review of the declaration of lantana species

Review of the declaration of

Lantana speciesin New South Wales

N S W D P I

Review of the declaration of

Lantana speciesin New South Wales

NewSouthWalesDepartmentofPrimaryIndustriesOrangeNSW2800

Frontispiece.AfloweringandfruitingbranchofthecommonpinkvarietyofLantanacamara,nearCopmanhurst(NSWnorthcoast,October2005)(Source:S.Johnson,NSWDPI).

©StateofNewSouthWalesthroughNSWDepartmentofPrimaryIndustries2007.Youmaycopy,distributeandotherwisefreelydealwiththispublicationforanypurpose,providedthatyouattributeNSWDepartmentofPrimaryIndustriesastheowner.

ISBN9780734718891

Disclaimer: The information contained in this publication is based on knowledge and understanding at the time of writing (December 2007). However, because of advances in knowledge, users are reminded of the need to ensure that information upon which they rely is up to date and to check currency of the information with the appropriate officer of New South Wales Department of Primary Industries or the user’s independent adviser. Job number 7262

ThisdocumentwaspreparedbyDrStephenJohnsonWeedEcologistWeedsUnitBiosecurity,ComplianceandMineSafety

Telephone:0263913146Facsimile:0263913206LockedBag21ORANGENSW 2800

Figure1.WhiteandpurplefloweringvarietiesoftheornamentalLantanamontevidensisplantedinamedianstrip,Griffith(southwesternNSW,September2005)(Source:S.Johnson,NSWDPI).

iv REVIEW OF THE DECLARATION OF LANTANA SPECIES IN NSW

EXECUTIVESUMMARY 1

SCOPEOFTHISREVIEW 3

REVIEWOFTHEDECLARATIONOFLANTANASPECIESINNSW 5

NOMENCLATURE 5

CONTENTS

Lantanacamara 5

Lantanamontevidensis 5

SPECIESDESCRIPTIONS 5Lantanacamara 5


TAXONOMY 9FamilyVerbenaceae 9

Lantanagenus 9

TheLantanacamaraspeciesaggregate 9

VarietiesofL.camarainAustralia 11

VarietiesofL.montevidensisinAustralia 14

ORIGIN 14

LIFECYCLE 16

DISPERSAL 19

GROWTHANDDEVELOPMENT 20

HABITAT 20Climaticrequirements 20

Soils 21

DISTRIBUTIONINAUSTRALIA 22Currentdistribution 22

Lantanacamara 22


Potentialdistribution 23

IMPORTANCE 24Detrimental 24

Pastures 25

Naturalecosystems 26

Forestry 28

Plantationandothercrops 28

Railwayandserviceproviders 28

Allelopathy 28

Hostsforotherpests 28

Humaneffects 29

Beneficial 29

Ornamental 29

REVIEW OF THE DECLARATION OF LANTANA SPECIES IN NSW v

Otherhumanuses 29

Alternativefoodandhabitatsourcesforwildlife 29

Otherbenefits 29

ThesaleofornamentalLantanaspecies 30

NegativeimpactsoftheremovalfromsaleofLantanaspecies 31

LEGISLATION 31Statedeclaration 31

AWeedofNationalSignificance(WoNS) 32

Benefitsthatmayaccruefromcontinuedlegislativecontrol 32

CONTROL 32Prevention 33

Herbicides 33

Othermanagement 34

Fire 34

Mechanicalclearing 35

Cultivation 35

Handremovalandflameweeding 36

Grazingmanagement 36

Revegetation 36

Biologicalcontrol 37

IntegratedmanagementofL.camara 37

Pasturesituations 38

Naturalecosystems 38

Controllingregrowth 39

Identifyingthecausesofinfestation 39

IntegratedmanagementofL.montevidensis 39

SOCIALLIMITATIONSTOCONTROL 40ThewidespreadacceptanceofL.camara 40

IssuesinvolvedwiththesaleofLantanaspecies 40

ConstraintstomanagingL.camara 41

Motivatinglandmanagerstoundertakemanagement 42

RECOMMENDATIONS

FURTHERRESEARCHNEEDS

INFORMATIONREQUIREDFROMCONSULTATION

42

43

44

CONCLUSIONS 44

REFERENCES

APPENDICES

48

59

vi REVIEW OF THE DECLARATION OF LANTANA SPECIES IN NSW

APPENDICESAppendix1 DeclarationsofLantanaspeciesacrossAustralia.Appendix2 RecommendationsfordeclarationsofLantanaspeciesinNSW.

ABBREVIATIONSACT AustralianCapitalTerritoryNSW NewSouthWalesNT NorthernTerritoryQld QueenslandSA SouthAustraliaVic VictoriaWA WesternAustralia

ACKNOWLEDGMENTSThisreviewhasbenefitedgreatlyfrominformationandcommentsprovidedbystafffromNSWDepartmentofPrimaryIndustries,BiosecurityQueensland,theUniversityofQueenslandandtheBotanicGardensTrust.

Figure2.A red and yellow flowering ornamental Lantana hybrid 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 vii

EXECUTIVESUMMARY aswellasotherareasreducingplantandanimalbiomass,abundanceandbiodiversity,generally

ThisdocumentexaminedthebasicbiologyandmanagementofLantanaspeciesinNSW.Twospecieshavenaturalised,thesebeingL.camara(lantana),apolyploidspeciesaggregatecomposedofatleast29differentweedyandornamentalvarietiesandL.montevidensis(creepinglantana)ofwhichoneweedyandatleastseveralornamentalvarietiesarepresent.VarietiesofL.camararesponddifferentlytoarangeofenvironmentalandmanagementconditionsandpractises.SpeciesofLantanaareabletohybridiseandmanysuchhybridsaregrownforhorticulturalpurposes.Despiteclaimstothecontrary,allvarietiesarefertiletosomeextent.AlthoughthereisextensivespreadofbothspeciesineasternAustralia,furtherspreadislikelyinbothcleanandinfestedareas.

Bothspeciesformdense,multibranchedthicketsmakingaccessandmanagementdifficult.Floweringandfruitsetmayoccurallyearroundwithdispersaloffruitbyarangeofbirdsandanimals.Bothspeciesareabletospreadvegetativelytosomedegree.

Lantanaspeciesinvadepastures,awiderangeofnaturalecosystems,forestryandplantationcrops,

causinglivestocktoxicityifeaten,oftenpreventingregenerationofnaturalareas,andaffectingtourism,recreationalandaestheticvalues.Alternatively,bothspecieshavebeenwidelyplantedasornamentalandhedgeplants,andasamenityplantings.Plantsmayprovidealternativehabitatforanimalsandprovideasourceofessentialoils.

LantanacamaraisaWeedofNationalSignificanceandhasbeenbannedfromtradeanddistributioninallstatesandterritoriesinAustralia.VarietiesofL.camaraaredeclarednoxiousinpartsofcoastalNSW,whilebothspeciesaredeclaredinQldandtheNT.ManagementofbothLantanaspeciesisdifficultwithintegratedcontrolprogramsneededincludingpreventative,chemical,mechanicalandbiologicalmeasureswhilefire,ploughing,handpulling,revegetationandgrazingmanagementareusefultools.

RecommendationsarisingfromthisdocumenthavebeenmadetotheNoxiousWeedsAdvisoryCommitteeandarecontainedinAppendix2.

REVIEW OF THE DECLARATION OF LANTANA SPECIES IN NSW 1

REVIEW OF THE DECLARATION OF LANTANA SPECIES IN NSW2 REVIEW OF THE DECLARATION OF LANTANA SPECIES IN NSW2

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 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


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).


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).


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


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.


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).


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

arie

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.


Spec

ies

Vari

ety

Bud

Flor

alc

hara

cter

isti

csM

iddl

eflo

ralr

ing

Out

erfl

oral

ring

Si

zea

ndC

olou

rofL

eave

s

L.ca

mar

aPi

nk(w

eed)

Pi

nk

Yello

wth

roat

,Pa

ley

ello

wp

etal

sO

rang

eth

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

ur

peta

ls(i

nner

dar

kert

han

oute

r)

Smal

lD

arke

rtha

npi

nkL

.cam

ara

L.ca

mar

aRe

d(w

eed)

Bl

ood

red

Yello

wth

roat

Pa

ley

ello

wp

etal

sRe

dth

roat

Re

dpe

tals

La

rge

Dar

kgr

een

L.ca

mar

aO

rang

e(w

eed)

O

rang

eYe

llow

too

rang

eth

roat

Ye

llow

pet

als

Ora

nge

thro

at

Ora

nge

peta

ls

Smal

lPa

leg

reen

L.ca

mar

aYe

llow

(o

rnam

enta

l)G

reen

ye

llow

Br

ight

yel

low

thro

at

Brig

hty

ello

wp

etal

sBr

ight

yel

low

thro

at

Brig

hty

ello

wp

etal

sSm

alle

rtha

nw

eedy

var

ietie

s

L.m

onte

vide

nsis

Purp

le

(wee

d)

Purp

le

Whi

teth

roat

Pu

rple

pet

als

Whi

teth

roat

Pu

rple

pet

als

Smal

lert

han

L.ca

mar

a

L.m

onte

vide

nsis

Lila

c(o

rnam

enta

l)Li

lac

Whi

teto

yel

low

thro

atLi

lac

peta

ls

Whi

teto

yel

low

thro

atLi

lac

peta

ls

Larg

erth

anw

eedy

var

iety

of

the

spec

ies

L.m

onte

vide

nsis

Whi

te(o

rnam

enta

l)W

hite

to

crea

m

Yello

wth

roat

Whi

tep

etal

sPa

ley

ello

wth

roat

Whi

tep

etal

sLa

rger

than

wee

dyv

arie

tyo

fth

esp

ecie

s

a b

c d

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.


SmithandSmith(1982)alsorecordedthreeothervarietiesofL.camarainNSW.ThefirstisaredfloweredvarietyknownaspaleStaffordredaroundKempsey,thesecondanorangeredvarietyknownasoblongredwestofGraftonalongtheGwydirhighway(apparentlynotabundantorwidespreadbutperhapsfoundelsewhereinthearea)andthethird,anorangefloweredvarietyknownastrueorangethatisoccasionalnearPalmBeach,northofSydney.Thislastvarietyiscommonlycultivated.

ThevalueofcategorisationofvarietiesofL.camaraintobroadflowercolourgroupshasbeenquestionedbyanumberofauthorsforvariousreasons.Thesereasonsincludewidevariabilityincolourwithinandbetweentaxamakingdifferentiationdifficult(Dayetal.2003;Sanders2006). Inaddition,therearedifferencesbetweensamecolouredvarietiesintheirtoxicitytolivestockandtheirsusceptibilitytobiologicalcontrolagentsorherbicidesbetweendifferentregions(DiatloffandHaseler1965;Seawright1965;Everist1981;Dayetal.2003). Furthermore,differentvarietiesofL.camarasurviveindifferentclimaticzonesandecosystemsinAustralia(Clarketal.2004).

Scottetal.(1997)analysedthegeneticrelationshipbetweenpinkandpinkedgedredweedyvarietiesofL.camarafromfourregionsalongtheeastcoastofAustraliaandfoundthatgeographicalproximitywasmoreimportantthanflowercolourindefininggeneticsimilaritybetweenpopulations.Whiletherewassomegeneticisolationbetweenvarietieswithinaregion,flowercolourhadlittlephylogeneticsignificancebetweenregions.TheoveralllevelofdifferentiationinL.camaravarietiesmakesitdifficultformanylandmanagersandscientistsaliketomanagethisweedandtopromoteconsistentweedmanagementmessages.

VarietiesofL. montevidensisinAustralia

Henderson(1969)recordedthattwovarietiesofL.montevidensisoccurredinAustralia,eachwithadifferentploidy.Thecommongardenvarietytestedwasatriploid(2n=36)whiletheweedyvarietywasatetraploid(2n=48).ItisnotknownifthefurtherintroductionofL.montevidensisvarietiesintoAustraliahasincreasedthenumberofploidylevels.Somedifferencesbetweentheweedyandtwoornamental

varietiesofL.montevidensis,onewithlilacflowersandtheotherwithwhiteflowers,areoutlined(Table1).Further,Neal(1999)indicatedthatcrossesbetweenL.montevidensisornamentalvarietiesmayhaveoccurredinAustralia.AvariegatedleafvarietyofL.montevidensiswithlilacflowersisalsocultivatedinthenewBotanicGardensinBrisbane(Swarbrick1986).

ORIGINThetwospeciesofLantananaturalisedinAustralia,L.camaraandL.montevidensisarebothnativetotropicalSouthAmerica.LantanacamarawasinitiallyintroducedintoEuropefromBrazilasanornamentalaround1636(Howard1969).FurtherintroductionsintoEuropecontinuedthroughoutthe17th,18thandespeciallythe19thcenturieswhereintroductions,ensuinghybridisationandvegetativereproductionofsomaticmutationsresultedintheregistrationof397newvarietalnamesinnurserycataloguesfrom1850to1900(Howard1969;Swarbrick1986).Inexcessof630varieties(not650asclaimedbymanyauthors)ofLantanahavebeendevelopedforhorticultureworldwideasaresultofhybridisation(Howard1969),althoughanumberofthesenamesmaybemisspellingsandsynonymsbecausedetailedbotanicaldescriptionsaregenerallyunavailable.

FromEurope,L.camarawasintroducedintoanumberofcountriesthatwereformerlypartofcolonialempires.Thespeciescannowbefoundgrowingasaweedinatleast60countriesorislandgroupsincludingtheUnitedStatesofAmerica,manycountriesinSouthAmerica,aroundthewesternMediterranean,throughoutAfrica,centralandsoutheastAsiaandvariouscountriesaroundthePacificoceanincludingNewZealandandAustralia(Holmetal.1979;Webbetal.1988;Dayetal.2003). AlthoughL.camaraiswidelygrownasanornamentalinothercountries,suitableterrestrialhabitatsfortheweedonlyoccurthroughouttropical,subtropicalandwarmtemperateareas(Figure9).

ThefirstrecordofL.camarainAustraliawasin1841intheoldAdelaideBotanicGardens(Bailey1841).LantanacamarawasfirstrecordedincultivationinNSWin1843nearSydney(Anon.1843,inMichael1972). Thespeciesquicklyspreadnorthwardsandwasrecordedasnaturalisedinthe1850’s,inBrisbane


in1861,andintheHastingsandClarencecatchmentsofNSWinthelate1860’s(Swarbrick1986).Bailey(1897)describedL.camaraas“ahugeramblingpricklybush...ithasspreadtoanalarmingextent,andformsimpenetrablethicketonthebanksofstreams,desertedfarms,andtheedgesofscrubs”inthePortJacksonandBrisbaneareas.Similarly,concernoverL.camararesultedinitbeinglistedasoneofthetenworstweedsinNSWin1895(Maiden1895)and1920(Maiden1920).

LantanacamarahasbeengrowninVicsince1852andWAsince1875,butmayhaveonlybeenintroducedtotheNTduringthe1930’sor1940’s(Swarbrick1986).Furtherexpansionintopreviouslyuninfestedareasislikelytostillbeoccurring,oftenasaresultoflandclearingandotherhumandisturbance(HumphriesandStanton1992;Swarbricketal.1998).

NumerousfurtherintroductionsofL.camarahaveoccurredinAustraliaasgardenplantsineasternAustralia(SmithandSmith1982),andprobablyintheNTandWA(Swarbricketal.1998). SmithandSmith(1982)consideredthat19varietiesweresufficientlycommonineasternAustraliatobeconsideredproblematicweedsortoxicplants.BothnurserystockandseedsofLantanaspecieswere,untillate2006,stillpermittedforimportintoAustralia(AustralianQuarantineandInspectionService2007).

Lantanamontevidensiswasalsowidelydistributedbymaninthe19thcentury(White1929;Swarbrick1986).Althoughmostauthorsgenerallyagreethatthenative

rangeofL.montevidensisincludessouthernBrazil,thereissomeconjectureiftherangealsoincludesUruguay(Everist1981)and/ornorthernArgentina(Dayetal.1999).

Swarbrick(1986)recordedthatthespecieswasintroducedintoEuropefromMontevideoin1822andtoAustraliaby1851(Johnson1872;Shepherd1851,bothinSwarbrick1986;Munir1996).Thespecies(L.sellowii)wasrecordedincultivationinsouthwestSydneyin1857(Anon.1857).Holmetal.(1979)indicatedthatL.montevidensishasonlybeenrecordedasaweedinAustralia,andperhapsinFlorida(Bailey1963)eventhoughitiswidelyplantedasanornamentalorhasnaturalisedinvariouspartsoftheworldincludingAustralia,NewZealand,AfricaandpartsofIndia(Everist1981;Webbetal.1988;Dayetal.2003;vanOosterhout2004).Suitableterrestrialhabitatsforthespeciesgenerallyoccurinsubhumidandsemiaridregionsofthetropicsandsubtropics(Everist1981;ParsonsandCuthbertson2001).

LantanamontevidensiswaswidelydispersedinAustraliaappearingintheMelbournebotanicgardensin1852andtheninmanybotanicgardensandnurserycatalogues(Swarbrick1986).ThatauthorstatedthatthespecieswasfirstfoundinAdelaidein1859andBrisbanein1875.ThefirstmentionofL.montevidensisaseitheragardenescapeorweedisbyBaileyandTenisonWoods(1879)whorecordedthespeciesintheBrisbaneriverarea.Swarbrick(1986)alsonotedaBrisbaneherbariumspecimenthatindicatedthatL.montevidensishadnaturalisednearIpswichin1888.

Figure9.Suitable terrestrial habitats for L. camara throughout the world. (Source: Swarbrick et al. (1998), used with permission).


Swarbrick(1986)recordedanumberofobservationsofL.montevidensisinsoutheasternQldfrom18831909, LIFECYCLEgenerallyofplantsthatweregardenescapesfromnearbytownsintoneighbouringpastures.Swarbrick(1986)notedthatrapidspreadoftheweedincoastalQldoccurredafter1900withherbariumrecordsfromGayndahin1913and1917,nearCairnsin1918andatRockhamptonin1925.The1917Gayndahrecordissignificantbecausethespeciesisdescribedasa‘verycommonweed’(O’Donnell2002).Bythe1950’sand1960’sL.montevidensishadbecomewidespreadthroughoutcoastalandsubcoastalQld,especiallyintheBurnettdistrict(O’Donnelletal.1999). O’Donnell(2002)recordsthatbythe1980’sand1990’sthespecieshadreachedlevelswheretheongoingviabilityofgrazingenterprisesinsomeareasofQldwassignificantlyaffected.

ItispresentlyunclearwhenL.montevidensiswasfirstnotedasaweedprobleminNSW.RecordsattheRoyalBotanicGardensinSydneyindicatedthatinfestationsofthespecieswerepresentasearlyas1952nearConcordinSydneyandaround1963nearCasinoontheNorthCoast.IncontrasttothewidespreaddistributionofL.montevidensisinQld,onlylimitednaturalisationsofthespecieshaveoccurredinNSWandtheseweregenerallyrestrictedtotheSydneybasin,aroundCasinoandMurwillumbahonthenorthcoastandtoTamworth.

AlthoughanumberofauthorsindicatethatonlyonevarietyofL.montevidensishasbeenintroducedintoAustraliaasanornamental(AuldandMedd1987;Conn1992;ParsonsandCuthbertson2001),vanOosterhout(2004)recordedornamentallilacandwhitevarietiesaswellasthecommonpurplefloweredweedyvariety(Table1;Figure1).

LantanacamaraxL.montevidensishybridshavebeendevelopedforuseinhorticulture.Forexample,Howard(1969)statedthattheL.montevidensishasbeenusedtoproducesuchhybridssincetheearly19thcenturyandHammer(2004)indicatedthatthispracticecontinues.Inparticular,L.montevidensisiscommonlyusedtoachievenewhorticulturalvarietieswithlow,moundingandtrailinggrowthhabits.Furthermoreanumberofauthorsnotedthehybridisationofbothpreviouslygeographicallyseparatedspecies(Sanders1989,inDayetal.2003)andvarieties(Spies1984b;Dayetal.2003)whengrowninthesamelocation.

SeedsofL.camaraandL.montevidensisgerminateatanytimeoftheyearprovidedthatthereissufficientsoilmoisture(ParsonsandCuthbertson2001;O’Donnell2002). Lantanacamaraseedsneedhighlightconditionsforgerminationandearlygrowth(GentleandDuggin1997b;DugginandGentle1998;Stock2004)withseedlingsunlikelytosurvivebeneathdenseinfestations.

ThelargestseedlingflushesofL.camaraoccurafterinitialsummerstorms,particularlyinhigherrainfallareas(ParsonsandCuthbertson2001;vanOosterhout2004). vanOosterhout(2004)indicatedthatincreasesinlightintensityandtemperaturearealsoimportantforgermination.Thesefactorsmayoccurasaresultofhumandisturbancee.g.clearing,burning,orpestanimalactivitysuchaspigrootingorrabbitburrowing.Germinationfromtheseeventswilloccurifsoakingrainsfollow,especiallyinareasthathavehighersoilmoisture,forexamplearoundcreeks,rivers,gulliesanddams(vanOosterhout2004).

Fieldgerminationratesrangebetween4and63%(GentleandDuggin1997b;Dayetal.2003).Germinationislikelytobeincreasedthroughanincreaseinnutrientsviaburning,andbybiomassremovalandsoilscarificationassociatedwithgrazingandfire(GentleandDuggin1997b;DugginandGentle1998).

SpecificstudiesontheconditionsrequiredforthegerminationofL.camaraseedsaresomewhatlimited.OnenotableexceptionisthestudyofVivianSmithetal.(2006)whichindicatedthattemperatureregimesbetween15and25oCweresuitableforgerminationofpinkandpinkedgedredvarietiesofL.camara. ThoseauthorsfoundthattheseedfeedingflyOphiomyialantanaedamagedtheseedofpinkedgedredvarietiesresultinginreducedgerminationandseedlingemergence.Incontrast,theemergenceofthemorecommonpinkvarietywasincreased,possiblyaresultofthedamagereducingdormancyinthisvariety.

O’DonnellandPanetta(2000)indicatedthatseedsofL.montevidensisrequiredlighttogerminateandthatseedburiedto12cmdidnotgerminate.O’Donnell(2002)presentedsomeevidencetosuggestgerminationflushesoccurredwhengoodsoilmoisture


conditionscoincidedwithcoldnighttemperaturesclosetoorbelowfreezing.Thatauthoralsofoundthatexposuretosmokefrompasturefiresfor5to15minutesalsostimulatedthegerminationofL.montevidensisseeds615times.Germinationandsubsequentgrowthwasenhancedinpasturegapsrangingfrom540cm(O’Donnell2002).

TheearlygrowthofbothL.camaraandL.montevidensisisslowwhilethetaprootandlateralrootsaredeveloped(ParsonsandCuthbertson2001;O’Donnell2002),forexample,seedlingsofL.montevidensismayonlygrowatratesof1cm/month(O’Donnell2002).StudiesbyStockandWild(2006)foundthatseedlingsofL.camaragrowtallerandwiderundertherelativeopencanopyoftheweedasopposedtothosegrowingunderneaththecanopyofnumberofsubtropicalrainforestspecies.Seedlinggrowthwasmostlimitedunderrainforesttreespeciesthatformedtheoriginalor‘oldgrowth’forests.

AlthoughSahuandPanda(1998)notedlowratesofseedlingandmatureplantmortalityinL.camara,similarstudiesdonotappeartohavebeenrepeatedunderAustralianconditions.O’Donnell(2002)indicatedthatmortalityofL.montevidensisseedlingsoccurredviaawiderangeoffactorsincludingmoisturestress,physicaldisturbance,fire,herbicides,cattletramplingandperhapsgrazing.Seedlingsthusestablishedmorereadilyinprotectedareassuchasstonyoutcropsandareasprotectedfromcattle,underneathfencesandfallentimber,withintussocksofgrassesthatwerenoteatenandinpasturesthatwerenotsubjecttohotfires.

ThestemsofL.camaraplantsbegintoentwineformingthicketswithintheirfirstseasonofgrowthbuttheydonotproduceflowerswithinthisseason.Theonsetofwintereitherreducesorstopsgrowthanddevelopment.AlthoughglasshouseplantsofaL.camaraxL.depressahybridandL.montevidensishavebeenshowntoproduceseedswithinayear(Neal1999;O’Donnell2002),thedevelopmentofseedlingsofL.camaraandL.montevidensisismuchslowerinthefield(ParsonsandCuthbertson2001;O’Donnell2002).Dormantseedlingsreshootinthefollowingspringandstarttoflowerinlatespring(L.montevidensis)orearlysummer(L.camara)inthesecondseasonofgrowth.ParsonsandCuthbertson(2001)recordedthatestablishedplantsofL.camaraflowerthroughout

summerandthenuntilMarchorApril.PlantsofL.camarawerereportedasfloweringbetweenSeptemberOctoberthroughtoMarchAprilinensuingyearsintemperateareasfromsouthernVictonorthernNSW(vanOosterhout).

Swarbricketal.(1998)summarisedvariousliteraturethatstatedthatfloweringinL.camaracanoccuryearroundunderconditionsofhighavailablesoilmoisture,airhumidityandtemperatureandwhenplantsgrewinwelllitsituations.ThisgenerallyresultsinfloweringandfruitsetallyearincoastalareasofQldandnorthernNSW,thatisinsubtropicalandtropicalareas(vanOosterhout2004).Incontrast,Webbetal.(1988)indicatedthatL.camaracanflowerallyearinNewZealand.Distinctflushesoffloweringfourtosixweeksafterrainfalleventsexceeding25mmfollowedbyfruitsetaremorecommonindrierinlandareas(Swarbricketal.1998;vanOosterhout2004).

AnumberofauthorsrecordthatestablishedplantsofL.montevidensisflowermostoftheyearinAustraliaandNewZealand(Conn1992;Webbetal.1988;ParsonsandCuthbertson2001).Morespecifically,O’Donnell(2002)recordedthatL.montevidensisfloweredinresponsetorainfallwithheaviestfloweringfollowingaprotracteddryperiod,forexampleafterthefirstspringrainfallbreak.

NewlyopenedflowersofL.camaraattractarangeofinsectpollinatorswhichresultinselfandcrosspollination.Theseinsectsincludebutterflies,moths,bumble,honeyandothertypesofbees,andthrips(Dronamraju1958;Schemske1976;Kugler1980inSwarbricketal.1998;Clemson1985;MathurandMohanRam1986).KhoshooandMahal(1967)statedthatbothpollenandseedviabilityresultedfromopenpollinationofplantsofallploidylevelstheyexamined(diploidspentaploids).Theysuggestedthatnormalsexualunionwasresponsibleindiploidsbutthatapomixis(reproductionwithoutfertilisationofgametes)wasresponsibleinplantsofotherploidylevels.HoweverthisisunlikelytobethecaseasthereviewofliteraturebyStirton(1977)foundnoevidenceforapomixis.Instead,SpiesandStirton(1982)foundnormalsexualembryosindiploid,triploidandtetraploidplantsbutnonormalsexualembryosineitherpentaploidorhexaploidvarieties.ThereareconflictingreportsabouttheabilityofL.camaratoselfpollinate(Dayetal.2003).


Spies(1984a)recordeddifferencesinpollenviabilitywhenstudyingdiploid,triploidandtetraploidplantsinSouthAfrica.Forexample,theaverageandrangeinpollenviabilitiesfordiploidplantswas59.95%(30.8585.25%),fortriploidplantswas36.21%(27.3044.38%)andfortetraploidplantswas63.3%(16.0183.22%). Neal(1999)recordedthat65%ofpollenfromtheweedycommonpinkvarietyofL.camarawasviableincontrasttothe2%ofpollenfromaL.camaraxL.depressahybrid.

Insummarisingseveralreferencesandotherobservations,Swarbricketal.(1998)statedthatanaverageof36.748%fruitsetoccurredinopenpollinatedL.camarainflorescences,witheachaveraging32.6flowers.Becausethereareusuallytwoinflorescencespernode,thetotalnumberoffruitproducedundergoodgrowingconditionsmaybeashighasseveralthousandpermetre.DatafromthePhilippinesindicatedthat24fruitswereproducedperinflorescenceandthattherewere511inflorescencesperplantresultingin12,264fruitsperplant.Thenumberoffruitthathavefullyformedembryosmaybeaslittleas36%however(Graaff1987). WhiletherehavebeenfewspecificstudiesontheinfluenceofthefruitpulponthegerminationofL.camaraseeds,bothGraaff(1987)andSwarbricketal.(1998)presentevidencethatadelayorinhibitionmechanismoccurs.Neal(1999)alsoindicatedthat0.162%offloretsproducedseedsinthesupposedlysterileL.camaraxL.depressahybridsheexamined.

TheseedbankdensitiesofL.camarahavebeenthesubjectoflimitedresearch.WhileGentleandDuggin(1997b,1998)indicatedthatlowdensitiesof1.43.4seeds/m2werefound,muchlargerdensitiesof5993674seeds/m2havebeendeterminedfromanotherstudy(VivianSmithetal.2006). Itisimportanttonotethatonlyasmallproportionoftheseseeds(616%)wereviableaseithergerminableordormantseeds.

vanOosterhout(2004)summarisedresearchthatindicatedupto50%ofL.camaraseedwillremainviableunderdryconditionsforuptotwoyearsafterdispersalwhileotherstudieshaveindicatedthatviableseedmaypersistforuptofiveyears(G.VivianSmithpers.comm.).Despitethesestudies,Dayetal.(2003)indicatedthatverylittlewasknownabouttheseedbankdynamicsofL.camara. Thesestudiesareimportantastheyplaceatimeframeforongoingcontrolfollowingtheremovalofdenseinfestations.

O’DonnellandPanetta(2000)indicatedthattheviabilityofL.montevidensisseedsdecreasedtobetween3044%afteroneyearofburialat12cmofdepthwhilethoseplantedonthesoilsurfacehadonly1018%viabilityafterasimilartime.O’Donnell(2002)addedthatallsurfacesownseedwasdeadaftertwoyearsbutthatupto20%ofburiedseedwasstillviable.TheseresultsindicatethattheseedofL.montevidensisisrelativelyshortlivedinthesoil.Inaddition,theseedpulpormesocarpinhibitedseedgerminationupto100daysaftersowingbuthadnoimpactafteroneyear,probablyasaresultofnaturaldecay(O’DonnellandPanetta2000).Aproportionofseedswillgerminatewiththepulpsurroundingthemhowever.

Plantsgrowrapidlyunderfavourableconditionsofsoil,humidity,temperatureandlightandmaygrowyearroundundertheseconditions.InmanyareasL.camaraisseasonallydefoliatedbybiologicalcontrolagentsbutplantsrecoveroncecoolweatherresultsinawaneininsectnumbers(DayandHannanJones1999;Dayetal.2003). Lantanacamaraisaperennialplantshootingorregrowingvigorouslyfromdormantbasalbudsifshootsareremovedbyfire,herbicide,physicalremoval,droughtoriffrostedoff.Inthesecasesplantscanflowerwithinseveralmonthsunderfavourableconditions(Swarbricketal.1998). PlantsofL.camaraplantsareverylonglivedwithconstantrenewalofstemsfromtheplantbase,especiallyifrainfalloccursafterstemdeath(vanOosterhout2004).

Plantswilldieunderprolongedstressfulconditions,forexample,intenseorprolongeddrought,orfromshading(Swarbricketal.1998). ParsonsandCuthbertson(2001)statedthatnewcanesofL.camaraareproducedfromthecrownofexistingplants,fromlateralrootsinearlyspringandthatshallowlateralssuckerifdamagedorbroken.Suckeringmayoccurafterplantsarephysicallyremoved,oftenbybulldozing,withlargepiecesoflateralrootsreshooting,especiallyundergoodsoilmoistureconditions(SaintSmith1964;Swarbrick1982;WaterhouseandNorris1987;ParsonsandCuthbertson2001). OneestimateplacesthenumberofnewshootsfoundafterclearingasiteinsclerophyllwoodlandinnorthernNSWthatarosefromsuckeringat30%(B.Johnsonpers.comm.).TheseobservationsareincontrasttoSwarbricketal.(1998)whostatedthatnosuckeringhasbeenobservedbythoseauthors.Furthermoretheyindicatedthatprostratestemswithadventitiousrootsmayhavebeenmistakenforrootsuckersaftertheyhad


producedvigorousshootsfromthenodes.Giventhedivergenceofopinionontheroleofsuckeringfromlateralroots,furtherresearchisrequired.Lantanacamaracanbepropagatedfromstemtiporhardwoodstemcuttingsorevenleafmaterialifplantedintomoistrootingmediaorsoil(Swarbricketal.1998;Neal1999).

Lantanamontevidensisreproducesbyseedandbylayeringrootingatthestemnodeswhencoveredinmoistleaflitterorsoil(ParsonsandCuthbertson2001). Henderson(1969)recordedapollenviabilityofapproximately65%inL.montevidensisincontrasttotheornamentalvarietyHenderson(1969)examinedwhichwaslessthan6%.Similarly,Neal(1999)indicatedthatpollenviabilitiesof82%fromtheweedyvarietypopulationsofL.montevidensissheassessedand1416%fromthepopulationsoftwoornamentalL.montevidensisvarieties.ThedifferencesbetweenthestudiesarelikelytobeduetotherestrictednumberofpopulationscollectedbytheHendersonstudy.

PlantsofL.montevidensisareabletoflowerandsetfruitthroughouttheyear,generallyfromearlysummermidwinterinQld(O’Donnell2002).SeedproductioninL.montevidensisgenerallyoccursfiveweeksafterflowering(O’Donnell2002).VariousauthorsincludingHenderson(1969)statedthattheweedyvarietyofL.montevidensishaveahighproductionoffertileseed.Forexample,O’Donnell(2002)foundthatyearlyseedproductionvariedbetween4,965and5,175seeds/m2andwhileeachdrupehadtwoseeds,generallyonly30%ofdrupesproducedasecondseedling.Attwater(1980)reportedthatthewoodyseedcoatofL.montevidensiswasimpermeabletocertaingassesandchemicalsandhencepreventedseedgermination.

Henderson(1969)andWebbetal.(1988)recordedthatornamentalvarietiesofL.montevidensisdidnotproducefruitinAustraliaandNewZealandrespectively.Incontrast,Neal(1999)demonstratedthat0.421%offloretsofthepurplefloweringornamentalvarietysheexaminedproducedfruitwhileleafmaterialofbothornamentalandweedyvarietiesofL.montevidensiswasabletogiverisetonewplants.PopulationsproducingfruitweregenerallyincloseproximitytoweedypopulationsofL.camaraand/orL.montevidensis.

ItisnotknownhowlongindividualplantsofL.camaramaylive(vanOosterhout2004)butO’Donnell(2002)suggestedthatplantsofL.montevidensismaysurviveforatleastfiveyears.

DISPERSALParsonsandCuthbertson(2001)notedthatL.camaraisdispersedbyseedsandbysuckers.Suckeringandseedlinggrowthincreasesthesizeanddensityofexistinginfestationswithinandontheedgesofthickets.Seedsareimportantinlongdistancedispersalofthisweed,especiallyafterbirdandmammalingestion(Swarbricketal.1998;ParsonsandCuthbertson2001).AlargenumberofnativeandexoticbirdshavebeenrecordedasfeedingonL.camarafruitsinAustralia.Theseincludedspeciessuchasthebrownpigeon,crow,cuckooandemeralddoves,emu,figbird,fruitdoves,Indianmynah,Lewin’shoneyeater,littlewattlebird,piedcurrawong,purplecrownedpigeon,rainbowlorikeet,redcrownedpigeon,regentandsatinbowerbird,silvereye,sparrow,spinycheekedhoneyeater,starling,variedtrillerandwongapigeon(Liddy1985;LoynandFrench1991;Swarbricketal.1998;Carter2000;StansburyandVivianSmith2003;vanOosterhout2004). SomeauthorsindicatedthatingestionincreasedthegerminationofL.camaraseedsoncetheyhadbeenexcreted(vanOosterhout2004).Initialinfestationsfrombirddispersedseedsarecommonunderperching,roostingandsheltertrees,fencelinesandotherperchsites(Swarbricketal.1998). Theseisolatedplantsformthefocioflaterinvasionsoftheweed.

LantanacamaraseedsareprobablyspreadbyawiderangeofotheranimalsinAustraliaincludingrodents,cattle,sheep,goats,horses,foxes,pigs,kangaroosandlizards(Swarbricketal.1998;Dayetal.2003;Stock2004).Whileingestionaidedspreadisgenerallylocalised,distancesofupto1kmormorearepossible(Swarbricketal.1998).

StudiesregardingtheroleofbirdsindispersingL.camaraseedsareneeded,inparticularstudiesonbehaviourandfeedingpreferencesandthedistancesdifferentbirdstravelaftereating(Dayetal.2003). OnenotableexceptionisthestudybyWillsonandCrome(1989)whichfoundthatseedsofmanyinternally


vertebratedispersedspeciessuchasLantanawerefounddispersedupto85metresintorainforest. GROWTHANDAnotherstudybyVivianSmithetal.(2006)indicatedthattheseedfeedingflyO.lantanaedamagedthefruitandseedofL.camaraensuringthatlessseedwasspreadbybirds.

Ensbey(2003)recordedthatL.camaracanbespreadinwater,incontaminatedsoilandonmachinery,aswellasaccidentallyonpeople.NewplantsariseviavegetativereproductionfromgardenwastecontainingplantmaterialofeitherL.camaraorL.montevidensiswhenitisnotdisposedofproperly(Neal1999;Ensbey2003). Thetradeofnurseryplantshas,andcontinuestoberesponsibleforthespreadofthisspecies.

TheseedsofL.montevidensisaredispersedbyanumberofmeans.ParsonsandCuthbertson(2001)notedthattheseedsarewidelyspreadbyfruiteatinganimalsandbirds,bywaterflowingacrossthesoilespeciallyafterheavyrainandinmudattachedtohooves,bootsandmachinery.O’Donnell(2002)alsonotedthatL.montevidensisseedsfloatandthatgulliesandwatercoursesweresusceptibletoinfestationasaresult.Inparticular,O’DonnellandPanetta(2000)recordedthatparrots,currawongs,whitecockatoos,crowsandemusatetheripefruitofthespecies.Paleheadedrosellashavealsobeenobservedeatingthefruitofthisspecies(S.Csurhespers.comm.).Initialinfestationsfrombirdswerecommonundernestingandroostingsites,especiallyaroundtreesandfences(O’Donnell2002).O’Donnell(2002)furtherindicatedthatantstransportandburyL.montevidensisseedsintheirnests.ThepresenceofpulpsurroundingseedsdoesnotprecludethegerminationofL.montevidensisseedsindicatingthatpriorbirdingestionisnotarequirementforsuccessfulgermination(O’DonnellandPanetta2000).Lantanamontevidensisisalsospreadwhenviableseedspassthroughthedigestivetractsofgrazingcattle(O’Donnell2002).

Swarbrick(1986)notedthattheweedyvarietyofL.montevidensismayalsoreproducebystemcuttingsandbythedivisionofestablishedplants.Incontrast,theornamentalvarietiesofL.montevidensisappeartohaveonlybeenpropagatedandspreadbystemcuttings(Swarbrick1986).ThesizeanddensityofexistingcoloniesofL.montevidensisincreasesasstemsrootatthenodesandasseedlingsdevelopwithinandnearexistingthickets.

DEVELOPMENTThefollowinginformationhasbeenextractedfromSwarbricketal.(1998)unlessotherwisenoted.Lantanacamaraisveryplasticinitsresponsetolightintensity.Seedlingdensitiesrangefromlessthanonetoseveralpersquaremetrewithplantsatlowerdensityproducingroundedplantswhilethoseathigherdensityproducingmoreuprightplantsthatcompeteforlight.Plantstendtodominateseveralsquaremetresasadults,growingthroughandoverotherplants,oftenproducingimpenetrablethickets,shadingoutotherplantsandoutcompetinganyL.camaraseedlingsthatmayemerge.vanOosterhout(2004)indicatedthatL.camarawilltoleratepartialbutnotcompleteshading.Newshootsemergefromthebaseofhealthyplantswhileoldershadedstemslosevigouranddie.Prostratestemsmayrootatthenodesifcoveredbymoistdebris,sometimesdevelopingintovigorousdaughterplants.

Lantanamontevidensisisabletopersistunderperiodsofextendeddroughtandhastheabilitytogrowandreproduceinsituationsfromfullsunlighttoshade(O’Donnell2002).Althoughthedroughtpersistenceofthisspeciesiswellestablished,evidencesuggeststhatthespeciesgrowsandspreadsatafasterrateunderwetterconditions(O’Donnell2002).

HABITATClimatic requirements Lantanacamaragrowswellinarangeofwarmerareasoftheworld,particularlytemperate,subtropicalandtropicalareas(Swarbricketal.1998). Growthdoesnotoccurbelow5oCandtheplantisfrostsensitive(Thaman1974;Stirton1977;Winder1980,inSwarbricketal.1998)withseverefrostskillingtheleavesandstems(vanOosterhout2004).Stirton(1977)notedthatL.camaraseldomoccurredwherethemeanannualsurfacetemperaturewasbelow12.5oCinSouthAfrica,whileGraaff(1986)recordedthatsomevarietiescouldwithstandminorfrostssolongasthesewereinfrequent.TheuppertemperaturelimitforgrowthforL.camarahasnotbeeninvestigated.


PopulationsofL.camaracoverawidegeographicandclimaticrangeinAustraliafromDarwininthenorthtoOrbostinthesouth(vanOosterhout2004)andfromareasreceivingmorethan3,500mm(DayandHannanJones1999)toareasreceiving650mmmeanannualrainfall(BartholomewandArmstrong1978). Lantanacamaragrowsbestunderconditionsofconstantrainfallorsoilmoisture,particularlyinareaswhichreceiveinexcessof900mmofrain(Swarbricketal.1998;Ensbey2003). The650mmisohyetisprobablythelimitofthespecieswestofwhichlowtemperaturesanddrysoilindependentlyrestrictthegrowthofthespecies(Swarbricketal.1998). Swarbricketal.(1998)postulatedthatbothofthesefactorswereresponsibleforlimitingthedistributionofthisspeciestoareaswestoftheGreatDividingRange.Whilethesefactorsarelikelytocontributetotherestricteddistributionoftheweedfurtherwestwards,lowtemperaturesanddrysoilarefactorsthatarecommonintheeasternpartsoftheGreatDividingRangeandincoastalareas.Furthermore,L.camaracantotoleratebothdrytohumidclimates(ParsonsandCuthbertson2001),althoughfloweringgenerallyoccursunderconditionsofhighsoilmoistureandairhumidity(Swarbricketal.1998). ItisthereforelikelythatotherfactorsareresponsibleforitscurrentdistributioninAustraliaandthesefactorsneedtobequantifiedbeforethepotentialdistributionofthisspeciescanbevalidated.

TherearetwocontrastingtheoriesastowhyL.camarahasnotbecomewidelynaturalisedinVicandsouthernAustralia.Severalauthorsindicatedthatthelimitingfactormaybemoreregularfrostsorlowertemperatures(Conn1999;ParsonsandCuthbertson2001;Dayetal.2003). Bywayofcontrast,Carr(1993)statedthatinVicneithertemperaturenormoisturewerelimitingtothegrowth,floweringandfruitingofestablishedL.camaraplants,butthatseedgerminationandseedlingestablishmentrarelyoccurredbecausesufficientmoisturewasnotavailablewhensummertemperaturesweresufficientforgermination.Thoseauthorsstatedthatoutsideofgardensituationsonlyonenaturalisedpopulationwasknowninasituationreceivingrunofffromroads.OtherweaklynaturalisedpopulationsinVicappeartohavebeenfoundmorerecentlyasoutlinedinMunir(1996)andConn(1999).EithertemperatureormoisturemayexplainwhythespeciesisnotmorewidelynaturalisedinWAalthoughalackofsuitablemoisturetoensureplantestablishmentisthemorelikely.

Dayetal.(2003)outlinedthattherewassomedifferenceintheenvironmentaltolerancesofdifferentvarietiesinAustraliawiththecommonpinkweedyvarietyfoundathigheraltitudesandlatitudeswhereasthepinkedgedredvarietywasrestrictedtowarmerareas. Incontrast,Ensbey(2005)indicatedthatthepinkweedyvarietyofL.camarawasfoundthroughouttheentirerangeofthespeciesthroughoutNSW,butthatthepinkedgedredvarietieswererelativelycommoninnortheasternNSWandwererarelyfoundsouthofthemidnorthcoastandHunterareas.Furthermore,Ensbey(2005)notedthattrueredvarietiesofthespecieswererestrictedtothenorthcoastofNSW(Clarence,CoffsHarbour,Bellinger,NambuccaandKempseycouncilareas)buthadthepotentialtospreadfurthernorthandsouth.TheorangeandwhitevarietiesofL.camarafoundinQldwererelativelyrareinNSW(A.Clarkpers.comm.)

Lantanacamaragrowsfromsealevelto1000metresinelevationineasternAustraliaandupto2000minotherplaces(Thaman1974;Munir1996;Swarbricketal.1998;ParsonsandCuthbertson2001).Swarbricketal.(1998)notedthatalthoughmostareassusceptibletoinfestationprobablyalreadyhadthespeciespresent,thatfurtherspreadofthisspeciescouldbeexpectedwithintheseareas,particularlyalongthefringesofexistinginfestationsandintotheriversystemsontheGulfofCarpentaria,theNTandWA.Webbetal.(1988)indicatedthatwhilebothL.camaraandL.montevidensisoccurredinthenorthernNewZealandL.montevidensiswasmoretolerantofcoldandisgrownfurthersouthasagardenplant.

Soils WhileL.camaragrowsbestonrichorganicsoilsitwillgrowonarangeofsoiltypesincludingstonyhillsidesandnearlypuresandsolongasthereisasourceofsoilmoisture(Winder1980,inSwarbricketal.1998;HumphriesandStanton1992;Munir1996).Thespeciesgrowswellondeeper,welldrainedclaysoilsofbasaltic,metamorphicorgraniticorigin(HumphriesandStanton1992).Thaman(1974)summarisedevidencethattherootsofthisspeciestendedtorotinwaterloggedsoilsandthatithadaverylowtolerancetosoilsalinity.

KleinschmidtandJohnson(1977)indicatedthatL.montevidensisiscommononshallow,stonysoils


inslopingareasthattendedtodryout.Incontrast,Munir(1996)statedthatL.montevidensisgrewinwelldrainedalluvialorloamsoils,anobservationalsomadebyO’Donnell(2002).

DISTRIBUTIONINAUSTRALIACurrent distribution

Lantana camara

LantanacamaracanbefoundincoastalandsubcoastalareasofAustralia(Figure10).AlongtheeastcoastofAustralia,thespeciescangenerallybefoundfromEden37o04’SinsouthernNSWtoCapeMelville(northofCooktown)14o12’SinQld.ThisnorthernmostrecordislikelytoincludeonlylargeinfestationsbecauseMunir(1996)statedthatherbariumrecordsofL.camarahavebeencollectedfromthetipofCapeYorkPeninsulaandfromtheTorresStrait.WhileClarketal.(2004)reportedthatinfestationsofL.camaracanbefoundontheNSW/Vicborder,otherinformationsuggeststhatveryfewinfestationsarefoundsouthoftheEurobodallaandMountDromedaryareasnearNarooma(36o13’S,Harding2005;M.Michelmorepers.comm.).TheseinfestationsarearoundBermagui,Tathra,PambulaandEden.ThesouthernmostinfestationontheeastcoastofAustraliaisprobablynearOrbost(37o42’S)incoastalVic(vanOosterhout2004).LantanacamaraappearstohavebecomeweaklyestablishedasagardenescapeinotherpartsofVic(Conn1999)andhasbeenrecordednearFrankston(38o09’S;Munir1996).

InNSW,Conn(1992)recordsthepresenceofL.camaraonthenorth,centralandsouthcoast,LordHoweandNorfolkIslands.Thespecieswasprobablyintroducedtotheseislandsinthemidtolate1800’s(Swarbrick1986). ThepinkfloweringvarietyofL.camaraisthemostwidespreadalongtheNSWandQldcoastwhiletheredfloweringvarietyiscommononthemidandfarnorthcoastofNSW(Ensbey2005).AredfloweringvarietyiscommonintheRichmondandKurrajongarea,northwestofSydney(AuldandMedd1987;McMillan1989;pers.obs.).

InQld,L.camaracanbechieflyfoundinallcoastalareasnorthtoCooktown(Figure10)andoftenbeyond.Inaddition,thespecieshasbeenrecordedonseveralislandsalongtheGreatBarrierReef(Munir1996).Seawright(1965)indicatedthatredfloweredvarietieswerecommoninfarnorthQueenslandaroundCairnsandbetweenKeppelBayandBroadSoundinCentralQueensland.TheremainderofcoastalQldareashadpinkfloweringvarietieswiththeexceptionofthewhitevarietythatoccurredintheeasternslopesoftheGreatDividingRangeinsoutheasternQld.AnumberofauthorsincludingSeawright(1965)havenotedthepresenceofanorangeyellowvarietyfoundgrowingaroundTownsvillebutnotelsewhere.vanOosterhout(2004)alsoindicatedthattheweedispresentinEmerald(centralQld)andWeipaonCapeYorkPeninsulawhileClark(2006)indicatedthespecieshasbeenfoundatvariousplacesininlandQldincludingSurat(Warrooshire,southernQld),inBelyandoandJerichoshiresincentralQldandinEtheridgeshireinnorthernQld.ScatteredinfestationsofL.camaracanbefoundintheTorresStraitIslands(Munir1996;Swarbricketal.1998,vanOosterhout2004).

LantanacamaracanbefoundintheNT,especiallyaroundDarwin,Maningrida(northcoastal),ontheGovePeninsula,AdelaideriverandonMelvilleIsland(Munir1996;JeffreyandReady1999;vanOosterhout2004). IsolatedoccurrencesalsooccurinotherstatesofAustralia(ParsonsandCuthbertson2001).Forexample,L.camaracanbefoundinsouthwestWA,especiallyinwetterareasaroundPerth,FremantleManjimupandAlbany,aroundBromeandGeraldton

Figure10.The current distribution of L. camara in Australia (Source: van Oosterhout (2004)).


andsouthofKununurra(Munir1996;Husseyetal.1997;vanOosterhout2004,Clark2006).Incontrast,Husseyetal.(1997)indicatedthattheKununurrainfestationwaseradicatedin1995.ScatteredinfestationsofL.camaracanalsobefoundintheSA,especiallyaroundAdelaide(JessopandToelken1986;Swarbricketal.1998;Blood2001;vanOosterhout2004). Swarbrick(1986)citedpersonalcommunicationthatindicatedthatalthoughLantanaspecieshavebeenwidelygrowningardensinTasmania,theyhaveneverescapedfromcultivation.

Lantana montevidensis

LantanamontevidensiscanalsobefoundincoastalandsubcoastalareasofAustralia(Figure11).AlongtheeastcoastofAustralia,thespeciescanbefoundfromNowra34o53’SinsouthernNSWtoCairns16o55’SinnorthernQld(Munir1996;Neal1999).AnumberofauthorsindicatedthatL.montevidensiscanbefoundonthenorthcoast(aroundCasino,MurwillumbahandByronBay)andcentralcoastofNSWnorthfromSydney(Everist1981;Swarbrick1986;AuldandMedd1987;Conn1992;O’Donnell2002).Thespeciesisalsofoundonthenorthwesternslopes(Tamworth)andsouthofSydney(Conn1992;Australia’sVirtualHerbarium2007).

LantanamontevidensisiscommoninsoutheastQld,mainlyfromRockhamptontotheNSW/Qldborder,againincoastalandsubcoastalareassuchasthe

Figure11.The current distribution of L. montevidensis in Australia (Source: Australia’s Virtual Herbarium (2007)).

Moreton,WideBay,BurnettandPortCurtispastoraldistricts(Seawright1965;KleinschmidtandJohnson1977;Everist1981;AuldandMedd1987;O’Donnelletal.1999;O’Donnell2002).Forexample,Seawright(1965)indicatedthatL.montevidensisgrewthicklyinpaddocksaroundBeenleighandIpswichinsoutheasternQld,O’Donnell(2002)recordedtheproblemsthespeciescausedintheBurnettwhileDayetal.(1999)recordedtheweedwasasevereproblemincentralQld.RecordsshownaturalisationinthecoastalpastoraldistrictsofSouthKennedy,NorthKennedy(aroundTownsville)andCook(aroundCairns)andtheinlanddistrictofLeichhardt(Hnatiuk1990;Munir1996;Australia’sVirtualHerbarium2007).

ThespeciesisusedasanornamentalinsometownsintheNT(Munir1996;ParsonsandCuthbertson2001)andhasbeenrecordedasaweedinothersituationsinDarwin(JeffreyandReady1999).O’Donnell(2002)notedthatornamentalvarietiesofthespeciescanbefoundfromTorresStraittoSydney,inMelbourneandAdelaide.

TherearetwootherherbariumrecordsofL.montevidensisinAustralia,oneinKununurra(WA)andtheothernorthofMelbourne(Vic)(Australia’sVirtualHerbarium2007).Inaddition,Conn(1999)indicatedthatthespecieshasbecomeweaklyestablishedawayfromgardensinVic.

Potential distribution LantanacamarahasthepotentialtospreadfurtherwithinAustralia.Itisprobablethatfurtherspreadwithinthecurrentdistributionofthisweedwilloccurasuninfestedareasareinvadedandasincreasesindensityoccurwithininfestedareas.Inthebroadestsense,Dayetal.(2003)statedthatL.camaracanbefoundbetweenthelatitudes35oNandS,eventhoughitismostcommonlyfoundintropical,subtropicalandtemperateareas.Incomparison,Australiastretchesfromapproximately10o30’SonCapeYorkto43o30’SatthebottomofTasmania.

TheecologicallimitationsofL.camaraarenotwellunderstoodandneedtobeinvestigated.TworeferencesindicatedthatL.camaraisnowpresentattheheadwatersofthemajorwesternflowingriversafterwestwardsexpansioninsoutheasternQldduringwetyearsinthe1970’s(A&RMCA&NZ,A&NZE&CCFM2001;vanOosterhout2004).Personal


communicationindicatedthattheseinfestationsarepresentintheupperCondaminecatchmentbutthere IMPORTANCEissomedoubtastothefurtherspreadofthespecieswestward(P.Blackmorepers.comm.;A.Clarkpers.comm.).ThespreadoftheweedintowesternNSWhasbeenpostulatedalongripariancorridorsandasaresultofpoorlandmanagementdecisions(vanOosterhout2004).Furthermore,Ensbey(2003)notedthatL.camarahasthepotentialtospreadintoVicwhiletheCooperativeResearchCentreforAustralianWeedManagement(2003)consideredthatthespeciesisabletospreadwestoftheGreatDividingRangeexpandingitsrangeintosouthernVic,SAandsouthwesternWA.Husseyetal.(1997)furtherindicatedthatL.camarahasthepotentialtobecomenaturalisedinWA.

Figure12outlinesthepotentialdistributionofL.camarabasedontemperature,rainfall,seasonalextremes,burningregimesandsoilmoisture.ThismodelleddatashowsthatL.camaramaybeabletospreadacrossCapeYorkPeninsula,andthroughoutthenorthernpartsoftheNT,theKimberleyregion,furtherinsouthwestWAandVicaswellasfurtherinlandinsouthernandeasternAustralia.Althoughthespecieshasnotyetspreadintotheseareasitspotentialasasleeperweedmayallowittospreadsometimeinthefuture.

Neal(1999)indicatedthatL.montevidensisisintheearlystagesofspreadinAustralia. ThatauthormappedthepotentialdistributionofL.montevidensisviaCLIMEXandconcludedthatatleast30%ofQldand10%ofNSWwereatthreatofseriousinvasion.O’Donnell(2002)notedthatsinceweedyandornamentalvarietiesofL.montevidensishavesimilarclimaticpreferencesthattheweedyvarietyofthespeciescouldbeexpectedtospreadbeyonditspresentrangeinAustralia.Itisimportanttonotehoweverthattheecologicallimitationsofthespeciesarenotknownandrequirefurtherresearchattention.HencegiventhecurrentsparsedistributionitwouldbereasonabletoexpectthatthespecieswillcontinuetoinvadecoastalandsubcoastalareasofNSWandQld,particularlyaroundareaswhereitcurrentlyoccurs.ExpansionontheDarlingDownsinQldandinthenorthandcentralwesternplainsofNSWmaybepossible,asmaybefurtherexpansionofthespeciesinsouthernNSWandVicsincethespeciesseemsmorecoldtolerantthanL.camarainNewZealand(Webbetal.1988).

Detrimental SincetheintroductionofL.camaratoAustralia,infestationshaveexpandedtooveratleastfourmillionhectares,2.5millionofthisinQldand1.5millionhectaresinNSW(Culvenor1985;Ensbey2005).Thecostsoftheseinfestationshavebeenconservativelyestimatedat$2.2millionperannum(Culvenor1985),althoughitislikelythatthisestimatedoesnotaccountfortheweedsmanyenvironmentalimpacts.AmoreuptodateestimateonthecostofcontrollingL.camarabyprimaryindustriesbasedinQldis$10millionperannum(A&RMCA&NZ,A&NZE&CCFM2001).

TheweedyvarietiesofL.camaraarewidespreadweeds,especiallynearhabitation,inpasturesandarableland,indisturbedgrasslands,woodlands,sclerophyllforestsandrainforest,alongroadsidesandfencelines,alongwaterwaysandinwasteland(AuldandMedd1987;Conn1992;Vranjic2000;ParsonsandCuthbertson2001).

Incontrast,Dayetal.(1999)outlinedevidencesuggestingthatL.montevidensisinfestedhundredsofthousandsofhectaresthroughoutcentralQld.InfestationsofL.montevidensiscurrentlyvaryintheirseverityandimpact.OneofthemostseverallyaffectedareasistheNorthBurnettwheresevereinfestationshavedecreasedgrazinganimalcarryingcapacitiessoastothreatentheviabilityoftheseenterprisesresultingindecreasesinlandvalues(O’Donnell2002).Neal(1999)alsostatedthatinfestationsofthisspecieshaveresultedinsignificantlanddevaluationandlossofincome.

Figure12.The potential distribution of L. camara in Australia (Source: van Oosterhout (2004)).


Lantanamontevidensisisanincreasinglyprominentweednearhabitation,inpasturesanddegraded

andguineapigs.

grasslands,inwoodlandsandsclerophyllforestsandonalluvialplains,inroadsidecuttings,alongfencelinesandriverbanksandonrockyoutcrops(Swarbrick1986;Munir1996;O’DonnellandPanetta2000;J.Hoskingpers.comm.).ItisimportanttonotethatinfestationsofL.montevidensisoftenoccurininaccessibleterrainandthatthisinaccessibilityresultsinlimitedweedmanagementoptions(O’Donnell2002).Theimpactofbothspecieswillbeexaminedinmoredetailbelow.

Pastures

LantanacamaraisatroublesomeweedofpasturethroughoutmanycountriesthroughoutthePacificandIndianoceanareas(Holmetal.1977). MuchofthefourmillionhaoutlinedaboveispasturecountryineasternAustralia.Inmanyinstancestheweedhasoverrunpastureecosystems,shadingoutmoredesirablespeciesandreducingtheirproduction,oftenproducingpurestandsthatareinaccessibletopeopleandlivestock.ThesetraitsarealsosharedbytherelatedspeciesL.montevidensis,whichin1997hadinfestedaround150,000haofpasturelandwithpotentialforfurtherexponentialspread(Flannery1997,inO’Donnell2002).Bothspecieshavebeenconsistentlyrecognisedascausingsignificanteconomiccosttograzingindustries(Grice2002;Barkeretal.2006).

LantanacamaraisgenerallyunpalatabletolivestockandallbutthreeofthevarietiespresentinAustraliaaretoxictocattleandsheep(Everist1981;ParsonsandCuthbertson2001).Inparticular,Everist(1981)statedthatallredfloweredvarietieswiththeexceptionofonefromMackayhavebeenfoundtobetoxic.OtherredfloweredvarietiesfromnorthQldarelesstoxicthanvarietiesfromcentralandsouthernQldandNSW.Onlytwosmallfloweredpinkvarietieswereshowntobenontoxic,thesebeingthecommonweedypinkvarietyandasimilarvarietyfoundaroundBundaberg.Alllargefloweringpinkvarietiesweretoxic.Everist(1981)presentedfurtherinformationthatindicatedthetoxicityofL.camarainparticularareasmaybearesultofgeneticfactorsnotenvironmentalfactorsandthattherandomintroductionofhorticulturalvarietieswasthemostprobableexplanationfortheregionaldistributionofvariousvarieties.Dayetal.(2003)alsosummarisedevidenceoftoxicitytootheranimalsincludinggoats,horses,dogs,buffalo,redkangaroos

IngestingplantmaterialoftoxicvarietiesofL.camarawillcausearangeofsymptomsoftenstartingwithin24hoursofasinglefeedingevent(Everist1981).Symptomsincludedepression,alossofappetite,sluggishness,constipation,frequenturination,diarrhoeaandjaundiceintheearlystagesdevelopingintophotosensitisation,inflammationofthemuzzleandmouth,damagetotheliver,kidneys,stomachandintestines,unthriftiness,internalparalysisandoftendeathinonetofourweeks(SeawrightandHrdlicka1977;Everist1981).Severefacialitchingcanresultininjuryandblindnessafterrubbing.LampandCollet(1989)indicatedthatwhitefacedcattlearemoresusceptibletopoisoningthanothers,perhapsasresultofincreasedphotosensitisation.EstimatesofcattlepoisoningduetoL.camaravarywidelyfromatleast1500cattleaffectedperannuminQld(Culvenor1985)toanestimated500600affectedperannuminonenorthernNSWcoastalRuralLandsProtectionBoardareaalone,Grafton,(Officer2002).

SincegrazingofL.camarainquantityonlyoccursinsituationsoflowfeedavailabilityorwhennewand/oryoungcattleareintroducedtograzinglandwherethespeciesispresent,poisoningcanusuallybeavoidedbyprovidingadequatefeedandspellingnewcattleinL.camarafreeareasafterpurchase.Inaddition,thesmellofherbicidetreatedL.camaramayattractgrazingfromlivestockwhenpastureisscarce(vanOosterhout2004).Livestockthatarebredinareasinfestedwiththeweedgenerallyavoidgrazingthespeciesbutlimitedtestfeedingmayoccurattimesandthisdoesnotappeartoresultinpoisoning(Everist1981;vanOosterhout2004;B.Johnsonpers.comm.).

Aswellasoutcompetingmanypasturespecies,L.camaraalsoimpedesthemovementoflivestockduringwateringandmustering(Swarbricketal.1998). Culvenor(1985)estimatedthattheannualcostofL.camaratopasturestobe$7.7millionmadeupof1500cattledeaths($0.5million),4.5%reducedperformance($2million),7.3%pastureloss($3million)and$2.2millionincontrolcosts.Theseestimateshaveundoubtedlybeensurpassed.

WhileL.montevidensisgrowsonshallowandoftenstonysoils,itcanbecomeanaggressiveweedinpasturesituations(O’DonnellandPanetta2000;Bray2002).O’Donnell(2002)statedthatwhilethe


abundanceofL.montevidensisisoftenlinkedwithovergrazing,evidencesuggestedthatthespeciesmorereadilyestablishedinundisturbedorunderlighttomediumgrazedconditions.OnlyafterL.montevidensisisestablisheddoesovergrazingappearfavouritsspread(O’Donnell2002).AnumberofauthorsindicatedthatL.montevidensisspreadsinnativeandunimprovedpastureswhentheseareweakenedunderdroughtconditions(Everist1981;AuldandMedd1987;Conn1992;O’Donnelletal.1999;O’Donnell2002). ParsonsandCuthbertson(2001)indicatedthatL.montevidensisisacompetitiveweedthatreducedavailablegrazingareasandhencepastureproductivity.

AlthoughsomeauthorsindicatedthatL.montevidensisisalsotoxictograzinganimals(AuldandMedd1987;Conn1992;CooperativeResearchCentreforAustralianWeedManagement2003),othersindicatedthatthespeciesisnottoxic(Seawright1965;DowlingandMcKenzie1993).O’Donnell(2002)postulatedthatifcattlehadbeengrazingpastureinfestedwithL.montevidensissincebirththenresistanceorimmunitymayhavedevelopedwhereasintroducedcattlemaynothavethisresistance.Everist(1981)notedthatthetoxicityofthespeciesrequiredfurtherstudy.

Naturalecosystems

Lantanacamara,andtoalesserextentL.montevidensis,poseasignificantthreattotheconservationofanumberofecosystemsfoundontheeastcoastofAustralia(Sindel2000;Randall2001).Forexample,BatianoffandButler(2001)rankedL.camaraasthemostinvasiveandfrequentweedonnaturalareasinsoutheastQueensland,whileL.montevidensiswasranked18th.EcosystemsthreatenedbyL.camaraincludefrontalduneandnearbycommunitytypessuchasmangroves,sedgeandheathlands,woodlandsassociatedwithmelaleucas,banksiasandcasuarinas,aswellopenwoodlandandforestcommunities(BensonandHowell1994;StockandWild2002;vanOosterhout2004).Inparticular,L.camaraisconsideredaseriousinvaderofdisturbedecosystemsthroughoutAustraliaandaroundtheworld(Swarbricketal.1998).Furthermore,thespeciesisaseriousthreattoseveralWorldHeritagelistedareasincludingtheWetTropicsofnorthernQld,FraserIslandandtheGreaterBlueMountainsregion(CooperativeResearchCentreforAustralianWeedManagement2003).

ThecapacityofL.camaratoinvadeAustralianforestecosystemshasbeenthesubjectofconsiderabledebate.Onreviewingtheavailableevidence,StockandWild(2002)concludedthattherewas“littleevidenceofitscapacitytodisplaceforestsintheabsenceofexternaldisturbances”. Indeed,theconverseappearedtobetrueinthatitwaslikelythatrainforestspeciesactuallysuppressedthefurtherexpansionofL.camarafromisolatedcanopygaps,inparticularbydensecanopyshading(Stock2004).ThiswouldparticularlybethecaseintropicalrainforestinnorthernQldwhereL.camarapersistsalongtheedges(includingroadsidesandcreeks)andincanopyopenings(HumphriesandStanton1992),andinsubtropicalrainforestsinsoutheastQld,bothhighrainfallenvironments(Stock2004).vanOosterhout(2004)notedthatL.camarathrivesalongwaterwaysbecauseoftheincreasedavailabilityofsoilmoistureandlight.

DisturbanceappearstobeoneofthekeyaspectsintheintroductionandestablishmentofL.camarainforeststhatreceivelessrainfall,forexampledryrainforest(Fenshametal.1994;GentleandDuggin1997b),warmtemperaterainforests,wetsclerophyllforest,andeucalyptforestsandwoodland(Lamb1988;HumphriesandStanton1992;BensonandHager1993;GentleandDuggin1998;DugginandGentle1998).Forexample,Fenshametal.(1994)correlatedL.camarainvasionwithsoildisturbancecausedbyferalpigs.Thisdisturbancekilledtrees,openingtheoverstoreycanopywithincreasedlightpenetrationoccurring.ThisincreasedlightlevelfavouredL.camaragrowthresultingintwooutcomes,thefirstthatpigsthenavoidedthesedenseinfestationscausingfurtherdisturbance,andtheseconditresultedinanincreaseinfireintensityduetoincreasedfuelloads.Thesemoreintensefiresthenkilltheremainingrainforestcanopy.StockandWild(2002)furtherpostulatedthatallstagesofthelifecycleofL.camaraincludinggermination,survivalandearlyseedlinggrowthwerestronglypositivelycorrelatedwiththeintensityofdisturbance,whichisinturnwaspositivelycorrelatedwithresourceavailability(DugginandGentle1998)andinparticulartolight.

TherehasbeensomedebateontheabilityofplantspeciestogerminateandgrowthroughL.camarainfestations(StockandWild2002),forexample,competitivespeciessuchasbroadleafprivet(Ligustrumlucidum)willgrowthroughinfestationsshadingthemout(Swarbricketal.1998). Studiesby


Stock(2004)showedthatwhilethegerminationofrainforestspeciesunderL.camaracanopieswassignificantlyreducedwhencomparedtogerminationunderrainforestcanopies,thesurvivalandgrowthofrainforestseedlingsunderL.camaracanopieswasnotsuppressed.ThisappearstobebroadlyconsistentwiththeobservationsofHumphriesandStanton(1992)thatrainforestspecieswilleventuallyestablishthroughL.camarainfestationsbutthattherateofregenerationislikelytobeafunctionofsoil,rainfallandrepeateddisturbancewithregenerationfavouredinareasofhigherrainfallanddeepersoils.Overall,L.camaraappearstohavethepotentialtoblocksuccession,displacenativespeciesandreducebiodiversity(Lamb1991;LoynandFrench1991;BensonandHager1993;Fenshametal.1994;Dayetal.2003;Stock2004;Vidler2004;CouttsSmithandDowney2006).

StockerandMott(1981)showedthatL.camaracanslowdownorblockgrassinvasionofdisturbedrainforest.GentleandDuggin(1998)indicatedthatL.camarawasaveryeffectivecompetitoragainstnativecolonisersunderhighlight,soilmoistureandsoilnutrientconditions.Lamb(1988)alsoshowedthatthecompetitivenatureofL.camarawillresultinanalterationtomanyoftheprocessesthatoccurnaturallyineucalyptwoodland,inparticularthatnativetreeslostvigourandthatnitrogencyclingwasalteredinfavourofL.camara. RegenerationofthesecommunitiesafterclearingL.camaramaybeslowbecausefewnativeplantpropagulesfromwhichregenerationcanoccurmaybeleft(Swarbricketal.1998).

ReductionsinbiodiversitywithL.camarainfestationsarecommon,particularlythatofthreatenedbiodiversity.Intheircomprehensivestudy,CouttsSmithandDowney(2006)foundthatL.camarawasathreat83threatenedplantspecies,twothreatenedanimalspecies(abutterflyandbird)and11threatenedecologicalcommunitiesinNSW,whereas15threatenedecologicalcommunitiesarelistedintheFinaldeterminationofL.camaraasaKeyThreateningProcess(DepartmentofEnvironmentandConservation2006).Thethreatenedecologicalcommunitiesincluderainforest,Eucalyptusandfloodplainforestsandwoodlands,andBanksiascrub.

Anationalstudyontheimpactofthespeciesonbiodiversitywillundoubtedlyidentifyfurtherthreats(P.Turnerpers.comm.).Incontrast,earlierbiodiversitystudiesoutlined20endangeredorthreatenedfloraspecieswhichwerethreatenedbyL.camarainfestationsintheuppernorthcoastofNSW(vanOosterhout2004),while80speciesandcommunitieswereidentifiedasthreatenedintheNorthernRiversCatchmentManagementAuthoritywhichencompassedthisarea(CouttsSmithandDowney2006).SixtyplantandanimalspeciesofconservationsignificancewereestimatedasthreatenedinQld(A&RMCA&NZ,A&NZE&CCFM2001).

Althoughlesscommonlyrecorded,itisalsolikelythatL.camarainfestationsreducethenumbersofnonthreatenedfloraandfaunaininfestedareas.ForexampleCummings(2004)reportedthatseveralfunctionalgroupsofantswerereducedinLantanainfestedareaswhencomparedtonearbyrainforestandsclerophyllvegetation.Again,Fernandesetal.(2001)indicatedthatarbuscularmycorrhizafungiarereducedwhenrainforestecosystemsinMadagascarareclearedandreplacedbyintroducedweedssuchasL.camara.

ThereissomeevidencetosuggestthatL.camarainfestationsalterfireregimesinnaturalecosystems(HumphriesandStanton1992;Fenshametal.1994;Swarbricketal.1998). Inparticular,L.camaraincreasedthefuelloadsingrassywoodlandsandforestsonrainforestmarginsallowingintensefiretopenetrateintorainforests(HumphriesandStanton1992;Fenshametal.1994). Incontrast,HumphriesandStanton(1992)recordedevidencethatL.camaraactedasaprotectivebarriertomildfires.Asidefromthesituationsoutlinedabove,L.camaramayrestrictaccesstospecificareasincludingforestsandothernaturalecosystemstherebyrestrictingecotourismandrecreationactivities(Clarketal.2004).

Lantanamontevidensisisconsideredaweedofnaturalecosystems,andinparticularnationalparks,becauseitisanefficientpioneerspeciesthatdisplacesnativevegetation(Flannery1997,inO’Donnell2002;Bray2002;CooperativeResearchCentreforAustralianWeedManagement2003).ItisacommonweedofopenwoodlandanddrysclerophyllforestinsoutheasternQldanddisturbedareasbehindmangroves(Munir1996;O’Donnell2002).Thereissomeevidence


tosuggestthatL.montevidensisreducedplantandanimalbiodiversityintheseandotherecosystemswhereitoccurred(Munir1996;O’Donnell2002;S.Csurhespers.comm.).

Forestry

Lantanacamaraisamajorweedoftimberplantationsincludinghooppine(Araucariacunninghamii)andmaybeproblematicintheearlyplantationstagesofexoticpinespecies(radiataandslashpine,PinusradiataandP.elliottiirespectively)inAustralia(Wells1984;Swarbricketal.1998;Hall2000;vanOosterhout2004).ThecostsofcontrollingL.camarainhooppineplantationsexceeded$200,000in1970(Waterhouse1970,inDayetal.2003)andarenowundoubtedlymuchhigher.Forexample,oneestimateplacedthesecostsinexcessof$500,000(A&RMCA&NZ,A&NZE&CCFM2001).InparticularL.camaravarietiesformimpenetrablethicketsundertheforestcanopy,competingstronglywithyoungtreesforlight,moistureandnutrients,decreasinggrowthrates,limitingaccesstothestandstherebyincreasingbothmanagementcostsandfirehazards(Swarbricketal.1998;Hall2000;Dayetal.2003).Incontrast,Swarbricketal.(1998)recordedthatL.camaraisrarelyaprobleminestablishedexoticpineplantationsbecauseitisshadedoutwhereaslightpenetrationismuchhigherinhooppineplantations(vanOosterhout2004).

vanOosterhout(2004)alsorecordedL.camaraasaseriousweedofhardwoodandcabinettimber(rainforestspecies)plantationswherethespeciesbecomesaprolificunderstoreymonocultureinhighlightsituations.

Plantationandothercrops

Lantanacamarahasbeenreportedasaweedofatleast14cropsthroughouttheworld(Holmetal.1977). Itisanimportantweedofbanana,coconut,coffee,copra,citrus,oilpalm,pineapple,rubberandteacropsparticularlyinsoutheastAsia,thePacificandAustralia,butalsoinothercountriessuchasIndia,NigeriaandTrinidad(Holmetal.1977;WaterhouseandNorris1987;SinghandAchhireddy1987;Swarbricketal.1998;ParsonsandCuthbertson2001;vanOosterhout2004).vanOosterhout(2004)alsorecordedL.camaraasaweedofalmond,

avocado,grape,guava,kiwifruit,lychee,macadamia,mango,nut,olive,papaya,pecan,pistachios,pomefruit,stonefruitandwalnutcrops.Holmetal.(1977)andvanOosterhout(2004)notedthatL.camaraisaweedofcottoninTurkeyandNicaragua,riceinIndonesia,sugarcaneinAustralia,IndiaandSouthAfricaandpeanutandsoybeancrops.

Railwayandserviceproviders

Lantanacamaraisoneofthemosttroublesomeweedsthataffectrailwaysandrailwaycorridorsduetoitssize,andtherapidrateofspreadofthespecies(Mahoney1967;vanOosterhout2004).Theweedisalsoproblematicalongclearedeasementsforelectricitylines,androadways(Swarbricketal.1998;vanOosterhout2004).

Allelopathy

SeveralauthorshavesummarisedstudieswhichillustratetheabilityofL.camaratoproducechemicalsthatinhibitthegerminationandgrowthofcertaincropse.g.wheatandsoybeanandryegrassandannualweedsunderlaboratoryconditions(GentleandDuggin1997a;Swarbricketal.1998;Stock2004). Theseinteractionshavenotbeenexaminedfurtherbecausetheyareoflimitedimportancetothisdiscussion.

GentleandDuggin(1997a)foundevidencethatL.camara“iscapableofinterruptingregenerationprocessesbydecreasinggermination,reducingearlygrowthratesandincreasingmortality”ofnativeAustraliantreespeciesthattheweedcooccurswith.SinghandAchhireddy(1987)haveshowntheL.camaraisallelopathictocitrusinFlorida.Swarbricketal.(1998)outlinedfurtherunpublisheddatathatsuggestedthatL.camaraisallelopathictowardsmanyendemicAustralianplants.FurtherresearchisneededtobetterunderstandtheallelopathicinteractionsthatL.camarahaswithvariousnativeandcropplants.

Hostsforotherpests

LantanacamarathicketsproviderefugiaforpestanimalsandotherplantpathogensandpestsinAustraliaandaroundtheworld(Holmetal.1977;Swarbricketal.1998;ParsonsandCuthbertson2001;Dayetal.2003;vanOosterhout2004).


Humaneffects

AsidefromthenuisanceaspectsofL.camararestrictingaccesstocertainareasandimpactingonotherhumanandfarmingactivities,thefruitofL.camaramayalsopoisonhumansifingested,occasionallyresultingindeath(Everist1981;Swarbricketal.1998). Shepherd(2004)alsoindicatedthattheleavesofL.camarawerepoisonous.InfestationsofL.camarareducetheoverallvisualamenityofvariousareas(Ensbey2003).

Beneficial

Ornamental

BothweedyandnonweedyvarietiesofL.camaraarewidelyplantedasornamentalplantsingardens,inparticularashedges(Swarbrick1986).Thenonpricklytriploidvarietiesthatproducelittleseedarebetterornamentalplantsthanthepricklyfruitingvarietiesbecauseoftheirshorter,denserplanthabitsandbecausetheflowerslastlongersincetheylargelyfailtofertilise(Swarbricketal.1998). BothLantanaspeciesarewidelyusedinlandscapedesign,publicandprivategardens,inparks,onroundabouts,inmedianstrips,inroadsidecuttingsandbesidefootpathsbecausetheyarecolourful,requirelittlemaintenanceandhavesomedroughttolerance(Neal1999;vanOosterhout2004).Everist(1981)andWebbetal.(1988)indicatedthatL.montevidensisiscommonlygrowninrockgardensandusedasagroundcoverplantonretainingwallsandbanksinNewZealand,andinotherpartsoftheworld.

Otherhumanuses

MuchofthefollowinginformationhasbeensummarisedfromMunir(1996)andDayetal.(2003),whicharethemselvesreviewsofawiderangeofinformation.ExtractsfromL.camaraareusedasherbalmedicines,especiallyinIndia,withextractsfromtheleavesshowingantimicrobial,fungicidal,insecticidalandnematicidalactivity,butnotantiviralactivity.Investigationsarecontinuingastowhetherextractsarealsousefulasherbicidesandnematicides.LantanamontevidensisisusedinherbalmedicineinSouthAmerica.

TheessentialoilspresentinL.camaraflowersandleavescanbeextractedforuseinperfumes.Productscontainingtheseessentialoilsareavailablefor

purchaseontheworldwideweb.ThetwigsandstemsofL.camaraarewidelyusedasfirewoodindevelopingcountries.

Alternativefoodandhabitatsourcesforwildlife

Swarbricketal.(1998)reviewedevidencethatL.camarawasbeneficialtowildlifeprovidingfeedingsitesforseed,leafandlitterfeedinginsectsandshelterforsmallbirdsandmammals.Theresultsofthatreviewwerenotconclusivehowever.BothLow(2001)andvanOosterhout(2004)contendedthatL.camarathicketsprovidedsuitablehabitatforbirdssuchasbrushturkeys,quail,whipbirdsandwrens,terrestrialanimalssuchasbandicootsandwallabies,frogsandreptilessuchassnakes,andinsectssuchasbirdwingbutterfliesandreedbeeswherethisweedhaddisplacednativevegetation.InmanyplacesL.camarainfestationsprovidedimportantreplacementhabitatwhennaturalhabitatwasnotavailable(Dayetal.2003). Clemson(1985)notedthatL.camarawasrarelyattractivetobeesbecausetheyobtainedverylittlenectarandpollenfromtheflowers.

Otherbenefits

Lantanacamaraisseenasusefulinpreventingsoilcompactionandsoilerosioninsteepriverinesituationsinsomeagriculturalsituations(Swarbrick1986;WaterhouseandNorris1987;vanOosterhout2004). InfestationsofL.camaramayalsobeavaluablesourceoforganicmaterforpastureimprovementandincreasenutrientmobilityineucalyptforestsandothersituations(Munir1996;Dayetal.2003;vanOosterhout2004).vanOosterhout(2004)indicatedthatL.camaracanalsobeusedtoprovidebarriersorbuffersagainstfurtherdisturbanceofvariousareasbyhumans,livestockandotherweeds,toprovideinterimbuffersonrainforestedgeswhileinfestationsinsidearemanaged,andtorestrictaccesstoecologicallysensitivearease.g.caveswherebatsbreed.Theseinfestationsneedtobemanagedtopreventfurtherspreadhowever.TheuseofL.camarafruitasastockfeedhasbeeninvestigatedinIndia(Lalletal.1983a,b). Lantanamontevidensismayalsopreventerosionofsteepslopes,althoughthisisonlybecauseithasreplacedmoredesirablespeciesthatformerlyprovidedsoilcoverage(O’Donnell2002).


ThesaleofornamentalLantanaspecies

ThepotentialsaleofthelargelysteriletriploidvarietiesofL.camarainSouthAfricahasbeencriticisedbySpiesandduPlessis(1987).Thoseauthorsshowedthatthetriploidvarietieswerecapableofproducingfertilepollen(27.344.4%)andviableseedsthushybridisingwithdiploidvarietiesoftheplant.Neal(1999)indicatedthatbetween2and16%ofpollenfromornamentalvarietiesofL.montevidensisandaL.camaraxL.depressahybrid,allsupposedlysterile,wasinfactviable.Furthermore20%ofBrisbanegardenssitessurveyedthatcontainedtheornamentalL.camaraxL.depressahybridandthepurpleornamentalvarietyofL.montevidensishad0.162%seedproduction/floret.PollinationstudiesbyNeal(1999)indicatedpollenfromweedyvarietiesofL.camaraandL.montevidensisresultedin26%and6%offloretsrespectivelyproducingseedinsupposedlysterileL.camaraxL.depressaplants.ThoseresultssuggestedthatwhenanornamentalvarietyofLantanaoccursinisolation,verylittleseedproductionoccurred.SeedproductiondoesoccurwhenornamentalvarietiesarefoundadjacenttoweedyL.camaraand/orL.montevidensisplants.

InadditionNeal(1999)demonstratedconsiderablepotentialforvegetativereproductionfromvariousLantanaspeciesandvarieties.Forexample,24%oftransplantedleavesfromtheornamentalL.camaraxL.depressahybridproducedrootmaterialafterfiveweekswhile4%alsoproducedshoots.Oneplantevenproducedflowersandseedaftertenweeks.Atotalof42%and34%ofornamentalwhiteandpurpleL.montevidensisvarietiesrespectivelyproducedrootmaterialafterfiveweeks,with2%and4%alsoproducingshoots.Incontrast,theweedyvarietiesofL.camaraandL.montevidensishad8%and12%rootproductionfromleavesafterfiveweeks.Thoseresultsindicatedthatthereisconsiderablepotentialforvegetativeproduction,indicatingthatthesevarietieshaveconsiderableweedypotentialquiteapartfromanythatoccursfromsexualreproductionanddispersal.

OneofthekeyaimsoftheWeedofNationalSignificancenationalstrategyforL.camaraisthephaseoutofsaleanddistributionofallnonsterilevarietiesofthespecies(A&RMCA&NZ,A&NZE&CCFM2001).ThetradeanddistributionofL.camaraisbannedinallstatesandterritoriesinAustralia(vanOosterhout2004;Appendix1). Despitethis,vanOosterhout(2004)

indicatedthatornamentalL.camaravarietieswerestillbeingplantedingardensandlandscapinginstatesandterritoriesofAustralia,particularlyinWA(D.Collopypers.comm.).Thetradeandplantingofthesevarietiesisgraduallydecreasinghowever.

vanOosterhout(2004)indicatedthatanumberofornamentalvarietiesofL.montevidensiswereavailableforsaleinAustralia.Inothercases,theweedypurplefloweredvarietywasgrowningardensasanornamental.Whiletheweedyvarietyproducesfertilepollenandseed,theornamentalvarietiesarelargelysterile,floweringprofuselyandrarelyproducingseed(Henderson1969;Everist1981).Unfortunatelypollenfromweedyvarietiescanpollinateornamentalvarietiesresultinginviableseedset.ThisaddstothegeneticdiversityofL.montevidensis. Inaddition,Hammer(2004)indicatedthatL.montevidensisisabletocrosswithL.camara.

AnysaleofornamentalvarietiesofL.camaraandL.montevidensishasthepotentialtoaddgeneticdiversitytotheweedyvarietiesalreadypresentinAustralia.Itislikelythatfurthergeneticdiversitywillhelpbothspeciestoexpandtheirdistributionintonewenvironmentsandtomakecontrolusingherbicidesandbiologicalagentsfarmoredifficult(vanOosterhout2004). BecauseofthediversegeneticbackgroundofL.camara(e.g.Dayetal.2003),thisdocumentrecommendstheremovalofallLantanaspeciesfromsaleinNSW.Inaddition,allplantsandseedssuppliedbynurseries,wholesalers,otherplantmarketerse.g.largesupermarketchainsandfromgardenclubexchangesshouldbebannedfromentryintothestateandaninformationprogramconductedtoalertandinformtheseplanttradeindustries/bodiesandthegeneralpublicofthesereasons.

vanOosterhout(2004)recommendedtheremovalofallplantingsofL.camaraandL.montevidensisfromprivateandpublicgardensandamenityareassothatgeneticmaterialdidnotmovefurtherintotheenvironment.WhilethisrecommendationissounditisprobablynotpracticalundercurrentNSWlegislation.Insteadtheinformationcampaignshouldencourageallgovernmentandprivatelandmanagerstoremoveallornamentalplantingsofbothspecies,toensuretheyareproperlykilled(solarisationunderorinplastichasbeensuggested),toreplanttheareaswithnativespecies(orlesspreferablynoninvasive


exoticspecies)thatarelowmaintenanceandthatdonothavethepotentialtobecomeweedyintheareafollowedbymonitoringofanyseedlingsorregrowth(A&RMCA&NZ,A&NZE&CCFM2001;vanOosterhout2004).

NegativeimpactsoftheremovalfromsaleofLantanaspecies

TherearelikelytobeanumberofminornegativeimpactsassociatedwiththeremovalofsaleofLantanaspecies.Thefirstisthecommercialimpactsthatsuchabanwillhaveonthenurseryindustryandotherplantsuppliers.Ensbey(2005)indicatedthatLantanaspeciesonlyfillasmallmarketnicheandthatthisnichecouldbereplacedbynativeornonweedyexoticspeciesthatfulfilledasimilarfunctione.g.speciesthatwerehardy,lowgrowing,requiredlowmaintenancelevels,weredroughttolerantandweresimilarlycolourful.AregulatoryimpactstatementassociatedwiththedeclarationandprohibitionfromsaleofLantanaspeciesinQldin2003indicatedthatthemarketforthespecieswasvaluedat$75,000/annum(Ensbey2005). ThecostsofcontrollingLantanaspeciesandassociatedimpactsfromtheweedsfaroutweighedtheminorpotentiallossestothenurseryindustry.TherehasonlybeenoneissuethathasariseninQldafterthesixmonthmoratoriumusedtoremovestocksofLantanaspeciesended,thisbeingwherealargeretailchainstoreofferedplantsforsalewhichweresourcedfromoutsideQld(A.Clarkpers.comm.).

LEGISLATIONState declaration PriortoMarch2006,L.camarawasdeclaredasanoxiousweedthroughthenorth,midnorthandsouthcoastareasofNSWandintheSydneyarea(Appendix1;NewSouthWalesDepartmentofPrimaryIndustries2005).Dependingonthelocalgovernmentareaandthevarietyoftheplant(pinkorredflowered),L.camarawasdeclaredasaW2weedinmostcasesmeaningthatitwastobefullyandcontinuouslysuppressedanddestroyed.LesscommonlyL.camarawasdeclaredasaW3plantwhichmeantthatitneededtobepreventedfromspreadinganditsnumbersanddistributionreduced.The

recommendationsfordeclarationofLantanaspeciesthatarisefromthisreviewareoutlinedinAppendix2.

InsummarisingtheinformationcontainedinAppendix1,Ensbey(2005)notedthatonlytheredfloweredvarietyofL.camarawasdeclaredasaW2orW3weedonthenorthandmidnorthcoastsofNSW,mainlyduetoitstoxicitytolivestock.Thepinkfloweredvariety,althoughwidespreadwasnotdeclaredintheseareaseventhoughitwascontrolledbylandholdersandmanagersasresourcesallowed.Movingfurthersouth,withtheexceptionoftheCessnockcouncil,L.camarawasnotdeclaredintheHunterandcentralcoastareaswhereasbothredandpinkfloweringvarietiesweredeclaredasW2weedsthroughoutSydney.Again,L.camarawasnotdeclaredsouthofSydneywiththeexceptionofthepinkfloweringvarietyinEurobodalla.Begavalleyhadanactivemonitoringanderadicationprogramforthespecieseventhoughitwasnotdeclared.

Ensbey(2005)recordedthatthenorth,midnorthandsouthcoast,aswellastheTablelandsRegionalWeedAdvisoryCommitteeshavedevelopedandareimplementingregionalcontrolprogramsforL.camara. Thesouthcoastplanincludedpriorityareaswithcontainmentzonesandbufferareastopreventthespreadofthespeciesfurthersouth.

AllLantanaspeciesincludingL.camaraandL.montevidensisaredeclaredasClass3pestplantsacrossQld(QNRW2007).AClass3pestplantisonethatiscommonthroughoutQldbutthatmayposeanenvironmental,socialoreconomicthreatin,oradjacenttoanenvironmentallysignificantarea.Thisclassificationmeansthatlandholdersneededtocontrolthisplantiftheylivednextto‘environmentallysignificantareas’suchasnationalparksandreserves,butonlyifthesewerefreefromtheweed.CertainlocalgovernmentareasinQldhavealsodeclaredbothweedsrequiringcontrolinareasthatarenotinoradjacenttoenvironmentallysignificantareas(Appendix1).Plantsofbothspeciescouldnotbesold.LandholderswerealsorequiredbylawtoreduceinfestationsofbothspeciesinsomeareasoftheNT(vanOosterhout2004).

ThetradeanddistributionofbothL.camaraandL.montevidensisisrestrictedinmanystates.


A Weed of National Significance (WoNS)

CONTROLLantanacamarahasbeendeclaredaWeedofNationalSignificance(WoNS)becauseofitswidespreaddistributionandimpactonprimaryindustries,conservationandbiodiversityareas(vanOosterhout2004).Amajornationalcontrolprogramhasbeenlaunchedfocussingonpreventingitsintroductionandspread(Ensbey2003).ContinuingdeclarationofthisweedwithinNSWwillgivevaluablesupporttotheseefforts.

Benefits that may accrue from continued legislative control ThereareanumberofotherbenefitsthatwillaccruefromongoinglegislativecontrolofL.camaraandtheimplementationoflegislationtocontrolallLantanaspecies.WhileEnsbey(2005)outlinedanumberofreasons,thesehavebeenexpandedwithinformationcontainedinthisdocumentasfollows:

n preventingthefurtherdistributionofL.camaraandL.montevidensis. ThereissomecontentionastowhetherL.camarahasspreadtoitsmaximumgeographicrangeornot.Onreviewingtheevidencethisdocumentoutlinesevidencethatfurtherspreadmayoccur,especiallyifnewgeneticmaterialmovesintotheexistingpopulation.Incontrast,itishighlyunlikelythatL.montevidensishasspreadtoitsmaximumgeographicrangeatthistime;

n preventingthefurtherspreadofL.camarawithincurrentdistributionareas.WhileL.camaraiswidelydistributedinNSW,thereareanumberofareaswithinthelimitsofitscurrentgeographicspreadthatarenotinfested.Continuedmanagementvialegislativecontrolmayhelptoreducetheinfillingofsuchareas;

n areductioninthenegativeimpactscausedbybothspeciesincludingpasture,forestry,cropandnaturalecosysteminvasionandareductioninlivestockandhumanpoisoning;and

n preventionoffurthergeneticmovementfromornamentalvarietiesintotheenvironment.Stabilisationofthegenomeofthisspeciesshouldaidinthefuturesuccessofbiologicalcontrolagents.

ThereareanumberofexcellentreviewsavailableexaminingthemanagementofL.camara,particularlyvanOosterhout(2004)andSwarbricketal.(1998).O’Donnell(2002)alsoproducedanexcellentreviewonthemanagementofL.montevidensis. ThepurposeofthissectionistohighlightthebasicprinciplesofmanagementofthesespeciesinAustraliaandinformationhasbeendrawnfromthesereferences.IngeneraltheprinciplesoutlinedforL.camarawillbeeffectiveforL.montevidensisunlessotherwisementioned.ItisimportanttonotethatthecontrolofL.montevidensisisoftenmoredifficultthanthatforL.camara(O’Donnell2002).Constantvigilanceisrequiredforthesuccessfulmanagementofbothspecies(Dayetal.2003).

ThereareanumberofreasonswhyinfestationsofL.camaraandL.montevidensiscanbedifficulttocontrolincludingthesizeofinfestations,pooraccesstoinvadedareas,thelowvaluesofinvadedlandandthecostoftheongoingcontrolthatisneeded(Dayetal.1999;Dayetal.2003;Ensbey2003). Anintegratedmanagementstrategyislikelytoprovidethemosteconomic,efficientandpracticalmeansofmanagingthisweed(Ensbey2003).Suchastrategywillincludebothphysicalandchemicalcontrol,followup,ongoingmonitoringandrevegetation(ParsonsandCuthbertson2001;Clarketal.2004;vanOosterhout2004)andneedstobetailoredtoeachsituation.Asummaryreviewofintegratedweedmanagementstrategieswillfollowtheinitialdiscussionofvarioustoolsthatmayformthesestrategies.

Itisalwayseasiesttoworkfromareaswherelightinfestationsoccurtowardsthosewithheavierinfestationsifpossible(CooperativeResearchCentreforAustralianWeedManagement2003).Incaseswhereextensiveanddenseinfestationsexist,initialcontrolmeasuresareusedtoeitherpromoteaccesstothesiteortoencourageregrowthsothatfuturecontroliseasier(vanOosterhout2004).Beforethisthough,itisalwayseasiesttopreventaweedfromenteringanarea.


Prevention PreventionofthespreadofL.camaraandL.

tebuthiuronandtriclopyr,andmixturesoffluroxypyrandaminopyralid,glyphosateandmetsulfuron

montevidensisintouninfestedareasisprobablythebestmeansofcontrol(CooperativeResearchCentreforAustralianWeedManagement2003).Therearethreemainmeansbywhichthiscanoccur:

n strategiccontrolofexistinginfestationssothattheydonotspreadfurther(toenablethisseethediscussionbelow);

n restrictingthesaleanduseofbothLantanaspeciesasgardenplantsbecausetheseplantscanactassourcesofnewinfestationsandintroducenewgeneticmaterialintoareaswhereexistinginfestationsoccurmakingcontrolmoredifficult(Randall2001;vanOosterhout2004).Noninvasivenativeorexoticspeciescouldberecommendedinplaceofthesespecies;and

n restrictingthefurtherimportationofanyLantanaspeciesintoNSW,notonlyL.camaraandL.montevidensis. Thiswilleventuallyresultinthepreventionofanynewvarietiesorspeciesescapingfromcultivation,naturalisingandhybridisingwiththeexistingLantanaspecies.

WhileL.camaraisrelativelywidespread,effortstopreventitspreadingtocurrentlyuninfestedareasarestilluseful.Incontrast,L.montevidensisisnotwidespreadinNSWandsomeareasofQldandvarioushygienemeasuresshouldbeobserved(O’Donnell2002). Theseincludefodderinspectionandrefusalofsuspectfodder,includingtheinspectionoffeedingsitesafterperiodsofdroughttoensurethespecieshasnotbeenspread.Quarantiningcattlethathavebeengrazingonplantsthathaveripefruitbeforemovingthemfrominfestedtocleanpasturesforfivetosixdaysmaybeaneffectivewayofreducingthespreadofviableseedsthroughthedigestivetractsofcattle.Vehiclehygienetopreventthemovementofseedsinmudorvegetativefragmentswillalsohelppreventspreadofthisspecies.TheuseofbufferstripsbetweeninfestedanduninfestedareaswillalsohelppreventthevegetativespreadoftheL.montevidensis.

Herbicides ThecurrentherbiciderecommendationsforL.camaracontrolinNSWinclude2,4Damine,dichlorprop,fluroxypyr,glyphosate,metsulfuronmethyl,

methyl,picloramand2,4Damineandtriclopyrandpicloram(AustralianPesticidesandVeterinaryMedicinesAuthority2007).Asidefromtriclopyrandtriclopyr/piclorammixtures,theseherbicidesworkbestbythoroughlywettingactivelygrowingfoliage(Ensbey2004;QNRM2004). Productscontainingtriclopyrareusefulforcutstumporbasalapplicationsandneedtobeappliedtoeachstemtoachieveeffectivecontrol.Thebestresultsfromcutstumpapplicationsoccurwhenthestumpiscutnohigherthan15cmfromthegroundandtheherbicideisappliedtothestumpwithin15seconds(vanOosterhout2004).Thefollowingdiscussionfocusesonfoliarherbicidesunlessotherwisenoted.

Thereareanumberoffactorsaffectingtheefficacyofthefoliarherbicidesoutlinedabove.Swarbricketal.(1998)summarisedarangeofAustralianandinternationalherbicidetrialsonL.camara. Thefollowinglistofregisteredherbicideswasderivedandarelistedinorderofdecreasingeffectiveness:fluroxypyr;glyphosate;piclorammixtures;dichlorprop;metsulfuronmethyl;and2,4Damine.Theseherbicideswereeffectiveagainstthecommonpinkandlesscommonredfloweringvarietyaswellasthe‘HelidonWhite’varietythatcommonlyoccursinsoutheastQueensland.Herbicidescontainingdicamba(notregistered)andfosamine(nowwithdrawnfromsale)weremoderatelyeffective.

Highvolumefoliarapplicationsoffluroxypyrweremoreeffectivethanlowvolumeapplications,especiallywhensurfactantsareadded(Love1989).Groundapplicationsof2,4Damine,glyphosate,dichlorpropandpicloramweremoreeffectivethanaerialapplications(Armstrongetal.1987).

Therewasamarkedseasonalityintheeffectivenessofherbicidecontrol.Inparticular,fluroxypyr,glyphosate,picloram,dichlorprop,metsulfuronmethyland2,4DamineweremoreeffectivewhenappliedbetweenDecemberandApril(HannanJones1998;Swarbricketal.1998). Ofthese,theactionofthegrowthregulatinghormonalherbicides(allherbicidesexceptglyphosateandmetsulfuronmethyl)waslinkedtothegrowthactivityoftheplantandmaypossiblybelinkedtoincreasedtranslocationoftheherbicide.Insummarisingothersresearch,Swarbricketal.(1998)


indicatedthatwaterstressandreducedfoliagecoverresultedinreducedcontrolduringautumn,althoughcoldweathermayalsobelinkedtothisreduction.Thebestresultsfromherbicideapplicationsweregenerallyachievedsixweeksaftergoodrains(rainfallexceeding35mm)andwhentemperaturesexceeded15oC(HannanJones1998).Theresultinggrowthflushresultedinagreateramountofherbicidebeingdepositedonyoungerleaftissue.

Graaff(1986)andHannanJones(1998)bothpostulatedthatthevariableresponseofL.camaratosimilarherbicidetreatmentsmaybeduetothelargevariabilitywithinthespecies.Forexample,therewasavariableresponseofthreedifferentvarietiesto2,4DamineinsoutheastQldinthatpinkandwhitefloweredvarietiesdiedwithinfourmonthsbutthatredfloweredvarietiesrecoveredwithinsixweeksandkeptgrowingdespitethefactthattherewerenodifferencesinleafwetabilityandsprayretention(DiatloffandHaseler1965). Variableherbicideresponsesto2,4DamineandfluroxypyrwerealsoobservedindifferentvarietiesfoundinnorthQld(VitelliandDorney1991).

Glyphosateappearedtogivedecreasingcontrolasplantsizeincreased(Wells1984).McMillan(1991)showedthatthevolumeofherbicideappliedneededtobeproportionaltothevolumeoftheplantratherthantheleafarea.vanOosterhout(2004)indicatedthatL.camararegrowthfrom30to100cmhighafterdroughtorfrostevents(orevenafterburning,cutting,slashingordozing)wasideallytreatedwithfoliarherbicidesbecauseitwasactivelygrowing,accesswasofteneasierandthereducedsurfaceareaoftheplantrequiredlessherbicide.

ThereareanumberofotherimportantconsiderationswhenapplyingherbicidestoL.camara(Ensbey2003).Inparticularthecostoftheinitialtreatmentsandfollowupherbicideapplications(orcostofothertreatments)onregrowthandonnewseedlingsthathaveemergedneedstobefactoredinasacostofanongoingcontrolprogram.Herbicidesarelikelytobeanexpensiveoptiontotreatlarge,denseinfestations(vanOosterhout2004)andwhileaerialapplicationsbyhelicoptermaybefeasibleforlargeinfestationsthatareinaccessibletoothermachineryorcannotbeburnt,thesearestillnotcosteffective.Herbicidechoicealsoneedstominimisepotentialofftargetimpactsondesirablenativeandpasturespecies,andwaterways

whererelevant(Dayetal.2003;vanOosterhout2004). Itisalwaysimportanttoreadandfollowlabelinstructionswiththeuseofanyherbicides.

O’Donnell(2002)recordedthatfluroxypyrgavethebestoverallcontrolofL.montevidensisandthatwhileglyphosatealsogavegoodresultsinautumnitdidnotachievegoodresultsinspring.Wettingagentsdidnotimprovetheefficacyoffluroxypyrorglyphosatetreatments.Inaddition,theuseoffluroxypyrallowedgrassproductionwithinthreemonthsoftreatment(O’Donnell2002).

KleinschmidtandJohnson(1977)recommendedtheuseofdichlorpropforthecontrolofL.montevidensisindicatingthatthoroughcoveragewasneededforactivelygrowingplants.O’Donnell(2002)recordedthatwhilethischemicalachievedcontrolofplantsfortwoyears,significantregrowthoccurredafterthistime.Herbicidescontaining2,4Daminesaltsandpicloram/2,4DmixtureswerelesseffectiveonL.montevidensisthanthosealreadymentioned.

O’Donnell(2002)recordedthatregrowthoccurredfromplantsthathadbeenapparentlydeadforperiodsofovertwoyearswithallherbicidetreatmentsusedandthatnoherbicideachieved100%control.RegrowthstimulatedfromaburningeventdidnotimprovetheperformanceofL.montevidensiscontroloverunburntareas(O’Donnell2002).

Inadditiontothoseherbicidesoutlinedabove,herbicidescontainingmetsulfuronmethyl,tebuthiuron,triclopyrandmixturesoffluroxypyrandaminopyralid,glyphosateandmetsulfuronmethyl,andtriclopyrandpicloramareallregisteredforthecontrolofL.montevidensis(AustralianPesticidesandVeterinaryMedicinesAuthority2007).HerbicidescontainingglufosinateandimazapyrhavebeenshowntobeeffectiveagainstL.montevidensisinresearchtrialsbutarenotregisteredagainstthisweed.

Other management

Fire

Althoughtheuseoffirehasbeenwidelyrecommendedinthepast(Goodchild1951;SaintSmith1964;BartholomewandArmstrong1978),Swarbricketal.(1998)commentedthatverylittleinformationwasavailableontheeffectsofseasonality,meteorologicalconditionsduringoraftertheburn,fuelloads,the


typeoffire(groundorcrownfire)andthetimingofthereintroductionoflivestockifapplicable.While

Mechanicalclearing

agreeingwiththesesentiments,vanOosterhout(2004)outlinedcertainconsiderationsthatarerequiredfortheuseoffiretohelpmanageL.camarainnaturalandpastureecosystems.

Thebestresultswithfireappearedtobeobtainedwhenthefireswerehotandtheweedwasactivelygrowing,inearlysummerinsoutheasternQldandinlatewinterandspringinNSW(CooperativeResearchCentreforAustralianWeedManagement2003). Exclusionofgrazinglivestockbeforeafirewasplannedhelpedbuildupfuelloads.BurningwasnotrecommendednearrainforestareasastheseecosystemsarehighlysensitivetofireandtheresultingburncouldprovidefurtheropportunitiesforL.camaratoinvadecanopygaps,ormaypromotetheregrowthofsclerophyllspeciesinsteadofmoredesirablerainforestspecies(Vranjicetal.2000;CooperativeResearchCentreforAustralianWeedManagement2003).Firemayalsodamageplantationtimberorothertreecropse.g.coconutsifused(Dayetal.2003).

FireisausefultoolinthemanagementofL.camaraclearingdensethicketsandreducingthenumberofplantswhencombinedinanintegratedmanagementprogram(ParsonsandCuthbertson2001),particularlypriorto,orasfollowuptotreatmentsformechanicalorchemicalcontrol(CooperativeResearchCentreforAustralianWeedManagement2003).Thedangersoffireescapinganddamagingpropertyandpeopleneedtobecarefullyassessedandfirepermitsmayneedtobeobtained(CooperativeResearchCentreforAustralianWeedManagement2003;vanOosterhout2004).

FirecanbeusefulintwowaysinmanagingL.montevidensisseedlings(O’Donnell2002).Burningtrialshaveindicatedthatsurfaceseediskilledbyhotpasturefiresbutthatburiedseed(especiallyinantnests)escapes.Anyseedthatescapesafirewasmorelikelytogerminateviasmokestimulusandtheseplantscouldbeeasilycontrolledbyherbicidesand/ortramplingasseedlings.Adultplantsarenotkilledbyfirebecauseregrowthoccurssoonafter,andwhileseedsetmaybedelayed,observationssuggestedthatflowerandseednumbersmaybehigheronpreviouslyburntplants.

AnumberofauthorshaverecommendedtheuseofmechanicalclearingtoremoveL.camarainpastures(Goodchild1951;SaintSmith1964;BartholomewandArmstrong1978).Thiscanbeeasilyperformedusingbulldozers,slashers,stickrakersorwithchainpullingwheretheterrainpermitsandwhereerosionriskswouldbeminimised(Ensbey2003;vanOosterhout2004). O’Donnell(2002)indicatedthatbulldozingtoadepthof15cmgavereasonablecontrolofL.montevidensis. Itwasimportanttofollowupanysuchclearingwithspotherbicideapplicationsorhandpullingofregrowthfromexistingplants,anynewseedlings,oranyotherweedsthatemergedduetosoildisturbance(O’Donnell2002;Ensbey2003;CooperativeResearchCentreforAustralianWeedManagement2003;vanOosterhout2004).Itwasalsoimportanttoconsiderthatreshootingfrombrokenstemsafterpushingorstickrakingwasalsolikelyandthatthisregrowthneededcontrol(vanOosterhout2004). Acombinedmulcher/harvesterhasbeenbuilttoclearwoodyweedsincludingL.camarawithinhooppineplantationsinQld(Swarbricketal.1998).

Cultivation

Whilecultivation,generallydiscploughing,givesveryeffectivecontrolofL.camara,itisimportanttoplantcompetitivepasturestopreventreestablishmentoftheweed(SaintSmith1964;Swarbricketal.1998;vanOosterhout2004).Variousfodderplantshavebeensuggestedincludingglycine(Neonotoniawightii),kikuyu(Pennisetumclandestinum),leucaena(Leucaenaleucocephala)andGuineaandRhodesgrass(PanicummaximumandChlorisgayanarespectively),(SaintSmith1964;Swarbricketal.1998;A&RMCA&NZ,A&NZE&CCFM2001).Itisimportanttoaccountforthepotentialweedinessofanynewspeciesintroducedtoensurethattheintroducedspeciesdoesnotbecomeaweed(e.g.Randall2001;Walton2003).Itmaybenecessarytoobtainapermitifnativevegetationisalsoclearedbythesemanagementmethods.Mechanicalgrubbingofindividualplantsinscatteredinfestationsmayalsobeuseful,especiallyifrevegetationiscarriedoutimmediately(vanOosterhout2004).

O’Donnell(2002)recordedthatmoreeffectivecontrolofL.montevidensisoccurredwithoffsetdiscsratherthanchiselploughsbutnotedthatmultiple


passesmaybeneeded.ChiselploughingcausedfragmentationofL.montevidensismaterialwithan

Grazingmanagement

increaseinplantnumbersresulting.

Handremovalandflameweeding

Lantanacamaraiswidelyremovedbyhandgrubbinginareasidentifiedforregenerationafterwhichselectedtreespeciesmaybeplantedandtendeduntilacanopyhasformedthatshadesoutanyfurtherinvasionsoftheweed(Swarbricketal.1998).Handgrubbingcanoccurwithamattockorhoe,whilea‘lantanalever’isunderdevelopmentinQld(vanOosterhout2004).Handcuttingusingabrushcutter,brushhookormacheteispopular,asistheuseoflargesecateursandhedgingtools,andevenchainsaws.Handcuttingallowsaccesstotheplantbaseforgrubbingorforthecutstumptobepaintedwithherbicide(vanOosterhout2004).Handpullingofseedlingsorevenlargerplantsafterrainfallispopularinsmallerinfestations(SaintSmith1964).FlameweedingalsocontrolledsmallL.camaraseedlings.ItisimportanttominimisesoilanddesirablevegetationdisturbancewithsuchactivitiestoreducetheriskoffurtherL.camaraseedgerminating(vanOosterhout2004).

ThemanagementofverysmallareasofL.montevidensismaybeachievedviapruningtogroundlevelbecausewhileregrowthoccurredafterthefirsttwopruningevents,plantsdiedafterthethird(O’Donnell2002).Asimilareffectmaybeachievedbychipping,mowingorslashing.Mulchingwithhaytoadepthofatleast20cmmayalsokilladultplants(O’Donnell2002).

GrazingmanagementasameansofmanagingLantanaspecieshasoftenbeenoverlookedbyanumberofreviews.Ensbey(2003)notedthatpropergrazingmanagementisausefulpreventativemeansofmanagementformanypastureweeds,notonlyL.camara.Conversely,theremovalofexcessivepasturebiomassthroughoverstockingandgrazingwillleadtopasturedegradationandtheinvasionofweedslikeL.camara.Replantingofdesirablepasturespeciesmayhelpreverseanyoverstocking(asmentionedabove).AvigorouspastureswardasopposedtograsstussockswillhelppreventtheestablishmentandgrowthofseedlingsofL.montevidensis(O’Donnell2002).

Revegetation

ThereplantingofdesirablenativespeciesalongwithcontinualmaintenanceoftheseareasisasimilarmeansofpreventingtheregrowthofL.camarainenvironmentallysensitiveareas(SaintSmith1964;CooperativeResearchCentreforAustralianWeedManagement2003;Ensbey2003).Revegetation,alongwithmonitoringandfollowupcontrolmeasures,willhelpensurethatsitesarekeptfreeofreinfestation(vanOosterhout2004).AfterstudyingtheresponseofL.camaratoshading,Stock(2004)recommendedthattreecanopydensitiesof75%preventedfurtherencroachmentbytheweed.Revegetationmayincluderesowingofpasturesaftermechanicaldisturbanceandeithertheactiveplantingoflocalnativeplantspecies,orallowingthesetoregeneratenaturally.Regenerationwillbemoresuccessfulifthenativevegetationisintactandifinfestationsbytheweedaremorerecent.Thisisbecausemanymorepropagulesofdesirablespeciesarelikelytoremain.Ifremnantvegetationispresent,regenerationcanbeencouragedbyremovingL.camaraontheinterfacesandthenextendingtheseinterfacesprogressivelyintotheinfestationsovertime(vanOosterhout2004)combinedwiththeplantingoffastgrowingtreespeciesifappropriate(Stock2004).


Biological control Therehasbeenwidespreadinterestandactivityin

aprogenitor(anoriginatoroftheselinesofdescent),incontrasttopopulationsfromtheSolomonIslands

thebiologicalcontrolofL.camaraforseveralreasonsincludingthecostoftreatinglargeinfestations,theinaccessibilityofdoingso,theneedforongoingtreatmentandthatinfestationsaregenerallyonlandofloweconomicvalue(Dayetal.2003).

ThefirstbiologicalcontrolagentswereintroducedintoQldin1914(Tyron1914)andanumberofotherinsectshavebeenintroducedsince.Although29insectspecieshavebeentrialledandreleasedforthecontrolofL.camarainAustralia,mostofthesebiologicalcontrolagentshaveprovided,atbest,onlyminororseasonalcontrol(TomleyandRiding2002;Dayetal.2003;Walton2004).Fourspecieshavemadethemostsignificantimpactofthosethathaveestablished,thesebeingasapsuckingbugTeleonemiascrupulosa,twoleafminingbeetlesUroplatagirardiandOctotomascabripennisandaseedfeedingflyO.lantanae(Dayetal.2003). FurtherinformationontherangeofbiologicalcontrolagentsreleasedinAustraliaandaroundtheworldforuseonL.camaraisavailableelsewhere(Dayetal.2003).

ItisimportanttorememberthatbiologicalcontrolisonlyonetoolthatmaybeusedaspartofanintegratedmanagementprogramagainstL.camara,andthatbiologicalcontrolagentsmayonlybeeffectiveatcertaintimesoftheyear(vanOosterhout2004). ThewidegeographicandclimaticrangeoverwhichL.camaraoccursinAustraliaalsomakestheestablishmentofpopulationsofbiologicalcontrolagentsthatareadaptedtothesevariableconditionsdifficult(Swarbricketal.1998;DayandHannanJones1999;Dayetal.2003). ThereisincreasingevidencethatinsectbiologicalcontrolagentsmayshowapreferenceforsomevarietiesofL.camaraoverothers(Harleyetal.1979;Cilliers1987;WaterhouseandNorris1987;Swarbricketal.1998;DayandHannanJones1999;Dayetal.2003). ThelargestproblemwiththedevelopmentandreleaseofnewornamentalvarietiesofL.camaraistheincreaseinthegeneticdiversitythatinevitablyresultsinweedyvarietiesmakingpotentialfuturebiologicalcontrolevenmoredifficult. Inothercases,planttaxonomymaybetheconfoundingissue.Forexample,thestrongaffinitybetweenpopulationsofthefiveweedyvarietiesofL.camarainAustralia(Table1),andinFijiandVanuatuindicatedthatthesepopulationsshareL.urticifoliaas

andMaui(Hawaii),(Scottetal.2002;Dayetal.2003).Thisknowledgehasnowbeenintegratedintothecontinuingsearchforeffectivebiologicalcontrolagents,particularlyagentsthathavesomespecificityforL.urticifolia(Dayetal.2003). Dayetal.(2003)suggestedfurtherresearchwasneededintotheclassificationandidentificationofweedyvarietiesofL.camarainAustraliaandaroundtheworldtosupporttheseefforts.Incontrast,therustProspodiumtuberculatumwhichwasreleasedin2001mayhavebetterefficacyagainstmanypinkfloweredvarietiesofthisweed(TomleyandRiding2002).

Careneedstobetakeninthereleaseofanybiologicalcontrolagentintotheenvironment,notonlytoensureitsefficacy,butalsotominimisenontargeteffects.NontargeteffectshavebeencausedbytheinsectAconophoracompressa(releasedagainstL.camarain1995)totheexoticornamentalVerbenaceaespeciesfiddlewood,(Citharexylumspinosum)andduranta(Durantaerecta),andanumberofothercommongardenspeciesincludingjacaranda(Jacarandamimosifolia)andyellowbells(Tecomastans)(Maheretal.2004;Palmeretal.2004,2005).

OnebiologicalcontrolagentforL.montevidensis,theleaffeedingbeetleCharidotispygmaeawasreleasedin1994(Dayetal.1999). PopulationsofthisagentarenotsustainedonL.camara. UnfortunatelythisbiologicalcontrolagenthashadnoimpactonL.montevidensis(Walton2004).Dayetal.(1999)notedthatthreeotheragentsreleasedforthecontrolofL.camaraalsoattackedL.montevidensis. TheseagentsarealeaffeedinginsectHypenalaceratalis,aleafblotchingflyCalycomyzalantanaeandaflowerfeedingmothLantanophagapusillidactylabutagain,theimpactoftheseagentsislimited.

Integrated management of L. camara AnintegratedmanagementprogramforL.camarawillrequirearangeofstrategiesincludingprevention,physicalandchemicalcontroltools,followup,monitoring,revegetationandperhapsbiologicalcontrolagentswherepresent(ParsonsandCuthbertson2001;vanOosterhout2004).Theuseofvarioustoolswilldependonalargenumberoffactors


includingthesizeandlocationofinfestations,andthetime,moneyandotherresourcesavailable.Anyintegratedmanagementstrategyneedstobespecifictothesituationencountered.

Pasturesituations

Physicalcontrolofsmallnumbersofplantsmayinvolvediggingplantsupbeforefloweringbyhand,handpullingseedlingsorclearingwithmachinery(oftenatractorandchain)andburning(ParsonsandCuthbertson2001;vanOosterhout2004).Careisneededwiththisapproachtoensurethatasmuchlateralrootmaterialisremovedaspossibletopreventreshooting.Thespotsprayingofherbicidesisgenerallyeffectiveinsmallbutdenseinfestations(vanOosterhout2004)whilecutstumpandbasalbarksprayingareeffectiveonsmallscatteredinfestations.

Onalargerscale,excludinglivestockpriortoaburningevent,bulldozingorslashingandthenstickrakingweedymaterialtogethercanaddtofuelloads(CooperativeResearchCentreforAustralianWeedManagement2003).Slashingcanbeusedreducetheheightanddensityofplantsandtocreateaccesssothattramplingbylivestockormachinerycanoccur,aswellasencouragingregrowth(vanOosterhout2004).Itisimportanttorememberthatregrowthfromlivestemscanoccurwhencontactismadewithmoistsoilevenifthesestemsarechoppedup,(vanOosterhout2004). BiologicalcontrolagentsmaybeusefulinsuppressingL.camaragrowthwhenpresentleadingtoimprovedaccessandmanagement(vanOosterhout2004).

Burningfollowedbydiscingonarableterrainandtheimmediateestablishmentofavigorousgrassandlegumepasturehasenabledsuitablecontrolinareassuitableforgrazing(Goodchild1951;SaintSmith1964;BartholomewandArmstrong1978;ParsonsandCuthbertson2001).Forexample,Goodchild(1951)andBartholomewandArmstrong(1978)recommendedtheuseofgreenpanic(P.maximumvar.trichoglume)orguineagrass(P.maximum)andsiratro(Macroptiliumatropurpureum)mixtures.Thismixtureensuresthatseasonroundgroundcoverageisachieved,siratroincreasessoilnitrogenfertility,whichisimportantforsubsequentgrassgrowth,andbothspeciesprovidedrymaterialforsubsequentburningifnecessary.BothBartholomewandArmstrong(1978)

andvanOosterhout(2004)notedthatpastureneededtoberesownatabovenormalratesbeforerainfalltoensuregoodestablishment.Theapplicationofsuperphosphatefertiliser,orotherfertilisersmaybenecessaryforseveralyearsalongwithlightgrazingmanagementwithinthefirst18monthstoallowgoodpastureestablishment(BartholomewandArmstrong1978).Ifburningiscontinuedthenitisimportanttoresowpasturesaftereachburnandthatlivestockareexcludeduntilthepastureisestablished(BartholomewandArmstrong1978).

Naturalecosystems

ManagementofL.camarainnaturalecosystemsandconservationareaswillrequireaslightlydifferentapproachtothestrategiesoutlinedforpasturesituationsabove.MuchofthefollowingdiscussionhasbeendrawnfromvanOosterhout(2004).Inmostcaseschemicalapplicationisnotappropriateandwhenusedcareisrequiredtoavoidofftargetimpacts.Cutstumpandlimitedfoliarsprayingmaybeuseful,asmayhandgrubbing.Slashingmayalsobeappropriatearoundforestedgeswhereasmechanicaldisturbanceisnotusefulinmostsituations.Forlargedenseinfestationsunderthecanopywhereaccessisrestricted,fireandbiologicalcontrolmaybetheonlypracticalmeansofmanagement.Lantanacamaracancreatehotterfireswhichmaythreatenrainforestvegetationandstructuremakingitaninappropriatetool.FireshouldnotbeusedtomanageL.camaraindryvinescrubvegetationwhichalsohasalimitedtolerancetofire.Eucalyptusforestsandwoodlandsaremoreadaptedtofireofvaryingintensityandfrequencysuchthatfiremaybeusedasatooltopromoteregrowthmanagement.Anymeansofmanagementneedstobefollowedbyrevegetation,eitherbynaturalmeansfromthegerminationofpropagulesinthesoilorbyactivereplantingafterL.camararemoval.Monitoringbirdroostingsitesshouldalsocontinueaccompaniedbythehandpullingofanyseedlings,whiledisturbanceshouldbeminimised.IfL.camaraisprovidingalternativehabitatorstabilisingsoilthensectionsratherthanentireareasneedtobeprogressivelycontrolledandrevegetated.


Controllingregrowth ControlofL.montevidensisisbestachievedbythephysicalremovalofadultplantsandtheuseof

Lantanacamararegrowthisbestspotsprayedinsummerandautumn,especiallyaftergoodrainfall.Thisisbestdonewhenitisvigorouslygrowingandbetween30and100cmtall(ParsonsandCuthbertson2001;QNRM2004;vanOosterhout2004).Variousherbicidesgiveeffectivecontrolincluding2,4Damine,2,4Damineandpiclorammixtures,dichlorprop,glyphosateandpicloram.Alternatively,2,4Damineortriclopyrappliedasabasalbarksprayorasadressingonacutstumpareeffective.Inaddition,fluroxypyrappliedtothefoliage(ParsonsandCuthbertson2001)andtebuthiuronappliedtothesoilatthebaseoftheplantarealsoeffective(GillettandWells1999;ParsonsandCuthbertson2001).Whenspotsprayingiscombinedwithseveralyearsofannualburning,infestationscanbereducedtolevelswherehandremovalbecomeseconomicallyfeasible.FollowupmanagementcombinedwithaprogramofcontinualmonitoringneedstooccurnotonlyforL.camarabutalsoforotherweedsandmaybenecessaryforatleasttwoyears(vanOosterhout2004).

Identifyingthecausesofinfestation

Finally,itisimportanttotryandidentifythecauseoftheinfestationsofL.camara.Forexample,weedinfestationsareoftensymptomaticofotherproblemsincludingdisturbance,overgrazing,inappropriateburningandclearing(vanOosterhout2004).Reducingthecausesofinfestationswillhelpreducetheinfestationsthemselves.

Integrated management of L. montevidensis AlthoughadultplantsofL.montevidensisareveryresilienttofire,drought,mechanicaldisturbanceandsomeherbicides,anintegratedweedmanagementprogramforL.montevidensisinvolvingmanyofthesameprinciplesasoutlinedabovewillbesuccessful(O’Donnell2002).MuchofthefollowingdiscussionhasbeenextractedfromO’Donnelletal.(1999),O’DonnellandPanetta(2000),ParsonsandCuthbertson(2001)andO’Donnell(2002),unlessotherwisenoted.

herbicides.Smallinfestationsoftheweedcanberemovedbyphysicallydiggingthemuptoremovetheplantsandbycultivation.Subsequenttooffsetdiscing,plantingcompetitivepasturescomposedofgrassspeciessuchasIndianbluegrass(Bothriochloapertusa)andwoollyfingergrass(Digitariaeriantha)willprovidegoodgrassbiomassandsomesuppressionofL.montevidensis(O’Donnell2002).Otherspeciessuchascreepingbluegrass(Bothriochloainsculpta)providedgoodbiomassbutgavenosuppressionwhereasbuffelgrass(Cenchrusciliaris)competedeffectivelywiththeweed.FewerL.montevidensisplantseventuatedwhenpasturespeciesweresownincombinationwiththelegumewynncassia(Chamaecristarotundifolia),whichmaybearesultofthesprawlinghabitofthelegumecombinedwiththeextranitrogenreleasedencouraginggrassproduction.Sincelightappearedtoberequiredforgermination,seedburialviaploughingshouldalsoreduceseedlingemergence.Seedviabilityfallsbyupto70%inthefirstyearofburialand80%inthesecondyear.

Wherecultivationisnotpracticable,repeatedherbicideapplicationsonactivelygrowingplantsinlatesummerandautumnshouldbeusedwherenecessary.Registeredherbicidesforthecontrolofthisspecieshavebeenoutlinedabove.Alternativelythenativeblackspeargrass(Heteropogoncontortus)canbecomeestablishedininfestationsofL.montevidensiswithoutgroundpreparationmakingitusefulonsteepslopeswherecultivationisnotpossible.ConservativestockingratescombinedwithpasturespellingtoallowregenerationofdesirablegrassesandthestrategicuseoffireisaneffectivemeansofmaintainingthevigourandcompetitiveabilityofnativepastureswhilereducingthegrowthofL.montevidensis.

Dayetal.(1999)outlinedthatwhilerepeatedherbicideapplicationonL.montevidensisregrowthandtheplantingofperennialpasturespecieswasausefulmeansofcontrol,thecostperunitareaoftenmadethisanunfeasibleoption.Burningwasalsoineffectivebecausetherewasgenerallyinadequatefuelloadstomaintainahotenoughfiretokilltheroots(Dayetal.1999;ParsonsandCuthbertson2001).


SOCIALLIMITATIONS Firstly,itiswellknownthatthespeciesLantanacamaraiscomposedofanumberofgeneticallyvariable

TOCONTROLThe widespread acceptance of L. camara Inthepast,thecommunityhaswidelyacceptedL.camaraasbeing‘partofthelandscape’(Clarketal.2004;vanOosterhout2004).Ineffortstoincreasethemotivationformanagingthisweed,aswellasprovidinginformation,managementtoolsanddirectionstoassistwithmanagement,Clarketal.(2004)outlinedfourneedstoensurethatincreasedmanagementoccurs,thesebeing:

n “areevaluationofattitudestoLantanaspeciestoensurereneweddiligence;

n improvedawarenessandexchangeofinformation;

n integrationofcontrolmethodsandprioritisationofactionstoachievebettercontrolresults;and

n strategicallycoordinatedmanagementtosecureongroundresults.”

ThereareanumberoffactorsthathaveresultedinthewidespreadacceptanceofL.camarabythecommunity(Clarketal.2004). Theseincludethelargesizeofinfestationsconfrontinglandmanagers,theinaccessibilityofmanysuchinfestationstoconventionalmethodsofmanagement,theconfusionaboutthecontinuedsaleofornamentalvarietiesandthewidespreadplantingofthesevarietiesbygardeners,councilsandbusinesses.Thesizeandinaccessibilityofinfestationsgenerallyreducestheimpetusforlandmanagerstobothstartandcontinuemanagingsuchinfestationsonanongoingbasis.

Issues involved with the sale of Lantana species ThereareanumberofissuesthatarisefromthecontinuedsaleofLantanavarieties. IthasbeenarguedthatbecauseofthewidespreadnatureofLantanavarietiesineasternAustralia,thecontinuedsaleofthesevarietiesintheseareasislikelytohaveaminorimpactonthefurtherspreadofthespeciesoutsideofitscurrentrange.Thisargumentcannotbesustainedonanumberofgrounds.

varietiesandthatthesevarietieseasilyhybridisewithoneanother(Spies1984a,b;SpiesandduPlessis1987;Neal1999;Dayetal.2003). Anyfurtheradditionstothegeneticdiversityofthespeciesarelikelytoresultinmoredifficultmanagementinthelongterm,especiallyforpotentialbiologicalandchemicalcontrol(Ensbey2003;Clarketal.2004),butmayalsoincreasetheadaptabilityofnewhybridsofthespeciestonewenvironments(A&RMCA&NZ,A&NZE&CCFM2001).Ensbey(2003)statedthisasareasonastowhyitwasnecessarytolimitnewintroductionsofL.camaratononinvasivevarieties.

Whiletheintroductionofnoninvasivevarieties,ifenacted,wouldbehelpful,itwouldnotentirelysolvetheproblem.Suchnoninvasivevarietiesarestilllikelytoproduceasmallproportionofviablepollenandseed,addingtothecurrentgenepoolintheenvironment(Neal1999;QNRM2004).Inaddition,thesevarietiesuponcrossingwithplantsintheexistinggenepoolwillresultinhybridsthatmayormaynotretainthenoninvasivecharacterofoneoftheirparents.Itismuchsaferinsteadtorecommendtheuseofalternativenativeplantsthatoffersimilargrowingandornamentalfeatures(Ensbey2003),butonlyifthenoninvasivenatureofthesenativespecieshasbeenwellestablished.ThisisimportantbecausethereareanumberofexamplesofnativeplantsfromoneareaofAustraliathatareinvasiveinanotherandthusconsideredasseriousenvironmentalweeds,forexampleCootamundrawattle(Acaciabaileyana)amongmanyotherAcaciaspecies,aswellassweetpittosporum(Pittosporumundulatum)(BennettandVirtue2004).

Secondly,thewidespreaddistributionofthisspeciesacrossanumberofclimatic,ecosystem,rainfall,topographic,elevationandrainfallzonesindicatesthatthisspeciesishighlyadaptivetoabroadrangeofclimaticconditions(Swarbricketal.1998;Dayetal.2003;vanOosterhout2004).Anumberofauthorspostulatethatthisspeciesmayspreadfurtheroutsideitscurrentrange,particularlyintothewarmertropicalnorthernAustraliancoastlineareasandperhapsintocoolersouthernNSWandViccoastalareas.Theintentionalsaleofthisspeciesintotheseareasislikelytoproducefurthergeneticmaterialfromwhichweedpopulationscanestablish.


Thethirdissuethatarisesisthatthesaleofthesevarietiesislikelytoprovideforcontinuedspreadofthespecies.EvenifL.camarahasreachedthelimitsofitspotentialrangeinNSW,continuedintroductionswillresultinthespeciesinvadingnewhabitatsandincreasingindensitywithinthisrange(Ensbey2003).Thiswillcontributetoincreasedmanagementneededforthecontrolofthespecies.

Thefourthissueisuniformityoflegislation.Forexample,ifLantanaspeciesarenotremovedfromsaleineverystatethentradeandmovementofthespeciesfromstatesthathavethespeciesforsalecanstilloccurintoareaswheretheyarerestrictedfromsale.Thiswillresultincontinuedspreadofthespecies.

Thefifthissueisoneofperception.Whilethevarietiessoldmaynotbe‘weedy’varietiesofL.camaraorL.montevidensis,landmanagersareunlikelytoeitherrecognisethisor,iftheydo,toplacethisknowledgeasidewhentheyconsidermanagementofinfestationsofL.camaraontheirland.Rather,landmanagerswillperceivethatthereislittlepointtomanagingLantana

Figure13.Purple flowering variety of the ornamental Lantana montevidensis planted in a median strip, Griffith (south western NSW, September 2005) (Source: S. Johnson, NSW DPI).

speciesifgardeners,councilsandbusinesscanplantit,therebyspreadingplantstothelandmanagers’landswhenvariousanimalseatandspreadit.Again,thereisaperceivedconflictofinterestifcouncilscontinuetoplantLantanavarietiesandthenenforcethemanagementofweedyvarietiesonprivateorpubliclands.ThisconflictofinterestextendstocouncilspermittingthesaleofLantanavarietiestothewidercommunitywhileenforcingcontrolonotherpartsofthecommunity.

Eveniflandmanagersrecognisethattherearesomedifferencesbetweenthevarietiesforsale(despitethefactthatthesevarietieswillresultinsomespreadandgeneflow)andthosethatareweedy,theyarelikelytosetthisknowledgeaside.Thismaybebecausetheyarenotconvincedofthetruthsoftheseclaimsortheyconvenientlyignorethembecausetheydonotwishtospendthemoneytocontrolexistinginfestations.Theseactionsmayprovidesuitableexcusestolandholdersthatclaimapotentialconflictofinterestwithlocalgovernment(asoutlinedabove).

Constraints to managing L. camara Aftersurveying1021landholdersandmanagersin2003,vanOosterhout(2004)indicatedthattimeandcostwerethetwolargestfactorsconstrainingL.camaracontrol.ThesefactorswerefollowedbythedifficultyinaccessingL.camarainfestationsparticularlyduetotheterrain.Economiesofscalewereachievedonlargerpropertiessuchthattheaveragecostperhectareonpropertiessmallerthan100hawas$31.70,decreasingto$8.40/haforproperties5001000haand$0.86forpropertieslargerthan10,000ha.Itwasencouragingtonotethatknowledgeofpotentialmanagementoptionswasconsideredaveryminorfactorlimitingcontrolandthatthevastmajorityofrespondentshadongoingfollowupatintervals.Ingeneralhowever,landholdersandmanagersneededtotrialmoreintegratedmanagementstrategiesinsteadofrelyingonasinglecontrolmethodsuchasamanual,mechanicalorchemicalremoval.


Motivating land managers to undertake management

n Committedenvironmentalattitudeandpriority.ThemanagementofLantanaspeciesislikelytoarise

Inconsideringhowtomotivatealllandholderstoundertakemanagement,Clarketal.(2004)notedthattherewereanumberofpositiveandnegativefactorsinfluencingthemanagementofthisspeciesandsummarisedtheseundersixbroadcategories.

n Availableinformation.LandmanagersmaybemoremotivatedtomanageLantanaspecieswhentheyhavesoundinformationavailabletothem.Thisinformationwouldincludebestmanagementpracticeandanintegratedprogramofcontrolcombiningmanagementandmonitoring.Suchinformationshouldhelppreventsinglecontrolmethodsandadhocmanagementwhichareoftenineffective.

n Resourceavailability.Variousresourcesincludingtime,money,equipmentandpersonnelwereneededtomanagethisweed.Positivemotivationmayarisefromfavourablecost/benefitratiosingrazingsystems,incentivesfromlocalgovernmentforlandholderstoundertakecontrolandtheavailabilityofpublicfundingtopreserveimportantareas.Fewresourcesaredevotedtounusedandvacantlandcreatinglittleactivecontrolandalackofmotivationformanagement.

n Access.Managementiseasierinreadilyaccessibleareaswhereasitmaybehamperedinrelativelyinaccessibleareassuchassteephillsidesorotherdenseareas.

n Activelandmanagement.AgriculturallandandnaturalareasthatareactivelymanagedarelikelytobeactivelymanagedforLantanaspeciesbecauseremovalwillincreaseproductivity,conservationandbiodiversityvalues.Incontrast,neglectedareasarelikelytoattractlesserattention.

fromthosewhohaveanincreasedenvironmentalawareness,forexample,landmanagerswhowishtofarminasustainablewayandlandholdersandenvironmentalvolunteerswhoaretryingtorestoreblocksoflandtoamore‘natural’speciescomposition.Theprioritiesofthesepeoplewillresultinincreasedcontrolofenvironmentalweedsaddingtotheconservationvalueandcommunityperceptionoftheselands.

n Legislationandcompliance.Whilelegislationcanbeusedasapowerfultooltoencouragegoodstewardshipofland,itisimportanttoencouragelandmanagerstovoluntarilyabidebyregulations,andontheotherhandforgovernmentbodiestoenforcethelegislationwhereitisappropriate.Goodstewardshipofthelandmaynotbepracticedwithoutenforcedcompliance.

RECOMMENDATIONSRecommendationsfromthisdocumenthavebeenmadetotheNoxiousWeedsAdvisoryCommittee.TheseareoutlinedinAppendix2.


FURTHERRESEARCH toparentalvarietiesviasomaticmutation.Suchchangeshaveimportantimplicationsinsuccessful

NEEDSThereareanumberoffurtherresearchneeds.Thesehavebeenbrieflymentionedbelow.

n AnaccurateassessmentofthenumberofvarietiesofL.camaraandL.montevidensisinAustralia,theiridentificationandpossibletoxicitiescombinedwithaccurateinformationonthebestmanagementofeachofthesevarieties.ThismaypartiallyinvolvefurtherDNAanalysisviaRAPDtechniques.

n TherateatwhichgeneticflowfromexistingornamentalvarietiesofLantanaspeciesoccursintonaturalisedpopulations.Thismayoccurviaspreadofpollen,seedorvegetativematerial.ThishasimportantimplicationsinthecontinuedmanagementofweedyvarietiesofbothL.camaraandL.montevidensis.

n FurtherquantificationofthefactorsresponsibleforL.camaraandL.montevidensisgermination,seedlingrecruitmentandmortality.Inparticular,nospecificstudieshavedeterminedtheinfluenceoffruitpulponthegerminationofL.camaraseedswhilefurtherstudiesontheseedbankdynamicsofthisspeciesareneeded.Thesestudiesareimportanttoaccuratelyplaceatimeframeonongoingmonitoringoncetheremovalofinfestationshasoccurred.

n FurtherstudiesintothebreedingsystemofLantanaspecies,inparticular,theabilityofL.camaratoselfpollinate.

n StudiesregardingtheroleofbirdsindispersingLantanaseedsareneeded,inparticularbehaviourandfeedingpreferencesandthedistancesdifferentbirdstravelaftereating.

n AnassessmentoftheabilityoflateralshootsofL.camaratoproducenewshootsoncebrokenordamagedinthesoil.Mechanicalorhandremovalproducesahighnumberofbrokenlateralrootsandsomeevidencesuggeststhattheserootsmaygiverisetonewplantsundersuitablegrowingconditions.

n AbetterunderstandingofthefactorsthatresultinoccasionalbranchesofL.camararevertingback

biologicalcontrolandtoxicitymanagement.

n InvestigationsintotheecologicallimitationsofbothLantanaspecies.Inparticular,studiesshouldinvestigatetheinfluenceoflowtemperatures/frostsandsoilmoistureasthereissomedisagreementabouttheimportanceofthesefactorsastheyrelatetothecurrentdistributionofbothspecies.

n QuantificationofthefactorsresponsiblefortheregenerationofrainforestandotherforestspeciesthroughLantanainfestations.Anunderstandingofthesefactorswillbeimportanttoensurerevegetationcanoccur.

n AccurateassessmentsontheimpactofLantanainfestationsonthreatenedplantandanimalspecies.ThepotentialofL.camaratoprovidealternativefoodandsheltertoinsectsandotheranimalsrequiresfurtherstudy.

nMoreaccurateassessmentoftheimpactofL.camaraonfireregimesinvariousnaturalecosystems.Giventhatfireisausefultoolinsomesituationsthisresearchneedstofocusontheeffectsseasonality,meteorologicalconditionsduringorrainfallaftertheburn,fuelloads,thetypeoffire(groundorcrownfire)andthetimingofthereintroductionoflivestockonthecontrolofbothspecies.

n ThecontinuedassessmentofpotentialbiologicalcontrolagentsagainstbothLantanaspecies.

n Furtherassessmentonthetoxicity,orotherwiseofL.montevidensis.

n FurtherproofofallelopathicaffectsofL.camaraonotherplantsunderfieldconditions.

n AmoreuptodateassessmentoftheeconomicimpactofbothL.camaraandL.montevidensisonprimaryproductionandtheenvironmentinAustralia.

n Collationofexistinginformationandresearchonalternativeornamentalspecies,growthhabitsandrequirementsincludinglikelysourcesofsuchplantingmaterial.


INFORMATION CONCLUSIONSREQUIREDFROMCONSULTATIONTheimpactsoftherecommendationsoutlinedinthisreportneedtobeevaluatedviaconsultationwithvariousstakeholders.Inparticularthisconsultationneedstoevaluatethefollowing:

n anaccurateassessmentoftheeconomicimpactontheremovalfromsaleofallLantanaspeciestothenurseryindustryandthecommunity;

n theaffectsofbothLantanaspeciesonpasturesandanimalproduction.Impactssuchasreductioninpasturebiomassandspeciesabundance,theimpactsofLantanatoxicityongrazinganimalsandtheimpedanceonlivestockandhusbandrymovementsneedtobeaccuratelyassessed;

n theaffectsofbothLantanaspeciesontheintegrityofnaturalecosystemswithparticularreferencetofloraandfaunabiodiversityandabundance,conservationandamenityvaluesandferalanimalcontrol;

n theimpactofL.camaraoncommercialforestryandotherplantationcropsincludingbanana,citrus,pineapple,andvariousfruitandnutcropsinAustralia;

n thecostsincurredbyLantanainvasionintorailway,electricityandroadcorridors;

n informationonthetoxicityorotherwisemedicinalvalueofLantanaspeciesonhumanbeings.TheextentthatL.camaraisusedasamedicinaloilsourceinAustraliaalsoneedstobeexplored.AbanonthesaleofLantanamaterialmayalsoresultinabanontheproductionprocessesandendproductsassociatedwithessentialoilsfromLantanaspeciesunlessanexemptionismadefortheseprocessesandproducts;and

n theintentoflocalgovernmenttocontinuetomanageL.camaraandundertakecontrolofL.montevidensis.

TherearetwonaturalisedspeciesofLantanainNSW,L.camara(lantana)andL.montevidensis(creepinglantana).Bothspecieshaveanumberofornamentalandweedyvarieties.

Lantanacamaraisamultibranched,perennialshrubthathastheabilitytoformdensethickets.Ifdamaged,shootsregrowvigorouslyfromtheplantcrownandfromdamagedstems.Thespeciesmayalsohavetheabilitytoreshootfrombrokenlateralroots.IncontrastL.montevidensisisacreepingperennialspeciesrootingatthestemnodesandproducinglowbutdensethickets.Thisspecieshasalargelignifiedtaprootthatallowstheplanttoresistshootdamage.Ornamentalvarietiesmayproducefewerfruitthanweedyvarieties.Thethicketformingnatureofthesespeciesmakesmanagementofinfestationsdifficult.Theapparentlowerfecundityofornamentalvarietiesmayreducetheirweedpotentialinsomesituations.

ThespeciesLantanacamaraisavariablepolyploidspeciesaggregatecomposedofatleast29differentvarietiesinAustralia.Existingvarietiesfreelyhybridiseresultingindifficultyincorrectlyascertainingtheiroriginandhencepotentialbiologicalcontrolagents.Althoughsomeornamentalvarietiesaresupposedlysterile,recentstudiesinSouthAfricaandAustraliaindicatethatthisisnotthecase.

Lantanacamaravarietiesvaryintermsoftheirploidylevels,plantmorphology,responsetoenvironmentalconditionsandnaturalenemies,chemicalcompositionandtoxicityandtheirresponsetoherbicides.ThelevelofdifferentiationinL.camaravarietiesalsomakesitdifficulttopromoteconsistentweedmanagementmessages.AlthoughitisunclearhowmanyvarietiesofL.montevidensisarepresentinAustralia,studiesindicatedifferencesexistinploidylevelsandplantmorphology.Thesedifferencesmayalsobeimportantinthemanagementofvarietiesofthisspecies.StudiesindicatethatdifferentLantanaspeciespreviouslyseparatedbygeographyhybridisefreelyandthatL.camaraxL.montevidensishybridsaregrownasgardenplants.

LantanacamarawasfirstintroducedintoAustraliain1841. ThespecieshasspreadextensivelythroughouttheeasterncoastalareasofNSWandQldintropical,


subtropicalandwarmtemperateareas.SmallerinfestationscanbefoundintheNT,WA,SAandVic.ThisindicatesthatL.camarahasawideclimaticrangeinAustralia,althoughspreadofthisspeciesinareasreceivinglessthan650mmofmeanannualrainfallhasnotyetoccurred.TheecologicallimitationsresponsibleforthedistributionofthisspeciesinAustraliaare,asyet,poorlyunderstood.ThereissomedisagreementastowhetherL.camarahasreachedthelimitofitsgeographicspreadinAustralia.Furtherinvasionofcurrentlyuninfestedareasisalsostilloccurring.AlthoughL.camaraisweedyinover60countries,ornamentalvarietiescanbefoundthroughouttheworld.

LantanamontevidensisappearstohavebeenintroducedintoAustraliain1851,wasrecordedasweedyinsoutheasternQldfromthe1880’sandhassincebecomeasignificantprobleminsubcoastalandcoastalareasofQld.ThespecieswasfirstrecordedasaweedinNSWinthe1950’sbuthasnotspreadsignificantlyinNSWasyet,despitetheclimaticsimilaritiesofNSWwithQld.SmallinfestationscanbefoundintheNT,VicandWA.ItislikelythatL.montevidensiswillspreadbeyonditspresentrangeinAustralia,particularlyincoastalandwesternareasofNSWandQld,andperhapsintocoolerareasinsouthernNSWandVic.WhileL.montevidensisiswidelyplantedasanornamentalaroundtheworld,ithasonlybeenrecordedasaweedinAustralia,andperhapsFlorida.BothnurserystockandseedsofLantanaspecieswerepermittedimportintoAustraliauntillate2006potentiallyincreasingthegeneticdiversityofbothspecies.

AlthoughgerminationofbothLantanaspeciesoccursthroughouttheyear,seedlingsaremorelikelytoestablishunderhighsoilmoisture,temperatureandlightconditions,particularlyafterdisturbance.Seedlinggrowthratesareslowandplantsgenerallydonotflowerintheirfirstyearofgrowthwithgrowthslowedorstoppedduringwinter.TheseedlingmortalityofL.montevidensismaybehighduringthisperiod.Thelongjuvenileperiodofbothspeciesallowsforconsiderablemanagementopportunities.

Afterspringregrowth,L.camaraplantsflowerandsetfruitallyearroundintropicalandsubtropicalareas,andthroughoutspring,summerandautumninmoretemperateareas,particularlyinresponseto

rainfall.FloweringandfruitsetinL.montevidensisalsotendstooccuryearround.Inexcessof12,000fruitmaybeproducedonlargeL.camaraplantswhileover2,500fruit/m2areproducedonL.montevidensisplants.Reductionsinseedviabilityofupto50%inL.camaraand80%inL.montevidensismayoccurwithintwoyearsofseedburial.Plantsofbothspeciesareperennialwithrapidshootgrowthoccurringaftertheremovalofshootsbyfire,herbicide,physicalremoval,droughtorfrost.Newplantsorcanesappeartobeproducedfromplantcrowns,fromintactordamagedlateralroots,andfromintactandcutstemmaterial.

Dispersalofbothspeciesoccursviaanumberofmeans.Alargenumberofnativeandexoticbirdspeciesspreadseed,aswellasawiderangeofferalanimalsandlivestock.Asidefromdeliberatespreadviathetradeofornamentalvarieties,spreadmayalsooccurinwater,insoil,onmachinery,onpeopleandvegetativelyondiscardedgardenwaste.Thewidevarietyofdispersalmechanismsindicatesthatspreadofthesespeciesisdifficulttocontain.

TwentyyearoldestimatesindicatethatinfestationsofL.camaracoveredinexcessof4millionhectaresandcostprimaryproductioninexcessof$10million/annum(amorerecentestimate).ThecostofL.montevidensisinfestationshasnotbeenestimatedbuthundredsofthousandsofhectaresareaffectedinQld.

BothLantanaspeciesoverrunpastureecosystems,shadingoutmoredesirablespeciesandreducepastureproductionwithinfestationsreducingaccessforpeopleandlivestock.AllbutthreevarietiesofL.camaraarepoisonoustograzinglivestockandotheranimals,producingarangeofsymptomsandoftendeath,forexampleinupto1500cattleperannuminQld.ThereissomedisagreementastowhetherL.montevidensisistoxictolivestock.Theconsiderablelossescausedtopasturebasedprimaryproductionindicatethereisconsiderableneedtocontinuetomanagethesespecies.

Lantanacamarathreatensalargenumberofecosystemsincludingfrontalduneandnearbycommunitytypessuchasmangroves,sedgeandheathlands,woodlandsassociatedwithmelaleucas,banksiasandcasuarinas,aswellopenwoodlands,tropical,subtropical,warmtemperateanddryrainforestsandwetanddrysclerophyllforestcommunities.ThereislittleevidencetosuggestthatL.camarainvadesforest


ecosystemsintheabsenceofdisturbanceeventssuchasofthesoil,increasedlightandfireintensitylevels.Infestationshavethepotentialtoblockorslowforestsuccession,displacenativespeciesandreducebiodiversity.ThereisincreasingevidencethatL.camarahasanegativeimpactonthreatenedanimalandplantspecies.Lantanacamaraalsoaltersfireregimes,commonlyallowingmoreintensefirestopenetrateintorainforestscausingextensiveandperhapsirreversibledamage.Ecotourism,recreationalandaestheticvaluesareaffectedbyL.camarainvasions.Lantanamontevidensisisalsoaweedofwoodland,forestandmangrovecommunitieswhereitdisplacesnativevegetationandreducesplantandanimalbiodiversity.AlthoughtheenvironmentalimpactsofLantanaspeciesarenotwellquantified,theyrepresentaseriousconcernandwarrantmanagementinaffectedecosystems.

Lantanacamaraisalsoamajorweedofmanyothercropsincludingforestry,particularlypine,hardwoodandrainforestspeciesplantations,andplantationsandorchardcrops.Itisaweedofroadway,railwayandutilitycorridors,providesrefugiaforpestanimalsandotherplantpathogensandmaybeallelopathic.Plantmaterialmayalsobepoisonoustohumansifingestedbutmayhavesomemedicinalusesinotherinstances.

BothLantanaspeciesarewidelyplantedasornamentalandhedgeplantsandarepopularinlandscapedesign,publicandprivategardens,inparks,onroundabouts,inmedianstrips,onroadsidecuttingsandbesidefootpathsbecausetheyarecolourful,requirelittlemaintenanceandhavesomedroughttolerance.AlternativebeneficialusesforLantanaspeciesincludetheharvestofessentialoils,useasfirewood,theprovisionofalternativeorreplacementhabitatsforanimalsandinthepreventionoferosion.

LantanacamaraisaWeedofNationalSignificance.Oneofthestrategiestoreducetheimpactsoftheseweedsistobanthesaleanddistributionofthesespecies.AllstatesandterritorieswithinAustraliahavebannedthetradeanddistributionofthisspecies.OnlyQldandtheNThaverestrictionsonthesaleandtradeofL.montevidensis. ThecontinuedsaleofornamentalvarietiesofeitherspecieshasthepotentialtoaddgeneticdiversitytotheweedyvarietiesalreadypresentinAustralia.Itislikelythatfurthergeneticdiversitywillhelpbothspeciesto

expandtheirdistributionintonewenvironmentsandtomakecontrolusingherbicidesandbiologicalagentsfarmoredifficult.Thenegativeimpactsofremovingthesespeciesfromsalearelikelytobelimitedbecausealternativespecieswhicharehardy,lowgrowing,requirelowmaintenancelevels,aredroughttolerantandsimilarlycolourfulareavailable.OneestimateindicatedthatthetotalmarketvalueofLantanaspeciesinQldwas$75,000/annum.

VariousvarietiesofL.camarahavebeendeclarednoxiousinNSWmidandnorthcoastallocalgovernmentcontrolareasandinSydney.Declarationsdonotgenerallyexistincentralandsoutherncoastalareas.Aregionalcontrolplanforthesouthcoastincludescontainment,bufferandmanagementplanstopreventthespreadofL.camarafurthersouth.

AlthoughthemanagementofbothspeciesofLantanaisproblematic,L.montevidensisisgenerallymoredifficulttocontrol.Themajorlimitingfactorsforthecontrolofbothspeciesarethesize,accessibilityandcostsofongoingcontrolofinfestationsandthelowlandvaluesthattheseinfestationsoccuron.Thesefactorsoftenseverallylimitcontroldirectedtowardsthesespecies.Anintegratedweedmanagementstrategyincludingtheuseofmanyofthefollowingstrategiesislikelytobesuccessfulhowever.

Preventionofthemovementofplantsintocleanareasviavarioushygienepracticesandabanonthesaleofthespeciesareeffectivemeansinpreventingoutbreaksofeitherspecies.Anumberofeffectiveherbicidesarecurrentlyregisteredforthecontrolofbothspecies.Theseherbicidesareeitherappliedtoactivelygrowingfoliage(includingregrowth)orascutstumporbasalapplications.Ofthefoliarherbicides,thefollowingdecreasingorderofeffectivenessisgenerallycorrect:fluroxypyr;glyphosate;piclorammixtures;dichlorprop;metsulfuronmethyl;and2,4Damine.Climaticandvarietaldifferencesmayaffectherbicideefficacy.Ongoingherbicideapplicationsonregrowthmaterialareneeded.Itisimportanttoconsiderofftargeteffectswithsomeherbicideapplications.

Fireisanotherusefulmanagementtooleventhoughanumberoffactorsaffectingitsperformancearenotwellunderstood.FireisusefulinclearingdensethicketsofL.camaraandinkillingseedsandseedlingsofL.montevidensis. Fireisalsoparticularlyuseful


whenusedpriorto,orasafollowuptochemicalormechanicalclearing.Again,offtargeteffectsneedtobeconsidered.

Mechanicalclearingusingarangeofmachineryincludingbulldozers,slashers,stickrakersorwithchainpullingequipmentisalsoeffectiveagainstL.camara.Regrowthneedstobecontrolledwithtoolssuchasherbicideapplicationsorwithhandpulling.Cultivation,generallydiscploughingfollowedbytheplantingofcompetitivepasturesisalsousefulinanumberofsituationsagainstbothLantanaspecies.TherearevariousmethodsofremovingL.camarabyhandortreatingplantswithflameweeding.Bothrevegetationandpropergrazingmanagementareimportanttoolsinpreventingthereinvasionofbothspecies.Although31differentbiologicalcontrolagentshavebeenreleasedagainstLantanaspeciesinAustralia,thesehaveprovided,atbest,onlyminororseasonalcontrolofthespecies.

RecommendationsarisingfromthisdocumenthavebeenmadetotheNoxiousWeedsAdvisoryCommitteeandarecontainedinAppendix2.TheserecommendationsconsiderallLantanaspeciesforanumberofreasons.ThemostimportantoftheseisthattheL.camaraisaspeciesaggregateandmanyweedyandornamentalvarietiesofitexist.ThesecondreasonisthatitisimportanttotryandreducethecurrentsizeoftheL.montevidensisproblemwhileinfestationsinNSWarestillsmall.ThepotentialformovementofgeneticmaterialfromvarietiesofornamentalL.camaraorL.camaraxL.montevidensishybridsispossibleifweedyvarietiesofL.camaracooccur.Inthesameway,geneticmovementispossibleifornamentalvarietiesofL.montevidensisorL.camaraxL.montevidensishybridscooccurwithweedyvarietiesofL.montevidensis. ThelistingofallLantanaspeciesalsocoversthepossibilityofcontinuedtradeofvarietiesofanyLantanaspeciesif,atthepointoftradeormovement,thespeciesnameiseithernotusedornotknown.

Avarietyoffurtherresearchneedswereoutlinedaswereinformationrequirementsfromconsultation.


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Figure14A flowering and fruiting plant of the common pink variety of Lantana camara, near Copmanhurst (NSW north coast, October 2005) (Source: S. Johnson, NSW DPI).


Figure15Vegetative naturalised plants of Lantana montevidensis (mid ground), near Oxley Lookout, Tamworth (April 2006) (Source: S. Johnson, NSW DPI).


APPENDICES Queensland

APPENDIX 1

DeclarationsofLantanaspeciesacrossAustralia

NewSouthWales

LantanacamaraisadeclaredweedundertheNoxiousWeedsAct1993inNSW.Priorto2006,twodeclarationsappliedtoweedypinkandredfloweringvarieties.TheseweretheW2NoxiousweeddeclarationwhichmeantthattheweedwastobefullyandcontinuouslysuppressedanddestroyedandaW3Noxiousweeddeclarationwhichmeantthattheweedwastobepreventedfromspreadinganditsnumbersanddistributionreduced.ThefollowinginformationwasdrawnfromNewSouthWalesDepartmentofPrimaryIndustries(2005).

ThepinkandredfloweredvarietyofL.camaraweredeclaredasW2weedsineachofthefollowingcontrolareas:Ashfield;Auburn;Bankstown;Botany;Burwood;Campbelltown;CanadaBay;Canterbury;Cessnock;Fairfield;Holroyd;Hornsby;HuntersHill;Hurstville;Kogarah;Kuringgai;LaneCove;Leichhardt;Liverpool;Manly;Marrickville;Mosman;NorthSydney;Parramatta;Pittwater;Randwick;Rockdale;Ryde;Strathfield;Sutherland;Sydney;Warringah;Waverley;Willoughby;andWoollahra;

ThepinkfloweringvarietyofL.camarawasdeclaredasaW2weedintheEurobodallalocalcontrolarea.

TheredfloweringvarietyofL.camarawasdeclaredasaW2weedinthefollowingcontrolareas:ClarenceValley(W2andW3declarationsappliedbecausethisareaincludedtheformerareasofCopmanhurst,Grafton,MacleanandPristineWaters,areaswhichhaddifferingdeclarations);andGreaterTaree,andaW3weedinthefollowingareas:Bellingen;CoffsHarbour;Hastings;Kempsey;andNambucca.

AllLantanaspeciesweredeclaredasW2weedsonLordHoweIsland.

AllcolouredvarietiesofL.camaraexceptthepinkfloweringvarietyweredeclaredasW3weedsintheFarNorthCoastCountyCouncil(includedtheformerareasofBallina,Byron,Kyogle,Lismore,RichmondvalleyandTweed).

AllLantanaspeciesaredeclaredClass3pestsinQldundertheLandProtection(PestandStockRouteManagement)Act2002. Class3pests,inthiscaseweeds,aredefinedasweedsthathaveestablishedinQldandhave,orcouldhave,adverseeconomic,environmentalandsocialeffects(includinginotherstates).Itisanoffencetointroduce,release,giveaway,sellorotherwisesupplyaClass3pest.ThesaleofallLantanaspeciesbecameillegalinNovember2003.LandholdersmayberequiredtocontrolaClass3pestifitisanenvironmental,socialoreconomicthreatinoradjacenttoanenvironmentallysignificantareasuchasanationalparkorreserves,butonlyifthesearefreefromtheweed.

Certainlocalgovernmentareashavealsodeclaredbothweedsunderlocallawrequiringcontrolinareasnotinoradjacenttoenvironmentallysignificantareas:Bowen;Cardwell;ChartersTowers;Dalby;Eacham;Gayndah;Johnstone;Kilkivan;Kingaroy;Maroochy;Maryborough;Mirani;Murgon;Pittsworth;Rockhampton;Sarina;Tara;andTownsville.

NorthernTerritory

BothL.camaraandL.montevidensisaredeclaredintheNTundertheWeedsManagementAct2001butunderdifferinglevels.TheseareaClassBNoxiousweed(regionaldeclaration)thatgrowthandspreadarecontrolledoutsidetownareasandasaClassCNoxiousweedwhichisnottobeintroducedintotheNT.DeclaredweedsarerestrictedfromsaleintheNT.

SouthAustralia

LantanacamarahasbeendeclaredasaClass11category3plantundertheNaturalResourceManagementAct2004. Thisdeclarationmeansthatthespeciesisrestrictedfromsalebutthatcontrolisnotrequired.

Tasmania

LantanacamarahasbeenprohibitedfromimportandsaleinTasmaniaundertheWeedManagementAct1999.Thespeciesmaynotbeotherwisesupplied.Landholdersmayberequiredtocontrolthespeciesontheirpropertyifinfestationsarefound.


WesternAustralia

LantanacamarahasbeendeclaredinWAundertheAgriculturalandRelatedResourcesProtectionAct1976andthePlantDiseasesAct1914. Thisdeclarationhasresultedinabanontheimportandtradeofthisspecies.ThedeclarationdoesnotincludeL.montevidensis.

Victoria

LantanacamarahasbeendeclaredanoxiousweedinVicundertheCatchmentandLandProtectionAct1994.Thetradeanddistributionofthespeciesisrestrictedunderthisdeclaration.

AustralianCapitalTerritory

LantanacamarahasbeendeclaredaprohibitedpestplantintheACTunderthePestPlantsandAnimalsAct2005. Thespeciesisnotabletobesuppliedorpropagatedasaresult.

Commonwealthlegislation

AllLantanaspeciesandmaterialincludingnurserystock,plantpartsandseedswereprohibitedentrytoAustraliaundertheQuarantineProclamation1998fromlate2006.


APPENDIX 2 geneticmaterialintotheenvironmentandsupportthenationalbanontheWoNSspecies.

RecommendationsfordeclarationsofLantanaspeciesinNSW

TherecommendationsfordeclarationsofLantanaspeciesinNSWthatarisefromthisreviewhavebeenoutlinedbelow(seealsoFigureA1overleaf).TheserecommendationsweremadetotheNoxiousWeedsAdvisoryCommitteeforconsiderationinWeedControlOrderNumber19oftheNoxiousWeedsAct1993.

ItisrecommendedthatallLantanaspeciesbedeclaredasClass3,Class4andClass5noxiousweedsinNSW.Thesedeclarationsshouldapplytolocalgovernmentcontrolareasasfollows:

n aClass3declarationinthelocalgovernmentcontrolareasofBegaandEurobodallatosupportmanagementeffortstoreduceLantanaspeciesintheproposedSoutherncontainmentzoneinNSW(Harding2005).AClass3declarationisalsoappropriateforLordHoweIslandtosupportcontrolefforts;

n aClass4declarationinallotherlocalgovernmentcontrolareaswhereeithervarietyofL.camarahasbeendeclaredpriorto2006.ThesedeclarationswouldincludethelocalgovernmentcontrolareasofAshfield;Auburn;Bankstown;Bellingen;Botany;Burwood;Campbelltown;CanadaBay;Canterbury;Cessnock;ClarenceValley;CoffsHarbour;Fairfield;FarNorthCoast;GreaterTaree;Holroyd;Hornsby;HuntersHill;Hurstville;Kempsey;Kiama;Kogarah;Kuringgai;LaneCove;Leichhardt;Liverpool;Manly;Marrickville;Mosman;Nambucca;NorthSydney;Parramatta;Pittwater;PortMacquarieHastings;Randwick;Rockdale;Ryde;Shellharbour;Shoalhaven;SouthSydney;Strathfield;Sutherland;Sydney;Warringah;Waverley;Willoughby;Wollongong;andWoollahra.Thesedeclarationswouldensurethatlocalgovernmentareascouldcontinuetomanageandreducetheincidenceofthisspeciessothatfurtherspreadandeconomicimpactsarereduced.ManagementplansneedtoincludeanintegratedprogramofmanagementmethodsasoutlinedinthisdocumentforthevarietiesofL.camaraorL.montevidensispresent;and

n aClass5declarationinalllocalgovernmentareasinNSWtopreventtradeanddistributionofLantanaspecieswillpreventthefurthermovementof

AClass3declarationisappropriatefor“plantsthatposeaseriousthreattoprimaryproductionortheenvironmentofanareatowhichtheorderapplies,arenotwidelydistributedintheareaandarelikelytospreadintheareaortoanotherarea”.

AClass4declarationisappropriatefor“plantsthatposeathreattoprimaryproduction,theenvironmentorhumanhealth,arewidelydistributedintheareatowhichtheorderappliesandarelikelytospreadintheareaortoanotherarea”.

AClass5declarationisappropriatefor“plantsthatarelikely,bytheirsaleorthesaleoftheirseedsormovementwithinthestateoranareaofthestate,tospreadinthestateoroutsidethestate”.

Thereareanumberofoperationalmattersthatneedtoaccompanythesedeclarationsasfollows:

n thedevelopmentofregionalLantanaweedmanagementplansencompassinganylocalgovernmentcontrolareasthathaveanyweedyLantanaspeciesoutsideofornamental,landscapeand/orgardenplantingswherethespeciesarenotdeclaredunderOrder19oftheNoxiousWeedsAct1993;

n theongoingsurveillanceandmonitoringofanyLantanaspeciesinallotherlocalgovernmentcontrolareasofNSW(wheretherearenooutbreaksoftheweed),whethertheseareashaveanyLantanaspeciesasornamental,landscapeand/orgardenplantingsornot;and

n thecontinuedprohibitiononthesaleormovementofplants,seedsorotherplantmaterialofLantanaspecies.

ConsiderationofaClass2declarationforLantanaspeciesmaybeconsideredforsouthcoastlocalgovernmentcontrolareasaftertheimplementationoftheSouthCoastRegionalLantanaManagementPlan(Harding2005)ortoLordHoweIslandtosupporteradicationattempts,ifappropriate.

Althoughdesirable,itisnotfeasibletorequirealllocalgovernmentcontrolareastoremoveallamenityplantingsofLantanamaterial.Astatewidelocal


governmentandpubliceducationcampaignshouldencouragerelevantstakeholderstoremovesuchplantingsandreplacethemwithalternativeplantspeciesthatareindigenoustothearea.Alternatively,acampaignsimilartothatencouragedbytheOrangecitycouncilfortheremovalofLigustrum(privet)maybeappropriatewhereeverLantanaspecieshavebeenplanted,thatis,peopletradingtheremovedweedfornewtubestockmaterialofmoredesirableornamentalspecies.

Postscript

DeclarationsarisingfromtheNoxiousWeedsAdvisoryCommitteesrecommendationscanbefoundinWeedControlOrderNumber19oftheNoxiousWeedsAct1993,publishedintheNewSouthWalesGovernmentGazetteNumber166,pp.1167111890.

FigureA1.Declarations of Lantana species in mainland NSW recommended to the Noxious Weeds Advisory Committee for consideration in Weed Control Order Number 19 of the Noxious Weeds Act 1993. Lantana species on Lord Howe Island are recommended to be declared as Class 3 weeds. Declaration in Class 3 and 4 areas also includes declaration at Class 5 (Source: A. Maguire, NSW DPI, used with permission).


REVIEW OF THE DECLARATION OF LANTANA SPECIES IN NSW 63 SPECIES IN NSW 63 REVIEW OF THE DECLARATION OF LANTANA SPECIES IN NSW SPECIES IN NSW LANTANALANTANALANTANA SPECIES IN NSW LANTANA

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