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JOURNALOFTHEANTHROPOLOGICALSOCIETYOF

SOUTHAUSTRALIA

SpecialEditionAboriginalRelationshipswithRiverCountry:

ArchaeologicalandAnthropologicalPerspectivesontheMurrayDarlingBasin

VOLUME41–DECEMBER2017

EDITORSAmyRoberts,MichaelMorrison,IanMoffatandHeatherBurke

JOURNALOFTHEANTHROPOLOGICALSOCIETYOFSOUTHAUSTRALIA

Volume41,December2017123

DIET,MOBILITYANDSUBSISTENCE-SETTLEMENTSYSTEMSINTHELATEHOLOCENELOWERMURRAYRIVERBASINOFSOUTHAUSTRALIA:TESTINGMODELSOFABORIGINALSEASONALMOBILITYANDSEDENTISMWITHISOTOPICANDARCHAEOLOGICALDATA___________________________________________________________________________F.DonaldPate1

1CollegeofHumanities,ArtsandSocialSciences,FlindersUniversity,GPOBox2100,Adelaide,SA5001,AustraliaAbstract___________________________________________________________________________Bone collagen stable carbon and nitrogen isotope data forarchaeological human remains suggest that the territorial behavioursreported in relation to Tindale’s ethnographic research for the lowerstretches of theMurray River of South Australia in the vicinity of therivermouthandadjacentcoastalregionscanbeextendedbackthroughthe late Holocene. Testing hypotheses regarding the presence of lateHolocene semi-sedentary and sedentary subsistence-settlementsystemsalongvariousregionsoftheLowerMurrayandadjacentcoastrequires comprehensive archaeological research, including improvedchronometricdating fora rangeofarchaeological sites,demonstrationof the use of a range of key plant and animal foods, and expansion ofstableisotopeapplicationstoincludestrontiumandoxygen.ThispaperprovidesanoverviewofthenaturalenvironmentoftheLowerMurrayRiverregionofSouthAustralia,includingtheavailabilityofwaterandarange of food resources. The importance of foods available in thevicinity of the riverine zone that could have provided reliable dietarysources is emphasised. Such ecological data regarding the abundanceand availability of key plant and animal foods are fundamental toimprovedunderstandingsofhunter-gathererbehaviouralvariability intheregion.



Introduction___________________________________________________________________________Aboriginal subsistence-settlement systems in the ecologicallydiverse Lower Murray River region of South Australia (SA)wouldhavebeeninfluencedsignificantlybythedistributionandabundanceofwater, floraand fauna (Lawrence1968;Peterson1976; Tindale 1981). This paper investigates the interaction offood and water resources available in the region and theexploitationofthoseresourcesbypastAboriginalpopulations.Itprovidesanoverviewofsomeofthekeyplantandanimalfoodsthat could potentially allowmore sedentary lifeways, includingyear-rounduseofbasecamps.

Previousresearchhasdemonstratedthatsedentaryandsemi-sedentary settlements are possible among hunter-gatherers in areas with abundant and reliable water and foodresources. Furthermore, in order to increase the reliability offood resources over longer periods of time, hunter-gatherersextended the availability of key foods by employing foodpreservation, storage, fire management, and technologicalinnovationsincludingfishweirs,netsandholdingareasforeelsand fish (Bettinger 1982; Builth 2002; Cohen and Armelagos1984; Fitzhugh and Habu 2002; Gamble 2008; Hiscock 2007;Kelly1983;LeeandDeVore1968;Lourandos1985;Owen2004;Pate1997, 1998, 2006;Pate andOwen2014;Price andBrown1985;Williams1987).

TheLowerMurrayregionwasoneof themostdenselypopulated geographic areas of Australia prior to Europeancolonisation (Clarke 1994; Mulvaney 1969:40; Radcliffe-Brown1918:230-231; Wilson 2017). On the basis of historical andethnographic accounts from the Lower Murray region of SA,Clarke (1994:75) reported that ‘Aboriginal descent groupterritories in this resource rich riverine-coastal zone, beingrelatively small, reflected amore sedentary life style’ (also seePeterson 1976). However, within these territories seasonalmovements of habitation sites occurred in relation to harshcoastal weather and limited food sources associated with thewintermonths.Somenon-coastalgroupsalsopracticedseasonalmovements from lakes and rivers to nearby sheltered inlandareasduringwinter(Clarke1994;Tindale1981).

Archaeologists have employed isotopic analyses ofhunter-gathererskeletalremainsfromvariousglobalregionsto



test ethnographic and historical models related to seasonalmobility versus more sedentary behaviours associated withyear-round use of base camps. Subsistence-settlement systemsbased on ethnographic and historical data cannot simply beextended to the archaeological past employing analogies.Variability in hunter-gatherer subsistence-settlementbehaviours across space and time must be investigated usingarchaeological data. In some cases, the isotopic data contradicttheethnographicandhistoricalmodelsandsuggestthepresenceof different behaviours in the past, while in other cases theisotopicdata support the long-termpersistenceof subsistence-settlement systems (cf. Roberts et al. 2013; Sealy and van derMerwe1986, 1988; Sealy 2006; Tafuri et al. 2017;Walker andDeNiro 1986;). Oxygen isotope analyses of shells fromarchaeological middens and mounds have also been used todemonstrateseasonaloryear-rounduseofcoastalsites(Jewetal.2014).

Bonecollagenstableisotopedata(PateandOwen2014)suggest that the territorial boundaries reported by Tindale(1974)fortheLowerMurrayandadjacentcoastalregionsofSAwere established by the late Holocene. Stable carbon andnitrogenisotopedataforcoastalskeletalsamplesdemonstrateaheavyrelianceonmarine-basedfoodsandlimitedmovementtoinland riverine and terrestrial habitats (Pate et al. 2002;Owen2004). Further research is required in order to determine theextenttowhichsedentaryandsemi-sedentarybehaviourswereassociated with hunter-gatherer populations along the upperinland stretches of the Murray River in SA (cf. Clarke 1985a,1985b,1994,1998,2009,2015;Hilletal.2016;Pate1997,2006;Littleton 1999; Littleton and Allen 2007; Martin 2006, 2011;WestellandWood2014).

StableisotopedatafortheinlandriverineSwanportandRoonka archaeological populations near Murray Bridge andBlanchetown, SA (Figure 1), respectively, indicate that lateHolocene inhabitants focusedonC3-basedplantananimal foodsourcesassociatedwiththeriverandadjacentplainsanddidnotinclude marine foods in their diets (Pate 1998a, 1998b, 2000,2006). Thus, marine foods from the southern coast were not



being imported to inland riverine regions and people living atSwanportandRoonkadidnotspendsignificantamountsoftimeinsoutherncoastalregionswheretheywouldhavehadaccesstomarine foods. However, the bone collagen stable carbon andnitrogen isotope data do allow formovement of the SwanportandRoonkainhabitantsalongtheinlandstretchesoftheriver,assimilar C3-based plant and animal foods would be associatedwiththeseregions.

The expansion of stable isotope research to includestrontiumandoxygenintoothenamel(Bentley2006;Buddetal.2004; Pate 2008a, 2008b) could provide more detailedinformationregardingrestrictionofhunter-gatherermovementsbetween various geographic localities and provide data toexaminehypothesesregardingsemi-sedentismandmobility forboth coastal and inland riverine regions. Westell and Wood(2014) andMartin (2006, 2011) argue that earthmound sitesalong the Lower Murray and Murray Riverine Plain wereassociated with repeat-use pit ovens for the intensiveexploitation of wetland resources for food and fibre. Theysuggest that the use of reliable root/tuber foods like bulrush(Typha spp.) allowed increased levels of residency andpopulationgrowthfromthemidtolateHolocene.

The reliable riverine water sources and diverse foodsassociated with the Lower Murray and adjacent mallee plainecosystems, provided the environmental conditions for thepotentialestablishmentoflong-termresidentialhunter-gathererbases associated with logistical mobility to access foods andother resources in the surrounding landscapes (Binford 1980,1982,1983;Bettinger1999;HabuandFitzhugh2002).

Pate (2006:238–239) provides an overview of theethnographic and historical accounts relating to thesocioeconomically complex Ngarrindjeri, who were organisedinto a number of territorial clans. The Ngarrindjeri occupied alargetriangularareaofcoastalandriverinelandstretchingfromjust aboveMurrayBridge in the north toEncounterBay in thesouthwest andKingston in the southeast. This region includesthe coastal Coorong, the Murray River mouth, and the LowerMurrayRiverinthevicinityofSwanport(SeeBell1998;Berndtand Berndt 1993; Jenkin 1979; Meyer 1846; Radcliffe-Brown1918;Salgado1994;Taplin1859–79,1874;Tindale1974;Pateetal.2003;Wilsonetal.2012;Wilson2017).



In addition, ethnographic and historical accountsidentifyarangeofkeyplantandanimalfoodsthatwereusedintheresourcerichLowerMurrayregionat thetimeofEuropeancontact. Coastal inhabitants had access to marine fish andshellfish,waterfowlandtheireggs,marinemammals(seals,sealions and stranded dolphins and whales), and vegetable foodslike berries. Further inland, key food sources includedfreshwater fish, shellfish, crayfish, yabbies, turtles, waterfowl,bird eggs, possums, rats, bandicoots, wombats, snakes, lizards,bardi(orwitchetty)grubs,andbulrushroot(Typhaspp.).Valuedmeat was obtained from larger mammals such as kangaroos,wallabies and emus (Angus 1847; Berndt and Berndt 1993:80;Clarke 1994, 2009, 2011, 2015; Pate 2006:237–238; Worsnop1897).Theavailabilityofarangeofreliableriverinefoodswouldhave provided an environment that allowed the potentialdevelopment of sedentary and semi-sedentary subsistence-settlementsystems.

Archaeologicalresearchsuggests thatamajorityof thelate Holocene earth mounds concentrated along the MurrayRiverine Plain of southeastern Australia were associated withthe baking/steaming of carbohydrate-rich wetland plant foodsuch as bulrush (Typha), club rush (Bolboschoenus) and waterribbons (Triglochin), as reported inethnographicandhistoricalaccounts for the region (Coutts et al. 1979; Gott 1999; Martin2006,2011;WestellandWood2014).Thesemoundsconsistofashysediments,charcoal,bakedclayheatretainers,bone,stoneand shell artefacts, a range of wetland faunal remains, andoccasionallyintrusivehumanburials(Martin2011:162–163).

Theseethnographicandhistoricalrecordsprovidedatato construct hypotheses about prehistoric diet and associatedsettlement systems in the region. A comprehensiveunderstanding of the distribution and abundance ofwater andfood resources in the Lower Murray will improve scientificprocesses involving the generation and testing of hypothesesemploying archaeological data. Detailed information regardingthe abundance and distribution of key plant and animal foodsassociated with the riverine ecosystem will provide importantdatatoaddresshypothesesregardingtheexistenceofsedentary



andsemi-sedentarylifewaysintheLowerMurrayregionduringthe late Holocene. Expanded archaeological research in theregionwillimproveourunderstandingofAboriginalbehaviouraldiversityacrossthediverseLowerMurrayecosystems.

Figure1Mapshowingkeyplacesreferredtointhetext(adaptedfromPate2006).



MurrayRiverEcology___________________________________________________________________________TheMurrayRiveristhemajortributaryoftheextensiveMurrayDarlingriversystemofsoutheasternAustralia.BoththeMurrayand the Darling rise in the higher rainfall areas of the GreatDividingRangeon theeastern coast and then flow through thedry, flat lowlands to the southwest. The rivers deliver largequantities ofwater to these semi-arid lands. TheMurray Riveroriginatesinthe2228mKosciuskoPlateauofNewSouthWalesand flows for approximately 2600 km to the Southern Oceansouthwest of Adelaide, South Australia (Gill 1973; Hills 1974;Jennings2009;Walkeretal.1986).

The Murray is the only major river in SA. Its watersenterthestateonthesoutheasternborder,flowwesterlytowardthe Mount Lofty Ranges and then southerly to the ocean. SAtopographyisgenerallylowinrelief.Approximately50percentof thestate is less than150mabovesea levelandover80percent is lessthan300m.TheMurraydropslessthan22minits642kmtrekbetweentheborderandtheocean(Aitchison1974).

SAisthemostaridstateinAustralia,with83percentofthelandmassreceivinglessthan250mmofrainfallperannum.The northeastern region of the state occurs in the arid core ofthe continentwheremean annual rainfall is less than 125mm(Gentilli 1971, 1972; Lee and Gaffney 1986; Williams 1979).Consequently, the Murray River, with its abundant watersupplies,providedanimportantecosystemforinlandsettlementbypastAboriginalpopulations.

The river morphology exhibits great variability withinthestate.FromthestatebordertoOverlandCorner(ontheriver75 km east of Roonka, Figure 1) the rivermeanders through awide alluvial valley bordered by tall limestone cliffs. BelowOverland Corner, the Murray has dissected the upliftedlimestone Pinnaroo Block producing a steep-sided gorgetopography(Twidaleetal.1978).TheLowerMurrayGorgewasincised during middle and late Pleistocene glacio-eustaticregressions (Twidale et al. 1978).Today, limestone cliffs tower30–40mabovetheriverbottom.Thisdeeplyentrenchedgorgehas restricted floodwater dispersal and swamp development



(Twidale et al. 1978). Thus, there is an abrupt transition fromthe surrounding semi-aridmallee eucalypt plain to the narrowalluvialriverflats.Thelimestonecliffsdecreaseinheight30kmsouth of Blanchetownnear SwanReach, andwide alluvial flatsemergeagain.Numerousswampsandbroad,shallowlakesoccurinthiscoastal lacustrinezone.Theseconditionscontinueforanadditional90kmtotherivermouthatLakeAlexandrina.(BrownandStephenson1991;Fenner1931;Firman1973,1985).

The modern river morphology became establishedbetween 8000 and 6000 years ago. Ancestral rivers in theMurrayDarlingsystemdatingfromca.14,000to7000BPweregenerally larger and straighter than theirmodern counterparts(Fluinetal.2009;Hilletal.2009;Pels1964;Prendergastetal.2009). The change of the Murray River to its present slower,meanderingmorphology resulted in increased floral and faunalproductivity associated with the increased representation ofshallow lagoons, billabongs, swamps, and lakes (Brown andStephenson 1991; Gingele et al. 2007; Pardoe 1995). LakeAlexandrina at the rivermouthwaswell-established by 7800–7600calyrsBP(Fluinetal.2009).

Priortohistoricalinterventions,likelocksanddams,thewater level of the Murray River was primarily affected byprecipitation in the eastern highlands. An examination ofsediments in a deep sea core (MDO3-2611) collected off thecoastofSA(Gingeleetal.2007)identifiedtwoperiodsofhigherriver water levels and discharge at 13,500–11,500 and 9500–7500 years BP associated with higher rainfall in easternAustralia. During the period from 17,000–5000 years BP, thesedimentary signature in the core was dominated by alluvialsedimentsfromtheMurrayDarlingBasin(MDB).

Water derived from monsoonal rains is transporteddown the Darling and spring snow melt down the Murray.Waters begin to rise in the Lower Murray in the late winterthrough early spring.However, the arrival of floodwaters fromthe two sources usually does not coincide and there are twopeaks during this period.When they do coincide, a large floodresults and the Murray floodplain can be entirely inundatedduring theseheavy floods.Regular floodingcannotbeexpecteddue to the unpredictability of droughts in the highlands. Theriver usually reaches its highest level inmid-December. In lateDecember the water begins to recede and reaches its lowest



level during the late summer and fall (Burton 1974; Gill 1973;Johnson 1974). Water temperature ranges from 8.5° C in thewinterto25.5°Cinthesummerwithameanof16.8°C(Shieletal. 1982). Even during extended periods of drought in thehighlands that significantly reduced river flow, there werereliable water supplies available in deep recesses of the riverbottom(Burton1974).

The Lower Murray region has distinct seasonalvariability, as does most of SA. The summers are hot and drywithmeanmaximumJanuarytemperaturesof28–32°C.Wintersare cool and wet with July minimum temperatures of 3–5° C.Two-thirds of the rainfall occurs fromMay toAugust as gentleshowers. Scattered convectional thunderstorms are commonduring the summer months. The Mount Lofty Ranges create adistinct rain shadow effect. Mean annual precipitation reaches1200mmintherangeseastofAdelaideanddecreaseseastwarduntil only 250–300 mm is received along the Murray River.There is considerable variation in precipitation from year toyear, andextensivedroughts lasting from two to five years arecommon. The inland areas experiencemore frequent droughtsthan the wetter coastal region. Evaporation exceedsprecipitation during all seasons. Due to the high evaporationrates and sandy texture of the soils, standing surface water israre and ephemeral. Water will accumulate on claypans andsoaks (claypans filled with sand) for short periods followingrainfall. Smaller streamson theadjacentplainsoriginate in thebetter-watered upland regions. The major ephemeral streams,theMarneRiverandBurraCreek,draintheMountLoftyRanges(Aitchison1974;Hills1974;Nix1981;Tisdall1974;).

Historicalaccountsdescribeariverineenvironmentrichinsubsistenceresourcesat the timeofEuropeanexploration inthe 1830s and 1840s (Angus 1847; Clarke 2002, 2003, 2009,2011; Eyre 1845; Lawrence 1968; Pate 2006; Sturt 1833). Theriverwithitsassociatedfloodplain,swamps,lagoons,creeksandflats,andadjacentmalleeplainsprovideddiversehabitatswhichsupported a wide range of floral and faunal species. The riverbank and floodplain supported stands of river red gums(Eucalyptuscamaldulensis). Bulrushes (Typha spp.), rushes and



sedges (Juncus,Bolboschoenus[formerly Scirpus],Cyperus spp.),andreeds(Phragmitescommunes)wereassociatedwithadjacentswampy areas. This aquatic environmentwould have providedvariousedibleroots,waterfowl,fish,turtles,frogs,molluscs,andcrustaceans.Thedenselywoodedflatsbackfromthefloodplainwere covered with river box (E. largiflorens), coolibah (E.microtheca), black oak (Casuarina cristata), false sandalwood(Myoporumplatycarpum),lignum(Muehlenbeckiacunninghamii),shrubs, and perennial grasses. Various fleshy fruits, seeds,tubers, greens, insects, and small game (possums, koalas,rodents) would be available. Lagoons and creeks were foundadjacenttothesesavannawoodlands.Sandflatswerestabilisedby native pines (Callitris spp.) and perennial tussock grasses(Angus 1847; Clarke 2002, 2003, 2009, 2011; Eyre 1845;Lawrence1968;Pate2006;Sturt1833).

The adjacent mallee plains were vegetated by variousdwarf eucalypts (2–12 m in height), including oil mallee (E.oleosa), bull mallee (E. beariana), blue mallee (E. polybroctea),white mallee (E. dumosa), and slender mallee (E. calycogona),saltbush (Atriplex, Rhagodia spp.), bluebush (Maireana spp.),mulga(Acaciaspp.),andgrasses(Danthonia,Stipa,Triodiaspp.).Larger game (kangaroos, wallabies, emus), birds, reptiles(snakesandlizards),andvariousplantfoodscouldbefoundonthemalleeplains.TheplainsextendhundredsofkilometresintoVictoria toward the east and are bordered by theMount LoftyRangestothewest(BowlerandMagee1978;Cleland1966;Lautetal.1977;LeighandCostin1974;Paton1983).

PollenevidencefromswampandlakecoresedimentsinsoutheasternAustralia indicates that theperiod from50,000 to10,000 years BP was significantly drier than the Holocene(Dodson 1974, 1975, 1977; Hope 1984; Kershaw 1981). Theshift from a dry Late Pleistocene to wetter Holocene in theregion is supported by a range of palaeoclimatic data (Dodson1989;Johnsonetal.1999;Mageeetal.2004).

Dodson (1975) provides an overview of climate andvegetationchangesforsoutheasternSouthAustraliaonthebasisofpollendata.Thedriestperiodwasfrom26,000to11,000BP.Anopeneucalyptwoodlandwithheathunderstoreywaspresentbetween50,000 and26,000BP.During the followingperiod ofpeakaridity,amoreopenvegetationdominatedbygrassesandherbs (Poaceae and Asteraceae) with scattered eucalypts and



heathhadexpanded toproduceamorediversevegetalmosaic.This mosaic was replaced by the modern eucalypt woodlandwith heath and savanna understoreys around 10,000 BP. Thepresence of eucalypts throughout this period suggests that themean annual minimum temperature for the region was above10° C and themean annual rainfall was greater than 200–250mm.

According to pollen evidence from Lake Leake andWyrie Swamp in southeastern South Australia (Dodson 1974,1977), theclimateof theHolocenehasbeenrelativelystable.Aslightly wetter period occurred between 6900 and 5000 BP.From5000to2000BPconditionsbecamedrier,awetterperiodreturned from 2000 to 1300 BP, and from 1300 BP to thepresent it has been relatively dry again (Dodson 1974, 1977).Climatic data suggest that the Holocene floral and faunaldistributionsof theLowerMurraywouldhaveresembledthosereportedatthetimeofEuropeancontactduringthedryperiods,whereasthewetterperiodsmayhaveallowedexpansionofplantandanimalspeciesandarelativelyrichersubsistencebase.

Recent palaeoenvironmental research (Fletcher andMoreno2012;Gliganicetal.2014;Moyetal.2002;Pethericketal. 2013) suggests that there were substantial changes in thepatternandspatialdistributionofwestern-drivenrainfallintheSouthernHemisphere after 6000–5000 years BP in associationwith increased variability in the El Niño Southern Oscillation(ENSO). In some regions, variations in these large-scaleatmospheric systems would have resulted in warmer, drierperiods associated with increased ENSO activity followed bycooler,wetterphases.However,pollendata fromAustralia andthe Pacific islands indicate little climate change during theHolocene(Pickettetal.2004;Wanneretal.2008).Gliganicetal.(2014) employed dated shoreline lake sediments, lacustrineshells and speleothems to examine the impact of these large-scaleatmosphericvariationsonregionalpalaeoenvironmentsinsouthern central Australia. Elevated lake levels in the FlindersRangesofSouthAustraliaassociatedwith increasedrainfallareidentified for the periods 5800–5200, 4500, 3500–2700 and1000 years BP. They argue that these lateHolocene periods of



increased precipitation would have increased the reliability ofresources for regional human populations during a time ofreducedwinterrainfall(Gliganicetal.2014).

Thus a range of palaeoenvironmental evidence suggestthat Aboriginal people living along the Lower Murray Riverwould have had access to a rich riverine ecosystem during atleast the last 7000 years. In contrast, the abundance andreliabilityofplantandanimalfoodresourcesassociatedwiththeadjacent semi-arid mallee plains would have fluctuated duringthe late Holocene in relation to alternating cooler, humid andwarmer, drier climatic phases related to variations in ENSOactivity.

The perennial water supply which originates in theeasternhighlandscreatedauniquehabitatwithinthesemi-aridplainsregionthatwouldhaveprovidedalarge,permanentwatersource for Aboriginal populations. According to historicaccounts,theLowerMurraywasoneofthemostdenselysettledAboriginal areas of Australia, along with the northern, easternandsoutherncoastalregionswhererainfallwashighest.

In this district the natives were very numerous, theirencampmentsbeingscatteredalongthenarrowstripofgroundbetween the limestone cliffs and the water's edge: there theyfindplentyoffoodfromthefish,mussels,crayfish,bullrushrootandotherproductsofthislargerriver.(Angas1847:58)

The greater availability and reliability of food and waterresources in these areas supported larger populations. Eyre(1845) estimated an Aboriginal population of three to fourpersonspermileofriverintheMoorunderegion,11kmsouthofRoonka.Henotedthatsettlementsizeschangedfromseasontoseasonaccordingtotheabundanceoffood.Duringthesummer,the river flooded and increased the habitat size of mussels,crayfishandfish.Summercampswerequitelargeandconsistedof simple brush windbreaks on the banks of the river. Eyre(1845)had encounteredAboriginal congregations of up to 600individuals.Inthewinter,thepopulationsdispersedintosmallercamps consisting of solid log huts covered with grass andvegetationorrocksheltersinresponsetothecold,wetweather.Sturt(1833)alsoencounteredAboriginalgroupsexceeding100personsashe travelledsouthwardalong theLowerMurray.On



thebasis of theobservationsofTaplin (1879),Radcliffe-Brown(1918:230) estimated that the Aboriginal population in theLowerMurraywasbetween1800and6000people,atadensityof between1–3.2 individuals per square kilometre prior to theEuropean introduced smallpox epidemics (Clarke 1994:57–60;Humphries2007).

Ahigherdegreeofsedentismassociatedwithlong-termresidential bases or year-round occupation of residential basecamps would be possible in the resource rich regions of theLowerMurrayenvironment(Owen2004;Pate2000,2006;Pateand Owen 2014). However, some mobility may have beennecessary due to the scheduling and seasonality involvedwithfloral and faunal procurement (Binford 1980; Flannery 1968;Poiner 1976) and the distribution of food resources and stonetoolrawmaterialsacrossthelandscape.Stratifiedarchaeologicalsites such as rockshelters and mounds accompanied bycemeteries could be related to increased sedentism and use oflong-term residential base sites, but such year-round use ofresidentialbasesitesmustbedemonstratedemployinga rangeofarchaeologicaldata.

Anexaminationoftheseasonalavailabilityofplantandanimal foods provides one means to address the temporalassociationofhunter-gathererswithvariousarchaeologicalsites(Bailey 1983; Jochim 2012; Kelly 1983; Rocek and Bar-Yosef1998; Rossignol andWandsnider 1992). Physical anthropologyandstable isotopeanalysesprovidepowerfulmethods that cansupplement conventional archaeological data to examinevariability in pre-contact Aboriginal diet, health, settlementpatterns and landscape use in the Lower Murray River Basin(Katzenberg 2000; Larsen 2003; Pate 1997, 2000, 2017; PateandOwen2014;PateandSchoeninger1993).



TheRiverineEcosystem___________________________________________________________________________SoilsandTerrestrialVegetationThesoilsandvegetationof the riverinezonearemoreuniformwhen compared to the adjacent mallee plain because they areless dependent on local rainfall. There is an abrupt vegetationtransitionfromthesemi-aridmalleetothebetter-watered,morefertile black silts of the river floodplain. The open malleewoodland with its sclerophyllous-chenopodiaceous shrubunderstorey isreplacedbydenselywooded flatsandasavannaunderstorey. The river banks are covered with bulrushes,rushes,andsedges(LeighandCostin1974;Specht1972).

Canopy composition changeswith reduced soil fertilityandwateravailabilityawayfromtheriverfront.Riverredgums(E.camaldulensis) are dominant along the river's edge and arereplacedbyriverbox(E.largiflorens)ontheflatsadjacenttothefloodplain. Eucalypts in these better-watered areas reach30minheightandhavea foliageprotectivecoverof10–30percent.Coolibah (E. microtheca), false sandalwood (Myoporumplatyearoum), black oak (Casuarina cristata), dry-land tea tree(Melaleuca lanceolata), sand pine (Callitris spp.), and mulga(Acacia spp.)arethedominantspecies inthe lowwoodlandsofthe drier peripheries. These trees are 5–10 m tall and have afoliage protective cover of less than 10 percent. The drought-resistant understorey expands as the canopy thins out (Specht1972).

The savanna understory is dominated by tussocks ofperennial kangaroo grass (Themeda triandra), wallaby grass(Danthoniaspp.),andspeargrass(Stipaspp.).Inthedrierareasaway fromtheriver, the topsof thesegrassesbecomedormantand wither and die in response to the hot, dry summerconditions.Theyachievetheirmaximumgrowthduringthelatespringfollowingwinterrainfall(Gott1982;Specht1972).

Orchids (Orchidaceae), liliaceous plants (Liliaceae andHypoxidaceae),andyamdaisies(Microserislanceolata)areotherdry-landplantsthatarewidespreadonthebanksadjacenttotheriver. These plants sprout green leaves from undergroundrhizomes inthecoolerdaysofautumnandusestarchstored inroots, tubers, corms, or bulbs until they can grow vegetativelywith winter rainfall. Flowers occur in spring and the plants



continue growth until water becomes limiting in the summer.During summer dormancy, the plants are reduced to theirundergroundpartsagain(Gott1982;Specht1972).

Shrubs, including golden wattle (Acacia pycantha),kangaroo thorn (A. paradoxa), daisy (Olearia spp.), hopbush(Dodonaea viscosa), and native box (Bursaria spinosa), aresparse, but will expand with repeated firing. Onion weed(Asphodelus fistulosus), onion grass (Romulea spp.), and otherannuals invade theunderstory in the spring after rainfall (Gott1982;Specht1972).

Wetlandplantsoccur indensestands in locationswithpermanentwateror inareas thatareseasonally floodedby theriver.Dominantrepresentativesincludebulrushes(Typhaspp.),rushes (Cyperaceae) and waterribbons (Triglochin procerum).Theseplantspropagate fromrhizomes inadditiontoproducingnumerous small seeds in the spring. Maximum growth occursduringthehotsummer.Theriveralsofloodsduringthisperiodand increases the area which will sustain growth. In the latesummerandearlyautumn,thetopsoftheplantsdieinresponseto reduced temperature and recession of floodwaters (Gott1982;Specht1972).TerrestrialFaunaA number of small mammals are associated with the riverinehabitat.Thenocturnalherbivorousphalangersareadaptedtoanarboreal life. This family includes possums (Trichosurusvulpecula, Pseudocheirus peregrinus) and the koala(Phascolarctus cinereus). These small mammals nest in thehollowsoftreesandfeedupontendershootsandleaves,flowers,nectar and fleshy fruits. The koala's highly specialised diet isrestricted to the leaves ofEucalyptus spp. (Barrett 1955; Jones1969;Ride1970).



AquaticFloraandFaunaThe freshwater biome includes two main ecosystem types:running waters or lotic habitats (streams and rivers) andstanding waters or lentic habitats (lakes, ponds, swamps,marshes).Swampsandmarshesaresomeofthemostproductivehabitats on earth (Simmons1979).Most of the flora and faunaareconcentratedintheshallowlittoralzone.Rootedplants,suchas cattails (Typha spp.), lilies (Nymphaea spp.) and pondweeds(Potamogeton spp.), and small floating plants, such asduckweeds (Lemna,Wolffiaspp.), are the dominant producers.Consumers include worms, insects, crustaceans, and molluscs.Waterfowl, fish, turtles,andamphibiansmove inandoutof thelittoralzone.Phytoplanktonbecomethedominantproducersintheopen-waterzone.Algaeanddinoflagellatesareconcentratednearthesurface.Zooplanktonincludecopepodsandcladocerans.Fish and fish-eating birds are the dominant larger animals.Bottomdwellersincludeworms,insects,andmussels(Simmons1979;Williams1981a,1983).

Thecombinedeffectsofannualdischargevalues,erraticchanges in water level, drought, and the aridity of thesurroundingmallee have produced a depauperate fauna in theMurray Darling system. Twelve families, 18 genera, and 19speciesoffishoccurintheMurrayDarling,whereas13families,52 species, and over 260 species are found in theMississippi-Missourisystem.LackofspeciationisalsoevidentintheMurrayDarling waterfowl, mammal, and invertebrate fauna. Riverinespecies have all adapted to the widely fluctuatingwater levelsand associated changes in productivity (Keast 1981a; Smith1978;Williams1981b).

The riverine fauna is dominated by invertebratesconsistingprimarilyofarthropods,e.g., insects,crustaceansandtheirallies.Themacro-invertebratefaunaisverylimitedduetotheunstablenatureoftheriverbottom(Williams1981a,1983).Two species of mussels (Mollusca) Velesunio ambiguus andAlathyria jacksoni and several crustaceans, including thefreshwater shrimp (Paratyaaustraliensis), river prawns (Macrobrachium) and crayfish (Cherax spp. and Eusastacus armatus),occurintheLowerMurray(Williams1981a,1983).



InsectsLarvae of the large rain moth Trictenaatripalpis(bardi grubs)escapedryseasonsanddroughtsinmoistundergroundburrowsat thebasesofeucalyptus treesalong the river.Eachmothwilllaythousandsofeggsatthebaseoflargeriverredgumtrees(E.camaldulensis).Whenthegrubshatchtheyburrowinto thesoilandfeedonthetreeroots.Followingrainfall,themothsemergefrom the burrows and disperse (Common 1990). These largepopulationsofbardigrubswouldhaveprovidedareliable foodsource for hunter-gatherers (See Grehan 1989; Tindale 1966;Yen2012;Yenetal.2017).FishAlthough the Murray Darling fish species diversity is low, theriver supports a large number of important food species,including catfish, perch, cod, and blackfish (Clarke 2002). TheMurray cod (Maccullochella macquariensis) is the largestfreshwaterfishinAustraliaandcanreachlengthsofupto1.8mandweightsofupto113kg.Theaveragespecimenweighs10kg.Golden perch (Plectroplites ambiguus) weigh up to 4 kg andaverage1.4to1.8kg.Breedinginmanyoftheselargerspeciesistriggered by increasing spring water levels and rising watertemperature. During these periods, both habitat size and foodsupplyareexpanded.Increasedplanktonanddetritusinputandgreater aquatic plant and insect productivity provide thisadditional food (Barrett 1955; Frith 1974; Keast 1981b;McDowall1996).Adultgoldenperchandsilverperch(Bidyanusbidyanus)migrate upstream for distances up to 1000 km afterspawning, whereas Murray cod and freshwater catfish(Tandanustandanus) are relatively sedentary (Reynolds 1983).Radio-trackedMurraycodintheMurrayRiverhavemigratedupto 120 km upstream to spawn before returning to exactly thesameriverchannelfromwhichtheydeparted(ToddandKoehn2009).



Thecontrolofriverflowbylockshashadadetrimentaleffect on the distribution and abundance of fish. Floodwatercontrol has decreased habitat size and reduced breedingopportunities. Winter flows have been reduced and latesummer/autumn levels increased. Macquaria australasica,Maccullochellamitchelli andGadopsismarmoratus are seriouslythreatened,whileT.tandanus,P.ambiguus,andM.macquariensisnumbershavebeendrastically reduced (Frith1974;Shiel et al.1982).WaterfowlTheMurray Darling is themost important waterfowl breedingarea inAustralia.Twelveof the19 species ofAustralianducks,geese and swans are common inhabitants of the river system,and two northern tropical species, the grass-whistle duck(Dendrocygnaeytoni) and water-whistle duck (D.arcuata), areoccasional visitors. These birds feed on aquatic plants,invertebratesandsmallfish(Frith1982).Waterfowlprovidedakey food source for Aboriginal people inhabiting the LowerMurrayregion(Clarke2016).

Waterfowl behaviour is also keyed to changes in riverlevel and food abundance. Species that inhabit shallow,temporary waters (Malacorhynchus membranaeeus, Anasgibberifrons)arehighlynomadic.Breedingoccursinresponsetorisingwater levels and increased food supplies, but as soon asthe floods recede the habitat can no longer support the entirelocal population and some migrate to better-watered coastalplains and estuaries. The remaining waterfowl species occupythe more permanent deeper swamps and lakes. They aresedentaryormigrateregularlybetweenlocalswampsandlakes.Shorebirds include plovers, dotterels, stilts, avocets, pratincole,gulls,andterns(Frith1974,1982;Pizzey2013).ReptilesThreespeciesofturtleinhabittheLowerMurray,theAustraliansnake-necked turtle (Chelodina longicollis), broad-shelled long-necked turtle (C. expansa), and Murray short-necked turtle(Emydura macquari). In Australia, freshwater turtles arereferred to as ‘tortoises’ to distinguish them from the largermarine forms. Freshwater turtles (‘tortoises’) are found in theriverandassociatedstreams, lagoons,andswamps.Adults feed



onfish,molluscs,crustaceans,frogs,tadpolesandaquaticinsects(Cogger2014;Goode1967;Worrell1970).

E. macquari and C. longicollis go ashore frequently tobaskinthesun,butC.expansaonlyleavesthewatertolayeggs.From 10–15 elongated, brittle-shelled eggs are laid in nestsexcavatedinmoistsoils.Theeggsareburiedandleftunattendedtohatchwith theheatof thesun.C.longicollus andE.macquarinestinearlysummerandeggshatchinlatesummerorearlyfall,whereasC.expansa layseggsinfallwhichoverwinterandhatchthe following spring. The young emerge fully developed andimmediatelyenterthewatertofeedoninsect larvae.C.expansaadults are the largest freshwater turtles in Australia. Theyachievecarapacelengthsupto50cmandweightsover5.5kg.C.longicollisandE.macquarireachcarapace lengthsof25cmand30cm,respectively(Cogger2014;Goode1967;Worrell1970).

Ectothermic reptiles are extremely sensitive totemperaturechanges(HeatwoleandTaylor1987).Turtlesentera state of winter dormancy (brumation) in response to coldtemperature.C.longicollisretreatstomoistearthenburrowsontheshore,whileC.expansaandE.macquariburrowunderwaterinbottommudorinadjacentbanks.

Awidevarietyofpredatorsposeathreattoturtlesatallstages of their life cycle (Goode 1967). Eggs are eaten bypossums,snakes, lizards,waterrats,anddingoes. Juvenilesareconsumedbywaterfowlandsnakes,andadultsbyhawks,eagles,waterfowl,waterratsanddingoes.AmphibiansFrogs were the only amphibians found in the pre-EuropeanLowerMurrayenvironment.Urodelesandcaecilians(newtsandsalamanders)andtoads(Bufonidae)didnotoccur inthenativeAustralianfauna.Mostfrogspeciesarenocturnalandterrestrial,butthegenusLitoriaalsocontainsarborealmembers.Alladultsarecarnivorousandwilleatanythingtheyareabletocatch,killandswallow.Insectsandsmallrodentsarecommonconstituentsof the diet. Adult size ranges from 13.8 mm in Ranidelladeserticola to 104.2 mm in Litoria caerulea (McFarland et al.1979;Tyler1978;Tyleretal.1981).



Waterlossisamajorproblemfacedbyamphibiansdueto the permeability of their skin. Moist microhabitats arerequired to regain water lost by evaporation and for thesuccessful development of eggs and larvae. Many terrestrialforms(Neobatrachus,Cycloranaspp.)burrowtolocatemoisture,while others (Limnodynastes spp.) are restricted to permanentwatersources.MostAustralianfrogsareopportunisticbreedersthatuseephemeralwatersforreproduction.L.rubellacompleteslarval development in as few as 14–15 days following fall andwinter rainfall, whereas species occupying permanent watersmayrequireupto5months.Riverinespeciesbreedyear-roundorinthelatewinterandspringinresponsetorisingriverlevelsand increased food supplies. Many terrestrial frogs remaindormantinburrowsorothermoistmicrohabitatsduringthehot,dry summer (McFarland et al. 1979; Tyler 1978; Tyler et al.1981).MammalsAquatic mammals include the platypus (Omithorkynchusanatinus)andwaterrat(Hydromyschrysogaster).Theseanimalsshelter in burrows excavated in the banks of the river andassociated creeks and lagoons. The diurnal platypus feeds onmussels, freshwater shrimp, insects, and worms at dawn andtowards dusk. Much of its time is spent in the deep burrow,whichmayextendupto30mfromthewater’sedge.Thefemalelays two to three soft-shelled eggs in a nest concealed in theburrow. The platypus remains in its burrow in hibernationduringwinter(Barrett1955;Jones1969;Morcombe1968).Thehighly specialised water rat is unique to Australia. It feeds oncrayfish, mussels, small fish, insects and aquatic plants. Thenocturnalanimal isextremelyshy.Adults reach lengthsofover50cm(Barrett1955;Jones1969).MolluscsFreshwater mussels are sessile bivalve filter feeders. Theyinhabitrivers,streams,lagoons,andbillabongsinwatersoflessthan 10m depth.Murray River species burrow in fine alluvialsilts or attach to large submerged dead trees. These shallowhabitats expose them to great temperature variations andfrequent desiccation (Burky 1983; Russell-Hunter 1983; Seed1983;Smith1978).



Hibernation in muddy bottom sediments occurs inresponse to cold winter temperatures. Velesunio ambiguusbecomesinactiveatwatertemperaturesbelow12°C(Hoffmann1983; Millington and Walker 1983; Walker 1981a, 1981b).Mussels also aestivate in mucus cocoons with falling latesummer/autumn river levels and the summer drying ofterrestrial habitats. Adults can survive in these cocoons foryears. Larger specimens survive up to 30 per cent longer thansmaller ones due to slower desiccation rates associated withlowersurfacearea/volumeratios(Dudgeon1982;Morton1979;Williams1983).

Great variations in mussel population size occur withfluctuatinghabitatsizeandfoodsupplies.Largenumbersofeggsareproducedinthespringashabitatareaisincreasedbyrisingriver levels and winter rainfall. The glochidial larvae attach toand are dispersed by fish hosts (Burky 1983). Velesunioambiguusadultsfromthemainriverstreamreachlengthsupto94 mm and heights to 55 mm. Lagoon forms are smaller andmoreflattened(Cotton1961;MilingtonandWalker1983).CrustaceansShrimp are simply smaller genera of prawns. The maximumlengthofthefreshwatershrimpParatayaaustraliensisis25mm,whileriverprawns(Macrobrachium spp.)growto178mmandweighupto400g.TheSAfreshwaterformsinhabitinlandriversand other permanent waters. They are active at warm watertemperatures. Optimum growth occurs at 28–30° C in M.rosenbergii. Their diet includes detritus, smaller crustaceans,molluscs, polychaetes, and other invertebrates (Hale 1927–1929;NealandMaris1985).

Femalescarryfromlessthan100toover100,000eggsontheirpleopodsforthreetofivemonths.InM.rosenbergii,theplanktonic larvaewashdownstream toestuarinewaterswheretheymaturewithinfourmonths.Thejuvenileswillonlysurviveinbrackishwaterswithsalinitiesfrom12–16ppt.Adultsmigrateupstream to fresherwaters (Neal andMaris 1985; Provenzano1985;VernbergandVernberg1983).



Freshwater crayfish are quite common throughoutinlandAustralia.Theyoccupy the shallowwatersofbillabongs,creeks, lagoons, and rivers (Frost 1975). Crayfish areomnivorous. Adults eat large quantities of aquatic vegetation.They are grazers in addition to taking phytoplankton anddetritus with filtratory setae. Animal foods include smallercrustaceans, molluscs, fish, annelids, arthropods, andzooplankton.Theyoung feedonplanktonanddetritus.Feedinggenerallyoccursduringtheearlymorningandevenings.Fisharethe primary predators (Cobb andWang 1985; Grahame 1983;MillsandMcCloud1983).

Theyabbie(Cheraxdestructor)isawarmwaterspeciesthat is extremely well-adapted to the dynamic Lower Murrayenvironment. It is active atwater temperatures above16° C inthespring,summer,andearlyautumn.BypossessingcombinedcharacteristicsofbothR-andK-reproductivestrategies(Pianka2011; Sastry 1983; Stearns 1977, 1980) yabbies can rapidlyincreasepopulationnumbersandcolonisenewhabitatscreatedbyspring floodwatersandwinterrains.Theybreedthroughouttheir active period, produce large numbers of young, provideparentalcare,matureearly,andexhibitrapidgrowthrates.

From300–400eggsarecarriedonthefemalepleopodsfrom October to March, and eggs hatch in about six weeks at20°C.Theyoungarebornaliveandclingtothemotheruntiltheyreach 3mm in length. This initial parental care results in veryhigh survival rates when compared to the pelagic larvae ofmarinecrustaceans(Provenzano1985).Youngare independentwithin 41–90 days of spawning, depending on watertemperature. The optimum temperature for growth is between20-25°C.Mostyoungyabbiesreach10cminlengthwithintwoyearsofhatching.Adultsreachtotallengthsof20cmandweighupto200g.Ediblemeatcomprisesabout25percentofthistotalweight. Life span is about three years (Faragher 1983; Frost1975;MillsandMcCloud,1983;Provenzano1985).

As water temperature drops below 16° C in the lateautumnandwinter,yabbiesretreattounderwaterburrowsandenterastateofdormancy.MostburrowsareoccupiedbyApril.Theyalsobecomedormantinresponsetodrought.Asephemeralwater holes dry out in the hot summer, the crayfish enter asealed burrow and follow the retreating water table. Tunnelsmayextendover3mdeep.Aestivationwillcontinueuntilwater



returns to the overlying ground. Dormant yabbies have beendiscovered beneath lake beds which have been dry for up toeight years, although yabbies attempt to avoid summerdormancybymigratingtomorestablehabitatsfollowingwinterrainfall(Frost1975).

TheMurrayRiver lobster (Euastacusarmatus) isacoldwater speciesactiveat temperatures in the5–10°C range. It ismuchmoreK-orientedthanC.destructor.Asinglematingoccursin late autumn and each female produces up to 400 eggs.DevelopmentissimilartothatofC.destructor,butgreatertimeisinvolved. At these colder temperatures eggs do not hatch untilfourmonthsafterspawning.Adultsreach50cmtotallengthandweighupto2.5kg.UnlikeC.destructor,thiscrayfishhasasmalltail-body ratio. Summerdormancy takesplace inmudburrowsintheriverbanks(Frost1975).RiverineTrophicStructure___________________________________________________________________________The primary limiting factors affecting the abundance anddistributionofaquatic floraand fauna in theLowerMurrayarefluctuating river level and water temperature. Flood watersbring inputs of nutrients, plankton and detritus, and increasehabitatsizeforbothfloraandfauna.Theactivity-breedingcyclesof crustaceans, molluscs, turtles, mammals and some fish arekeyedtochangesinwatertemperature.

Because large aquaticplants are scarce, phytoplanktonand detritus are themajor sources of primary productivity. Inaddition, wetland plants provide abundant supplies of smallseeds during the spring and early summer. Insects are thedominant riverine herbivores. Omnivores include crustaceans,molluscs,fish,andwaterfowl.Theplatypus,riverrat,andturtlesareprimarilycarnivorous(Walkeretal.1986).

In relation toAboriginal food supply, animals aremostabundantinthespringandearlysummerwhentheriverlevelisrising and water temperatures are warm. In the winter, thecrayfish Cherax destructor, shrimp, mussels, turtles, mammals(platypus,riverrat)andsomeinsectsbecomedormant,andfishandwaterfowlnumbersaregreatlyreducedinresponsetocold



watertemperatures,lowriverlevels,andreducedfoodsupplies.Thecold-adaptedcrayfishEusastacusarmatus ismostabundantduringtheseperiods(Walkeretal.1986). DuetotheerraticdischargevaluesoftheMurrayRiverthatexistedpriortohistoricalwatermanagementpractices,themagnitude of these seasonal changes in floral and faunaldistribution and abundance would have been less predictable.Furthermore, major non-seasonal droughts in the easternhighlands have disrupted the regular cycle in the past andreducedthelowerreachesoftherivertoaseriesofwaterholes(FrithandSawer1974;Walkeretal.1986).TheMalleeEcosystem___________________________________________________________________________VegetationThe major part of the mallee occurs between the 200 and450 mmannual isohyets.Thecanopyof themalleeopenscrubconsistsofsmall treesand tall shrubsdominatedbyEucalyptusspp. Mallee eucalypts are sclerophyllous evergreen perennialswhich sprout from subsurface lignotubers. They have deep taproots and shallow extensive lateral root systems. The diversesclerophyll heathland understorey contains over 130 plantgenera (Beadle 1981; Specht 1972, 1979, 1981a, 1981b;Williams1979). Theamountandseasonalityofrainfallhasagreateffecton the availability of mallee plant foods. Regular autumn andwinter rainfall produces lush periods of vegetation growthduring which both perennial and annual plant foods areabundant. Groundwater reserves accumulated during the rainyseason allow plants to remain productive during the annualsummerdrought.However,plantfoodsaregreatlyreducediftheregular rainfall does not arrive. During these dry periods onlythe stems, leaves, roots, fruits and seeds of drought-resistantplants are available (Cloudsley-Thompson and Chadwick 1964;Mayhew1968).



FaunaBirdsSpecies density surveys (Foulkes and Gillen 2000; Pianka andSchall 1981; Simpson 1973a, 1973b) indicate that birds are byfar the most diverse vertebrate class inhabiting the malleeregion. Foulkes and Gillen (2000) provide an overview of themaximum total number of species recorded from various pastsurveys. Birds (257 spp.) are followed by reptiles (75 spp.),mammals(66spp.),andamphibians(fourspp).Mobilityallowsbirds to successfully exploit patchy distributions of food andwater.Nectar,flowers,pollen,fleshyfruits,seeds,andinsectsareprimary sources of food. Abundant nectar-producing plantsprovide a concentrated high energy source in an otherwisemarginal environment (Recher 1981). Numerous species ofpasserine birds feed primarily on nectar. The Meliphagidae(honey-eaters)contributegreatlytothehighspeciesdiversityinthemallee.

Many of these small birds conserve preformed andmetabolicwater anddonotneedtodrink.Water conservation,combined with an ability to tolerate high body temperatures,allow these birds to remain active during the heat of the day(Dawson1981;Keast1981a,1981b;Kikkawaetal.1979).Sincemanybirdsarenomadictheyopportunisticallyenterthemalleefor food and then return to better-watered surrounding areas,such as the riverine and coastal zones. The emu (Dromaiusnovaehollandiae)onlyentersthemalleefollowingwinterrainfalland is active during the evening and cooler parts of the day.Others, including the territorial mound-building mallee fowl(Leipoa ocellata) and bustard or plains turkey (Ardeotisaustralis),areresidentspecies.Additionalgamebirdsinhabitingthe mallee include quails, pigeons and doves (Barrett 1955;Pizzey2013;Kikkawaetal.1979).



ReptilesReptilesarewell-adaptedtoaridregions.Severalcharacteristics,including a thick skin with few dermal glands, excretion ofconcentrateduricacid,nocturnalactivity,andtheuseofburrowsduringthehottestperiodoftheday,minimisewaterlosstotheenvironment. The almost exclusive carnivorous diet, includinginsects,frogs,birdsandsmallmammals,providesanappreciablepercentageofthereptilianwaterrequirement.Additionalwateris obtained by opportunistic drinking and from metabolicsources.Breedingusuallyoccurs in the spring followingwinterrainfall when a greater quantity of higher quality food isavailable(Mayhew1968).

Fat storageallowsreptiles to reduceactivityor remaindormant in their burrows during colder months or periods offood shortage. Such daily and seasonal metabolic inactivityallows reptiles to capitalise on limited and unpredictable foodsupplies.Itisamajorfactorcontributingtotheirrelativesuccessover mammals and birds in the arid interior of Australia(Mayhew1968;Pianka2011).

Lizardsandsnakesarethedominantreptiliangroupsofthe mallee. Rawlinson's (1966) literature review of reptilesrecordedintheVictorianmalleereports73speciesrepresenting34generaandninefamilies.Theseinclude45speciesof lizardsand25speciesofsnakes.

Microhabitat specialisation and variability in times ofactivity have contributed to the great lizard diversity. Manyspeciesarerestrictedtosandridges,sandplain-Triodiaorshrub-Acaciahabitats.Alargenumberarenocturnal.Nocturnallizardsinclude geckos, pygopodids, and skinks. Most agamids andvaranids are diurnal. The majority of lizards in regions withMediterranean climates breed only during the spring wheninsects becomeabundant (Barrett 1955;Mayhew1968; Pianka1981).



Theelapidsarethemostcommonsnakesofthemallee.Snakeshavearelativelylimitedheattoleranceandconsequentlymostarid-landsnakesarenocturnal.Daysarespentinburrows,hollowlogsorcoolriverinehabitats.Theirdietincludesrodents,frogs, lizards, birds, bird eggs, other snakes, insects, andadditional small invertebrates. Snakes are capable of goingwithout food for periods of six to twelve months without anyseriousconsequences.Breedingoccursinthespringandautumn(Barrett1955;Mayhew1968;Pianka1981).MammalsSmall rodents are the dominant mammal group of the mallee.Malleerodentsincluderats,mice,rat-kangaroos,andbandicoots.The burrowing rodent-like wombats (Lasiorhinus latifrons) arealso common (Cloudsley-Thompson and Chadwick 1964; Jones1969;Leeetal.1981;Ride1970).

Nocturnal activity, combined with the use of burrowsandnestsduringtheheatofthedayand/ordailytorpor,allowsrodents to evade arid-land conditions and reduce evaporativewater loss. Evaporative water loss is a critical problem due totheir small size and large surface area/volume ratio. Most oftheirwater isobtained from foodandmetabolic sources.Manyrodents enter a prolonged state of dormancy or aestivationduring the annual summer droughts. Aestivation is similar towinterhibernation,butitoccursinresponsetoextendedperiodsofheatandaridityrather thancold (BartholomewandDawson1968;Cloudsley-ThompsonandChadwick1964;Leeetal.1981).

Rodent populations and territorial ranges weredrasticallyreducedbythe1890–1920sduetocompetitionwithintroduced European species (Lee et al. 1981). Rabbits(Oryctolaqus cuniculus) and house mice (Mus musculus)competedforburrowspaceandfood,whilefoxes(Vulpesvulpes),dogs(Canisfamiliaris)andcats(Feliscatus)increasedpredationpressures (Keast 1981b; Lee et al. 1981). Other nocturnalburrowers include the echidna or spiny anteater (Echidnaaculeatus)and thenumbatormarsupialanteater (Myrmecobiusfasciatus). These mammals feed on a variety of small insects,includingantsandtermites.



Large herbivores are represented by the familyMacropodidae,whichincludeskangaroosandwallabies(Dawson1995;Hume1982;MainandBarker1981).Thelowermetabolicratesandruminant-likedigestionofkangaroosallowanefficientuse of poor quality fibrous vegetation (Janis 1976; Tyndale-Biscoe2005).

During drought periods small 2.5 kg hare-wallabies(Lagorchestes spp.)cansurviveontheopenmalleeusingwaterobtained directly from vegetation (preformed water) and viametabolicoxidation(metabolicwater),whereaskangaroosover25 kg must move toward areas where free water is available(Main and Barker 1981). Large kangaroos move toward theMurrayRiver in thesummerandthenreturn to themalleeandarid grasslands following winter rainfall (Archer 1984a; FrithandCalaby1969).

Distributions of macropods are also related to habitatand diet. Themajor proportion of the diet consists of grasses.Grasses are supplemented with herbs. The red kangaroo(Macropus rufus) occupies open plains and remains highlymobile during the rainy season in search of newly sproutedgrasses and ephemeral dicots which have greater water andprotein contents. Red kangaroos occur on the arid grasslandsbordering the mallee. Grey kangaroos (Macropus fuliginosus)rarelyventurebeyondthedensemalleescrubandwoodedareasadjacent to theMurrayRiver. They consume greater quantitiesof grass than red kangaroos and are less selective. The greykangaroo remains closer to the river during drought periods,whileredkangaroosmove toward theriver front fromthearidgrasslands(FrithandCalaby1969;Jones1969;Newsome1965a,1965b;Tyndale-Biscoe2005).

Solitary macropods include the euro (Macropusrobustus) and wallabies. Euros and rock wallabies (Petrogalespp.) seek shade in rocky outcrops, gullies, and beneathmulga(Acaciaaneura). Euros do not normally venture further than a1km radius of their shelters. They feed on lower qualityperennial grasses such as spinifex. Other wallabies arewidespread throughout all inland woodland and desertcommunities (Frith and Calaby 1969; Jones 1969; Newsome1965a,1965b;Tyndale-Biscoe2005).



The native carnivorous mammals that have inhabitedtheSAmalleeregionareconfinedtotwofamilies,theDasyuridaeandtheThylacinidae.TheTasmaniandevil(Sacrophilusharrisii)and the native cats (Dasyurus spp.) are included in theDasyuridae, and the dog-like Tasmanian wolf or thylacine(Thylacinus cynocephalus) is the only member of the familyThylanicidae.Allofthesecarnivoreswerenocturnal.

TheTasmaniandevilandTasmanianwolfarenolongerpresent on the mainland. Mainland extinction in southeastAustralia probably dates back no more than a century. TheTasmaniandevilsurvivedinWesternAustraliauntil430±160BPand in the Northern Territory until at least 3120±100BP.Mainland evidence for the Tasmanian wolf comes from theKimberleyregionat0±80BPandthesouthwestat3090±90BP(Archer 1981). Native cats still survive today, but theirpopulationsandrangeshavebeenreducedsubstantially(Archer1981;Barrett1955).

Competition associated with the introduction of thedingo (Canisfamiliarisdingo) between 4000 and 3000 BPmayhave been responsible for the mainland extinction of thethylacineandtheTasmaniandevilsincethesenativevertebratefauna survived in Tasmania in the absence of the dingo.CompetitionwithdingoesandEuropeanintroduceddogs(Canisfamiliaris), red foxes (Vulpes vulpes) and cats (Felis catus),accompanied by reductions in rodent populations, led toTasmanian devil and native cat declines (Archer 1984b; Gollan1984; Johnson andWroe 2003; Oskarsson et al. 2011). Nativecatswerealsokilledbyfarmersbecauseoftheirincessantraidsonhenhouses(Archer1981;Barrett1955;Jones1969).



Aboriginal Subsistence-Settlement Systems in the LowerMurrayEnvironment___________________________________________________________________________Thepalaeoenvironmentaldataindicatethattherehasbeenlittlechange in the climate and vegetation of southeastern Australiaduring the last 10,000 years. Major vegetation changes areconfined to the period following European colonisation. Themodern productive Murray River system was established by7000 BP. Therefore, Aboriginal populations living along theLowerMurrayRiverexperiencedasimilarnaturalenvironmentthroughout thepast7000–8000years (Fluinetal.2009;Hill etal. 2009). Coastal groups near the mouth of the Murray Riverwouldhavehadagreateraccess tomarine resources followingsea-level stabilisation at 6000 BP (Lampert and Hughes 1974;Perlman1980).

Consequently,modern ecological data can complementarchaeological and historical records by providing estimates ofthe floral and faunal resources available to Aboriginal riverinepopulations during the majority of the Holocene. Althoughavailableatdifferentseasons,theemploymentofarangeofplantfoods from the riverine andmallee ecosystems would provideyear-round access to fruits and seeds (Clarke 1985b, 1994,2015).Plantstorageorgans(tubers,roots,bulbs),aquaticplants,and naturally desiccated fruits, such as Santalum andSolanumspp.,wouldremainavailableduringperiodsofdrought(Clarke1985a,2015;Gott1982,1999;Martin2011;WestellandWood2014).However,manydrought-escapingseedsandgreensfrom the mallee habitat would not be available (Specht 1972,1981b;Pate1986).

Storagecouldbeusedtoextendtheavailabilityofdriedfruits and seeds. Techniques for the drying of plants wereemployed in the Lower Murray. In relation to basketmaking,rushes were sun dried or placed in a heated oven and thenbundledupandstored for futureuse(Clarke2015:234).Fruitsof Kunzea pomifera were dried in the form of cakes (Tindale1981:1879).

Faunal variety ensured that animal foods were alsoavailable throughout the year. Furthermore, buried turtle andmallee hen eggs and dormant crayfish and mussels could beobtainedbydigging.Variousmeatscouldbepreservedbydryingandthenstoredfor futureuse.AccordingtoClarke(1994:165–



170, 2009), fishing techniques employed in the LowerMurrayregion ranged from netting, spear fishing and trapping, toopportunistic harvesting and storage (Eyre 1845; Krefft 1862–1865;Tindale1930–1952).Ethnographicaccountsindicatethatfish were stored in small dams, pounds or mud pools (Clarke1994:169;Sturt1833,Vol2:165).Inaddition,theavailabilityoffish was extended via drying on racks (Berndt and Berndt1951:29). Bird eggs (e.g., emu, waterfowl) and freshwatermusselswere stored formonthsbyburying them indamp soil(Clarke 1994; Simpson and Blackwood 1973;). Non-localdroughts that reduced river flowwould have had the greatestimpactonfaunalabundance.Manyanimals,includingwaterfowl,fish and turtles, could migrate to better-watered areas duringthese stressful periods. On the other hand, local droughtsresultedinincreasedfaunaldensitiesneartheriver.

Ifhumanbehaviour isviewedasanadaptive responseto specific environmental conditions, then pre-contactAboriginal population size in the Lower Murray should belimitedbytheperiodofgreatestresourcestress.Unpredictableextended droughts that reduced both riverine and malleeresources would have provided the ultimate test of survivalskills. Food storagewouldprovide an importantmechanism tosustain populations during extended periods of drought, but iffood reserves were depleted, alternative strategies would berequired.Access tomore reliable foodsources in the resource-rich Murray River mouth and nearby coastal regions wouldprovideinlandriverinepopulationswithameanstoreducetherisksassociatedwithextendeddroughts.

IfAboriginalgroupsinhabitingtheLowerMurraywereto obtain non-local resources with the least amount ofoppositionandeffort,someformofsocialtieswouldberequiredin these areas. Exogamous marriage between regions aspracticed in themarginal desert habitats ofAustraliawouldbeonemechanism to facilitate such relations (Berndt and Berndt1993; Stanner 1965; Strehlow 1965; Yengoyan 1968, 1976).Individuals socialised in one area would carry the essentialresource procurement knowledge of their homelands to theirspouse'speople.Tradenetworkswouldprovideaccesstoother



resourcesthatwerenotequallyavailabletoallpopulations,suchas stone, marine and riverine shell, ochre, and local foods(McBryde1984;McCarthy1939).Alternatively, the use of foodstorage,fishtraps,andothertechnologicalmeanstoincreasethereliability of local foods (intensification) could be employed toreducethedependenceonneighbouringgroupsandfavourmoreclosed social relations associated with territoriality (Clarke1994;Lourandos1985,1988,1993,1997).

HistoricalAboriginalpopulationnumberswerereducedsubstantially by introduced European diseases and violentconfrontationswith settlers. Smallpox spread rapidly along thecoastsanddowntherivervalleys(Butlin1993;Campbell2002).The first smallpox outbreak among Aboriginal populationsoccurredinSydneyin1789,withinoneyearoftheestablishmentof the colony. Itwas again reported along theMurray River atSwan Hill in the early 1800s (Stirling 1911). The disease wascommon among Aboriginal groups of the Murray Darlingbetween 1830 and 1845 (Sturt 1833; Stirling 1911; Clarke1994).

Disputes over trespassing and the hunting of livestockresultedinnumerousAboriginaldeaths.ManyAboriginalpeoplewere either poisoned or shot on sight (Eyre 1845, II; Smith1880). Burke et al. (2016) provide a detailed overview ofhistorical accounts of frontier violence in the western CentralMurraybetween1830and1841withafocusonfive ‘hotspots’where repeated evidence of conflict occurred. This researchinvestigatedthesignificantincreaseinviolentconflictassociatedwith the opening of theOverland StockRoute between Sydneyand Adelaide in 1838. Campbell (1939) estimated that theAboriginal population of southeastern South Australia declinedby50percenteveryfiveyearsfrom1840through1900.

Consequently, historic and ethnographic accounts ofAboriginalbehaviourmost likelydonotaccuratelyportraypre-contact lifestyles. Rapid population reductions andenvironmental degradation could completely disrupt formersocial relations and produce new adaptive requirements andopportunities.TheremnantsofformerAboriginalgroupswouldbefacedwiththechallengesofsurvivalunderavastlydifferentsetof conditions than thoseexperiencedby theirpredecessors.Access to food and blanket rations,medical assistance, alcohol,andemploymentresultedinconcentrationsoflargenumbersof



Aboriginal people from various groups around Europeanmissions,governmentposts,homesteads,andpopulationcentres(Foster 2000; Foster et al. 2001;Rowley1971). Eyre (1845, II:445) stated that only 150 of the 600 to 800 Aboriginal peoplewhofrequentedAdelaidewereoflocalorigin.Conclusions___________________________________________________________________________Accesstoreliablewaterandthewiderangeofplantandanimalfoods available in the Lower Murray riverine and adjacentmallee ecosystems provides environmental data to generatehypotheses about past variability in hunter-gatherersubsistence-settlementsystemsintheregion.Ethnographicandhistorical data relating to diet and landscape use in variousregions of the LowerMurray can contribute tomodel buildingandhypothesistesting.However,modelsrelatingtopre-contactseasonalmobilityor increased sedentismassociatedwith long-termuseofresidentialbasesmustbetestedemployingarangeofarchaeologicalandenvironmentaldata inordertoassessthevalidityoftheirapplicationacrossspaceandtime.

In relation to support for hypotheses regarding thepresence of semi-sedentary or sedentary hunter-gathererlifeways along the Lower Murray during the late Holocene,archaeological research must establish multiple lines ofindependent evidence that suggest the long-term use of basecamps associated with particular riverine landscapes orterritories. Collaborative research projects involvingarchaeologistsandvariousAboriginalcommunitiesintheregionwillbeessentialtothesuccessfulachievementofthesegoals.

In order to test these hypotheses across the LowerMurray region, improved data associated with the followingareasisrequired:



Ø Expanded excavation and chronometric dating for a range

ofsite types, includingrockshelters,middens,moundsandcemeteries to demonstrate long-term use particulararchaeological sites (cf. Littleton et al. 2017; Pate et al.1998,2003;Wilsonetal.2012);

Ø Comprehensiveandsystematicanalysesoffaunalandfloralremains and artefacts associated with variousarchaeological sites to demonstrate past use of a range ofkeyanimalandplantfoods;

Ø Expanded analyses of earth mounds to providearchaeological and environmental evidence to supportethnographicmodelsregardingtheprocessinganduseofarange of key plant and animal foods, including faunalanalyses and residue analyses (chemical, phytolith) ofgrindstonestoestablishassociationwithplantslikeTyphaspp.;

Ø Expandedstable isotopebaselinevalues inboneand teethfor a range of animals associatedwith different diets andhabitats;

Ø Expanded stable isotope data base in bone and teeth forhuman remains associated with a range of cemetery andother burial sites across different habitats to demonstratelong-termtiestoparticularsitesandregions, i.e.,presenceofbasecampsorterritories;and

Ø Comprehensive and systematic employment of physicalanthropology associated with a range of cemetery andotherburialsitesacrossdifferenthabitats.

Acknowledgements___________________________________________________________________________Over thepast35years,my researchhasbenefited significantlyfrom collaborative involvement and support from a number ofAboriginal communities in South Australia. I acknowledgecontributionsfromtheGerardCommunityCouncil,theMannumAboriginal Community Association Inc. and the NgarrindjeriHeritage Committee. In addition, the South AustralianMuseumhasprovidedinvaluablesupportforarangeofresearchprojects.I thank Mick Morrison and Philip Clarke for their invaluableadviceregardingtherevisionofthemanuscript.



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