Review of the declaration of Lantana species

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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

Lantanamontevidensis 7

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

Lantanamontevidensis 23

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).

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

REVIEWOFTHE usesthetaxonomicnameL.camaratodescribethisaggregatespecies.

DECLARATIONOFLANTANASPECIESINNSWNOMENCLATURE

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

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

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

AbriefdiscussionofthevalidityofthetaxonomicrevisionofL.camaratoL.strigocamaraiscontainedinthetaxonomysection(Sanders2006).Thisdocument

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

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

ThesetwoLantanaspeciesaretheonlyspeciesrecordedasbeingnaturalisedinNSW.Theyhavebeenreferredtobyscientificnamethroughoutthisdocument.

SPECIES DESCRIPTIONS

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

REVIEW OF THE DECLARATION OF LANTANA SPECIES IN NSW 5

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

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

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

flowercoloursinornamentalvarieties.Ornamentalvarietiesaregenerallymorecompactplantswithsmallerleaves(vanOosterhout2004).

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

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

REVIEW OF THE DECLARATION OF LANTANA SPECIES IN NSW 6

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

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

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

REVIEW OF THE DECLARATION OF LANTANA SPECIES IN NSW 7

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

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

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

8 REVIEW OF THE DECLARATION OF LANTANA SPECIES IN NSW

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

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

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

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

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

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

REVIEW OF THE DECLARATION OF LANTANA SPECIES IN NSW 9

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

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

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

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

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

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

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

10 REVIEW OF THE DECLARATION OF LANTANA SPECIES IN NSW

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

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

FourorfivemajorgroupsofL.camaracanbeseparatedviatheirflowercolourasfollows:­

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

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

nwhiteorpale-pinkfloweredvarieties;and

n orangefloweredvarietieswithflowersthatremaindeepyellow(similartoFigure7)toorangethroughouttheirlife.

ThesevarietieshavebeendocumentedinvanOosterhout(2004),pg.6.AsummaryofthemostdistinguishingcharacteristicsisoutlinedinTable1andFigure8.

Ensbey(2003)outlinesfourofthemostcommonL.camaravarietiesthatoccurinNSW.Theseincludethe:­

n commonpinkfloweredvarietythatisspreadthroughoutNSW(includedinthepinkfloweringvarietiesabove);

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

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

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

REVIEW OF THE DECLARATION OF LANTANA SPECIES IN NSW 11

Tab

le1

.D

istin

guis

hing

flor

ala

ndle

afc

hara

cter

istic

sof

L.c

amar

aan

dL.

mon

tevi

dens

isv

arie

ties

outli

ned

byv

anO

oste

rhou

t(20

04).

The

flow

erh

eads

oft

hew

eedy

var

ietie

sof

L.c

amar

aar

eill

ustr

ated

inF

igur

e8.

Alth

ough

ther

ear

eor

nam

enta

lvar

ietie

sof

L.c

amar

aw

itho

ther

flow

erc

olou

rs,o

nly

the

yello

wv

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.

12 REVIEW OF THE DECLARATION OF LANTANA SPECIES IN NSW

Spec

ies

Vari

ety

Bud

Flor

alc

hara

cter

isti

csM

iddl

eflo

ralr

ing

Out

erfl

oral

ring

Si

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.

REVIEW OF THE DECLARATION OF LANTANA SPECIES IN NSW 13

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

14 REVIEW OF THE DECLARATION OF LANTANA SPECIES IN NSW

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).

REVIEW OF THE DECLARATION OF LANTANA SPECIES IN NSW 15

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

16 REVIEW OF THE DECLARATION OF LANTANA SPECIES IN NSW

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).

REVIEW OF THE DECLARATION OF LANTANA SPECIES IN NSW 17

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

18 REVIEW OF THE DECLARATION OF LANTANA SPECIES IN NSW

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

REVIEW OF THE DECLARATION OF LANTANA SPECIES IN NSW 19

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.

20 REVIEW OF THE DECLARATION OF LANTANA SPECIES IN NSW

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

REVIEW OF THE DECLARATION OF LANTANA SPECIES IN NSW 21

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)).

22 REVIEW OF THE DECLARATION OF LANTANA SPECIES IN NSW

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

REVIEW OF THE DECLARATION OF LANTANA SPECIES IN NSW 23

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)).

24 REVIEW OF THE DECLARATION OF LANTANA SPECIES IN NSW

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

REVIEW OF THE DECLARATION OF LANTANA SPECIES IN NSW 25

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

26 REVIEW OF THE DECLARATION OF LANTANA SPECIES IN NSW

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

REVIEW OF THE DECLARATION OF LANTANA SPECIES IN NSW 27

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).

28 REVIEW OF THE DECLARATION OF LANTANA SPECIES IN NSW

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).

REVIEW OF THE DECLARATION OF LANTANA SPECIES IN NSW 29

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

30 REVIEW OF THE DECLARATION OF LANTANA SPECIES IN NSW

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.

REVIEW OF THE DECLARATION OF LANTANA SPECIES IN NSW 31

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.

32 REVIEW OF THE DECLARATION OF LANTANA SPECIES IN NSW

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)

REVIEW OF THE DECLARATION OF LANTANA SPECIES IN NSW 33

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

34 REVIEW OF THE DECLARATION OF LANTANA SPECIES IN NSW

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

REVIEW OF THE DECLARATION OF LANTANA SPECIES IN NSW 35

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).

36 REVIEW OF THE DECLARATION OF LANTANA SPECIES IN NSW

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

REVIEW OF THE DECLARATION OF LANTANA SPECIES IN NSW 37

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.

38 REVIEW OF THE DECLARATION OF LANTANA SPECIES IN NSW

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).

REVIEW OF THE DECLARATION OF LANTANA SPECIES IN NSW 39

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.

40 REVIEW OF THE DECLARATION OF LANTANA SPECIES IN NSW

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.

REVIEW OF THE DECLARATION OF LANTANA SPECIES IN NSW 41

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.

42 REVIEW OF THE DECLARATION OF LANTANA SPECIES IN NSW

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.

REVIEW OF THE DECLARATION OF LANTANA SPECIES IN NSW 43

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,

44 REVIEW OF THE DECLARATION OF LANTANA SPECIES IN NSW

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

REVIEW OF THE DECLARATION OF LANTANA SPECIES IN NSW 45

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

46 REVIEW OF THE DECLARATION OF LANTANA SPECIES IN NSW

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.

REVIEW OF THE DECLARATION OF LANTANA SPECIES IN NSW 47

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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).

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

58 REVIEW OF THE DECLARATION OF LANTANA SPECIES IN NSW

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.

REVIEW OF THE DECLARATION OF LANTANA SPECIES IN NSW 59

WesternAustralia

LantanacamarahasbeendeclaredinWAundertheAgriculturalandRelatedResourcesProtectionAct1976andthePlantDiseasesAct1914. Thisdeclarationhasresultedinabanontheimportandtradeofthisspecies.ThedeclarationdoesnotincludeL.montevidensis.

Victoria

LantanacamarahasbeendeclaredanoxiousweedinVicundertheCatchmentandLandProtectionAct1994.Thetradeanddistributionofthespeciesisrestrictedunderthisdeclaration.

AustralianCapitalTerritory

LantanacamarahasbeendeclaredaprohibitedpestplantintheACTunderthePestPlantsandAnimalsAct2005. Thespeciesisnotabletobesuppliedorpropagatedasaresult.

Commonwealthlegislation

AllLantanaspeciesandmaterialincludingnurserystock,plantpartsandseedswereprohibitedentrytoAustraliaundertheQuarantineProclamation1998fromlate2006.

60 REVIEW OF THE DECLARATION OF LANTANA SPECIES IN NSW

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

REVIEW OF THE DECLARATION OF LANTANA SPECIES IN NSW 61

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).

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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