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SCOPEOFTHISREVIEWThisdocumentoutlinesthebasicbiologyandmanagementofspeciesofthegenusLantanathatarepresentinNewSouthWales.ThereviewwasundertakentoascertainifthecurrentdeclarationofthenoxiousweedspeciesLantanacamarawasappropriate.AnassessmentofLantanamontevidensisandthelargenumberofornamentalLantanavarietieswasalsomadetodetermineifnewdeclarationswereappropriate.

Informationfromtheliteraturehasbeengroupedintothefollowinggeneralareasthroughoutthisdocument:­nomenclature,speciesdescriptions,taxonomy,origin,lifecycle,dispersal,growthanddevelopment,habitat,distribution,importance,legislation,controlandrecommendations.Furtherresearchneedsandinformationrequiredfromconsultationhavebeenhighlighted.

Figure3.A flowering plant of the red variety of Lantana camara, near Seeview (Grafton, NSW north coast, October 2005) (Source: S. Johnson, NSW DPI).

REVIEW OF THE DECLARATION OF LANTANA SPECIES IN NSW 3

REVIEW OF THE DECLARATION OF LANTANA SPECIES IN NSW4 REVIEW OF THE DECLARATION OF LANTANA SPECIES IN NSW4

REVIEWOFTHE usesthetaxonomicnameL.camaratodescribethisaggregatespecies.

DECLARATIONOFLANTANASPECIESINNSWNOMENCLATURE

Lantana camara ThereareavarietyofcommonnamesusedforLantanacamaraL.inAustralia.Theseinclude:Lantana,Commonlantana,Kamaralantana,Large­leaflantana,Red­floweredsage,WhitesageandWildsage(Shepherdetal.2001).

Internationalcommonnamesinclude:Achmann(Cambodia),Bahug­bahug(Philippines),Bands(India),Boengapagar(Indonesia),Bungatahiayam(Malaysia),Cambaradeespinto(Brazil),Cariaquillo(PuertoRico),Chiponiwe(Zimbabwe),Cuasquito(Nicaragua),Guphul(India),Kauboica(Fiji),KembangsatikandKembangtelek(Indonesia),Largeleaflantana(USA),Latoramoa(Tahiti),Nagaairi(India),Pha­ka­krong(Thailand),Phullaki(India),Pricklylantana(Malaysia),Putus(India),Red­floweredsage(Barbados,ThailandandTrinidad),Saliara(Indonesia),Sapinit(Philippines),Tahiagam(Indonesia),Tantbi(India),Taturamoa(Tahiti),Telekan(Indonesia),Thomoi(Vietnam),Tickberry(easternAfricaandZimbabwe),Vieillefille(Mauritius),Wildsage(Jamaica)andWhitesage(ThailandandTrinidad).

Synonymsthatareusedtodescribethespeciesinclude:­ CamaravulgarisBenth.,LantanaaculeataL.,Lantanacamarasubsp.aculeata(L.)R.W.Sanders,Lantanacamaravar.aculeata(L.)Moldenke,LantanacamaraL.var.camara,Lantanacamaravar.crocea(Jacq.)L.H.Bailey,Lantanacamaravar.sanguineaL.H.Bailey(pink­edgedredfloweringvariety),Lantanacamaravar.splendensMoldenke,LantanacroceaJacq.(pink­edgedredfloweringvariety),LantanatiliifoliaCham.andLantanascabridaSol.(Holmetal.1977;Everist1981;JessopandToelken1986;ParsonsandCuthbertson2001;APNI2007).

AbriefdiscussionofthevalidityofthetaxonomicrevisionofL.camaratoL.strigocamaraiscontainedinthetaxonomysection(Sanders2006).Thisdocument

Lantana montevidensis ThereareanumberofcommonnamesusedforLantanamontevidensis(Spreng.)Briq.inAustralia.Theseinclude:Creepinglantana,Lantana,Polecatgeranium,Purplelantana,Sellow’slantana,Smalllantana,Trailinglantana,WeepinglantanaandWildverbena(Shepherdetal.2001;O’Donnell2002).

TherearetwosynonymsusedforthisspeciesthesebeingLantanasellowianaLink&OttoandLippiamontevidensisSpreng.(White1929;StanleyandRoss1986;Shepherdetal.2001;APNI2007).LantanamontevidensiswasknownasL.sellowianafromitsintroductionintoAustraliauntilthe1930’s(Swarbrick1986). PlantsformerlyidentifiedasL.sellowianahavenowbeenclassifiedasL.montevidensis. OtherspeciesnamesthathavebeenusedinvariousliteratureincludeL.sellowii,L.selowianaandL.delicatissima(Anon.1857;O’Donnell2002).ThesenameshavenotbeenrecordedintheAustralianPlantNamesIndex(APNI2007).

ThesetwoLantanaspeciesaretheonlyspeciesrecordedasbeingnaturalisedinNSW.Theyhavebeenreferredtobyscientificnamethroughoutthisdocument.

SPECIES DESCRIPTIONS

Lantana camara Lantanacamaraplantsaremulti­branchedshrubsthatgenerallygrowfrom2­4minheight(AuldandMedd1987;Conn1992,Figures3­5and14).Thebranchesclimbovereachother,formingdensethickets(Conn1992),to15minheightifsupportedbysurroundingvegetation(Swarbricketal.1998). Thebranchesaresquareincross­sectionand2­4mmindiameterwhenyoung,becomingmorerounded,grey/brownandupto150mmindiameterwhenmature(Swarbricketal.1998;ParsonsandCuthbertson2001).Theyoungstemsoftheweedyvarietiesarehairyandhaveshortrecurvedprickleswhilethoseonthenon­weedyvarietiesarerounder,moreslenderanddonothaveprickles(Swarbricketal.1998). Thebranchesmaybe

REVIEW OF THE DECLARATION OF LANTANA SPECIES IN NSW 5

woodyorbrittleandoftenhavepithycentres(Everist1981;Swarbricketal.1998;ParsonsandCuthbertson2001;vanOosterhout2004).

Theoval­shapedleavesareborneoppositeeachother,are20­120mmlongand15­80mmwide,onpetioles5­30mmlong(Holmetal.1977;StanleyandRoss1986;Conn1992;Munir1996;Swarbricketal.1998;ParsonsandCuthbertson2001).Theleafbasesarerounded,pointedorheart­shapedwhilethemarginsareroundlytoothed(Conn1992;Swarbricketal.1998). TheleavesofthepinkfloweringvarietyofL.camaraarepalegreenwhilethoseoftheredfloweringvarietyaredarker(AuldandMedd1987).LeafsizeandshapeisdependantonthevarietyofL.camaraandtheavailabilityofmoisture(vanOosterhout2004).Theleavesmaysometimesbeglossy(ParsonsandCuthbertson2001).Theuppersurfaceoftheleavesisoftenwrinkledandcoveredinsharp,rigidhairswhilethelowersurfacemayormaynotbecoveredinshort,softhairs(Conn1992).Theleafveinsareprominentonthelowersurfacewhiletheuppersurfacecontainsveryfewstomata(Swarbricketal.1998). Theleaveshaveastrongodourwhencrushed(Swarbricketal.1998).

Theflattoppedtodome­shapedflowerheadsareproducedinpairsintheaxilsofoppositeyoungleaves(Swarbricketal.1998;Dayetal.2003;Figure4)andare10­30mmindiameter,whilethepeduncleorflowerheadstemis20­95mmlong(JessopandToelken1986;Conn1992;Munir1996;Swarbricketal.1998).Lance­shapedbractsarepresentunderneatheachflowerandare3­8mmlong(Munir1996).Thereare20­40stalkless,tubularflowersineachheadwitheachcorolla(flower)9­14mmlongwithfourspreadingroundedlobes(AuldandMedd1987;Conn1992;Swarbricketal.1998). Theflowerbudsareangularandtightlypacked,openinginitiallyfromtheoutsideoftheflowerheadandmovinginwardstowardsthecentre(Swarbricketal.1998). Newlyopenedflowersnormallyhaveyellowthroatswithflowerheadscolouredincombinationsofwhite,cream,yellow,orange,red,purpleandpink.Thesecolourstendtochangewithage(Conn1992;Swarbricketal.1998;ParsonsandCuthbertson2001).ThemostcommonvarietyofL.camarainNSWhasflowerheadsto25mmwideandindividualflowersthatarepalecreamtodarkyellowatfirst,changingtopinkandlilacorpurple(AuldandMedd1987).Thereareanumberof

flowercoloursinornamentalvarieties.Ornamentalvarietiesaregenerallymorecompactplantswithsmallerleaves(vanOosterhout2004).

ThefruitofL.camaraisadrupe(incorrectlyknownasaberry),4­8mmindiameter,greenandhardwhenimmature,turningashinypurple/blackwhenripe(AuldandMedd1987;ParsonsandCuthbertson2001;Figure5).Fruitareborneinclustersofupto20witheachfruitcontainingone‘seed’thatispear­shapedandpalestrawincolour,hardand1.5­4mmlongandwide(Holmetal.1977;Swarbricketal.1998;ParsonsandCuthbertson2001).Thisseedisinfacttwofusedpyreneseachwithasingleembryo,bothofwhichareviable(Swarbricketal.1998;Vivian­Smithetal.2006).Swarbricketal.(1998)furthernotedthatnon­weedyvarietiesofL.camaratendtoretaintheirflowerslongerafterpollinationincontrasttotheflowersofweedyvarietieswhichdarkenincolourafterpollination,losingtheiryellowcentreandthenfallfromtheplant. Non­weedyvarietiesproduceveryfewfruit,mostlyfailingtosetseedafterpollination.

Figure4.Lantana camara shoot showing leaf and flower head morphology (Source: Botanic Gardens Trust (2007), used with permission).

REVIEW OF THE DECLARATION OF LANTANA SPECIES IN NSW 6

Figure5.A branch with flowers, green and mature fruit of the common pink variety of Lantana camara, near Copmanhurst (NSW north coast, October 2005) (Source: S. Johnson, NSW DPI).

TherootsystemofL.camaraisbrownandwoodyandhasashorttaprootwithmanyshallowlateralbranches(Swarbricketal.1998;ParsonsandCuthbertson2001).Theselateralsdividerepeatedlytoformadenserootmat(Swarbricketal.1998). Lantanacamaraisabletovigorouslyregrowfromthebaseofthestem,fromtheplantcrownifdefoliatedbyfireorherbicide,orafterdroughtorfrostdamage,fromlateralrootfragmentswhenbrokenandslowlyfromrootedhorizontalstemsthatcomeintocontactwithmoistsoil(Saint­Smith1964;Swarbrick1982;WaterhouseandNorris1987;Swarbricketal.1998;ParsonsandCuthbertson2001).

Lantana montevidensis Lantanamontevidensisisashrubwithhorizontallygrowingbranchesthatmayrootatthenodes(Conn1992;Munir1996,Figures1,6,13and15).Thebranchesalsotrailoverrocks,banksandclimbalongtreebranchesforsupport(White1929;Everist1981;Conn1992).Theyoungbranchesare1­2mmwide,squareincross­sectionandwithageupto5mmindiameter,becomingroundedastheymature(White1929;Everist1981;AuldandMedd1987;ParsonsandCuthbertson2001).Thestemsgrowfrom1­4minlengthbutrarelygainmorethan0.5metresinheightwiththeendsofbranchesgrowingupwards(Munir1996;ParsonsandCuthbertson2001;CooperativeResearchCentreforAustralianWeedManagement2003).

REVIEW OF THE DECLARATION OF LANTANA SPECIES IN NSW 7

Figure6.Lantana montevidensis shoots showing leaf and flower head morphology (Source: S Johnson NSW DPI. Line drawing source: Botanic Gardens Trust (2007), used with permission.).

Thesebranchesformlowdensethicketsormatsandarefrequentlytrimmedtoformhedgesoverexistingstructures(StanleyandRoss1986;Swarbrick1986;CooperativeResearchCentreforAustralianWeedManagement2003).Thebranchesarealsoroughtotouchwithshortrigidhairs,havenoprickles,mayormaynothavehairsandareoftenglandularontheyoungerparts(Conn1992).

ThebrightgreenleavesofL.montevidensisareborneoppositeeachother,areoval­shaped,generally8­30mmlong,5­16mmwide,onpetioles20­40mmlong(Conn1992),althoughoccasionallylarger(White1929).Theleavesoftheornamentalvarietiesofthisspeciesmaybeslightlylarger,from25­40mmlongand6­18mmwide(Everist1981).Theleafbasesmayberoundedortruncate(cutoffinappearance)whiletheleafmarginsaretoothed(StanleyandRoss1986;Conn1992).Theuppersurfaceoftheleavesiswrinkledandcoveredinsharprigidhairswhilethelowersurfacemayormaynotbecoveredinshortsoftwhitehairsandiscommonlyslightlypaler(Everist1981;Conn1992).Thelowerleafsurfacehasyellowandorangeglandsonit.Theleavesarestronglyaromaticwhencrushed(AuldandMedd1987;ParsonsandCuthbertson2001),andthisismostmarkedintheweedyvariety(Everist1981).

8 REVIEW OF THE DECLARATION OF LANTANA SPECIES IN NSW

Theflowerheadsofthisspeciesis10­40mmindiameterwhilethepeduncleis15­100mmlongand TAXONOMYborneintheleafstalk(KleinschmidtandJohnson1977;Conn1992;Munir1996).Thereareupto20flowersineachheadalthoughtheweedyvarietyhasfewerflowers(Everist1981;ParsonsandCuthbertson2001).Broadlyoval­shapedbractsarepresentunderneatheachflowerheadandare4­7mmlong.Eachcorolla(flower)is8­12mmlongand4­10mmindiameter(Conn1992;Munir1996;ParsonsandCuthbertson2001). Theflowersarepalepurple,mauveorlilacincolourwithapaleyelloworwhitecentrewhenyoung,becomingpurpleonmaturity(White1929;Everist1981;AuldandMedd1987;Conn1992;ParsonsandCuthbertson2001;O’Donnell2002).Munir(1996)statedthatflowersofL.montevidensisarealsopink,roseormagenta.OrnamentalvarietiesofL.montevidensishavelilacandwhiteflowersandaremorecompactplantswithsmallerleaves(vanOosterhout2004).Munir(1996)alsostatedthatyellowfloweringvarietieswerecommonincultivationbutthesearemisnamedastheyareahybndofL.camaraxL.depressa

TheellipsoidfruitofL.montevidensisisadrupe(incorrectlyknownasaberry),greenatfirst,2­8mmindiameterandpurple/blackorreddish/brownwhenmature(White1929;Everist1981;Conn1992;Munir1996;Conn1999;ParsonsandCuthbertson2001).Severalauthorsnotethatonlytheweedyvarietyofthisspeciesproducedfruitwhereastheornamentalvarietiesrarely,ifever,producedfruit(AuldandMedd1987;Everist1981;Swarbrick1986;O’Donnell2002).O’DonnellandPanetta(2000)indicatedthateachfruithasuptotwoseeds.Theseedsofthisspeciesareapalestrawcolourandupto4mmlong(ParsonsandCuthbertson2001).Thisspecieshasabrownwoodytaproot,stronglateralrootsandfinewhiteroots(ParsonsandCuthbertson2001).Thislargelignifiedtaprootisalsoknownasaxylopodiumanditactsasacarbohydratestorageorganallowingthespeciestoresistfire,droughtandherbicidedamage(O’DonnellandPanetta2000).

ThetwospeciesareeasilydifferentiatedwithL.montevidensishavingahorizontalgrowthhabit,branchesthatreadilyrootatthenodes,purplishflowers,leavesthatarelessthen25mmlongandthelowerleafsurfacethathasyellowtoorangeglands.Lantanacamaradoesnothaveanyofthesecharacteristics.

Family Verbenaceae TheVerbenaceaefamilyincludesaround75generaand3000speciesofherbs,shrubsandtreesoftropicalandsubtropicalpartsoftheworld(Conn1992).Conn(1992)recordedthepresenceof17generaand62speciesinAustraliaandninegeneraand19speciesarefoundinNSW.Hosking(inprep.)recognisedoneadditionalgenusandatleastfivemorespecies.AsidefromLantana,thereareanumberofAustraliangenerathatcontainweedyspeciesincludingPhyla(lippia),Verbena(purpletop/verbena)andStachytarpheta(snakeweed)(ParsonsandCuthbertson2001).

Lantana genus Dayetal.(2003)summarisedthecomplextaxonomyinherentinthefamilyVerbenaceaeandgenusLantana.TheLantanageneraiscomposedof150herbandshrubspecies,nativetotropicalAmericawithseveralnativetoAfricaandAsia(Conn1992;Dayetal.2003). Holmetal.(1979)recordedthatnineofthesespeciesareweedsinvarioustropicalandsubtropicalareas.FourdistinctsectionsinthegenusLantanaarerecognised(Munir1996). TwoofthesearesectionCalliorheaswhichincludesL.montevidensisandsectionCamarawhichincludesL.camara. ThehaploidchromosomenumbersforthesectionCalliorheasaren=12whilethoseinthesectionCamaraaren=11.

The Lantana camara species aggregate TheaggregatespeciesknownasL.camaraisa“variablepolyploidcomplexofinterbreedingtaxa”(Dayetal.2003;Sanders2006). Itcontainsawidediversityofvarietiesarisingfromhorticulturalandnaturalhybridisation,selectionandsomaticmutation(Swarbricketal.1998;Dayetal.2003,e.g.Figure2).IthasbeenwidelystatedthatthespeciesL.camaraarosefromhybridisationofanumberofsimilarorcloselyrelatedbutspatiallydistincttropicalAmericanspecies,orindeedfromcomplexesderivedfromthesespecies.Forexample,RAPD(RandomAmplifiedPolymorphicDNA)studieson30differentpopulationsofthepinkvarietyofL.camarafromtheeastcoastofAustralia(16.9­35.4oS)indicatedthatthesepopulationsprobably

REVIEW OF THE DECLARATION OF LANTANA SPECIES IN NSW 9

arosefromL.urticifolia(Scottetal.2002). ItislikelythatvarietiesofL.camarainothercountrieshavedifferentorigins.

Inreviewingthesituationoutlined,Sanders(2006)statedthat“currentusageofL.camaraincludesawidelycultivatedandnaturalisedcultigenspeciesofhybridoriginthatistaxonomicallydistinctfromL.camara”. ThatauthordescribedtheweedymaterialheexaminedbelongingtothiscultigenasanewspeciesLantanastrigocamara. Itisimportanttonotethatthisdoesnotimplythat“allweedyLantanainallcountriesisL.strigocamaraduetothemultipleimportationsofLantanatoeachcountry”(W.Palmerpers.comm.).Palmerfurtherstatesthatover250AustralianLantanaspecimenshavebeensenttoDrSanderstoascertainthecorrectspeciesnameofthismaterial.Inlieuofthatwork,thisdocumentusesthenameL.camara.

TheaggregatespeciesL.camarafreelyhybridiseswithinthespeciesitselfproducingfertilehybridsthatmayexhibitsomemorphologicaldifferenceswhencomparedtotheadultplants(Spies1984a).Membersofthisspeciescanalsohybridisebacktotheparentspeciesthatthecomplexwasderivedfrom(Swarbricketal.1998). Thetaxathathaveresultedfromthishybridisationhavebeengivenspecies,form,cultivar,biotype,subspeciesandvarietalstatus(Dayetal.2003).Despitethefactthatthebotanicalrankvarietas(variety)hasbeenoverusedforplantsthatareactuallycultivarsandthatvarietalnamesaresomewhatmisapplied(Sanders2006),thisreviewusesthewordvarietytodenotedifferencesinL.camara.

ThelargenumbersofvarietiesofL.camarafoundthroughouttheworldvaryintermsofploidy,bushshape,flowercolour,prickliness,leafshape,responsetoenvironmentalconditions,naturalenemies,herbicides,chemicalcompositionandtoxicitytoanimals(Swarbricketal.1998;Dayetal.2003). Everist(1981)recordedthatmostweedyvarietiesofL.camarainAustraliaweretetraploids(2n=44),butthatseveralweretriploids(2n=33),onewasadiploid(2n=22),whileanotherwasapentaploid(2n=55).Swarbricketal.(1998)concludedthatmostweedyvarietiesinSouthAfricaandIndiaweretetraploidswhilemostornamentalornon­weedyvarietiesweretriploids,andconcludedthatthiswasprobablyalsothecaseinAustralia.HexaploidvarietiesofL.camarahavealsobeenrecorded(NatarajanandAhuja1957;Spies1984a,b;Swarbrick

etal.1998). Thoseauthorssummariseinformationindicatingthattheseploidylevelsarisefromabreedingsystemthatissexual,semi­sexualandasexualorapomictic(notinvolvingthefusionofmaleandfemalegametesinreproduction),(KhoshooandMahal1967)andfrombothauto­andallo­polyploidywithinL.camaraandotherLantanaspecies(NatarajanandAhuja1957). Auto­polyploidyarisesfromhavingtwoormoresetsofchromosomesderivedfromthesamespecieswhileallo­polyploidyarisesfromhavingtwoormoresetsofchromosomesderivedfromdifferentspecies.

Someauthorshaveindicatedthatthereappearedtobenodirectrelationshipbetweenploidylevelormorphology,flowercolourandseedset(Swarbricketal.1998). Incontrast,TandonandBali(1955)notedthattriploidvarietiesweremorevigorous,hadlarger,thickeranddarkerleavesandproducedlargerandmorenumerousflowersincontrasttothediploidvarietiestheyexamined.Spies(1984a)alsostatedthatincreasesinseedmasswerealsoobservedasploidylevelincreasedfromdiploidtopentaploid.Incontrast,Swarbricketal.(1998)comparedthemorphologyof13weedyandornamentalvarietiesfromsoutheastQldandfoundsimilarityintheweedyvarietieswhichallhadlargerbushes,longerinternodelengthsandleafblades,pricklystemsandhighlevelsoffruitproduction.Fiveofthesixornamentalvarietieshadsmallerbushes,shorterinternodelengthsandleaves,lackedpricklesandwhiletheyproducedsimilarflowernumbers,theysetveryfewfruit. ThesixthornamentalvarietyDrapd‘orhadmuchlargerbushes,internodesandleavesbutlackedpricklesandhadlittleseedproductionindicatingitwasderivedfromadifferentsourcetotheornamentalandweedyvarietiesexamined.

BranchesofcertainL.camaraplantsoccasionallymorphologicallyreverttoothervarietiesandbredtruetothatvarietywhencuttingsaretaken(SmithandSmith1982;Dayetal.2003). Thereasonswhythesesomaticmutationsoccurarepoorlyunderstoodeventhoughthebrancheshavebeenrecognisedasareversiontoparentvarietiesthattheplantwasderivedfrom(SmithandSmith1982).Dayetal.(2003)postulatedthatenvironmentalswitcheschangedtheexpressionofchromosomesinthesebranchesandthatsincethesechangeshadimportantimplicationsinsuccessfulbiologicalcontrolandtoxicitymanagement,thattheyneededtobeinvestigated.

10 REVIEW OF THE DECLARATION OF LANTANA SPECIES IN NSW

ThecontinuedsaleofsupposedlysteriletriploidornamentalvarietiesofL.camarawillfurthercomplicatethetaxonomyofthisspecieswhereverweedyvarietiesarealsopresent(Dayetal.2003).Forexample,thereisevidencetosuggestthatthesesterilevarietiesarecapableofhybridisingwithfertileweedyvarietiesinAustraliaandSouthAfrica(SpiesandduPlessis1987;Neal1999).Therateatwhichthesenewgenecombinationsareintegratedintoweedypopulationsisnotknown(Neal1999;Dayetal.2003).

Varieties of L. camara in Australia AlthoughSmithandSmith(1982)recordedatleast29differentvarietiesofL.camarahadnaturalisedinAustralia,itishighlylikelythatthisnumberhaslongagobeenexceeded.Whilenaturalisedvarietiescanbepartiallydifferentiatedbyflowercolour,completedifferentiationreliesonarangeofcharacteristicsincludingflowercolourandsize,flowerlimbmorphologyandothervegetativecharacteristicssuchasthesize,shapeandcolouroftheleaves,thestructureofleafhairs,thorninessandthelengthofbracts(Everist1981;SmithandSmith1982;ParsonsandCuthbertson2001;vanOosterhout2004).Swarbrick(1986)notedthatmostweedyvarietiesofL.camarainAustraliahadlong,rambling,thornystemsandfreelysetseedwhiletheornamentalvarietiesatthattimetendedtobethornlessornearlysoandsetcomparativelylittleseed.

FourorfivemajorgroupsofL.camaracanbeseparatedviatheirflowercolourasfollows:­

n redfloweredvarietiesthatareorangeoryellowafteropeningbutchangetoashadeofredwhenmature.Therearetwosubgroupswithinthesevarieties,thosewithpinkedgesandthosethataredarkred.Everist(1981)andvanOosterhout(2004)givethesetwosubgroupsvarietystatus,thatistheredandpink-edgedredvarieties;

n pinkfloweredvarietiesthataredividedintotwosubgroupsbasedonflowersizethatissmallandlargeflowervarieties.ThetypicalweedyvarietyofL.camaraisasmallflowervariety,withflowersstartingaspaleyelloworwhiteandmaturingintopink(adifferentcolourtothatoutlinedabove);

nwhiteorpale-pinkfloweredvarieties;and

n orangefloweredvarietieswithflowersthatremaindeepyellow(similartoFigure7)toorangethroughouttheirlife.

ThesevarietieshavebeendocumentedinvanOosterhout(2004),pg.6.AsummaryofthemostdistinguishingcharacteristicsisoutlinedinTable1andFigure8.

Ensbey(2003)outlinesfourofthemostcommonL.camaravarietiesthatoccurinNSW.Theseincludethe:­

n commonpinkfloweredvarietythatisspreadthroughoutNSW(includedinthepinkfloweringvarietiesabove);

n commonpink-edgedredfloweringvariety(e.g.Figure3)thatisfoundonthenorthcoast,aroundKempsey,Dorrigo,Bellingen,CoffsHarbourandGraftonandonthecentralcoast(includedintheredfloweringvarietiesabove);and

n roundredandStaffordredvarietiesthatarefoundonthenorthcoast,aroundKempsey,Bellingen,CoffsHarbour(againincludedinthered­floweredvarietiesabove).

Figure7.A yellow flowering ornamental hybrid variety of Lantana camara planted in a median strip, Griffith (south western NSW, September 2005) (Source: S. Johnson, NSW DPI).

REVIEW OF THE DECLARATION OF LANTANA SPECIES IN NSW 11

Tab

le1

.D

istin

guis

hing

flor

ala

ndle

afc

hara

cter

istic

sof

L.c

amar

aan

dL.

mon

tevi

dens

isv

arie

ties

outli

ned

byv

anO

oste

rhou

t(20

04).

The

flow

erh

eads

oft

hew

eedy

var

ietie

sof

L.c

amar

aar

eill

ustr

ated

inF

igur

e8.

Alth

ough

ther

ear

eor

nam

enta

lvar

ietie

sof

L.c

amar

aw

itho

ther

flow

erc

olou

rs,o

nly

the

yello

wv

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tyh

asb

een

outli

ned

here

(sim

ilart

oFi

gure

7).

Info

rmat

ion

from

Eve

rist(

1981

)has

als

obe

enin

clud

edin

this

tabl

e.

12 REVIEW OF THE DECLARATION OF LANTANA SPECIES IN NSW

Spec

ies

Vari

ety

Bud

Flor

alc

hara

cter

isti

csM

iddl

eflo

ralr

ing

Out

erfl

oral

ring

Si

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aPi

nk(w

eed)

Pi

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Yello

wth

roat

,Pa

ley

ello

wp

etal

sO

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roat

,Pa

leo

rdar

kpi

nkp

etal

sLa

rge

Pale

gre

en

L.ca

mar

aW

hite

(wee

d)

Crea

m

Yello

wth

roat

Pa

ley

ello

wp

etal

sO

rang

eor

yel

low

thro

atLi

lac

peta

ls

Smal

l

L.ca

mar

aPi

nk-e

dged

Re

d(w

eed)

Pink

to

dull

red

Ora

nge

thro

at

Pale

yel

low

too

rang

epe

tals

Ora

nge

thro

at

Two­

tone

dpi

nkto

red

colo

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ls(i

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dar

kert

han

oute

r)

Smal

lD

arke

rtha

npi

nkL

.cam

ara

L.ca

mar

aRe

d(w

eed)

Bl

ood

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Yello

wth

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Pa

ley

ello

wp

etal

sRe

dth

roat

Re

dpe

tals

La

rge

Dar

kgr

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O

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pet

als

Ora

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thro

at

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nge

peta

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Smal

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leg

reen

L.ca

mar

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llow

(o

rnam

enta

l)G

reen

ye

llow

Br

ight

yel

low

thro

at

Brig

hty

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wp

etal

sBr

ight

yel

low

thro

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Brig

hty

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wp

etal

sSm

alle

rtha

nw

eedy

var

ietie

s

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onte

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nsis

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le

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le

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als

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als

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lert

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a

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atLi

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iety

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the

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ies

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a b

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e

Figure8.Schematic representation of the flower heads of the weedy varieties of Lantana camara (extracted from van Oosterhout 2004, used with permission). ThevarietiesareasfollowsFig.8apinkflowered,Fig.8bwhiteflowered,Fig.8cpink­edgedredflowered,Fig.8dredfloweredandFig.8eorangeflowered.

REVIEW OF THE DECLARATION OF LANTANA SPECIES IN NSW 13