolia ft; pappus Platycarpha glomerata P ft; . . parviflora cus in occurs and

Copyright (c) American Society for Plant Taxonomists. All rights reserved. Delivered by Ingenta to IP: 192.168.39.151 on: Fri, 24 Sep 2021 18:12:04 01 FN N OKME:PAYAPEE 197 FUNK AND KOEKEMOER:PLATYCARPHEAE 2011] by aclusteringof heads onthecrown ofthe rhizome. [Allphotosby M. Koekemoer(PRE)] deeply dividedand rolled; D.Closeupoftwo secondaryheads,notethatthedelimitation oftheprimaryheadsis obvious.Seco tion ofthepinnatesect ,notethespinosemargins andtherugose surface,C.Secondary head,notethelongstylesand Fig. . Iae o of Images 4. Platycarpha glomerata Less. A. A populationoftightlypackedplants, notethattheleavesare arching upwards, B. An enlarged por- long corolla lobesthatare ndary headsare formed Copyright (c) American Society for Plant Taxonomists. All rights reserved. Delivered by Ingenta to IP: 192.168.39.151 on: Fri, 24 Sep 2021 18:12:04 largest leaves30cmlong; comparison tooneofthelargest (secondaryheads9cmand (ca. 3cm)andleaves(2–5cm; absence. focused collectingeffort isneededtoestablishitspresence or veys. Thehabitatinthisarea wouldsuit new areas oritcanbeasignofinadequatebotanicalsur- tribution range,whichcouldindicateactive‘invasion’of plant. Manynew‘recent’ records are notedwithinthedis- road infrastructure inthearea andthespinynature ofthe suspect thisisduetoinsufficient collectionsbecauseofpoor the crowns are closetogetherandtheleavesarch upwards. crowded areas suchasdepressions ordenselyvegetatedplaces of therhizome. [pencilillustrationby Alice Tangerini (US)] [Volume 36 SYSTEMATIC BOTANY 198 (e.g. barium labelsasanuisancein pastures andonsportsgrounds ulations thatcantakeoveran area. Itisoftenlistedon her- they spread bymeansofstoutrhizomes andformlarge pop- measurements for but forthemostparttheywere simplyattheshortendof the measured partswere alittlesmallerinPenthercollection corolla aslight purpleorwhitishnearthe basewithadark one recent collection( ple, violet,mauve,and‘turning whitewithage.’Photosfrom (e.g. The specimenwithoneofthesmallestsecondaryheads The distributionseemstobeintwodistinctareas butwe Fig. Ecology— Starke s.n. Theron 1039 . Ilsrto o of Illustration 5. Once plantsofthis species becomeestablished ). Corolla colorislistedvariouslyaspink,pur- ). A fewsheetslistitashavingasweet scent Koekemoer 3466 Platycarpha glomerata Koekemoer no. 3466 Koekemoer 3466 Penther 10622 . Less.showingthehabitwitharching leavesandthegroup oflarge headsnestedtogetheronthecrown ). Ingeneralsomeof ; Fig. 4 ) showthe P. glomerata ) wasexaminedin anda River, 21Nov1894, 1973, [3324] Humansdorp,1kmW ofKabeljousStation,19May Cape: [3228]MouthofKeiRiver, s.d., square cell( Larsen etal.2009 )]. SOUTH AFRICA. Eastern identify theNWcornerof longitude (S)latitude(E)degree able plantshavebeenconsumed. that itcanbefoundinpastures androadsides where allpalat- Brandwijk 1962 ) andthisagrees withourfieldobservations livestock andfeedingtestswere negative( Watt andBreyer- evil spirits( Arnold etal.2002 ). Itisapparently noteatenby infusions are knowntobeusedassprinklingcharmsagainst as ‘imboziza’,and Pooley (1998) citesitas‘usiphahluka.’ Plant Raymond (2002) and Hutchings (1996) report theZuluname Mkhipheni Ngwenya,ZuluBotanicalKnowledgeProject). a homesteadtoprotect againstlightningstrikes(pers.comm. whole plant(called‘intelezi’)issprinkledintheyard around believed tohavesomemagicalpowers;aconcoctionofthe Gray 2003 ), ‘inyathelo’and‘imbozisayabesuthu’itis several commonnamesinKwaZulu-Natal:‘imbozisa’( Gordon- species. Little isknownaboutthepollinationorseeddispersalofthis area showlilaccolored florets butthismaybeafactorofage. purple glandattheapexwhileotherphotosfrom thesame Humansdorp Rd, 27 Apr 1947, Adtoa Seies Examined— Specimens Additional Value— Cultural and Economic Leibold 45211 (PRE);[3325]5milesfrom Thornhillon Schlechter TRV-1418 Story 2409 Platycarphaglomerata nmes n brackets in [numbers (PRE); Addo Elephant Flanagan 2882 (PRE); Gamtoos (PRE); has Copyright (c) American Society for Plant Taxonomists. All rights reserved. Delivered by Ingenta to IP: 192.168.39.151 on: Fri, 24 Sep 2021 18:12:04 01 FN N OKME:PAYAPEE 199 FUNK AND KOEKEMOER:PLATYCARPHEAE 2011] n pappus. and margin at theapexbutismore slender; D.Floret showingthe longtubeandstylebranches;E. Longitudinal sectionoffloret s of thepalesfrom theouterpartsofaheadit issimilartotheinvolucralbracts; C.Oneofthepalesfrom theinnerpartof Fig. . Ilsrto o of Illustration 6. Platycarpha glomerata Less. A. Partofasecondary headshowingtwoheadsandafewleavesaswellthethickrhizome; B.One a headitstillretains theflared howing anthers;F. Achene Copyright (c) American Society for Plant Taxonomists. All rights reserved. Delivered by Ingenta to IP: 192.168.39.151 on: Fri, 24 Sep 2021 18:12:04 from silk-wooly crown, lyingflaton theground, inseveral times widely spaced ( Fig. 7 (ca. 1cmindiameter);forming matsbutindividualssome- ). Leavesprostrate, radiating Platycarphella carlinoides without bridgeacross pore, noncaveate.Species2. nate, tricolporate,withoutelaboratedborder oncolpiand truncated andoverlappingwhitescales.Pollengrainsechi- papillae orhairsbecomingobsoletetoward tips.Pappusof papillose upperpartofstyleshaft;branchesslightlytapered, 8–13 mmlong;stylebranchesca.1long,shorterthanthe rets, notdeeplyembeddedinthesurfaceofcrown; corollas heads onthecrown oftherhizome; headswith3–14(–20)flo- Inflorescences ofsecondary headsformedbyatightclusterof lating orminutelydentate,sometimesnarrowly pinnatisect. oblanceolate, 8–60mmlong,margins entire, somewhatundu- prostrate inrosettes, radiatingfrom awoolycrown, blades rescences, proliferating byrhizomes; withoutlatex.Leaves [Volume 36 SYSTEMATIC BOTANY National Park,NofCamp,27Jan1966, 200 1894, [ Mar 1951, (PRE); s.d., & Zeyher191 Zwartkops River, 2.1Villa PaulMare toUitenhage,Jan, Redhouse, PortElizabeth,Nov1914, Feb 2007, 11569 Albany Divison,Grahamstown,13Dec1911, Mouth ofKariegaRiver, 19Jan1947, Alexandria, Grahamstown,9May1981, Platycarphella Rd toPontoon,7Nov1974, 2005, (PRE); Vernon Crooks Nature Reserve, border fence,7Dec along thefenceofLakeElandGameReserve,2006, PRE-45193 Rd, 19Feb1962, Hilliard2119 Menne 1366 Harold JohnsonNature Reserve,23Oct1996, (PRE); EshoweDistrict,Mandini,c.1kmNofturnoff to a tribalarea atMapumulo inlandofStanger, 2006, Stanger andSouthofTugela, 2006, Dist., s.d., Feb 1974, Nichols1411 Natal: [2831]ImpaphalanearFort Yolland, 25Jan2006, Meeting Nature Reserve,3Jan1984, 942 Grahamstown onRdtoPortElizabeth,18Jan1955, Nov 1895, unused gravelpit,Jan1939, TRV-10622 Perennial herbs,prostrate, acaulescent,rhizomes slender Perennial, rosettiform, acaulescentplantswithsessileinflo- (PRE);9Dec1971, K!, scannedimageatUS). 1864”], S latitude,[withoutdatebut markedas“received 8IV Africa 3:430,1877.—TYPE: South Africa. 23degrees Platycarphacarlinoides & H.Rob.CompositaeNewsletter47:24–27,2009. noides Newsletter 47:24–27,2009.—TYPE: (PRE);1kmfrom Bathurstonrd toGrahamstown,5 Ward &Swanepoel15894 Penther 1157 Ward 8571 (Oliv. &Hiern)V. A. Funk&H.Rob. Koekemoer 3466 Forest Off.45190 Galpin 3027 (PRE); Isingolweni,inlandfrom PortShepstone, Theron 1039 ] (PRE);Nov1925, (PRE);DoornkoponStangerRd,24 Nov1963, (PRE);[2930]Westville, UniversityofDurban,27 James ChapmanandThomasBaines s.n. (M);[3029]25miWofPortEdward onBizana (PRE,S,US);Uitenhage,Jan1908, Bolus s.n. Noel 2394 V. A. Funk&H.Rob.,Compositae (M);Dec1907, (K,PRE,photoatUS);Pietermaritzburg (PRE); Albany Divison,ca.37kmfrom Starke s.n. (K,PRE);25Nov1894, (S);12Jan1814 (PRE,US);Grahamstown,Kowie, Oliver&Hiern,FloraofTropical (PRE);[2931]Mapumulo,inlandof (K,PRE);[3030]Ixopo,s.d., Stirton 1406 Simon 29774 (PRE);[3130]PortEdward, on ( (MO,PRE);Bathurst,Waters Oliver &Hiern)V. A. Funk Thode A656 Tennant 4218. Styles s.n Gobez 2 Liebenberg 7703 (PRE). Paterson 1889 (PRE);Port Alfred, 9 Burchell 4423 Acocks 13275 Wilson s.n. Platycarphella carli- (PRE).KwaZulu- (PRE);Mvozane, (PRE);15Oct Schonland 1689 (PRE);[3326] Penther 1182 Burtt Davy Williams & (holotype: Styles s.n. Styles s.n. (K);10 (PRE); (PRE); (MO); Comins (PRE); Ecklon Webb

ary headwasin floweratthesametime( Fig. 7B, C ). Older when itstarted tofloweritappeared as ifthewholesecond- it hadsomanysmallheads, eachwithonlyafewflowers, mately 400heads,eachonewith onlyafewflorets. Because and Steenkamp6357 one secondaryheadof Platycarphaglomerata hundreds ofsmallheadsinthelarge secondaryheadwhile large secondaryheads,inthat from heads inthesecondaryhead( Fig. 7 , 8A than theotherspecies)anditstendencytohavemanysmall ). This speciesdiffers originating from thecrown (sometimesasmanysix,more pinnatisect leaveswithoutspines,multiplerows ofleaves species inthetribebyitsshortstylebranches,entire todeeply irregular thanthoseoftherugose achenes. Figures 7–8 . sclaes ofthesmoothacheneslonger, more narrow, andmore margins irregularly dentate;apexacutetoacuminate,stiff, late, persistent,whitescalesofvaryinglengths,2–3×1mm; widths, smoothorrugose. Pappusofca.10lineartolanceo- types, oblong,2.50×1.25mm,darkbrown, 3sidesofunequal apex, stylebranches<1mm,apexrounded. Achenes oftwo len area alongbackside of stylebranchesallthewayto below thestylebranches,shortstiff hairsfrom baseofswol- purple tolavender, 9–10×<0.5mm,slightswollenarea just 0.5 mmlong,filamentsca.1longandpaleincolor;styles white, subequallysagittateatbase,apicalappendagesacute< mm long,thecaeca.3purpleoroccasionallypink down ontothethroat and tube( Fig. 8D ); stamensca.4.0–4.5 ial surfaceofapexeachlobe,glandssometimesextending lobes 3mmlong,spreading withdarkcolored glandsonabax- 3.5–4.0 ×1mm,glabrous to sparselyhispidonproximal end, tubes 4–5×<0.5mm,sparselyhispidatdistalend,throats occasionally 11); corollas purpletopink-purple, 7–9mmlong, altogether. Florets gobletshaped,ca.3–7(frequently 3–4, bracts andsomeheadswithonly3florets appeartolackthem are fewflorets theyare difficult todistinguishfrom involucral to oblanceolate,ca.(5–)7–9mm×<0.5–1.0mm,whenthere a shortstiff tip.Pales(receptacular bracts)ifpresent, linear ance; margins entire, apicesacuminateoracuteanderect with the adaxialsurfaceandapexgivingheadawoolyappear- gins ordistalportion,glabrous orwithlongwhitehairson with strawcolored margins sometimescanbepurple onmar- difficult totellwhichbractsbelongaspecifichead],green inner ones[organization ofheadissuchthatitsometimes 5–10 ×1mm,smallestouterbractsabouthalfthelengthof Involucral bracts7–25,lanceolatetolinearin3–4series; or lesscylindricaloblong,3–4mmindiameter( Fig. 8B ). face sometimeswithwoolywhitehairs.Headsdiscoid,more but someyoungheadssubtendingolderheads,crown sur- heads are mostlyofthesameageinsecondaryhead, the rhizome, secondaryhead2–10cmindiameter( Fig. 8A small headsgrouped inasecondaryheadonthecrown of ); with densewhitetomentum.Inflorescence sessilewithmany rugose surfacewithsmallglands,abaxialcovered stiff spine-likeprojection; adaxialsurfacepalegreen witha natisect ( Fig. 7 ); apicesrounded withaslightpointorsmall lating, minutelydentate,oroccasionallydeeplypin- where itmeetsthecrown, margins entire, somewhatundu- cm, petiole-likebaseoflarge leavesbroadens to 5–15mm larger lowerleaves8–20×1–3cm,smallerupper1.50.5 layers; bladesoblanceolatetoovatemore orlesslinear, Platycarphellacarlinoides Platycarpha glomerata ) was6.5cmindiameterand hadapproxi- hasfewerlarger heads.Forinstance, Platycarphella carlinoides canbedistinguishedfrom theother , theotherspeciesthatproduces Platycarphella parvifolia ( Oliver, Muller has Copyright (c) American Society for Plant Taxonomists. All rights reserved. Delivered by Ingenta to IP: 192.168.39.151 on: Fri, 24 Sep 2021 18:12:04 01 FN N OKME:PAYAPEE 201 FUNK AND KOEKEMOER:PLATYCARPHEAE 2011] in asomewhatwetter environment, theplants are ina“peak”flowering stage. southern Namibia inaverydryenvironment, theplantsare inalate flowering stage;B&Ewere takenbyC. A. Mannheimer(WIN est andlargest onthebottomandfactthatinnersmaller leavesare arching upward somewhat. A, C,&Dwere takenbyM located atthecrown oftherhizome; D.Smallersecondaryhead,notethewhitepollen; andE.Individualplantwithseveralrin a relatively wetarea, notethenearlyentire leavesandlarge secondary head;C.Plantinadryarea, notethedividedleaves small sizeoftheheads: A. Habitandhabitat,notethe dryrocky soilandtheupward thrusting oftheplantwhenthey are clos Fig. . Iae o of Images 7. Platycarphella carlinoides (Oliv. &Hiern.)V. A. Funk&H.Rob.note thehemisphericappearence ofthesecondaryheadsand and thelarge secondaryhead e toanotherone;B.Plantin gs ofleaveswiththeold- D) southofWindhoek . Koekemoer(PRE)in Copyright (c) American Society for Plant Taxonomists. All rights reserved. Delivered by Ingenta to IP: 192.168.39.151 on: Fri, 24 Sep 2021 18:12:04 F. style withveryshortbranches;G.rugose acheneand pappus;H.smoothacheneandpappus. [pencilillustrationby Alice Tang heads havefewif any pales;D.floret showing atubeandstylebranchesthatare shorterthan leaves beinglarger, this large clusterofsmallheads isinfullflower;B.headwiththree florets; C.pale thatisverysimi [Volume 36 SYSTEMATIC BOTANY 202 Fig. . Ilsrto o of Illustration 8. Platycarphella carlinoides (Oliv. &Hiern.)V. A. Funk&H.Rob. A. Habit,notethesizevariationinleaveswith lower/oldest P. glomerata ; E.longitudinal sectionoffloret showing anthers; lar toinvolucralbracts, somesmall rn (US)] erini Copyright (c) American Society for Plant Taxonomists. All rights reserved. Delivered by Ingenta to IP: 192.168.39.151 on: Fri, 24 Sep 2021 18:12:04 (PRE); Ca45km NofMaltehöheontherd toWalvis Bay, 11 852 (WIND); [2416]NaukluftPlateau, 16May1978, 1987, Giess 3594 WIND); FarmBergland REH264,10 Aug 1963, on rd toRehoboth,11 Jun1955, 1983, (M); [2317]40kmfrom Rehobothonrd toKalkrand,27Feb Vlok &Viviers 1805 [2316] betweenKlein Aub andNauchas,19 Apr 1987, Brukkaros andvicinity, 1Jun1931, (PRE); 24Jun1984 Zebra Pan,35milesEofGobabeb,2Jul1969, S-48 [2620] Welkom, 5kmSofTwee Rivieren, 21Mar1976, square cell( Larsen etal.2009 )]. BOTSWANA. Kcalagadi: identify theNWcorneroflongitude(S)latitude(E)degree leaves are heavilygrazedearlyintheseason. in pastures; intheKalahari itwasobserved(MK)thatyoung sional notesonherbariumsheetsthatitissometimesfound the economicorculturalusesofthisplantexceptforocca- heads (pers.obser.). cies butblackantswere frequently seenvisitingmanyofthe is knownaboutthepollinationorseeddispersalofthisspe- anther thecaeare purpleand thepolleniswhite( Fig. 7 ). Little glands ontheabaxialsurfaceofapexcorolla; the lection ( ary headsthatare pastfruiting. Photos from onerecent col- by meansofrhizomes which seemtoemanatefrom second- further northitisalsofoundonstonysoilsandhillslopes. sources andindrainagelines, whileinthehigherrainfallareas tends togrow inshallow depressions, around seasonalwater areas towards thesouthernendofitsdistributionrange,it /granite. Inareas withless rainfallthatismore unpredictable, hard soil,drymargin ofdam,sandydrysoil,andingravel pan, grassland,rocky flats, rocky hillsides,drysandysoil,red 203 grow inavarietyofhabitats basedonlabelinformation:drysalt 1,800 m.Itisoftencollectedafterrainsanditseemstobeable vious discussionand Fig. 1 ). Itisfoundatelevationsfrom 700– it israrely foundinthesouthwestcornerofBotswana(seepre- and intheKalahariregion oftheNorthernCapeSouth Africa; spread inthesouthernandcentralhighlandarea of Namibia lector atthesamelocationthree dayslaterhadnormalleaves. leaves (lessthan1cm)althoughanotherplantbythesamecol- (Fig. 8B).Onecollection( the leavesandinvolucralbractswhere theyattachtothecrown white hairs,eitherscattered ordense,are foundatthe basesof age oftheplantandamountwaterthatisavailable.Long FUNK AND KOEKEMOER:PLATYCARPHEAE the charactersofheadandleavessoitmaybeafactor There doesnotappeartobeanygeographiccorrelation between ondary headsandthesizemargins ofleavesvarygreatly. short timeuntiltheoldheadispressed flat.Thesizesofthesec- older secondaryheadandsowillriseabovethefirstonefora 7A ). Also, onesecondaryheadsometimesdevelopsoutofan together andpushupwards foracoupleofcentimeters( Fig. if twosecondaryheadsare closetooneanothertheymaygrow ( Fig. 7A,D ). This speciesseemstoalwaysbeprostrate although secondary headshaveafewflowersscattered here andthere 2011] Adtoa Seies Examined— Specimens Additional Value— Cultural and Economic Ecology— Habitat— and Distribution (PRE,WIND); FarmOuKamkas,3Jan 1995, (PRE). NAMIBIA.Erongo: [2315] NamibDesertPark, Sievers 109 Immelman 430 Koekemoer 2696 (PRE,WIND);OanobDanNature Reserve,14Mar Once thisspeciesbecomesestablisheditspreads (WIND);Rehoboth,22Feb1979, (MO,PRE);Damas,3Jun1939, (PRE,WIND);50.2milesNofKalkrand Phillipson 866 ) showthecorolla aspurple withdark Volk 12100 Platycarphellacarlinoides De Winter 3522 (K). Gibeon:[2517]Mount There isnoinformationon , M)hadextremely narrow Sordahl 114 nmes n brackets in [numbers (US).Hardap: Muller &Tilson Merxmüller & (K,M,PRE, Rauh 48953 Dreyer 466 Jensen 284 Volk 2219 s wide- is Skarpe Bean,

2 Apr 1963, and surroundings, 9Mar1988, mi NEofWindhoek, Jun-Jul1916, Keres, Dist.Windhoeck, 19 Apr 1939, Apr 1960, Karasburg, 21Sep1989, Aug 1961, Galpin14168 6 May1976, to RoshPinah,Vogelstruisskluft (probably Vogelstrausskluft), mi NofHelmeringhausen,19 Apr 1963, Farm Helmeringhausin,20 Apr 1949, Calitz&Sheuyange115 Grensplaas 490,2kmSofHomesteadinriver, 19 Apr 1997, Mar 1995, Bergland, FarmFinkenstein,11 Apr 1966, the waytopumpstation,10Jun1963, 11 Mar1995, 2077 1962, Jun 1939, 434, 14Mar2001, 2004, [2819] Ca.75kmfrom Karasburg onrd to Ariamsvlei, 6Jun miles WNWof Aroab, 14Oct1949, 1955, 677 Aroab atKlipfonteinonrd toKarasburg, 30 Aug 1974, of Aroab, 14Oct1949, Keetmanshoop, 3May1955, [2619] Dist.Keetmanshoop,7miWof Aroab onroad to 1976, Unknowns.n. [2617] betweenHelmeringhausenandBethanien,21Jun1987, Tonpfanne auf Plateau,26Jun1974, 3675 WIND); [2518]FarmKleinDaberas19,11 Feb1998, of Twee River, 18May1956, Kalahari GemsbokNationalPark,EindpaalNtipofpark,NW Proefplaas Rohrbeck,11 Oct1960, 154, 23Mar1998, 5 Jul1947, 695 (M);59kmSWofMaltahöheonrd toHelmeringhausen, between FarmSchwarzkuppeandOsis,28Dec1974,Friedrich the rd toMaltehöhe,29Feb1996, (K, PRE,WIND);[2516]±45kmNofHelmeringhausenon miles Eof Aranos, 26 Apr 1960, 1956, 6 Apr PRE, WIND);FarmHaribes, Apr 1956, [2417] 18kmSofKalkrand,29May1974, Barod 15385 28080 Schelpe162 Barod 15595 Jul 1962, Jun 1963, Pan onRHS,10 Mar1983, Rousseau2 1780 [2316] Naos,FarmReh46,29 Aug 1972, 139 [2218] BetweenGobabisandWindhoek, Feb1921, (PRE); Windhoek, Neudamm,7Jul1937, (PRE);7milesWof Aroab onrd toKeetmanshoop,3May (PRE);FarmHelen231,9Mar2001, (WIND);hillswestofMaltahöhe ongravelrd toSolitaire, (WIND);[2616]Farm Aar LU16;HäufigaufBrauner (M);[2317]FarmStolzenfeld442,SchaapRiver, s.d., Oliver, Muller&Steenkamp6352 (PRE,WIND);NearNauchas,8 Apr 1977, Hanekom 62 De Winter 3399 Koekemoer 2696 Hanekom 99 Seydel 3549 Volk 2266 (WIND);34kmNofRehobothonrd toWindhoek, Nordenstam &Lundgren Burgoyne 3459 Schweickerdt 2532 Seydel 2353 (M);Windhoek, 15May1954, (WIND); FarmNamibgrens, 13 Mar1953, (MO, WIND);FarmPlateau,s.d., Kers 1616 (PRE);between Ariamsvlei andKarasburg, 14 Volk 12229a Oliver, Muller&Steenkamp6357 (WIND);Feldschuhhorn,WofSeeheim,6May Germishuizen 7824 (WIND);12miEofWindhoek, 10Jul1954, Strohback &Dauth2075 Klaassen 239 (M);MunisipalegebiedWindhoek, 7Jul (K,US);Finkenstein Region,Bergland, (K,PRE,WIND);Keetmanshoop,21.5 (PRE); Ariamsvlei Station,4Jul1937, (PRE,WIND);22Mar1910, (S);Bergland, FarmBodenhausen,21 (M,WIND);FarmLichtensteinWest (WIND).Karas:[2516]HelmeRiver, Dean 688 Unknown s.n. (PRE);FarmFriedland,s.d., (M);[2419]FarmLekkerwater, 32 Immelman 560 Story 5532 (PRE);betweenOnseepkansand deWinter 3399 (WIND);FarmOngeama,20 Bohlmann 8826 (PRE,WIND);Bergland on Burgoyne &Snow5049 (PRE).Khomas:[2216]Farm Pearsoon 9879 603(S);[2518]Farm Aukam Van Vuuren &Giess1098 Freyer 88 Gassner 139 (PRE);Rdfrom Seeheim (PRE);Mariental,Farm Acocks 15583 (WIND);5kmSWof Volk 12100 Kinges 2188 Giess 13386 Klaassen 223 (PRE);Bergland, 10 Nordenstam 2296 Galpin 14167 Goldblatt 1993 Merxmüller &Giess (M);21.5miWNW (WIND);Kalahari Brain 2 Seydel 4410 (WIND);[2519] Seydel 3549 (K,MO,PRE); (K);Windhoek (ID; Aris, (WIND); (M);[2217]35 (M,WIND); Lavranos & Lavranos & (PRE);16 (K,PRE); (PRE);22 (PRE,S); Mucke 37 (WIND); Strohbach Gillman (PRE); (PRE); Walter Walter Clarke (MO, (M); (M); (S,

Copyright (c) American Society for Plant Taxonomists. All rights reserved. Delivered by Ingenta to IP: 192.168.39.151 on: Fri, 24 Sep 2021 18:12:04 subglobose, 3–4 mmindiameter( Fig. 10C ). Involucral bracts ing olderheads andoftenneartheperimeter. Headsdiscoid, the samesecondaryhead,some veryyoungheadssubtend- ondary head1–2cmindiameter; headsare different agesin heads grouped onthecrown oftherhizome forming asec- with densewhitetomentum. Inflorescence sessile, few-20 a rugose surfacewithsmallglands,abaxial surfacecovered rounded withaslightpoint;adaxialsurfacepalegreen with ( Fig. 10B broadens to4–7mmwhere itmeetsthecrown, margins entire ), somewhatundulatingorminutelydentate;apices proximally turningred inmost,petiole-likebaseoflarge leaves ( Fig. 9 radiating from silk-woolycrown, lyingflatontheground A, D);bladesoblanceolatetolinear, 12–60×3–10mm, (ca. 1cmindiameter);formingdensemats.Leavesprostrate, Platycarphella parvifolia Upington, 21 Apr 1928, Park, 22Mar2008, (PRE); justSofTwee Rivieren outside ofGemsbokNational around MelkvleiPicnic site, 22Mar2008, Gemsbok NationalParkinthefloodplainofNossobRiver 15 kmNofBokspits,12Jun1990, Twee Rivieren, 16May1956, Brynard216 Kalahari GemsbokNationalPark, Auob Riverbed,15Jul1953, [VolumeRiver, KwangPan,17 36 Apr 1960, Rooyen 3675 National Parks,Dikbaardskolk Afdraai, 15Jun1984, “Okohandje”, 20Mar1928, 1983, s. n. Woortman 208 Farm Okawari,Wolfkalkveld, 4Jun1974, 1927(28?), SYSTEMATICon rt1800–1808,s.d., BOTANY Mason&Boshoff2594 Klaassen223 May 1949, 23 Feb1965, 204 1960, Jun 1927, S ofOtjiwarongo, Jul1953, Omatjiune ExperimentalFarm,s.d., on Steinhausenrd, 5Mar1997, 1 May1963, (S-2 sheets);[2119] FarmLausitz,EofSteinhausen,Epukiro, Zyl 29 Schwurdsfeger4149 Volk &Leistner6720 [2118] FarmLausitzinEpukiro Omuramba,1May1963, Krone 136,11 Mar1968, 1886, 11704a Gravenstein, 2Mar1963, Perennial herbs,prostrate, acaulescent,rhizomes slender SOUTH AFRICA. NorthernCape:[2520]KalahariGemsbok (M);[2117] FarmHortensia,45kmfrom Hochveld,6Mar at Ksays“MissBarrett-Hamilton, 21Oct1901”]. type: K!)[specimenatBMwasdonatedin1902,specimen Hamilton s.n. Orange RiverColony, [withoutdate], AFRICA. Western Transvaal: Wolmaransstad district, in JournalofBotany41:136,1903.—TYPE:SOUTH Rob. CompositaeNewsletter47:24–27,2009.S.Moore Germishuizen 2613 Seydel 2231 Marloth 1342 (WIND);48milesNofWitvlei, 1May1963, (M).Omaheke: [2116] Okahandja, Hereroland, May v.Wernsdorff s.n. Marloth PRE-45173 Liebenberg 4544 (PRE);[2620]KalahariGemsbokNationalPark, (PRE);KalahariGemsbokNationalPark,Nossob (WIND);Okatjirute West, Witvlei, 24 Apr 1969, Seydel 4211 Bleissner 224 (PRE,WIND);FarmErichsfelde,s.d.1956, (K,MO,US);Omatako-Sicht,28 Apr 1976, (holotype:BM,scannedimageatUS;iso- (WIND);FarmOkaruako, 26 Apr 1984, Koekemoer 3593 (PRE);7miNofSeeis,FarmDeutsch (WIND);FarmOkasondana,23Jun1953, (PRE);Site161,30kmfrom Witviel Bester &Van Eck4 (PRE,WIND);Quickborn,PostOffice (MO,US);s.d., Pole Evans2162 (PRE);FarmHelen231,9Mar2001 (M);[2218] At turn-off toWitvley Leippert 4265 (M,2sheets).Otjozondjupa:[2016] Wanntorp &Wanntorp 219 Basson 16 Bradfield 247 Story 5485 (S.Moore) V. A. Funk&H. (PRE);Omongongua,31Mar Barnard 512 (PRE);[2820]41milesNof Germishuizen 9420 Pfeiffer s.n. (PRE);[2116] Okahandja, Leistner 1841 (M);7Mar1956, (PRE). (PRE). Seydel 4410 (PRE);SWKalahari, Jacobsen 4603 (ID; Gobabis, (WIND); Capt. G.E.Barrett- (PRE);Kalahari, Koekemoer 3591 (WIND);23mi (K,PRE); (K);Farm Kers 1040 (PRE); (K,S); (PRE); Giess, Volk Volk Van van

1968, RD toHamiltonsRustNR.drainage Linebelowdam,1Sep Vals river, Sep1929, ( ary headsthatare pastfruiting. Photos from recent collections by meansofrhizomes which seemtoemanatefrom second- dams andstreams inshort grasslandsoronfloodplains. m (seeprevious discussion and Fig. 1 ). Itisoftenfoundnear in thecentralpartofSouth Africa atelevations1,200–1,500 to thecrown are thickwithlongwhitehairs(Figs.10B,H). older, lowerleaves.Thebasesoftheleaveswhere theyattach leaves, whichare justabove theothers,are smallerthanthe be foundinasinglepopulation.Onlarger plantstheyounger leaves donotvarytoomuchandallofthesizedifferences can a coupleofcentimeters.Thesizesthesecondaryheadsand to oneanothertheymaygrow togetherandpushupwards for always beprostrate although iftwosecondaryheadsare close consistently smallsecondaryheadsandleaves.Itseemsto species inthegenusbyitsentire ornearlyentire leavesand to generaandspecies),minutelydenticulate. Figures 9–10 . × 1mmwithoverlappingmargins; apextruncate (seekey lanceolate, persistent,whitescalesofvaryinglengths,2.5–3.0 sides ofunequalwidths,rugose. Pappusof10–12linearto ( Figs. 9C ). Achenes oblong,2.0–2.5×1mm,darkbrown, 3 all thewaytoapex,stylebranches1mm,apexrounded from baseofswollenarea alongbacksideofstyle branches mm, swollenarea justbelowthestylebranches,shortstiff hairs ca. 1mmlongandpaleincolor;styleslavender, 11–13 ×<0.5 tate atbase,apicalappendagesacute<0.5mmlong,filaments mm long,thecae3.5–4.0purple,subequallysagit- on abaxialsurfaceofapexeachlobe( Fig. 9C ); stamensca.5 near base,lobes3mmlong,spreading withdarkcolored glands hispid atdistalend,throats 4.5–5.0-×1mm,sparselyhispid to pink-purple,10–13mmlong,tubes7–8×<0.5mm,sparsely (counted) orpossiblyupto20(estimated);corollas lightpurple every floret hasacorresponding pale.Florets slender, ca.5–14 for apexwhichissomewhatstiff, glabrous; itis not clearif lar bracts)linear, ca.8–9mm×<1ephemeral,soft,except slightly more narrow andresembling pales.Pales(receptacu- and erect, withashortstiff tip;innerbractssimilar becoming at distalend,stiff atapex,glabrous, entire, apicesacuminate mm, bractsofimmature heads5×1mm,green becoming red 10–15 lanceolatetolinearinca.3series;outerbracts8–10×1.5 Rd, Leeuwspruit floodplain, 31 Aug 1997, North from R59intersectionofParysRd andWonderfontein Smith 6295 of Vaal River, oppositeTheSmall IslandNR,Sep1932, Hoeveld, 6Sep1927, [2926] Above railwayline4miWofHomestead on Farm square cell( Larsen etal.2009 )]. SOUTH AFRICA. Free State: identify theNWcorneroflongitude(S)latitude(E)degree it isprobably noteatenbylivestock. sional noteonherbariumsheetsthatitisfoundinpastures so the economicorculturalusesofthisplantexceptforocca- well indisturbedovergrazed areas (pers.obs.). pollination orseeddispersalofthisspecies.Itseemsto do surface oftheapexcorolla. Littleisknownaboutthe the corolla aslightpurplewithdarkglandsontheabaxial Koekemoer &Funk2866 Ecology— Habitat— and Distribution Platycarphellaparvifolia Adtoa Seies Examined— Specimens Additional Value— Cultural and Economic Scheepers 1751 (PRE);Boschbank12,Rossouw’s Farm,extending Once thisspeciesbecomesestablisheditspreads (K,PRE);[2627]Parys, aboverightbank Pont 618 C. A.Smith4456 and canbedistinguishedfrom other Funk &Koekemoer12564 (PRE);FarmRietgat73,SWof Platycarphellaparvifolia There isnoinformationon nmes n brackets in [numbers (PRE);[2727]Near Kroon 15266 occurs ) show ) (PRE); C. A. Copyright (c) American Society for Plant Taxonomists. All rights reserved. Delivered by Ingenta to IP: 192.168.39.151 on: Fri, 24 Sep 2021 18:12:04 C &DbyM.Koekemoer (PRE)] colorful stylesand anthers;D.Secondaryheadswith abutterflyvisiting,notethesmall sizeoftheprimaryheads,[Photos A, in Ethelongslender throat ofthecorolla (aperson’shandcanbeseen inbothofthesephotos);C.Close upofflorets showi surface ofleavesthatare growing flatontheground; B&E.Clustersofsecondaryheads,notethesmallsize both theprim 205 FUNK AND KOEKEMOER:PLATYCARPHEAE 2011] Fig. . Iae o of Images 9. Platycarphella parvifolia (S.Moore) V. A. Funk&H.Rob. A. A densepopulationfoundona secondaryfloodplain,notewhiteunder- ary andsecondaryheads ng shortstylebranches and B&EbyV. A. Funk(US); Copyright (c) American Society for Plant Taxonomists. All rights reserved. Delivered by Ingenta to IP: 192.168.39.151 on: Fri, 24 Sep 2021 18:12:04 amount ofpubescence; H. Achene and pappus.[Pencilillustrationby Alice Tangerini (US)] E. Floret; F. Longitudinalsection ofafloret showingtheanthers; G.Style,notetheshortstyle branchesthatare open only showing theplacement ofthepubescenceoncrown surfaceandthefewflorets withslendertubes; D.Palewhichissimilart small headsonthecrown oftherhizome; B.Leafshowingtheexpandedleafbase, pubescence ontheundersurfaceandhai [Volume 36 SYSTEMATIC BOTANY 206 Fig. 0 lutain f of Illustration 10. Platycarphella parvifolia (S.Moore) V. A. Funk&H.Rob.: A. Habitshowingthree secondaryheadseachformedbyaclusteringof slightly andthedecreased o theinvolucralbracts; rs atthebase;C.Head Copyright (c) American Society for Plant Taxonomists. All rights reserved. Delivered by Ingenta to IP: 192.168.39.151 on: Fri, 24 Sep 2021 18:12:04 Bavr, 95 osets eeu seirmu Arctotidinearum . specierumque generum Conspectus 1915b. G. Beauverd, Bavr, 95 otiuin à ’td ds opse X A L place La A. X. Composées des l’étude Tomalin , à Contributions M. 1915a. Snyman, G. Beauverd, E. E. Hawker , C. L. Prentice , A. C. H., T. Arnold, on theSIDepartmentofBotanywebsite http://botany.si.edu/botart/ . the beautifulpencilillustrations.Manyofherillustrationsare available We are especially gratefulforallthetime Alice Tangerini (US)spenton Ortiz, ananonymousreviewer, andtheeditorsfortheirhelpful comments. tion onpossibleexplanationsofthedistributiontaxaandSantiago thank PeterLinderandRichard Cowlingwhoprovided helpfulinforma- We appreciate the photographstakenbyC.Mannheimer(WIND). Also, we tions inNamibia,andH.Steyn(PRE)forgeneratingthedistributionmap. E. KlaassenandC.Mannheimer(WIND)whokindlyhelpedwiththeloca- supplying anelectronic list of funding forfieldwork;andSANBIassistanceinwork for Smithsonian Institution’sRestrictedEndowmentProgram forprovided images ofthecollectionshousedattheirinstitutions.Thanksto A. Anderberg (S)andF. Schuhwerk(M)fortakingthetimetoscan their respective institutionsandforsendingthescannedimages;alsoto (K), N.BiggsandM.Hjertson(UPS),forhelpingtrackdowntypesat 207 1893, FUNK AND KOEKEMOER:PLATYCARPHEAE Dyer3199 Vereeniging Govt.Ex,towards drainagelevel,30 Aug 1935, Gouda, 27 Apr 1978, Willem Pretorius GameReserve-Wildtuin, langsvleiNaby Sep 2004, and Sasolburg nearbridgeoverKromelmboogspruit, 7 7 Sep2004, of Hwy59onRDfrom “TheBarrage”,SEofPotchefstroom, plain, 3Sep1996, Farm Uitkomst,5–6kmW-NW ofSasolburg, Rietspruit flood- 2011] (by MK). As aresult thematerialwasnotusedforthisstudy. no plantswere founddespite anextensivesearch ofthearea out destroying thespecimens. Duringarecent triptothearea condition ofthecollectionmadeidentificationdifficult with- ( July and August bythesamecollectorbutindifferent years lia. side therangeofeither 10874 Hafström 936 bankar vidflodenVaal mittemotWindsorton, 13Oct1963, Windsorton, 13Oct1936, Aug 1936, Esterhuysen 3633 9096 (M, PRE);[2824]Warrenton, Vaal River, 31 Aug 1921, [2824] Kimberley, ModderRiver, Sep1946, Sutton248 1376 (PRE);[2725]Wolmaransstad, Boskuil,22Oct1929, Hanekom1835 1948, (PRE); Lichtenburg Dist.,Hakboslaagte,around dam, 1 Aug Lichtenburg Dist.,Hakboslaagte,12Sep1947, near Barberspan,2Oct1921, Britz GameFarm,10 Apr 2003, (PRE). NorthernCape:[2823]Postmasburg. “Lime Acres”, Galpin 8480 Acknowledgments. Abgos Specimens— Ambiguous Theywere collectedfrom LimpoponearNaboomspruit in BulletindelaSociété BotaniquedeGenève(2sér.) fians. africaines . australiens des Arctotidées etleursrapportsavecles Arctotidenées . . ruh ad . otsBrhr . 20 eiia ad magical des and Medicinal 2002 . Pottas-Bircher . C. and ofsouthern Africa: anannotatedchecklist . Crouch , R. N. (PRE);Fourteenstreams, banksofVaal River, Aug 1936, (PRE);[2922]Prieska,13Oct1936, Kinges 1766 Schlechter 1293 Ursinia (PRE);Wolmaransstad, 23Jul1902, (PRE).Mpumalanga:[2629]Standerton,16Oct Esterhuysen TRV35857 Funk &Koekemoer12564 Koekemoer 2866 in1913and (S); [2825]Smitskraal,21Sep1911, Bulletin delaSociété BotaniquedeGenève(2sér.) parmilesComposées-Tubuliflores. B.Lesrepresentants (K,PRE);Klerksdorp,3Sep1905, (PRE);Klerksdorp,Wolwerand, 4Sep1972, (PRE);Warrenton, BanksofVaal River, Kroon 11935 (PRE).NorthWest: [2625]Hieromdraai, Our thanksgotoJ.Gregson (BM),D.J.N.Hind ieaue Cited Literature Muller 2133 Platycarphella carlinoides 534 Acocks H936 There are twocollectionsfarout- (PRE);2kmNofR59btwParys Platycarpha in1921;bothare atPRE).The (PRE);NearParys,1.5kmN Pole Evans207 Zietsman &Preece 221 (PRE);Vaal Riveropposite (PRE).Gauteng:[2627] specimens.Thanksalsoto (US);[2827]Winburg, (PRE);Kapkolonien, Bryant 332 : 41–56. 7: Strelitzia Hamilton 10354 (PRE);[2626] Brüeckner 764 r or Kinges 1491 Burtt Davy Burtt Davy : 21–40. 7: (PRE). P. parvifo- 3 1–203. 13: (PRE); Mogg

Bee, . 94. 1994. K. Bremer, Lne, . . 01. To e seis f of species new Two 2001 . P. H. Linder, Lne, . 95. 1995. P . H. Linder, Bhmn, . n C Zeo . 17 ee emcoie u aus Germacrolide Neue 1977. Zdero . C. and in F. 536–537 Bohlmann, 163–533, Pp. Compositae. 1873. Hooker . D. J. and G. Bentham, geographical and history classification, the on Notes 1873. G. Bentham, Gro-ry K D. e. 20 2003. ed. Cape D., the K. Gordon-Gray, of conspectus A plants: Cape 2000. Manning. of J. and evolution P. the Goldblatt, into Insights 2004. Keeley . S. and Chan, R. A., V. Funk, Bonifacino, M. J. Bayer , J. R. Baldwin, G. B. Anderberg , A. A. A., V. Funk, new a and Platycarpheae tribe new A 2009. Robinson. H. and A. V. Funk, in 335–383 Pp. Cichorioideae. 2009. Chan. R. and A. V. cit- Funk, for system reference degree A 1971. Leistner . A. O. and D. Edwards, in 999–1015 Pp. review . Cynareae-systematic 1977. M. Dittrich, in 1836. P. A. Candolle, 364–366 thicket De subtropical African southern Pp. of origin the On (Arctotideae) . 2005. M. R. Cowling, Arctotidées Les 1821. H. in 118 –119 Cassini, Pp. Arctotidées . 1816. H. Cassini, Knel J. 20 ae poie f aii Namibia. of profile Water 2007. J. Kundell, Lsig C F 13 e yatees israin ura. quarta. dissertation Synanthereis De 1831. F. C. Lessing, Lre, . T Hlen S D Pae , . aii ad . okf 09. Using 2009. Roskaft. E. and Maliti, H. Prager , D. S. Holmern , T. R., Larsen, the of analysis Phylogenetic 1992. Bremer . K. and Källersjö, M. O., P. Karis, in 200–208, Pp. Arctotidieae . 2007. O. P. Karis, 1996. A. Hutchings, 1977. eds. 2000. Turner , L. Welman B. . and G. Harborne, B. W. J. H., V. and Heywood, Koekemoer , M. Retief, E. J., P. P. Herman, Lite, . , d 00. Se pat o suhr Arc: aiis and families Africa: southern of plants Seed 2000. ed. A., O. Leistner, Bulletin Bredasdorp plains . GeneraPlantarum 335–577. distribution ofCompositae . ern seaboard flora ofSouth Africa . 637–655. the tribe Arctoteae (Compositae)using IAPT . Vienna : gen- next the Compositae J. metatrees: Wagstaff , Compositae S. 2009. Vogt , Watsoneration R. . Vallès E. , . Pp. J. L. N. Urtubey , and Robinson, Susanna, 747–780 E. Ward A. , Stuessy H. Urbatsch, , F. in T. L. Randle, Serrano, P. Unwin, M. M. Semple, C. C. Pelser , J. B. Sancho, Nordenstam , P. G. B. Roque, Ortiz, Mort, Lowrey , E. S. K. M. T. Oberprieler , Tadesse , Kimball, M. C. T. McKenzie, R. J. Kilian, R. N. Lundberg , Keeley O. , J. P. C. S. Karaman-Castro , V. Katinas, Källersjö, L. Karis, M. S. Kadereit , Hansen, W. V. J. H. Galbany-Casals, Gruenstaeudl , M. Himmelreich , Freire , M. Gemeinholzer E. , S. B. D. Dillon, Garcia-Jacas , Coutinho, O. P. N. M. X. Crisci, A. V. Chan, J. R. Crawford , J. Carbajal, R. Brouillet , L. Breitwieser , I. CompositaeNewsletter genus IAPT . Vienna : Bayer . R. and Stuessy , T. evolution andbiogeographyofCompositae Staatssammlung München ing biologicalrecords insouthern Africa . Press . Academic London: Turner . L. B. and Harborne, ogy andchemistryoftheCompositae Würtz . and Treuttel Paris: vegetation . Normant . Dictionnaire desSciencesNaturelles Oregon . Portland, glomerata Naturelles sharing ofbiodiversitydata . the extendedquarterdegree gridcellsystemtounifymapping and eoearth.org/article/water_profile_of_Namibia . of theMissouriBotanicalGarden Cichorioideae (Asteraceae),withemphasisontheMutisieae . Verlag . cular plants University ofNatalPress . chemistry oftheCompositae Leistner . Asteraceae (Compositae) . in Natal . eea. genera . Platycarphella 6 465–477. 56: Strelitzia . Strelitzia , vl 2 Spl), d . . uir . Prs: L Normant. Le Paris: Cuvier . L. G. ed. (Suppl.), 2 vol. , Phytochemistry . d. V A Fn , . uan , . . tes , n R Bayer . R. and Stuessy , F. T. Susanna, A. Funk, A. V. eds. . o. es . . aeet n C Jfry . Bri Springer- Berlin: Jeffrey . C. and Kadereit W. J. eds. 8. vol. . Durban : Ethekwini ParksDepartmentandUniversityof South AfricanJournalofBotany Asteraceae: cladisticsandclassification Zulu medicinalplants:Aninventory

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