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Stone tool assemblage variability is considered a reliable proxy measure of adaptive variability. Raw material richness,
transportdistances, and the character of transportedtechnologies are thought to signal (I) variation in raw material selectivity based on material quality and abundance, (2) optimizationof time and energy costs associated with procurementof
stonefrom spatially dispersed sources, (3) planning depth that weaves raw material procurementforaysintoforaging activities, and (4) risk minimizationthat sees materials transported in quantities and forms that are energetically economical
and least likely to fail. Thispaper dispenses with assumptions that raw material type and abundance play any role in the
organizationof mobilityand raw materialprocurementstrategies. Rather,a behaviorallyneutralagent-based model is developed involving a forager engaged in a randomwalk within a uniformenvironment.Raw material procurementin the model
is dependentonly upon randomencounters with stone sources and the amountof available space in the mobile toolkit. Simulated richness-sample size relationships,frequencies of raw material transfers as a function of distance from source, and
both quantity-distanceand reductionintensity-distancerelationshipsare qualitativelysimilar to commonlyobservedarchaeological patterns. In some archaeological cases it may be difficult to reject the neutral model. At best, failure to reject the
neutral model may mean that interveningprocesses (e.g., depositional time-averaging) have erased high-frequencyadaptive signals in the data. At worst, we may have to admit the possibility that Paleolithic behavioral adaptations were sometimes not responsive to differences between stone raw material types in the ways implied by currentarchaeological theory.
Se considera la variabilidadde las colecciones de litica como una medidaconfiablede la variabilidadde las adaptacionesal
medio ambiente.La diversidadde materiasprimas, la distancia a sus yacimientosy la tecnologia empleada reflejarian(1) la
selecci6n de la materiaprima con base en su calidad y abundancia,(2) la optimizacionde gastos de tiempoy energiaempleados en la obtencion de materia litica de yacimientosdispersos; (3) la integracionanticipadade las visitas a los yacimientos
con actividadesde caza y recoleccion,y (4) una estrategiade reduccionde riesgos que consiste en transportarla litica en las
cantidadesyformas mas eficientesdel punto de vista energeticoy menos susceptibles al desgaste. En este trabajoprescindimos de suponer que el tipo y la abundanciade materiaprima hayanjugado un papel en la organizacion de estrategias de
obtencion de esta y de los desplazamientosen general. En su lugar,partimos de un modelo conductualmenteneutralbasado
en el agente (individuo),como seria un cazadorque se desplaza al azar en un medio ambienteuniforme.La obtencionde materiaprima dependeentoncesunicamentede hallazgosfortuitosde yacimientosde litica y la cantidadde materialque el cazador
pueda agregar a su equipaje.La simulaci6n de la relaci6n entre la diversidady el tamanode la muestra,asi como de la frecuencia de uso, la cantidady la reducciondel volumenenfuncion de la distancia al yacimiento,revelanpatronesque se asemejan, de manera cualitativa,a los que arrojaa menudoel registroarqueologico. En algunos casos arqueol6gicos resultaria
entonces dificil descartartal modelo. En el mejorde los casos la imposibilidadde descartarlosenialarfaque los procesospostdeposicion (p.ej. la combinaci6nde artefactosde distintas epocas en una sola coleccion) han borrado todos los indicadores
de la adaptacion.En el peor de los casos, nos veriamosobligados a reconocerque las adaptacionesdel comportamientopaleolftico a veces no obedecian a las diferencias entre los tipos de materiasprimas de la litica de la manera que sugieren las
teorias vigentes en arqueologia.
geo-
titative.
MotooKimura(1983:xiv)
P. Jeffrey Brantingham * Departmentof Anthropology,Universityof California,Los Angeles, 341 Haines Hall, Box
951553, Los Angeles, CA 90095-1553. Email: branting@ucla.edu
AmericanAntiquity,68(3), 2003, pp. 487-509
CopyrightO2003 by the Society for AmericanArchaeology
487
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Figure 1. Raw material richness in assemblages from the Middle Paleolithic site of Grotte Vaufrey, approximately
204,000-74,000 B.P. Richness changes through time (a) but is heavily dependent upon sample size (b). Procurement probabilities estimated from the observed raw material proportions (c) illustrate the structure of the distribution underlying
procurement behaviors. The primary question is whether this probability distribution is derived from the environmental densities of different raw materials, or whether a biased behavioral strategy is responsible. Data from Geneste (1988)
with revisions.
P. Jeffrey Brantingham]
PROCUREMENT
NEUTRALMODELOF STONERAWMATERIAL
489
Geneste 1988, 1989). Two assumptionsarenecessary to infer geographic range from these data,
namely that raw materialprocurementis embeddedwithinotherforagingactivities(Binford1979;
Rensink et al. 1991), and that the maximum
recordedtransportdistancefor a materialtranslates
approximatelyinto themaximumradiusof the foraging area(Roebroekset al. 1988:30).Thus,in the
CentralEuropeanMiddle Paleolithic, the maximum recordeddistancefor raw materialtransfers
is around300 km, while in the Middle Paleolithic
of Aquitaine Basin the maximum distance is
approximately 100 km (Figure 2) (FeblotAugustins1993). The differencesbetweenregions
arepresumedto reflectrangesizes adaptedto different ecological settings. Nonetheless, in both
cases the decline in the frequencies of transfers
from greaterdistances is taken to indicate minimizationof the costs associatedwith rawmaterial
procurementand transport(see below).
More interestingperhapsis Feblot-Augustins'
of "interal modes"
(1993:243-245) interpretation
for rawmatefrom
the
trend
decay-like
deviating
rial transfersoverall (Figure2a) (see also FeblotAugustins 1997b, 1997c). She views
"unexpectedly"high frequenciesof raw material
transfersfrom distantsourcesas indicatingeither
logistical(sensu Binford1980) use of distantecological patchesby specializedtask groups,or seasonal residentialmoves to non-core areas in the
searchfor migratoryprey (see also Blades 1999;
Feblot-Augustins1997a,1997b,1997c;Rensinket
al. 1991). This inferenceimplies extensive depth
of planningin landscapeuse.
A finalfamilyof modelsfocusesin greaterdetail
on the decay-likerelationshipbetweenthe quantities of specific stone raw materialsfound in an
assemblageand the distancefrom the sourcesof
those materials.In general,the closest stone raw
materialsourcescontributethe greatestquantities
to an assemblage(60-80 percentfrom sources<5
km away),whilethemostdistantsourcescontribute
diminishinglysmall quantities(1-2 percentfrom
sources >20 km away) (Feblot-Augustins1993;
Geneste 1988). In CentralEurope,Middle Paleolithicassemblagesshowa dramaticdeclinein the
percentage frequency of stone materials from
sourcesbeyond50 km fromthe site (Figure3). At
Kulna(Layer 11), a MoravianMiddle Paleolithic
cave site, 87 percent of the assemblageis com-
AMERICAN ANTIQUITY
490
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formalriskmanagementstrategies(Geneste1989).
DistanceFromSource(km)
In particular,technologies are expectedto be formally designed from the outset to be small, light
gh
to
to
Figure 2. Frequencyhistogramsof raw materialtransfer may designed r
are to be
and resstant o flure
lure ithe
in (a) Central we
distaiicesforMiddlePaleolithicassemblages
Eurolpe and (b) the Aquitaine Basin. High frequencies of
short -distancetransfers(A) are interpretedin terms of
optimlization of time and energy costs associated with raw
materrialprocurement.
Internalmodes(B) are interpreted
in terims of logistical or seasonal use of distant ecological
patchles. Raw material transfers from very distant sources
(C) alre thoughtto markthe maximumextentof the geographic range of the populations in question. Data from
Feblo}t-Augustins(1993).
P. Jeffrey Brantingham]
ment,consumption,anddiscardareindependentof
rawmaterialtype designationsandthereforecomprise an unbiasedstone raw materialprocurement
strategy.The model providesa baseline for comparisonwherewe can be certainthatadaptationis
not responsiblefor observedpatternsin raw material richness,transportdistances,and both quantity-distance and reduction intensity-distance
relationships(see Gotelli and Graves1996:6).
Elements of a Neutral Model
491
Hubbell 2001). Such models thus call into question the ultimate role of ecological assembly
rules-especially interspecific competition-in
establishingandmaintainingcommunity-levelecological diversity (Connell 1980; MacArthurand
Levins 1967;Roughgarden1983; Schoener1983;
Weiherand Keddy 1999).
A neutralmodel of lithic technologicalorganizationbasedon similarfirstprinciplesoffersa radical pointof departurefromtraditionalapproaches
(butsee Neiman [1995] and ShennanandWilkinson [2001] for related discussions of "cultural
drift").Theneutralmodelof stonerawmaterialprocurementdevelopedherejettisonsthe assumption
thatdifferencesbetween stone raw materialsnecessarilyinfluencebothprocurementdecisions and
how stone toolkits are maintainedand materials
ultimatelydiscarded.The alternative,neutralstarting point is to assumethat all stone raw materials
areequivalenton aper unitbasis.Forexample,each
unitweight of rawmaterial,regardlessof whether
we classify it as chert, obsidian, or quartzite,is
assumedto be equivalentin termsof probabilities
of procurement,consumption,and discard. The
dynamicsof stone procurement,use, and discard
in thisneutralframeworkarethereforeindependent
of how we mightlabel raw materialtypes and any
unique functionalqualities we might attributeto
them.
It is pertinentto ask whetherthe patternsof raw
materialdiversityobserved in the archaeological
recordare consistentwith procurementstrategies
modeledusing such extremeneutralassumptions.
If they are consistent,then we must confrontthe
difficultpossibilitythatempiricallyobservablefeaturesof thearchaeologicalrecordsuchasrawmaterial richness and transportdistances may not be
telling us muchof anythingaboutoptimizationof
procurementbehaviors, depth of planning, risk
managementstrategies,and, ultimately,adaptive
variability.Of course, one may reject the neutral
modelif observedarchaeologicalpatternsarefound
to be inconsistentwith its expectations.Interpretations linking patternsof raw materialdiversityto
specializedbehavioraladaptationswouldbe more
robustas a result.
492
AMERICAN ANTIQUITY
modelingplatform(http://repast.sourceforge.net).
The simulated"world"consists of a two-dimensional grid (500 x 500 cells) that holds all of the
rawmaterialsourcesanda single foragerengaged
in a randomwalk throughthe environment.Each
grid cell is assumedto containa uniform,infinite
food supply,ensuringthatthereareno patchchoice
decisions to be made in forager movement (see
below). The worldcan containa variablenumber
of raw materialsources(Figure4). In most simulations,theworldis seededwith5,000 pointsources
of stone raw materialand each of these sourcesis
arbitrarilyassigneda uniquetype label i = 1, 2,..
.5,000. The coordinatelocationof each rawmaterial sourceon the grid is chosen at randomfrom a
uniform distribution (x, y) = [1, 500], without
replacement.There are 250,000 cells that could
potentiallyhold a uniquerawmaterialsource.The
probabilitythat any one cell contains stone raw
materialis approximately.02 (i.e., 5,000 sources/
250,000 cells) and, since each point sourcerepresents an arbitrarilyuniqueraw materialtype, the
probabilitythatanyone cell containsa specificraw
materialtype is approximately4.0 x 106 (i.e., 1
type / 250,000 cells). Alternatively,these numbers
may be thoughtof as the environmentaldensities
of all stone raw materialsand specific raw material types, respectively.
100
:
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100
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*-
66
80
90
1001
Figure 4. A snapshot of a 100-x-100 cell area of the simulation world (total size 500 x 500 cells) showing the random
distribution of raw material sources in the environment.
The mean distance between nearest neighboring raw
material sources is 3.72 grid cells (standard deviation =
1.85 cells;
minimum
= 1.0 cells;
maximum
= 8.25
cells).
P. Jeffrey Brantingham]
The initialcoordinatepositionof the single foragerin the environmentis chosenin a mannersimilar to the positioning of raw material sources,
namely from a uniform distribution(x, y) = [1,
500]. At each time step, the foragermoves to one
of the nearest eight neighboring cells (i.e., the
Moore neighborhood)or stays in the presentcell,
with equal probability(p = 1/9). This movement
rule defines a randomwalk whereinthere are no
first-orhigher-order
correlationsin movedirections
(Turchin1998:78).
Whena rawmaterialsourceis encountered,the
forager evaluates the present size of the mobile
toolkit and collects only as much raw materialas
is necessaryto provisionthe toolkitup to the maximumsize. Themobiletoolkitis simulatedas a vectorviwhereeach elementrepresentsthe amountof
stone raw materialin the toolkit of uniquetype i.
Inmostsimulations,themaximumamountof materialthatcan be carriedin the mobiletoolkitis arbitrarily set at 100 units. Accordingly,the mobile
toolkit must always meet the constraint:
v < 100
(1)
i(2)
Yi,Vi(3)
i
493
494
AMERICAN ANTIQUITY
6000 .
n = 22,830
5000
4000
.E
3000
2000
1000
__1__
I.
0
10
11
P. Jeffrey Brantingham]
NEUTRALMODELOF STONERAWMATERIAL
PROCUREMENT
495
496
AMERICAN ANTIQUITY
definesthemaximumpossiblepackingof resources
into a local foraging area. When the simulated
world contains 5,000 unique sources-a global
density of .02 sources/gridcell-the set of minimumnearestneighborswill on averageinclude 11
sourceswithinthe boundariesof a foragingareaof
P. Jeffrey Brantingham]
PROCUREMENT
NEUTRALMODELOF STONERAWMATERIAL
497
Toolkit Richness
1
2
3
4
5
6
7
Total
Number of Occurrences
160
104
93
49
54
18
3
481
Mean
63.6
30.3
29.7
26.6
14.9
20.8
6.7
38.7
d=10
d=20
d=30
20.7
9.0
3.9
Note: Data are for one simulationrun using baseline parameters.
given in Figure 11.
"Expectedamount (arbitraryunits) calculatedfrom the equationy = 47.726e-0'0834X
Maximum
100
95
95
95
70
66
12
100
d=40
1.7
AMERICAN
ANTIQUITY
498
12
10
2'
10000
11000
12000
13000
14000
15000
Time Step
Log Quantity of Material in Mobile Toolkit (arbitraryunits)
mentpoint for each uniqueraw materialtype presentin the mobile toolkit.This procedureprovides
aneffectivemeasureof how fara rawmaterialtype
has been transportedfromits source.Figure9 presents frequency histograms of distances from
sourcefor materialsin themobiletoolkitmeasured
in two differentspatialcontexts.Figure9a shows
a randomsample of approximately10 percentof
all raw materialoccurrencesin the mobile toolkit
for one simulationrun lasting 24,294 time steps.
Representedarethe numberof time stepsin which
a materialpresentin the mobile toolkitis found a
specifiednumberof gridcells fromthepointsource,
regardlessof whetherthe forageris presentlyat a
source of material.Figure 9b, in contrast,shows
thedistancefromsourceformaterialsin themobile
toolkit observed only at the points of encounter
withrawmaterialsources.Thetwo spatialcontexts
displaysome interestingsimilarities.
When observationsof the mobile toolkit are
madeindependentof spatiallocation,thefrequency
distribution of distances displays an "internal
mode" with a long right skew (Figure 9a). The
internalmodeindicatesthatthemobiletoolkitmost
often containsmaterialtypes from sourcesin relativelyclose proximityto the spatiallocationwhere
the toolkit is observed.The mediantransportdistance (9 cells) represents1.2 percentof the maxiTransportDistance
mumpossiblelineardistancebetweentwo sources,
At each time step in a simulation,it is possible to the diagonaldistanceof 707 grid cells across the
evaluatehow far the forageris from the procure- simulationworld.The modaltransportdistance(5
P. Jeffrey Brantingham]
PROCUREMENT
NEUTRALMODELOF STONERAWMATERIAL
800
600
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400
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200
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21
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200
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100
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16
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28
32
36
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AMERICAN ANTIQUITY
500
225
400
625
900
0.35
0.3
0.25
0
L,
a
0.2
P.
o
0.15
0.1
to d.4
0.05
10
15
20
25
30
35
Technologyof TransportedMaterials
The simulationalso providesneutralexpectations
for the quantity and technological characterof
materialsin the mobile toolkit transportedfrom
sourcesat differentdistances.Figure11 illustrates
thatthe quantityof materiali in the mobile toolkit
as a functionof distancefrom the source of i follows an exponentialdeclineof the formy = b*e-ax.
Materialsfromthe closest sourcesareusuallyrepresentedin thegreatestquantities,while thosefrom
the most distant sources are representedin the
smallest quantities.Note, however,that variation
aroundthe mean increaseswith increasingproximity to the sourceof the material.Thus,materials
from nearbysources are frequentlyprocuredand
transportedeitherboth in high andlow quantities.
At greaterdistancesfromsource,rawmaterialsare
more uniformlyrepresentedby low quantities.
The high variancein the quantitiesof materials
transportedover short distances stems from the
influencethatmaterialsalreadyin the toolkithave
on the size of rawmaterialpackagesprocured.For
example,if raw materialsources are close to one
anotherin space, then thereis a high probability
that the toolkit will be nearlyfull when a unique
sourceis encountered.Becauseprecedenceis given
to materialsalreadyin the toolkit-there is no discard of excess material-the quantityof material
procuredfrom the newly encounteredsourcewill
be relativelylow. If materialsourcesarespacedfar
apart, however, then the mobile toolkit will be
nearlyor completelyemptywhen a uniquesource
is encountered.In this case, the size of the raw
materialpackageprocuredfromthenewly encounteredsourcewill approachmaximumtoolkit size.
The low variancein materialquantitiestransported from distant sources reflects a different
mechanism:materialstransported
fromgreaterdistancesdisplaythe effects of repeatedconsumption
events,dampeningout anyeffectof initialpackage
P. Jeffrey Brantingham]
100
90
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cc
.s
?-
80
70
60
50
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ct
40
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30
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Cy
20
501
10
0
1
10
13
16
19
22
25
28
31
34
37
40
size. Inessence,rawmaterialstransportedfromthe
most distantsources have been "atrisk"for consumptionthe longest and thereforeoccur consistently in the smallestquantitieswithinthe mobile
toolkit. Table 3 presentsdata on the sizes of raw
materialpackagesprocuredwhenthemobiletoolkit
is at differentrichnesslevels. Mean package size
procureddecreasesas richnessincreases,illustrating thatmaterialsalreadyin themobiletoolkitplace
certainlimitson the amountof materialthatcanbe
procuredfrom a newly encounteredsource.
Table 3 also illustrates,as alludedto above, a
fundamentalrelationshipbetween distance from
source and intensity of material consumption.
Quantitiesof materialsin the mobile toolkit from
nearby sources generally have spent the least
amountof time in the toolkit and have therefore
been exposedto limitedriskof consumption.Predictably, these materials are not intensively
reduced.Risk of consumptionincreaseswith the
amount of time a materialtype has spent in the
mobile toolkit, which equations 4-6 show is
directlyrelatedto raw materialtransportdistance.
Materialsfrom more distantsourcesare thus predictablymore intensivelyreduced.This patternis
independentof rawmaterialtypedesignations.For
example,for a startingmobiletoolkitof richnessk
Archaeological Parallels
Severalimportantobservationsandpredictionsare
derivedfrom the neutralmodel developedabove.
First, raw material richness within the mobile
toolkit shouldbe expectedto varydynamicallyas
a functionof both spatiallocationandtime, butin
allcasesrichnessis constrainedto be muchless than
themaximumrichnesstheoreticallyattainable.The
signatureof this process is a dependenceof maximum toolkitrichnesson samplesize. Remembering thatall materialsin the simulatedenvironment
occur at equal densities, it is clear thatthe biases
in the representationof raw materialswithin the
toolkit arise solely from a neutral,non-adaptive
raw materialprocurementstrategy.The fact that
some materialspresentin theenvironmentarenever
procured,othersare only rarelyprocured,while a
few are commonly procuredneed not imply raw
materialselectivityon the partof the forager.
Second,thefrequencydistributionof rawmaterial transportdistancesdisplays an internalmode
and a long right skew. Maximum transportdistances are expected to be three to four times the
distancerepresentedby theinternalmode.Theratio
of the maximumto the modalraw materialtransport distanceis largely unaffectedby changes in
mobile toolkit size, consumptionrate,or the naturaldensitiesof rawmaterialsin the environment.
Maximumtransport
distancesmaytranslateintothe
"utilizationrange"of a givenrawmaterialtype,but
are equivocallyrelatedto the "geographicrange"
of the forager.
Finally,the quantityof materialof a given type
withinthemobiletoolkitgenerallyfollowsanexponential decline with increasing distance from
source.As an extension of this pattern,materials
from the most distantsources are expected to be
representedconsistently(i.e., withlow variance)in
small quantitiesandbe heavilyreducedrelativeto
502
AMERICAN ANTIQUITY
P. Jeffrey Brantingham]
503
504
AMERICAN ANTIQUITY
imply anydepthof planning,optimization,orrisk- ham et al. 2003; Brantinghamet al. 2001; Turchin
1998)-that random-walk foraging and a rawmanagementstrategies.
materialprocurementstrategythatis indifferentto
Discussion
stone types couldbe optimal.The conclusionhere
an
common
sense-tells
would
be thatthe absence of both planningdepth
Many ethnography-and
us thatforagersemploycomplexoptimalforaging andexplicitriskmanagementstrategieswouldyield
strategies(Binford2001; Kelly 1995;Smith1991). higherfitnessthantheirmorecomplexalternatives.
Itis perhapsmorelikelythatotheraspectsof forIt is assumedthatoptimalforagingstrategiesmust
influenceand,therefore,be diagnosedby stoneraw agingbehaviorwere subjectto optimization,depth
materialprocurementpatterns.Thepresentmodel, of planning,andriskmanagementin the ways sughowever,is basedon assumptionsthatforagersdo gestedby contemporaryarchaeologicaltheory,but
not optimizeany specificcurrencyassociatedwith thatstone raw materialprocurementandtransport
movement,do not dependon anyformof planning were not, at leastas reflectedin the empiricalmeadepth,andarerisk insensitivein all of theirmove- suresdiscussedhere.9Foragingstrategiesoptimized
ment and procurementdecisions. Surprisingly, with respectto mobile prey,or seasonallydistribthese extremeassumptionslead to patternsin raw utedplantresources,maybe entirelystochasticwith
materialrichness and transportthat are qualita- respectto rawmaterialsourceencountersandraw
tively similarto commonlyobservedarchaeologi- materialprocurement.Moreover,whetheror not
cal patterns.Whatshouldwe concludefromthese stone procurementbehaviorswere optimized(or
sensitiveto risk) may have had little or no impact
results?
Thereare at least two answersto this question. on the optimalityof food procurementstrategies.
First, it is possible thatthe processes of archaeo- The conclusionherewould be thata random-walk
strategyandindifferenceto stone
logical site formation-especially time-averag- stoneprocurement
of
discrete
eradicate
the
fine
details
raw
material
ing-may
type may not entailthe fitnesspenalof
behavties
a
events
commonlyassumedand, importantly,thatthe
leaving palimpsest
procurement
ioral traces indistinguishable from the neutral morecomplexalternatives
maynotnecessarilyyield
model. In general, time-averagingoccurs when- higherpayoffs.This is a logical extensionof arguevereventsthathappenedatdifferentpointsin time ments suggestingthatstone raw materialprocureappear synchronous in the geological record ment was completely embedded within other
(Kowalewski1996). It influencesourunderstand- foragingactivities(Binford1979).Absentanydirect
ing of behavioral, ecological, or evolutionary fitnessconsequences,stoneprocurementandtransrateis slower port behaviorswould be neutralwith respect to
processeswheneverthesedimentation
thanthetime scale of theprocessin question(Bush selection andwouldbe free to varyin a stochastic
et al. 2002; Kowalewskiet al. 1998; Stem 1994). manner.
The presentmodel,like manyneutralmodelsin
Stone procurementstrategieswere implemented
on time scalesof minutesto perhapsmonths,if sea- community ecology (Gotelli and Graves
sonalplanningwas in play,while sedimentationis 1996:5-6), raises questions about how we infer
generally a much slower process. Moreover, adaptationfromempiricalpatterns.Thissaid,there
geochronologicalcontrolsrarelyoffer such fine- will be temptationforcriticsto rejectthemodeloutscale resolution.Thereis thus little questionthat right by claiming that foragers "would" never
time averagingwill be a concern in interpreting engage in a randomwalk and"would"nevermake
most cases of stonerawmaterialprocurementand raw-materialprocurementdecisions withoutcontransport(but see Close 2000). It may be the case sidering raw materialtype. Similarly,motivated
thatdiscretestoneprocurementeventswerehighly objectionshave been voiced over neutralmodels
adaptivein all of the ways suggestedby current discountingthe importanceof interspecificcomecologicalcommunities(see
archaeologicaltheory,butin some cases it maynot petitionin structuring
be possibleto distinguishthe aggregatepatternsof Connell 1980; Conner and Simberloff 1979;
procurementbehaviorfromthe neutralalternative. Roughgarden1983; Simberloff1983). In this latSecond, it is conceivable-though perhaps ter case, conventionalwisdom suggeststhatcomunlikelyin manyenvironments(butsee Branting- petition is the critical determinant of whether
P. Jeffrey Brantingham]
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Notes
1. A two-dimensionalrandomwalk on a lattice is commonly used as a discrete approximationof spatial diffusion.
N - (d/l)2 is the mean squareddistance of spread(the variance) after N time steps for an ensemble of particlesundergoing diffusion. Denny and Gaines (2002) provide a
straightforwardderivationof this formulafor the one-dimensional case. The derivationsfor the two- and three-dimensional cases yield the same fundamentalrelationship(Denny
and Gaines 2002:163-167).
2. This analysis is based on the limiting case where raw
material sources are encounteredonly within the 100 time
steps following a firstencounter,and not subsequently(i.e., d
= 10 using equation[4]). Of course,raw materialsourcesmay
be encounteredover time intervalsgreaterthanN = 100. The
resultsarecomparableto this limitingcase becauseof a tradeoff between time spent at different richness states and the
amountof materialrepresentedby the most abundanttype in
the toolkit (see discussion in text).
3. The discrepancybetween the observed and expected
maximumtransportdistancesreflectsthe pathdependenceof
a randomwalk. The analyticalform N = d2may substantially
under- or over-estimatethe actual distance traveledon any
c!(l-c)Nj!(N- j)!
P. Jeffrey Brantingham]
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