Forgotten impacts of European land‐use on riparian and savanna … · 2018. 1. 24. · Simon E. Connor, School of Geography, University of Melbourne, Melbourne, VIC, Australia.

J Veg Sci. 2017;1–11. wileyonlinelibrary.com/journal/jvs  | 1

Journal of Vegetation Science

© 2017 International Association for Vegetation Science

Received:5June2017  |  Accepted:29October2017DOI:10.1111/jvs.12591

S P E C I A L F E A T U R E : P A L A E O E C O L O G Y

Forgotten impacts of European land- use on riparian and savanna vegetation in northwest Australia

Simon E. Connor1,2  | Larissa Schneider3 | Jessica Trezise3 | Susan Rule3 |  Russell L. Barrett4,5 | Atun Zawadzki6 | Simon G. Haberle3

1SchoolofGeography,UniversityofMelbourne,Melbourne,VIC,Australia2CIMA-FCT,UniversityoftheAlgarve,Faro,Portugal3CentreofExcellenceinAustralianBiodiversityandHeritage,andDepartmentofArchaeologyandNaturalHistory,AustralianNationalUniversity,Canberra,ACT,Australia4NationalHerbariumofNewSouthWales,RoyalBotanicGardensandDomainTrust,Sydney,NSW,Australia5CollegeofMedicine,BiologyandEnvironment,ResearchSchoolofBiology,AustralianNationalUniversity,Canberra,ACT,Australia6InstituteforEnvironmentalResearch,AustralianNuclearScienceandTechnologyOrganisation,Menai,NSW,Australia

CorrespondenceSimonE.Connor,SchoolofGeography,UniversityofMelbourne,Melbourne,VIC,Australia.Email:[email protected]

Funding informationKimberleyFoundationAustralia;AustralianResearchCouncil,Grant/AwardNumber:CE170100015andDP140103591

Co-ordinatingEditor:PetrKuneš

AbstractQuestions:FireandlivestockgrazingareregardedascurrentthreatstobiodiversityandlandscapeintegrityinnorthernAustralia,yetitremainsunclearwhatbiodiversitylossesandhabitatchangesoccurredinthe19–20thcenturiesaslivestockandnovelfireregimeswereintroducedbyEuropeans.Whatbaselineisappropriateforassessingcurrentandfutureenvironmentalchange?Location:Australia’sKimberleyregionisinternationallyrecognizedforitsuniquebio-diversityandculturalheritage.Theregionishometosomeoftheworld’smostexten-siveandancientrockartgalleries,createdbyAboriginalpeoplessincetheirarrivalonthecontinent65,000yearsago.TheKimberleyisconsideredoneofAustralia’smostintactlandscapesanditsassumednaturalvegetationhasbeenmappedindetail.Methods:InterpretationsarebasedonacontinuoussedimentrecordobtainedfromawaterholeontheMitchellRiverfloodplain.Sedimentswereanalysedforgeochemicaland palynological proxies of environmental change and dated using 210Pb and 14Ctechniques.Results:Weshowthatthepresent-dayvegetationinandaroundthewaterholeisverydifferenttoitspre-Europeancounterpart.Pre-Europeanriparianvegetationwasdom-inatedbyAntidesma ghaesembilla and Banksia dentata,bothofwhichdeclinedrapidlyatthebeginningofthe20thcentury.Soonafter,savannadensityaroundthesitede-clinedandgrassesbecamemoreprevalent.Thesevegetationshiftswereaccompaniedbygeochemicalandbiologicalevidencefor increasedgrazing, localburning,erosionandeutrophication.Conclusions:WesuggestthattheKimberleyregion’svegetation,whilemaintaininga‘natural’appearance,hasbeenaltereddramaticallyduringthelast100yearsthroughgrazingandfireregimechanges.Landscapemanagementshouldconsiderwhetherthecurrent(impacted)vegetationisadesirableorrealisticbaselinetargetforbiodiversityconservation.

K E Y W O R D S

Australianmonsoontropics,fireregimes,geochemistry,grazing,humanimpact,palaeoecology,Potentialnaturalvegetation,riverinevegetation,savanna

Nomenclature:WesternAustralianHerbarium(2017);Mucina&Daniel(2013).

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1  | INTRODUCTION

Human impacts associatedwithEuropean colonizationbeginning inthe 15th century profoundly altered ecosystems around the globe.Impacts includedthe introductionofexoticspecies,manipulationoffireregimes,deforestation,major land-usechanges,displacementofindigenouslandmanagementpracticesandalterationstobiogeochem-icalcycles (Ireland&Booth,2012;Johnsonetal.,2017;Kirkpatrick,1999).Astheseimpactsprecededawidespreadunderstandingofeco-systemecologyandresearch,inmanypartsoftheworlditisdifficulttounderstandhowecosystemsfunctionedpriortoEuropeancoloniza-tion.Thisraisesquestionsabouthowtobestmanagealteredecologiesinaneraofrapidenvironmentalchange(Johnsonetal.,2017).

Acommonbaselineforlarge-scaleecosystemmanagementistheconceptofPotentialNaturalVegetation(PNV).PNVtakesinformationfromareasofassumednaturalvegetation(latesuccessional)andex-trapolatestoother (impacted)areaswithsimilarenvironmentalcon-ditions (Loidi&Fernández-González, 2012).ThePNVapproachhasbeen criticized because its predictions sometimes conflictwith thefossilrecord(Abrahametal.,2016;Carrión&Fernández,2009;Rull,2015).PNValsofailstoaccountforecosystemdynamics (Chiarucci,Araújo, Decocq, Beierkuhnlein, & Fernández-Palacios, 2010) and isunabletosimulateecologicalstructureinculturallandscapes(Stronaetal., 2016). Despite these constraints, PNV remains an accessiblebaselineforconservationandmanagementdecisions.

NorthernAustraliaisregardedasoneofthelargestareasofintacttropicalsavannaworldwide(Bowmanetal.,2010;Woinarski,Mackey,Nix,&Traill,2007;Ziembickietal.,2015). If this is true,PNVmapsfortheregionshouldconstituteavaluablebaseline(Beard,Beeston,Harvey,Hopkins,&Shepherd,2013;Mucina&Daniel,2013).Yettheecosystems of northern Australia have been extensively modifiedsinceEuropeancolonization through livestockgrazing,disruptionofIndigenous land curation and the introduction of invasive species,among other impacts (Douglas, Setterfield, McGuinness, & Lake,2015;Hnatiuk&Kenneally,1981;Lonsdale,1994;Radford,Gibson,Corey,Carnes,&Fairman,2015;Vigilante&Bowman,2004).Impactsarelinkedtoextinctions,decliningnativemammalpopulations,alteredstream ecology, eutrophication, erosion and catchment instability(Payne,Watson, &Novelly, 2004;Woinarski etal., 2007; Ziembickietal.,2015).

Taking a differentview, recent research links the recent declineoffire-sensitivespeciesacrossnorthernAustraliatointeractionsbe-tween fire and flammable grasses (Bowman,MacDermott, Nichols,&Murphy, 2014;Trauernicht,Murphy,Tangalin,&Bowman, 2013).Uncertaintiessurroundingthecausesofrecentspecieslossesremainobscure in the absence of reliable historical evidence for 19th andearly20th centuryenvironmental change in this sparselypopulatedregion.Suchevidencecouldprovideacriticaltestofecosysteminteg-rityandreveallong-termdriversofspeciesdecline.

NorthernAustraliaishometohighlevelsofbiodiversity,muchofwhichremainsundocumented(Barrett,2013,2015;Barrett&Barrett,2011,2015;Maslin,Barrett,&Barrett,2013;Moritz,Ens,Potter,&Catullo, 2013). Phylogenetic research reveals extraordinary levels

ofgeneticdiversityandendemism,perhapsequal totherecognizedbiodiversity hotspots of easternAustralia (Moritz etal., 2013). TheKimberleyregioninNWAustraliaisinternationallyrecognizedforitsextensiverockartgalleries,whicharesomeofthemostancientexam-plesofhumanartisticexpressionglobally(Aubert,2012)andaresitu-atedinanancientculturallandscapecreatedbyindigenousAustralians(Hiscock,O’Connor, Balme, &Maloney, 2016; Rangan etal., 2015).Thereisapressingneedtounderstandanddocumenttheregion’sbi-ologicalandculturalheritageaseconomicdevelopmentpressuresontheNorthernAustralianenvironmentintensify.

Inthispaper,weconfronttheassumednaturalvegetationoftheKimberley regionwith fossil evidence,aiming toassess theecologi-cal integrityof the region’s tropical savannaand riparianvegetationtypes.Ourdatapertaintothesavannawoodlandsandriparianthick-ets (Mucina&Daniel,2013)of theMitchellPlateau,oneof the lastareasoftheAustraliancontinenttoexperienceEuropeancolonization(McGonigal,1990).Wecomparepollendatawith independentgeo-chemicaldatatounderstandthetiming,directionanddriversoftwocenturiesofenvironmentalchange.

2  | METHODS

2.1 | Study area

TheKimberleyRegion issituated inNWAustraliaandconstitutesabiogeographicallyandgeologicallydistinctentitywithintheAustralianMonsoonalTropics(Pepper&Keogh,2014).Theregionischaracter-izedbycomplexandancientgeology,whichhascombinedwithlongperiodsofsub-aerialexposureandweatheringtocreateuniqueland-forms and topography (Pillans, 2007; Tyler, 2016). The Kimberleyregion isbroadlydivided into threegeologicalbasins: theextensiveKimberleyBasininthenorth,theOrdBasintothesouthandeast,andtheCanningBasintothesouthandwest(AppendixS1).Thesebasinsareseparatedbytwoorogenicbeltsofmetamorphosedandintrusiverocks.Thewesternportionof theKimberleyBasin isdominatedbytheKingLeopoldSandstone (erosion-resistant arenites) andCarsonVolcanics (weathered basalts), while the Warton and PentecostSandstonesdominatetheeasternportion.Theserockshavebeensub-jectedtoonlyminorfaultinganddeformationsincetheirdepositionsome1840–1800millionyearsagoandthick,erosion-resistantlater-itesdevelopedoverthebasalts70–50millionyearsago(Tyler,2016).Geologyhasadefininginfluenceonthedistributionofsoiltypesandvegetationunits,aswellasbiogeographicandphylogeneticpatterns(Barrett,2015;Barrett&Barrett,2015;Hnatiuk&Kenneally,1981;Moritzetal.,2013;Mucina&Daniel,2013;Pepper&Keogh,2014).

On theMitchell Plateau, savannawoodlands on basalt tend tohave a ground layer of Allopteropsis semialata, Chrysopogon fallax,Heteropogon contortus, Sehima nervosum, Sorghum plumosum and manyothergrasses,withatreelayerofeucalypts(Eucalyptus bigaler-ita, E. tectifera, Corymbia greeniana, C. bella, C. disjuncta), ironwoodtrees (Erythrophloemum aff. chlorostachys) and terminalia (Terminalia canescens, T. fitzgeraldii),dependingontopography(Mucina&Daniel,2013).Savannawoodlandsonsandstone-derivedsoilshavedominant

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Sorghum and Triodia grasses andadominant tree layerof eucalypts(Eucalyptus tetradonta, E. miniata, E. apodophylla, E. houseana, Corymbia latifolia, C. nesophila, C. torta)andpalms(Livistona eastonii;Hnatiuk&Kenneally,1981;Mucina&Daniel,2013).Sandstoneoutcropshaveanaltogetherdifferentflora,havingahighproportionofspeciesconsid-eredtohaveGondwananheritage,aswellasmanyendemics(Dunlop&Webb,1991).Keyspeciesofthesescrubs/heathsincludeAcacia gono-carpa, A. kelleri, A. arida, A. tumida, Bossiaea arenitensis, Calytrix brownii, C. exstipulata, C. achaeta, Corymbia polycarpa, Gardenia pyriformis and Grevillea agrifolia(Mucina&Daniel,2013).Rainforestremnants(vinethickets)occurinfire-protectedlocationswheremoistureisavailableallyear(Hnatiuk&Kenneally,1981;Ondei,Prior,Williamson,Vigilante,& Bowman, 2017). Albizia lebbeck, Atalaya variifolia, Cochlospermum fraseri, Sersalisia sericea and Wrightia pubescens are important can-opyspeciesintheserainforests(Beard,1976).WetlandsandriparianzonesaredominatedbyPandanus and Melaleucaspecies(P. aquaticus, P. spiralis, M. leucadendron, M. nervosa, M. viridiflora).Nauclea orientalis, Timonius timon and thedeciduous shrubAntidesma ghaesembilla are alsocommonintheseareas.

TheKimberley regionhasadry tropicalmonsoonal climate,withrainfall seasonality driven by the Indo-Australian SummerMonsoon.DoonganStation,nearourstudysite,receivesanaverageof1,205mmannualprecipitation,95%ofwhichfallsfromNovembertoApril(BureauofMeteorology,http://www.bom.gov.au/climate/data/accessed5Jun2017).Rainfalleventsusuallyoccurintheformofprolongedtropicalstorms and/or cyclones during the humid ‘wet’ season, flooding theregion’swatercourses (Mucina&Daniel,2013).Asteep rainfallgra-dientexistsacrosstheKimberleyregion–onaverage,over1,400mmof precipitation falls annually on the seaward slopes of theMitchellPlateau,whilethesouthernpartsoftheKimberleyadjoiningtheGreatSandyDesertreceivelessthan400mm.Maximumdailytemperaturesaverageover30°CinmostpartsoftheKimberleyRegionthroughouttheyear(BureauofMeteorology).

2.2 | Sample collection and dating

A sediment core was collected from a small (~100m2) ellipticalwaterhole on the floodplain of the Mitchell River (15°10′30.5″S,125°53′29.3″E;Figure1,AppendixS1)usingamud–waterinterfacecoreroperatedfromafloatingplatform.Thesite (MP11A) is150mfrom themain river channel and forms part of a complex of lakesand swampswhere the floodplainwidensas it follows thegeologi-calcontactbetweenbasaltandsandstone.Thewaterhole is fringedby Melaleuca leucadendron to thesouthandsurroundedbysavannawoodland on all sides. Nymphaea violacea and Eleocharisgrowinthewater,whichwas~50-cmdeepatthetimeofcorecollection.

Thecorewasdatedusingacombinationof210Pb and 14Ctech-niques(Wrightetal.,2017).Eight210Pbsampleswereselectedfromthe upper 35cm of the core. Macroscopic plant remains were re-moved before the sampleswere freeze-dried, ground andweighed.Subsamples of >1g were dated at the Australian Nuclear Scienceand Technology Organisation (ANSTO) using alpha spectrometry.Radiocarbon(14C)datingwasperformedonthreesamples>5gusing

AcceleratorMass Spectrometry atDirectAMS Laboratories (Seattle,OR,USA).Macroscopicplantmaterialwas removedprior to furtherpre-treatment. An age–depth model was constructed using cubicsplineregressioninClam2.2(Blaauw,2010).

Surface sediment samples are routinely collected in palaeoeco-logical research to calibrate pollen–vegetation relationships (Morris,Higuera,Haberle,&Whitlock,2017).IntheKimberleyRegion,surfacesampleshavebeenanalysedtoaidtheinterpretationofpollenrecords(Field,McGowan,Moss, &Marx, 2017; Proske, Heslop, &Haberle,2014),butmulti-sitecalibrationhasnotbeenattempted.Wecollectedsurface sediment samples from ten small lakesandwetlandsacrosstheregion(Figure1).Multiplerepresentativestandsofthesurround-ingvegetationweresurveyedusingaHaglöfHEC-Relectronicclinom-eter (Haglöf,Avesta, Sweden) and aGRS densitometer (GeographicResourceSolutions,Arcata,CA,USA)toestimatecanopyheight,treevolumeanddensity.CompositionwasmeasuredontheDAFORscale(Kent,2012).Satelliteimagerywasusedtoestimatetheproportionofeachvegetationtypewithina100-mradiusofthecoringsite.Stand-basedestimateswereweightedusingtheseproportionstoobtainveg-etationestimatesforcomparisonwithpollenassemblages.

2.3 | Pollen and charcoal analysis

Seasonally variable environments often have less suitable condi-tionsforpollenpreservationthanmorestableenvironments(Head&Fullager,1992),requiringrelativelylargesedimentsamples.Samples

F IGURE  1 Locationofthemainstudysite(MP11A-triangle)andsurfacesamplingsites(circles)inrelationtoregionalvegetationstructure,NWKimberleyRegion,WesternAustralia.VegetationunitsafterMucinaandDaniel(2013)

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of 5ml sedimentwere treated according to standard palynologicaltechniques(Jones&Rowe,1999),includingremovalofclaysthroughsettlinganddecantation,sievingtoremovelargeparticles(>120μm),treatmentwith10%KOHat90°Cfor20mintoremovehumicmate-rial,heavyliquidseparation(s.g.2.0)toremoveminerogenicmaterial,acetolysis(9:1mixtureofC4H6O3toH2SO4),carbonateremovalwith10%HCl,HF(50%)toremovefinesilica,dehydrationin100%etha-nol,andmountinginglycerolonglassslides.Lycopodiumsporetabletswereaddedearlyinpre-treatmenttoenablecalculationofpollencon-centrationsandaccumulationrates.Pollen,sporesandselectedfun-galandalgalremainswereidentifiedat400×magnification,usingtheextensiveKimberleycollection in theAustralasianPollenandSporeAtlas(Rowe,2006)andpublishedguides(vanGeel&Aptroot,2006;Macphail & Stevenson, 2004). Identificationswere agreed upon bythreeanalysts(JT,SRandSH).

Charcoalparticles arean indicatorof fireoccurrence,withmac-roscopicparticlesconsideredindicativeoflocalfiresandmicroscopicparticles indicative of regional burning (Whitlock & Larsen, 2001).Microscopic charcoal was quantified on pollen slides by countingopaqueblackparticles10–120μm.Macroscopiccharcoalwassievedfromaknownvolumeofsedimentusinga120-μmmeshandparticlesidentifiedusingabinocularmicroscope(Stevenson&Haberle,2005).Resultswere converted to accumulation rates for comparisonwithpollendata.

The pollen sequence was divided into relatively homogenoustime spansusing zonation (CONISS), implemented inTilia (v2.0.41,EricGrimm, http://www.TiliaIT.com). Pollen–vegetation relationshipswere assessedby comparing the abundanceof eachpollen type totheabundanceofpollen-producingplantswithina100-mradius(themostcommonlyusedsamplingradiusincomparablestudies:Buntingetal.,2013).

2.4 | Geochemical analysis

Toexaminetherelationshipbetweenpastvegetationandchangesinthe floodplain environment, we analysed elemental concentrationsof34geochemicalelements.Weusedthealkalineelementssodium(Na),potassium(K),magnesium(Mg),calcium(Ca)andbarium(Ba)totrackerosion.Theseelementsare relativelywater-solubleand theirmobilityduringerosionprocessesresults inelevatedconcentrationsinwater-bodysediments(Davies,Lamb,&Roberts,2015).Thelitho-genicelementtitanium(Ti)wasusedasacontrolforthemobileele-mentsas it isgeochemicallystableandhostedbyresistantminerals(Daviesetal.,2015).Theelementphosphorus (P)wasusedtotracklivestockgrazing.CattledirectlyaffectthelocalPcyclebecausethephosphorus they consume is returned to the environment as dung(Parham,Deng,Raun,&Johnson,2002).ThisincreasesPconcentra-tionsinthesoilandultimatelycontributestoPenrichmentinwaterbodiesandsediments.

Sedimentsamples(33fromthecoreandtensurfacesamples)of0.2geachweredigestedusing1mlHNO3(Aristar,BDH,AU)and2mlHCl (MerckSuprapur,Darmstadt,Germany)andanalysedformetalsusing an inductively coupled plasma mass spectrometer (ICP-MS;

Perkin-Elmer,Waltham,MA,US;modelDRC-e)withanAS-90auto-sampler(Telford,Maher,Krikowa,&Foster,2008).Certifiedreferencematerials (NRC-BCSS1, NRC-Mess1, NIST-1646, NRC-PCAS1) andblankswereusedas tocheck thequalityand traceabilityofmetals.PCAwasusedtolinkgeochemicalvariablestopollenassemblages.

3  | RESULTS

Figure2showstheadoptedage–depthmodel for theMP11Asedi-mentrecordwithdatedlevelsandassociatederrors.Two14Cdateswereexcludedfromthemodel(30.5-and40.5-cmdepth).Thesesam-ples yielded ages thatwere the same or older than the lowermost14Cageinthecore.Re-depositionofolderCbyfloodsisacommonproblemwhendatingrecentfloodplaindeposits(Ely,Webb,&Enzel,1992).Thechronologyestablishedby210Pb has low errors and indi-catesrelativelyconstantsedimentationrates.Theapparentincreasein sedimentation toward the top of the core is likely to reflect therelativelyuncompactednatureofthemorerecentsediments.

Figure3showskeychangesinthevegetationoftheMP11Asiteanditssurroundingsthroughthelast200years(seeAppendixS2forcompletepollenrecord).Fourmainphasesweredefinedbyzonation,describedbelowinrelationtochangesincharcoalandgeochemistry:

1. Phase 1 (AD 1823–1907) is characterized by relatively highproportions of Corymbia, other Myrtaceae, Callitris, Terminalia,Banksia and Antidesma ghaesembilla pollen. Phase 1 pollen as-semblagesareverydifferenttothoseofsurfacesamplescollectedin modern waterholes (see PCA, Appendix S3). Microscopic

F IGURE  2 Age-depthmodelfortheMP11Asedimentcore,showingcalibratedagesandassociatederrorsfrom210Pb and 14Cisotopicdating,withsmoothsplineinterpolation

210Pb210Pb

210Pb

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charcoal accumulation rates were relatively high in Phase 1compared with later phases, whilst macroscopic charcoal accu-mulationrateswerelow(average4%totalcharcoal).Geochemicalindicators show a dip in concentrations of weathering resistantTi later in this phase and an increase in silicon (Si).

2. Phase 2 (AD 1907–1948) exhibits a steep decline in Antidesma ghaesembilla and Banksia pollen.Pollenof the canopydominantsfluctuated,withCorymbia and Callitrisattainingtheirhighestpro-portions intheentiresequencebeforerapidlydecreasingtowardtheendofthephase.Botryococcusdecreasedsubstantiallyatthesame time and Sordaria fungal spores increased in abundance.Charcoaldeclinedsubstantiallytowardtheendofthisphase,reach-ingitslowestconcentrationsintheentirerecord.Incontrast,themobileelements(Na,KandBa)increasedwhileTiconcentrationsdeclined.TheendofthisphaseismarkedbyasharpincreaseinP,reachinglevelsovertentimesbackgroundconcentrations.

3. Phase3(AD1948–1993)beginswithmajorincreasesinPoaceaeand Pandanuspollen.Thesechangeswerefollowedbyamajorin-creaseinbothmacroscopicandmicroscopiccharcoal,reachingthehighestconcentrationsintherecord.Treepollendecreasedtoan

averageof34%(comparedto50%inthepreviouszone).Phase3pollen assemblages are similar to those of surface samples frommodern waterholes (see PCA, Appendix S3). Geochemical datashowatemporarydeclineinconcentrationsofNa,KandBaandanincreaseinTi,MgandCa.

4. Phase4(AD1993–2016)isdifferentiatedfromthepreviousphasebyafurtherdeclineintreepollen(average:30%)andthehighestlevelsofPoaceae,Acacia and Dodonaeapollenfortheentirerecord.Macroscopiccharcoalisabundantcomparedtomicroscopicchar-coal (average 30% total charcoal). Sordaria remained abundantthrough this phase. Geochemical indicators show relatively highconcentrationsforallelementsexceptP,whichdecreased.

Figure4 shows the relationships betweenmajor pollen taxa andvegetationparametersforsurfacesamples(AppendixS4).Corymbiapol-lencorrelatesstronglywitharea-weightedplantabundance (r2=.87),savanna basal area (r2=.74) and savanna canopy cover (r2=.48).Pandanuspollenwas lessstronglyrelatedtoabundance (r2=.37)andhadno relationship tosavannabasalarea (i.e., treedensity).Poaceaeproduced a short abundance gradient (being dominant in vegetationsurveys), but displayed an inverse relationship with savanna density(r2=.42) and canopy cover (r2=.27). Corymbia:Poaceae ratios cor-relatedpositivelywithbothCorymbiaabundance(r2=.64)andsavannadensity(r2=.67).

4  | DISCUSSION

Ourmulti-proxyresultsprovideaglimpseofhistoricalchangesthatoccurred in the savannas and inland floodplains of the KimberleyRegion. A longer record of palaeovegetation change comes fromBlack Springs, 75km SW of site MP11A (15°38′S, 126°23′23″E),showing PandanusexpansionduringHolocenewetphasesandaridi-ficationcycleslinkedtoweakeningmonsoons(Fieldetal.,2017).Thisrecord,however,has lowtemporal resolution (centennial scale)anduncertain dating for recent centuries. TheMP11A record providesdecadalresolution,revealingasequenceofenvironmentalchangesintheupperMitchellRiverareasinceEuropeancolonization.The210Pb chronologyshowsnosignsofflooddisturbance(Arnaudetal.,2002).Nevertheless,theassignedagesshouldbeviewedasapproximate.

The sequence of events is summarized in Figure3. An ini-tial decline of the riparian shrub Antidesma ghaesembilla began around1900(Phase1),followedbyBanksiadeclineadecadelater.Corymbia,indicativeofsavannadensity(Figure4),increasedaround1912andCallitrisfollowedinthe1920s(Phase2).Atthesametime,regional fires decreased (indicated by lower levels ofmicroscopiccharcoal)andgrazingintensified(indicatedarapidincreaseindungfungalspores,suchasSordaria).The1930sbroughthighergrazingintensity,increasingerosion(indicatedbyrisingBaandNaconcen-trations)andamajordeclineinsavannatrees(Corymbia and Callitris)and regional burning. Eutrophication followed in the 1940s,witha pronounced peak in P lasting until the 1970s (Phase 3).Duringthis interval,grassabundance increasedandPandanus replacedA.

F IGURE  3 SummaryofmajorchangesinenvironmentalproxiesatsiteMP11A,MitchellPlateau,WesternAustralia(seeAppendixS2forcompletepalynologicalandgeochemicalproxydiagrams)

1850 1900 1950 2000 AD

Micro-charcoalMacro-charcoalNeurospora

Dung fungiBarium

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ghaesembillaasthedominantriparianelement.Fireoccurrencerosesubstantiallyinthelate1970s,particularlythelocalfiresindicatedby macroscopic charcoal andNeurospora fungal spores (van Geel&Aptroot,2006;Whitlock&Larsen,2001).This increase in localburningwasaccompaniedbyevidenceofincreasedgrazingintensity(dungfungi),peakingaround1990.Thelasttwodecadeshaveseenfurther expansion of grasses and a decline in savanna density, aswellasthedisappearanceofPandanusfromthewaterholemargins(Phase4).

Observed changes in savanna fire regime, savanna vegetationstructureandriparianzonevegetationarediscussedinthefollowingsectionsinrelationtomajordriversofenvironmentalchange.

4.1 | Changes in savanna fire regime

On a global scale, savannas are thought to exist as an alterna-tive stable state to forests, maintained by frequent fire (Bond,Woodward, & Midgley, 2005). Changing fire regimes are impli-cated in major vegetation shifts across the Northern Australiansavanna zone (Bowman, 2003; O’Neill, Head, &Marthick, 1993;Russell-Smithetal.,2003), including thesimplificationofvegeta-tionstructure(Craig,1997),compositionalchange(Bowmanetal.,2014;Trauernichtetal.,2013),shiftingforest–savannaboundaries(Banfai&Bowman,2005;Hnatiuk&Kenneally,1981) andnega-tive impacts on nativemammals (Radford etal., 2015; Ziembickietal.,2015).TheeffectsoffireregimechangeinAustraliaarenotcompletely understood, given a lack of information on historicalfireregimesandfireecology(Silcock,Witt,&Fensham,2016).Fireregime change is often linked to the dispossession of Aboriginalpeoples of their land and consequent loss of traditional burningpractices,followedbythedifferentburningpracticesofEuropeancolonists (Ritchie,2009;Russell-Smithetal.,2003).Alternatively,fire regime change may stem from the invasion of fire promot-ing grass species, creating a grass–fire positive feedback cycle(Bowmanetal.,2014).

Our data demonstrate profound changes in fire regime aroundtheMP11Asiteover200years.Phase1samplesaredominatedbymicroscopiccharcoal.Thissuggeststhatregionalsavannafireswerecommon,whilelocalfiresintheriparianzonewereapparentlyrare.Thisagreeswithethnographicevidence(Russell-Smithetal.,1997;Vigilante, 2001). The major reduction in regional burning in the1930sand1940s(Phase2)coincideswiththedeclineofcontinuousAboriginaloccupationoftheMitchellPlateau,ending inthe1950s(Wilson, 1981). Data from the 1960s onwards (Phases 3–4) showthatlocalfiresbecamemoreprevalent,impactingbothsavannaandriparian zones (Figure3). Historically, Aboriginal fire managementisassociatedwith light/early season firescompared tounmanagedareas,whereseveredryseasonfiresaremorefrequent(O’Neilletal.,1993).

ThenatureoftheMP11Asitelimitstheconclusionsthatmaybedrawn fromcharcoal analysis. Fluvial transport is amajor sourceofprimarycharcoal(directproductsoffire)andsecondarycharcoal(re-depositedbyerosion:Patterson,Edwards,&Maguire,1987;Whitlock&Larsen,2001).Overlappingradiocarbondatesatdifferentsedimentdepths could indicate contamination by older (secondary) charcoal(Figure2).However,themacroscopiccharcoalrecordisvalidatedbyasimilartrendinNeurospora,abiologicalproxyforlocalfire(vanGeel&Aptroot,2006).Charcoalabundanceisnotclearlyrelatedtoerosionproxies(Figure3)andthereisnoindicationofamajorshiftinsedimentsource that explains the dramatic increase inmacroscopic charcoalsincethe1960s.Hence,theMP11Acharcoalrecordiscautiouslyin-terpretedasareflectionoffireoccurrenceintheriparianzone(macro-scopiccharcoal)andsurroundingsavanna(microscopiccharcoal).ThechangesareconsistentwithatransitionfromAboriginaltoEuropeanfire management (Haberle, Tibby, Dimitriadis, & Heijnis, 2006), butremaintobecorroboratedatothersites.

F IGURE  4 Scatterplotsofplantabundanceandsavannadensityparametersversuspollenabundancesinsurfacesamples

aib

myro

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nadn

aPea

ecao

Pai

bmy

roC

eaec

aoP:

Savanna density4 5 6 71 2 3 4 5

0

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0

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25

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0

1

Plant abundance

Pol

len

(%)

r2: .64

(basal area: m2/ha)(weighted DAFOR)

Pol

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ratio

r2: .67

r2: .00 r2: .42

r2: .07r2: .37

r2: .87 r2: .74

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4.2 | Changes in savanna structure

Acriticalquestion inunderstandingsavannavegetationdynamics ishow global-scale factors interact with the local environment (Case& Staver, 2017). Previous research has debated the role of fire vslocaledaphicfactorsindeterminingvegetationstructureinnorthernAustralia (Bowman, 1988; Craig, 1997;Douglas etal., 2015;Ondeietal., 2017). These studies often used space-for-time substitutionor repeataerialphotography toanalysevegetationchanges.Space-for-timesubstitutionislimitedbyconfoundingvariablessuchasdif-ferencesinsoiltype,climate,speciespoolsandlanduseatdifferentsamplinglocations.Aerialphotographyprovideslong-termdataonabroadgeographicscale,buthas limitedtemporalscope (usually late20th century) and provides little detail on changes in understoreyvegetationbeneaththecanopywheretheimpactsofsavannafirearepresumablygreatest.Palaeoecologicaldatafromsmallbasins,suchastheMP11Arecord,revealvegetationchangesatasmallspatialscaleoverlongtimescalesandareideallyplacedtoinformdebatesonveg-etationstructure(Marianietal.,2017;Morrisetal.,2017).

Ourresultspointtoa long-termdecline insavannatreedensity.Thedecline is registered in decreasingCorymbia pollen abundancesand Corymbia:Poaceaeratios,proxiesstronglyassociatedwithsavannabasalarea(Figure4).Treedensitydeclinewasparticularlyacuteinthe1930s.Thisevidence,albeitfromasinglesite,contrastswithobser-vationsof20thcentury‘woodythickening’acrossnorthernAustralia(Bowmanetal.,2010;Tngetal.,2011;cf.Murphy,Lehmann,Russell-Smith,&Lawes,2014).Observedchanges invegetationdensityaredependentonthebaselineused.Theearliestaerialphotographyformost of northernAustralia dates to the1940s and1950s. Savannadensity change at MP11A since the 1940s has been negligible(Figure3).Comparedtoabaselineofthe19thorearly20thcentury,however,losshasbeensubstantial.

AroundMP11A,grasscoverincreasedassavannadensitydeclined(Figure3).Historically,livestockhavecausedmajorchangesingrass-land species composition in the Kimberley Region (Petheram, Kok,&Bartlett-Torr,1986).Cattle, sheep,pigs,horsesanddonkeyswereintroducedbyEuropeangraziersfromthe1880s(McGonigal,1990).Livestocknumbersincreasedrapidlyfromlessthan100,000cattleinthe1890s tomore than600,000 in1914 (Bolton,1953), andwerelinked to environmental degradation from the 1930s (Payne etal.,2004).Cattlegrazingcantriggeratransitionfromperennialtoannualgrasses inAustraliansavannas (Ash,McIvor,Mott,&Andrew,1997;Fensham&Skull,1999).Thesametypeof transition is linkedto in-creasingfirefrequency(Bowmanetal.,2014).ThehistoriesofgrazingandfireatMP11Aaresolongandintertwinedthatasinglecauseofrecentsavannastructuralchangeisunlikelytoemerge.

Perennial Triodiagrasses,ifleftunburnedforlongperiods,cangainsignificantsizeanddensity,providinghabitattoarangeofvertebratesand invertebrates.Old tussocks, often exceeding2m in height and5m inbreadth,werefrequentalongtheFitzroyCrossingRoaduntilthemid-1980s.Recent changes in fire regimehavegreatly reducedtheextentofunburnedTriodiatussocks,achangethatmaybelinkedtothedeclineofsmallmammalsintheregion.Pollendatacannotbe

usedtointerrogatethehistoryofspecificgrassspeciesbecausediffer-entspeciesproducepollenthat ismorphologically indistinguishable.AnalysesofphytolithsorancientDNAmayhelpaddressthisshortfall.

Callitris intratropicahasbeen the targetofconsiderable researchintofireimpactsinnorthernAustraliansavannas(Bowman&Panton,1993;Bowmanetal.,2014).Callitrispollenismorphologicallydistinct,andtheMP11Apollenrecordprovidesinsightintotherecenthistoryof this fire-sensitiveconifer.Callitrisapparentlyexperiencedamajordecline inthe1930sandhassincerecoveredslightly.C. intratropica treeswerenotsufficientlyabundantinourvegetationsurveystoper-mit a quantitative test of the grass–fire cycle hypothesis (Bowmanetal., 2014). Theoretically, if the expansion of grasses is linked toCallitrisdecline, the ratioofCallitris:Poaceaepollenshoulddecreaseovertime.Instead,thereisanincreasesincethemid-20thcenturyatMP11A (Figure3).Thepurported linkbetween fire andgrass coverfindslittlesupportinourdata,withcharcoalandPoaceaepollenfol-lowingdifferenttrajectories.Ourdatacannotanswerquestionsaboutgrazing–fire–grassinteractionsonaregionalscale,butpointtoapro-foundlegacyof20thcenturygrazingandfireimpactsintheMitchellPlateausavannas.

4.3 | Changes in the riparian zone

DatafromtheMP11Awaterholeprovidestrongevidenceforthe20thcenturydeteriorationofariparianshrublayercomprisingBanksia and Antidesma. BanksiapollenrepresentsB. dentata,theonlyBanksiaspe-ciesextant intheKimberleyregion.Thisnon-serotinousspeciesoc-cursinseasonallyinundatedareasandswampmarginsonsandstonebedrock, where it grows alongside Eucalyptus apodophylla (Hnatiuk& Kenneally, 1981). Extensive aerial surveys across the Kimberleyconfirm this almost exclusive habitat restriction (R. Barrett, per-sonalobservation).OurdatasuggestthatBanksia dentata also occu-pied riparian zones in the past (i.e.,Melaleuca leucadendron alliance ofHnatiuk&Kenneally,1981;RiparianThicketsgroupofMucina&Daniel,2013).

Antidesma ghaesembillaisafire-sensitiveriparianshrubthatbearsedible fruits and is known to expand rapidly in areas of unburnedwoodland (Bowman, Wilson, & Hooper, 1988; Russell-Smith etal.,2003).ItoccursfrequentlyalongtheMitchellRiverfloodplaintoday.The MP11A pollen record suggests A. ghaesembilla was dominantaroundthissiteuntiltheearly20thcentury.Itsdeclinefollowsapeakinmacroscopiccharcoal,implicatinglocalfiresasthecause.Accordingtoethnographic sources,Aboriginalpeopleavoidedburning riparianvegetation (Russell-Smith etal., 1997;Vigilante, 2001). Post-fire re-coverymayhavebeenimpededbygrazing,asAntidesmaspeciesaremorepalatabletocattlethanbrowse-andburning-resistantPandanus.

Riparianandwetlandzonesarewherecattleandferalanimalim-pacts in theKimberley aremost acute (Legge,Murphy,Kingswood,Maher,&Swan,2011).Grazingandtramplinghavehadsignificantdet-rimentalimpactsonriparian‘bamboo’(Phragmites karka),oncecommonintheKimberley,butnowlistedasaspeciesofconservationconcerninwesternAustralia.Enhancederosion isattestedgeochemicallybyincreased concentrations of mobile vs immobile elements (Bierman

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etal.,1998).AtMP11A,mobileNa,KandBaincreasedasrelativelyimmobile Ti decreased during the 20th century. Erosion is furtherindicated byPseudoschizaea palynomorphs (López-Marino,MartínezCortizas,&LópezSáez,2010),whichpeakinPhases3–4.GrazingmaycausePenrichmentfromfaecesandlossesofCaandMgfromurinepatches(Early,Cameron,&Fraser,1998;Parhametal.,2002).IntheMP11Arecord,theearly–mid20thcenturyischaracterizedbyanin-creaseinP(Figure3)andadecreaseinCaandMg(AppendixS5).Withtheadditionalsupportofproxiesfor livestockdung(Figure3),thesegeochemicalchangesmaybelinkedtograzingimpacts.

SomeofthechangeswitnessedintheMP11Arecordcouldbeex-plainedbyotherenvironmentalshiftsoccurringintheMitchellRivercatchment.Larsen,May,Moss,andHacker(2016)demonstratedhowthegeomorphicprocessofknick-pointmigrationcausedriparianfor-est loss inAustralia’sNorthernTerritory.Sandslugs releasedbyup-streamerosioncauselong-lastingimpactsondownstreamecosystemsin many Australian rivers (Prosser etal., 2001). These explanationsseem inadequate in thecaseofMP11A–aerialphotographsof thestudy area in 1949 and 2017 show no obvious change in riparianforestandfloodplaingeomorphology(AppendixS6).

Therearenomajor floodingevents corresponding to the timeofpeakimpactatMP11A(Gillieson,Smith,Greenaway,&Ellaway,1991; Wohl, Fuertsch, & Baker, 1994). Meteorological recordsindicate some higher-than-average rainfall years in the 1940sand 1950s, butmuch higher values in recent decades (Bureau ofMeteorology), part of a wetting trend across northern Australia(Bowmanetal.,2010).Thedecades-longenvironmentaltransforma-tionevidentatMP11Acannotbeexplainedbyasingledisturbanceevent.Agreementbetweengeochemicalandbiologicalproxies forgrazing and fire, alongside clear evidence for the loss of savannatreedensityandtheshrubstratumwithintheriparianzone,suggestgeomorphicprocessesareanunlikely causeofhistoricvegetationchangesinthestudyarea.

4.4 | Implications for conservation and vegetation mapping

Our evidence casts doubt on the notion that northern Australia’slandscapesandecosystemsareintactandunmodified(Bowmanetal.,2010;Woinarskietal.,2007;Ziembickietal.,2015).Justastheintactcanopyof‘Australian’eucalyptsdisguisesasavannaflorawithstrongSEAsianaffinities(Haynes,Ridpath,&Williams,1991),thesamecan-opymaydisguisethepervasiveecologicaltransformationsthathaveoccurredoverthelast100years.Theforgottenimpactsofearly20thcenturyfireandgrazinghaveprofoundlyalteredsavannaandriparianecosystemsontheMitchellPlateau,andperhapsmorewidely.

Ourdatacallforcautioninadoptingmid-20thcenturybaselinesforassessingecologicalchange,giventhattheimpactsofEuropeancolonization often manifested themselves much earlier. PotentialNatural Vegetation (PNV) maps have similar limitations. BecausePNVmaps are populated from data collected in the current (im-pacted)environment,therearelikelytobeknowledgeshortfallsinreconstructingpre-Europeanvegetation(Beardetal.,2013).Given

thedurationandmagnitudeofhistorical impact, itcouldbeaskedwhether restoration to pre-European conditions is desirable orrealistic.

Thereisaclearneedforreplicationbeyondthesinglepalaeoeco-logicalrecordpresentedheretobetterunderstandspatialandtempo-ralvariationintheKimberleyregion’secosystems.Itisacknowledgedthat obtaining palaeoecological data for vegetation types existingundermorearidconditionsmaybedifficult(Head&Fullager,1992).HencePNVprovidesauseful‘nullmodel’(Somodi,Molnar,&Ewald,2012)thatcanonlybe improvedasbetterdataandnewsourcesofinformationcometohand.

Failure to recognize the Kimberley Region’s historically alteredecologicalstatecouldplacetheremainingbiodiversityandecosystemsatgreaterriskthancouldbeexpectedundertruly ‘intact’ecologicalconditions.

ACKNOWLEDGEMENTS

We acknowledge the traditional owners of the land on which ourresearch was conducted. Sincere thanks to Cecelia Myers, SusanBradleyandDoonganStationstaffforfacilitatingandsupportingourresearch.ThankstoJanelleStevenson,PeterKershawandananony-mous reviewer for constructive reviews. Funding was provided bytheKimberleyFoundationofAustraliaandtheARC(DP140103591)project ‘Environmental Transformations linked to Early HumanOccupationinNorthernAustralia’.ResearchisalsosupportedbytheARC Centre of Excellence for Australian Biodiversity and Heritage(CE170100015).SECledthewritingandconductedfieldwork;LSledthegeochemicalandstatisticalanalysesandcontributedtothemanu-script;JTanalysedthepalynologicaldataandledtheliteraturereview;SRanalysedadditionalpalynologicaldataandoversawpollentaxon-omy;RLBcontributedtothemanuscript,particularlyits“Introduction”and “Discussion” sections; AZ provided 210Pb dating and expertise;SGHcoordinatedtheresearch,conductedfieldworkandcontributedtothemanuscript.

ORCID

Simon E. Connor http://orcid.org/0000-0001-5685-2390

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SUPPORTING INFORMATION

Additional Supporting Information may be found online in thesupportinginformationtabforthisarticle.

Appendix S1.Mapofsamplinglocations,NWKimberleyRegionAppendix S2. Complete pollen and geochemical diagrams, MP11ArecordAppendix S3.PrincipalComponentsAnalysisresultAppendix S4.PollenandvegetationdatafromsurfacesamplesAppendix S5.GeochemicaldatafromsurfacesamplesAppendix S6.Aerialphotographsofstudyareain1949and2017

How to cite this article:ConnorSE,SchneiderL,TreziseJ,etal.ForgottenimpactsofEuropeanland-useonriparianandsavannavegetationinnorthwestAustralia.J Veg Sci. 2017;00:1–11. https://doi.org/10.1111/jvs.12591