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Journal of Archaeological Science (1997) 24, 3963

Contemporary Shellfish Gathering Strategies among theMeriam of the Torres Strait Islands, Australia: Testing

Predictions of a Central Place Foraging ModelDouglas W. Bird and Rebecca L. Bliege Bird

Department of Anthropology, University of Utah, Salt Lake City, UT84112, U.S.A.

(Received 1 September 1995, revised manuscript accepted 15 December 1995)

This paper explores a central place foraging model with data on Meriam intertidal shellfish gathering strategies, fieldprocessing practices, patterns of resource transport, and consequences of these factors for introducing variability inshell assemblage composition among these Islanders of the eastern Torres Strait, Australia. As a result of differentialfield processing and shell material transport, we show that some species are likely to be consistently over-representedin shell assemblages while others are likely to be under-represented relative to their actual dietary importance.

Explanations for this behavior are tested with the predictions of a central place forging model derived from behavioralecology. We find consistent relationships between the anticipated and observed patterns of transport for five species.These results have implications for current thought about the relationship between faunal assemblages and transportbehavior in a wide array of contexts, interpretations of site function and settlement patterns, and arguments about roleof shellfish in prehistoric diets. 1997 Academic Press Limited

Keywords: SHELLFISH GATHERING, DIFFERENTIAL TRANSPORT, SHELL MIDDENS, CENTRALPLACE FORAGING THEORY, MERIAM.

Introduction

Archaeologists often base their arguments about

prehistoric subsistence strategies, food sharingbehavior, the sexual division of labor, and

settlement patterns on inferences drawn from patternvariability in faunal assemblage composition. Suchpatterns of variability can be influenced by numerousprocesses, including decisions that humans make aboutresource transport and processing (Perkins & Daly,1986; White, 1952, 1953). When differential process-ing and transport have been examined in enthno-graphic contexts, analyses of the types and amountsof subsistence remains found at butchery sites andresidential sites provide evidence that variability inbone accumulation is influenced by the value of

particular anatomical parts (Yellen, 1977; Binford,1978, 1981; Bunn, 1983; Bunn et al., 1988; OConnellet al., 1988, 1990; OConnell & Marshall, 1989; andcontributors to Hudson, 1993). Much of thiscurrent work focuses on contemporary butchery andcarcass transport, and analyses their effects onskeletal part profiles produced in the context of largemammal hunting. This work also provides insightsinto processes that might affect faunal variability inshell middens. If foragers differentially transport theshells of certain shellfish prey, simple analyses ofmidden composition may incorrectly estimate theimportance of particular shellfish species or the rela-

tive contribution of shellfish to the wider prehistoricdiet.

Recently, archaeologists have begun to apply the

logic of economic decision-making to understand whyforagers choose to discard some resource parts atthe procurement site and transport others (Jones &Metcalfe, 1988; OConnell et al., 1988, 1990; Jones &Madsen, 1989; OConnell & Marshall, 1989; Elston,1989, 1990; Rhode, 1990; Metcalfe & Barlow, 1992;Barlow & Metcalfe, 1996). The model presented byMetcalfe and Barlow in particular draws our attentionto a small set of cost/benefit variables (foraging rangeand processing efficiency) that influence processing andtransport decisions where foragers are interested in therate at which they deliver resources to a central locale.This model is derived from central place foraging

theory (sensu Orians & Pearson, 1979; Stephens &Krebs, 1986), and has merit for archaeological prob-lems in its reliance on the deductive methodology ofbehavioral ecology (see OConnell, 1995).

This paper investigates the utility and applicabilityof the MetcalfeBarlow central place foraging modelfor generating predictions about human transportbehavior and its depositional consequences. We testpredictions of the model with ethnographic data oncontemporary intertidal gathering strategies among theMeriam of the eastern Torres Strait Islands, Australia.In particular, we focus on (1) dissimilarities betweenthe relative contribution of some types of intertidal

390305-4403/97/010039+ 25 $25.00/0/as950095 1997 Academic Press Limited


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resources to the Meriam diet and the contemporarydeposition of remains at residential sites, and moreimportantly, (2) testing the hypothesis that locations ofprocessing activities can be predicted by attributesof the resources. Our approach to ethnoarchaeologydiffers from research that seeks out ethnographicanalogies with widely applicable archaeologicalimplications. We seek rather, to test with ethnographicdata a formal hypothesis derived not from ethno-graphic observations, but from a well establishedtheoretical framework. Although there are importantarchaeological lessons presented by these observations,

it is the hypothesis, not necessarily the case study,that has wide application in studies of prehistoricsubsistence.

The Study Site

The Meriam are indigenous inhabitants of the eastern-most Torres Strait in tropical northeastern Australia.Their three islands (Mer, Dauar and Waier) lie 6 kmfrom the outer Great Barrier Reef, about 250 kmnortheast from the Australian mainland and 140 kmsoutheast from Papua New Guineas southern coast.

Mer, Dauar and Waier (Figure 1) are of volcanic originand are forested with deciduous vine thicket. TheMeriam community, consisting of about 400 people, istoday nucleated into a single village on the northwestforeshore of Mer. Dauar and Waier are used exten-sively for temporary residence, gardening, fishing andgathering.

The Meriam are of Melanesian descent and probablyoriginated from the region near New Guineas FlyRiver delta. Their traditional language, Meriam Mir, isclosely related to those spoken in the Oriomo low-lands of southern Papua New Guinea (Wurm, 1972).

Although the islands have been part of the state ofQueensland since 1879, much of their oral history andculture has strong ties to southern coastal and riverineNew Guinea.

Traditionally, the Meriam were intensive horticul-turists. Since the 1970s extensive gardening activitieshave been somewhat curtailed. Flour, rice and othergoods now arrive on a fortnightly barge. Most house-holds maintain kitchen gardens, but the large swiddenplots in the interior of the islands often remain fallow.

Despite this decrease in horticultural activities, re-sources acquired from traditional marine subsistencepractices remain a substantial part of the Meriam diet.

2k m

50km

Me r

D a u a r

Waier

NeKeper

WaierPit

Ormi

Giar Pit

Teg

N

10

50mcont our int er val

Main resid ent ialar ea

Fr ingin greef

Meria mIslan ds

Werbadu

Eger

Las Keper

Las

Ulag

Australia

Mer

P a pua NewGui nea FlyRiver

Figure 1. Map of Mer, Dauar and Waier. Intertidal gathering activities occur on most parts of the fringing reefs that surround the MeriamIslands except the subtidal lagoons Las Keper and Ne Keper.

40 D. W. Bird and R. L. Bliege Bird


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Fish and marine turtle comprise the majority of proteinconsumed on Mer. Fin-fish and squid are caughtusing hand lines, nets, and spears from the foreshore,fringing reefs, and submerged atolls near the islands.Sea turtles are hunted in dinghies between May andSeptember, while during October through April they

are usually taken at nesting sites on the islands [seeBliege Bird et al. (1995); and Bliege Bird & Bird(in press), for details on Meriam fishing, turtleacquisition, and food sharing strategies]. Intertidalgathering, primarily of shellfish, also remains animportant part of Meriam subsistence. We focus ontwo types of intertidal gathering in detail: reef flatcollecting and rocky shore harvesting.

Data collection

Intertidal gathering activities were monitored throughfocal individual foraging follows. Focal individual fol-

lows consist of sampling all occurrences of behaviors ofindividuals, recording episode length, time spent ineach activity, and measuring the individuals yieldduring the sample period (Altmann, 1974). We con-ducted a total of 91 opportunistic and systematicintertidal focal follows over 14 months betweenJanuary 1993 and March 1995. Opportunistic follows(N=32) occurred when we were participating in forag-ing episodes when intertidal gathering took place.Systematic follows (N=59) were recorded duringobservational sampling of time allocation to foreshoreand reef subsistence activities. We attempted to samplewidely the intertidal gathering activities of adultwomen and men, as well as children. Thirty-three

childrens (15 and younger) focal follows, 16 mensfollows, and 42 follows of women were completed.Fifty individuals are represented in the follows.The follows amounted to more than 137 foragerobservation hours.

Meriam Intertidal Gathering

Reef flat collecting

Reef flat collecting of molluscan species constitutes90% of intertidal gathering effort. Reef flat environ-ments are structured into midlittoral zones consisting

of sea grass beds, sandy flats, micro-atolls, soft corals,branch coral flats, and a sublittoral fringe composed oflive corals and algal pavement. Reef flat resources arerandomly and sparsely distributed within zones, al-though species densities are variable. Gastropodprey commonly encountered are spider conch (Lambislambis), trochus shell (Trochus niloticus), tiger cowry(Cypraea tigris), red lipped conch (Strombus luhuanus),and less commonly vase shell (Vasum turbinellus),turban shell (Turbo petholatus), baler shell (Meloamphora), Australian trumpet (Syrinx aruanas), andabalone (Haliotis varia). The bivalves collected includemostly tridacnids (large, algal symbiotic clams): bear-

paw clam (Hippopus hippopus), fluted clam (Tridacnasquamosa), elongated clam (Tridacna maxima), giantclam (Tridacna gigas), and boring clam (Tridacnacrocea). Other bivalves rarely collected includepearl oyster (Pinctada margaritifera), hammer oyster(Malleus malleus) and comb oyster (Lopha cristagalli).

Occasionally other invertebrates such as Octopus vul-garis are taken if opportunistically encountered on thereef while collecting other molluscs.

Collecting and field processing strategies.Most collect-ing begins as foragers enter the reef flat from theforeshore. Foragers travel through the midlittoral,harvesting prey, and when the tide recedes, move intothe mid-sublittoral margin of the reef. Collecting tech-nologies are relatively simple: 10 litre plastic buckets, along-bladed knife, and a hammer are commonly car-ried, but foragers may also carry spears and handlines.While children form relatively large foraging groups

(three to seven individuals), adult men and womentend to forage separately and on their own.Appendix Table A presents data from 48 focal

follows during which Hippopus were collected. Loadsof Hippopus ranged from 50 g of edible flesh to 82 kg.An average 122 kg (S.D.=156) of edible flesh washarvested per focal individual follow. Each specimentook an average of 138 min to field process(S.D.=15), and each processed Hippopus weighed180 gm (S.D.=120). Foragers processed immediatelyupon encounter by spearing the mantle with a knife toprevent the valves from closing, cutting the flesh fromthe shell and removing the kidney.

Appendix Table B presents data recorded during 33

focal follows when Tridacna spp. were harvested.When harvested, these species made up on average113 kg (S.D.=174) of edible flesh in a load of reef flatresources, ranging from 45 g to 85 kg. Tridacna spp.required an average of 26 min per specimen to fieldprocess (S.D.=191 min). The average flesh weight perharvested animal was 300 g, but specimens variedwidely (S.D.=420 g, ranging from 40 g to 17 kg).Unlike Hippopus, Tridacna squamosa, T. maxima, andT. crocea are all attached to the reef by strong byssalthreads and require extraction before processing.Foragers spear the mantle with a knife before thevalves close and use the knife as a lever to pry the

animal free of its byssus. T. crocea, the most abundantshellfish species on the reef, is completely embedded incoral and its handling costs inevitably cause foragers toignore it. T. gigas are often too large to be picked up(let alone transported) when they are encountered.These were exploited only when the tide fully exposedthe reef edge where these huge bivalves are found.

Data recorded on 66 focal follows during whichLambis were collected are presented in Appendix TableC. Foragers harvested an average of 490 gm(S.D.=573) of edible Lambis flesh per follow, rangingfrom about 25 g to just over 3 kg. Each Lambisrequires on average 092 min (S.D.=084) to field

Contemporary Shellfish Gathering Strategies 41


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process and yields 25 g (S.D.=6) of edible flesh.Foragers process each animal singly, often as it iscollected. Animals are processed by cracking the shellswith a hammer and removing and discarding theentrails, retaining only the muscle and small claw-likeoperculum.

Shell disposal, cooking and dinner-time camps.When shellfish are field-processed, foragers deposit theshells at processing sites on the reef or at dinner-timecamps (sensu Meehan, 1982) on the foreshoressupratidal fringe. Processing sites on the reef are verysmall and very ephemeral. They are marked by thepresence of one species in very small numbers (in thecase of tridacnids, only one shell is left behind at eachprocessing location). Concentrations of processedLambis shell on the reef flat remained for no more thantwo tidal cycles.

During four focal follows, foragers in groups

processed and consumed Lambis at dinner-time campsnear the procurement area. Foragers processed and ateLambis after roasting them in-shell in a small hearthprepared on the foreshore. Heating the shell causesit to become brittle and makes it easier to crack.Dinner-time sites during our study were marked by thepresence of only one or two species (Lambis and/orCypraea) and a single ephemeral hearth. If foragerstransport shell remains to residences, they oftendeposit them in heaps covered with sand to avoidstepping on sharp shell fragments.

Constraints on reef flat collecting.Shellfish collecting is most productive during the low-

est daily tide: any water on the reef makes preyrecognition difficult. All collection that we observedtook place during the 24 h window of opportunitycreated by a diurnal low spring tide. There are alsoseasonal tidal cycles which fully expose the reef justafter midnight during half of the year and just afternoon in the other half. Most shellfish gathering duringour study occurred between 1 May and the end of July1994 when there were 50 days during which at least10% of the fringing reef was exposed during daylighthours. From November 1994 through January 1995diurnal ebbing tides exposed 10% of the reef on only 16days and shellfish collecting was far less common.

Foragers on the reef flat are also constrained by thevolume and weight of a load of unprocessed shellfish.Hippopus average 18 kg (gross weight) per specimen,Tridacna spp. average 26 kg per specimen, and Lambis03 kg per specimen. On average, an individual adultforager can collect about 55 Hippopus, 3 Tridacnaspp., and 17 Lambis specimens (or about 20 kg grossweight) per hour (see Appendix Tables AC). Reef flatcollecting generally lasts longer than an hour andwithout field processing a forager would quickly exceedthe weight she could comfortably carry and still con-tinue collection. Furthermore, one 10 litre bucket willhold only four or five medium sized unprocessed

tridacnids or about 30 medium sized unprocessedLambis. Given their collecting rate, foragers will al-most always fill up a container within an hour if theydo not field process. Foragers must then decidewhether they should trade-off foraging time with timespent field processing to decrease inedible bulk, or

whether they should trade-off an increase in theamount of edible flesh in a load for more time to collectand transport more resources.

Rocky shore harvesting

Rocky shore harvesting is a true central place foragingactivity. Foragers travel to known patches of rockyshore resources, spend time harvesting within thepatch, and transport shellfish back to a central place.Rocky shore harvesting takes place just out from theforeshore in the upper intertidal. Productive patchesare marked by a matrix of fine sands, intermixed

with gravels and overlain by aggregations of basalticcobbles and boulders. No such areas exist immediatelyadjacent to the village foreshore on Mer: foragers walkto patches near Ulag and Eger on Mer, or while onDauar, to locations near Waier Pit and Giar Pit (seeFigure 1). Two types of shellfish are exploited in thispatch: sunset clams (Asaphis violascens) and wavednerites (Nerita undata). Foragers never collect smalllimpets (Patelloida sp.) and ox-palate nerites (Neritaalbicilla), which are also encountered while harvesting.The average amount of edible flesh harvested per loadwhile rocky shore harvesting is 719 g (N=10,S.D.=407).

Asaphis are a small sand bivalve (mean edible flesh

weight per harvested animal is 93 g, Appendix Table D:N=7 trials, 446 specimens, S.D.= 12 g) found in thesilts and gravels beneath large stones. To harvest them,foragers overturn stones and look for the small holescreated by the Asaphis siphon. As foragers dig forAsaphis they also encounter Nerita undata in the crevicesof rocks overlaying the clam bed. These small gastro-pods average 22 g edible flesh per animal (N=6 trials,939 specimens, S.D.=02 g, see Appendix Table E).

While we never observed field processing of either ofthese two prey types prior to transport, foragersinformed us that they do occasionally roast, processand consume Asaphis violascens at dinner-time camps.

When foragers process Asaphis at home, they may cutthe flesh from the valves and remove the stomachcontents, but Asaphis may also be roasted in shell. Thiscooking method involves clearing a hearth of its coalsand placing the shellfish, with their lips facing downand their hinges uppermost, on a bed of hot sand. Thehot coals are placed on the phalanx of the shells. Whenthe juices cease to boil, the cook removes the shellsfrom the hearth and allows them to cool. She thenextracts the meat and cleans the stomach contents.Nerita undata are usually boiled. After the shells cool,diners pull the flesh from the shell with a small pin,consuming the entrails along with the muscle. Nerita



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can also be cooked on a fire but because of their smallsize they are difficult to separate from the coals.

Tidal conditions do not seem to constrain rockyshore harvesting to the same extent as reef flat collect-ing. Patches are located higher in the littoral andcan be exploited even when the reef is not completelydry. Nevertheless, there is still only a 4 h window ofopportunity created by diurnal low tides during whichharvesting can occur. If foragers do not field process,they collect about 306 kg of rocky shore resourcesper hour (Appendix Tables D and E: total weightharvested per total time harvesting). In 4 h, a forager

could harvest about 12 kg of unprocessed resources, aload that would be difficult to transport much morethan 1 km, especially if the forager is accompanied bya small child. As with reef flat collecting, container sizeconstrains harvesting. A 10 litre bucket of Asaphisweighs 910 kg and can be filled easily in about 3 h ofcollecting. Foragers face the familiar trade-off: once abucket is full or if they exceed the weight they cancarry, they must either field process to make room formore resources or cease harvesting to carry the loadback home.

Shellfish Remains at Residences and RelativeDietary Importance

In order to test the assumption that the relative dietaryimportance of shellfish types in the Meriam diet can beinferred from the types and amounts of shellfish thatare deposited at a residential site, we sampled the shellaccumulations of six households and compared thesedata with observed dietary contribution of shellfish.Between 24 March and 4 July 1994 we asked sixrandomly selected households to deposit the valves ofall shellfish transported home whole into containers.We checked the containers once a week and counted

and weighed all shells according to species. We thenconverted shell weights to edible flesh weights andcompared these with observational data. Table 1presents summary data of shell accumulations and therelative contribution of species to the diet. Estimates ofthe relative dietary importance based on residentialshell accumulations were calculated by multiplyingshell weights of each prey type by its correspondingexperimental mean edible flesh to shell ratio (seeTable 2). The focal follow sample in Table 1 gives theedible flesh weights collected by members of the samplehouseholds during those intertidal gathering follows

that occurred between 24 March and 4 July. Table 1also presents the total edible flesh collected of eachspecies during the entire study period (see AppendixTables AE).

Figure 2 illustrates the relationship between the threeedible flesh figures from Table 1. Although proportion-ally the edible flesh weight collected by householdsduring the sample period differs little from the totalflesh weight collected across all follows, discrepanciesbetween these figures and the edible flesh weight rep-resented in the shell accumulations are obvious. This isespecially the case for Hippopus and Tridacna spp.whose dietary contribution is under-represented in the

shell accumulations, and the rocky shore resourcesAsaphis and Nerita whose remains are over-representedin the accumulations. The tridacnids (Hippopus andTridacna genera) represented less than 10% of thereconstructed edible flesh weight and yet in the samplefollows and across all follows they made up 62% and67% of the edible flesh weight collected respectively.No Tridacna spp. shells were deposited at the resi-dences and yet during the sample period householdmembers collected 615 kg ofTridacna flesh, making upalmost 20% of the total sample flesh weight. Con-versely, rocky shore resources represented over 30% ofthe flesh weight in the household shell remains but less

Table 1. Shellfish remains, reconstructed dietary importance, and observed dietary importance (ranked in order ofreconstructed dietary importance)

SpeciesFlesh:shell

ratio

Household samples Sample follows All follows

kg fleshreconstructed

kg shellobserved kg flesh # follows kg flesh # follows

Lambis 0092 366 3978 814 11 3237 66Asaphis 0338 350 1036 221 4 434 7Cypraea 0278 346 1055 038 5 223 21Trochus 0208 157 755 057 3 414 12Hippopus 0115 148 1287 1356 9 5855 48Nerita 0301 105 349 072 3 207 6Strombus 0148 023 155 005 2 183 13Vasum 0148 015 101 0012 2 008 7Tridacna 0170 000 000 615 7 3714 33Total 1510 8716 31788 14275

Flesh:shell ratio was obtained experimentally (Table 2) (no data on Vasum are available, flesh:shell ratio forStrombus which is similar in size and shape was used).Household samples include shell accumulation of six households between 24 March and 4 July 1994. Samplefollows are focal follows of individuals from the household sample during the sample period. All follows includeall 91 focal individual follows (kg flesh is total w1 in Appendix Tables AE).



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than 10% of the sample-household edible shellfish

weight and less than 5% of the total flesh weight. In theMeriam case, unlike other studies comparing archaeo-logical and ethnographic evidence (e.g. Meehan, 1982),we can eliminate the possible hypothesis that thesediscrepancies are the result of changes in prey choice:the patterns of predation and deposition are contem-poraneous. We know that these differences resultedfrom differential field processing and transport of shell-fish prey types. The task now is to provide explanationsfor this behavior to account for biases in shell accumu-lation composition.

Central Place Foraging Models

We are just beginning to investigate the special oppor-tunities and constraints that human foragers mightface in traveling to collect resources and transportingthem back to a central locale (see Schoener, 1971;Orians & Pearson, 1979; Kacelnik, 1984; Stephens &Krebs, 1986; for development of central place forag-ing models in studies of animal behavior). Modelsin animal studies have addressed questions aboutresource load size and its relationship to variability inthe rate of food delivery to a central place whendelivery rate is often critical to a foragers overallreproductive success. Metcalfe & Barlows (1992) cen-

tral place foraging model may be especially useful forexploring the solution to trade-offs that human for-agers are liable to face in decisions about processingand transporting resources that have parts of both lowand high value. Shellfish often have parts of relativelylittle utility and we might expect intertidal gatherers (ifthey are interested in foraging efficiently and are con-strained by the size of a load that they can carry) toface decisions about what will maximize their deliveryrate of high quality material per unit time spent

foraging: spending time to cull parts of little value nearthe site of procurement (field processing) to increasethe value of the transported load, or forgoing thebenefits of field processing to spend more timeforaging. The MetcalfeBarlow model predicts thatcollection and field processing time will vary with (1)distance to and from the procurement site, and (2)expected variation in the time spent collecting andprocessing (costs), and increases in the quality of a loadif it is processed prior to transport (benefits).

Figure 3 illustrates the relationship between the costsof field processing and benefits of increased load qual-ity [the utility function, U(t)]. The slope of this func-

tion predicts the time traveled when field processingbecomes economically profitable. Higher field process-ing costs associated with smaller increases in loadquality will increase the minimum distance from acentral locale at which field processing will maximizethe overall efficiency of foraging (collecting, process-ing, and transporting) and vice versa. If foragers trans-port the optimal load size, the travel time beyondwhich field processing will increase foraging efficiency(i.e. the rate-maximizing solution to trade-offs betweenfield processing and transport) can be calculated withthe following equation (after Metcalfe & Barlow, 1992:344):

Lambis

Tridacna

0.1

Pr opor t ion oft ota led ible fles h weig h t

Preytype

0. 50.2 0.4

Asaphis

Cypraea

Trochus

Hippopus

Nerita

Strombus

Vasum

0.30.0

Figure 2. Relative Nutritional Importance of Meriam Shellfish.Taken from the relative proportions of edible flesh presented inTable 1. Reconstructed proportion is the edible flesh represented inhousehold shell samples between 24 March and 4 July 1994, samplefollow proportion is the edible flesh harvested by sample householdmembers during the sample period, and all follows include theproportions of edible flesh harvested during 91 focal individualfollows throughout the entire study period. : Reconstructed fromhousehold shell accumulations;: sample follows; : all follows.

Roun d-tr iptr aveltime

Ut

x1

0.4

1.0

0.8

0.6

0.2

0

x0z

y1

y0

Collectin gan dfieldpre cessingtim e

Loadutil

ity

Figure 3. Graph of the relationship between the costs (x0 and x1)and benefits (y0 and y1) of field processing [the utility function U(t)]

and the point at which field processing before transport will increasenet yield per unit time spent foraging (z).



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z =y0x1 y1x0

y1 y0(1)

where z=the minimum round trip travel time at whichfield processing will increase the delivery rate of highquality material, x0 is the cost of collecting a load ofunprocessed resources, x1 is the cost of collecting andfield processing a load of resources, y0 (benefit without

field processing) is the proportion of an unprocessedload of resources consisting of high quality material,and y1 (benefit with field processing) is the proportionof a processed load of resources consisting of highquality material.

Estimating the models parameters and predictions

Costs.Equations (2) and (3) monitor variability in theanticipated costs associated with different shellfishtypes by calculating the time required to collect andfield process a load of resources. Here:

x0 =tc

w0(L) (2)

and,

x1 =tc + tfp

w1(L) (3)

where for each shellfish species, x0 is the cost (in min)of collecting 1 kg of gross (unprocessed) shellfishmultiplied by the total weight of the load transported(L), x1 is the cost (in min) of collecting and fieldprocessing 1 kg of edible (processed) shellfish flesh,again multiplied by the transported load size (L), tc is

the time spent collecting a resource during a focalfollow, tfp is the time spent field processing, w0 is theload weight of a resource if no field processing were tooccur, and w1 is the load weight of a resource if fieldprocessed.

The Appendix Tables A through E present data onthe times, weights, and costs associated with collectingand field processing each shellfish species. In Table 2we present data on processing experiments when

foragers were asked to process loads of shellfish. Bothobserved and experimental data were used to calculateintertidal gathering costs.

For reef flat collecting follows during which some ofthe resources were not field processed, increases in xwith tfp were calculated with average field processingrates. For Hippopus and Tridacna we calculated tfpfor unprocessed specimens by multiplying w1 by theaverage time a child (age 15 or under) or adult requiresto obtain 1 kg of edible flesh by field processing [seeAppendix Tables A and B; time fp per kg fp: Hippopus:children=195 min, adults=62 min (there were nosignificant differences between adult females andmales, t-test, df=32, P=083); Tridacna: children=333min, adult females=122 min, adult males=62 min.]Unlike the tridacnids, there were no significant differ-ences between the time required to process a large orsmall sized Lambis [correlation comparing the averageflesh weight (kg fp per # fp) and the average timerequired to field process a specimen (time fp per # fp),r2=004, F-test, P=028]. Therefore, to calculate tfpfor Lambis specimens that were not field processedwe multiplied the number harvested but not fieldprocessed by the average time a child or an adult wouldrequire to process one Lambis specimen (children=16 min, adults=043 min, no significant differences

Table 2. Experimental data summary: processing reef flat and rocky shore resources

Prey typeN

(experiments)#

(specimens)Flesh per gross

(kg)Flesh per shell

(kg)tp per flesh(min kg1)

Flesh tp(g min1)

Hippopus 13 21 0098 0115 S.D. = 002 S.D. = 001

Tridacna spp.* 8 8 0117 0170 S.D. = 002 S.D. = 001

Lambis 6 98 0078 0092 S.D. = 001 S.D. = 002

Cypraea 4 51 0193 0278 264 388S.D.=002 S.D.=52 S.D.=73

Strombus 4 174 0124 0148 1162 86S.D.=001 S.D.=08 S.D.=01

Asaphis 5 394 0183 0338 1216 92S.D. = 002 S.D. = 002 S.D. = 519 S.D. =30

Nerita 3 205 0301 0301 1683 59S.D. =002 S.D. =002 S.D. =42 S.D. =02

tp: Time processing.*T. maxima and T. squamosa.



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between female and male adults, t-test, df=21,P=023).

Estimating tfp for rocky shore resources (Asaphisviolascens and Nerita undata), which were never fieldprocessed during the focal follows, required exper-imental data. We asked three adult female foragers at

the conclusion of five foraging trips to process loads ofeach of these shellfish types as they would if theresources had been processed away from home (with-out cooking). Asaphis were processed using the sametechniques generally employed for home processing:prying open the shell, removing the flesh and cleaningthe stomach of its contents. Processing Nerita involvedinserting a small pin past the operculum and pullingout the flesh. Neither the stomach contents nor the tinyoperculum of this prey type were removed prior toconsumption. We weighed and counted the animalsprior to processing, measured the total amount of timerequired to process the load of each shellfish type, and

weighed the edible flesh after processing. Increasesin collecting costs with field processing were thencalculated by multiplying w1 by the experimentalaverage time required to process 1 kg of edible flesh(Table 2; minimum processing kg1 flesh: Asaphis=1216, Nerita=1683).

Benefits.Quantifying benefits for field processingspecies loads can be expressed as:

y0 = 0 (L) (4)

and,

y0 = 1 (L) (5)

where y0 and y1 measure of the proportion of edibleflesh that makes up the transported load before andafter field processing respectively, 0 and 1 are esti-mates of the proportion of edible flesh per kilogram ofunprocessed and processed shellfish respectively, and Lis the total load weight transported.

The benefits of field processing were also calculatedwith a mix of observational and experimental data.Because the Meriam usually field process H. hippopus,Tridacna spp., and Lambis on-encounter, we wereoften able to weigh only the field processed flesh.Similarly, if the shellfish were not field processed, weonly had the opportunity to weigh only unprocessedshellfish. If we measured only gross weight or edibleflesh weight during a follow, the average experimentalproportions of each species (see Table 2; kg flesh kggross1) were used as an estimate of 0. Because afully processed load contains only parts of high quality(i.e. edible flesh), 1 was always 10.

Predictions and assumptions.The Appendix Tables AEalso present for each follow the predicted valuesfor travel times and distances beyond which fieldprocessing is expected to occur (zt and zd1 respectively).The times were calculated using equation (1) and

distances (one way) were estimated at a walking speedof three km h1. Again, graphically, z is thex-intercept of a line tangent with the utility functionU(t) (Figure 3).

With reliable estimates of these variables, we can testthe following prediction for intertidal gathering among

the Meriam: If Meriam intertidal gatherers fieldprocess to maximize the rate of edible flesh deliveredto a central locale, then we expect the relationshipbetween distance and transported waste material to bedescribed by a dichotomous function in which theproportion of waste in a load falls immediately to zero(100% edible flesh) as the maximum terminal foragingdistance (MTFD) exceeds the z value. The analysispresented below compares the MTFD and the pro-portion of the transported load made up of wastematerial for each follow. For each shellfish species,there should be no waste material transported to acentral locale (either a residential or dinner-time camp)

during those follows where the maximum terminalforaging distance exceeds the z value. We have modi-fied the models prediction to reflect the amount ofshell that will be transported and, because resourcesare transported while foraging within the reef flatpatch, we used the MTFD (m) from the centralbase during the follow rather than time spent in travelto the patch. For rocky shore harvesting, no transportoccurred within the patch, and thus the MTFD is alsothe one way travel distance to the patch.

This model works under several simplifyingassumptions. First, it assumes that foragers have al-ready decided to carry efficient load sizes and exploitefficient patches and prey. Second, it assumes that

travel, collection, and processing are mutuallyexclusive activities. Third, it presumes that shellfishhave two components, a valuable one (edible flesh) anda worthless one (primarily shell). And finally, theprediction specifies edible flesh weight rather thanenergetic gain as the currency to be maximized. In theanalysis below we discuss the extent to which theseassumptions might be violated by Meriam intertidalgathering and how this might affect differences betweenpredicted values and observations.

Analysis

Reef flat collecting

Tridacnids.As the z values in the Appendix Tables Athrough E demonstrate, the range of predicted dis-tances at which field processing becomes profitable issubstantial. Although there is considerable variabilitybetween follows, the mean of z for each prey type issignificantly different from the others. At one extremeare the two giant clam genera Hippopus and Tridacnawith relatively small mean z one way distance values(zd1) of 746 and 137 m respectively. These prey typesare typified by relatively low processing costs and largeincreases in load quality if they are field processed. If



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foraging occurs beyond about 150 m from the resi-dence, no shells of either of these prey types areexpected to be transported to a central place. Figures 4and 5 show the relationship between (1) the differencebetween the MTFD and the one way distance atwhich field processing will be economically profitable(distance minus zd1 on the x-axis) and (2) the pro-portion of waste material in loads of Hippopus and

Tridacna on the y-axis. Each point represents a singlefocal follow during which these two prey types wereexploited. The model predicts that all positive x-axisvalues (when the MTFD exceeded the point at whichfield processing would increase the amount of edibleflesh returned home) should have no waste material intransported loads, while negative x-axis values shouldbe associated with y-axis values that are greater than 0.

3500

1

0.2500

MTFD minusz

d1 (m)

Proportionofloadconsistingofw

astematerial

2000

0

0.8

0.6

0.4

0.2

0 500 1000 1500 2500 3000

Outliers

Figure 4. Relationship between Hippopus hippopus proportional waste transported to a central locale and the difference between the maximumterminal foraging distance (MTFD, in m) and the predicted point (in one way m) at which field processing will increase the weight of edibleflesh harvested and transported to a central locale ( zd1). Follows during which the MTFD exceeded the z value (positive points on the x-axis)and with no waste transported or follows that did not exceed the z value (negative points on the y-axis) and with some waste transported fitthe models prediction (43 follows, five outliers).

3500

1

0.2500

MTFD minus zd1 (m)

Proportionofloadconsistingo

fwastematerial

2000

0

0.8

0.6

0.4

0.2

0 500 1000 1500 2500 3000

Outliers

Figure 5. Relationship between Tridacna spp. proportional waste transported to a central locale and the difference between the MTFD (in m)and the predicted point (in one way m) at which field processing will increase the weight of edible flesh harvested and transported to a centrallocale (zd1). Explanation as in Figure 4 (30 follows, three outliers).



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We were able to estimate z values for 43 of the 48follows during which Hippopus were harvested. TheMTFD in all cases except one exceeded the distance atwhich field processing is predicted to increase the rateof edible flesh delivered to a central locale. During thisfollow, no shellfish were field processed. On five of the

remaining 42 follows during which Hippopus wereexploited and we were able to estimate the z value, atleast some shell and other waste material was trans-ported home. In other words, 37 of these 42 followswere predicted by the model. As such, 38 of the 43 datapoints (88%) lie within our predictions. If we usemean z values according to age (children=869 m,adults=714 m) for those follows for which we wereunable to calculate z, 43 of the 48 data points (90%) fitthe predictions.

Tridacna spp. were harvested on 33 follows, 30 ofwhich we were able to calculate predictive values.During four of the 33 follows at least some shell was

transported home. Three of these four follows wereassociated with MTFDs that exceed their z values andthus lie beyond the anticipated distance at which fieldprocessing should have occurred. Of the follows forwhich we have z values, 90% (27 of 30) of the vari-ability is accounted for by the models predictions. Ifwe include mean Tridacna z values for children(1729 m), men (819 m), and women (1418 m) forthe three follows that we were unable to estimate theassociated z, 29 of all 33 Tridacna follows (88%) fit theprediction.

These figures demonstrate a good fit between thepredicted and observed results, and illuminate somekey factors influencing variability in large clam field

processing. For adult female foragers, the fit for bothHippopus and Tridacna improves to 92% for Hippopus(23 of 25 follows with z values) and 100% for Tridacna(all 13 follows). Women transported only two loads ofHippopus and one of Tridacna spp. that consisted ofany low quality parts. Thus, adult females are slightlymore likely to field process the tridacnids than arechildren and adult males.

This discrepancy may be the result of a number offactors, particularly the intensity of intertidal gather-ing. For men and children, reef flat foraging can oftenbe embedded in other activities. Men occasionallycollect substantial amounts of shellfish while spear

fishing at the edge of the fringing reef, while childrenmay harvest clams opportunistically after reef edgefishing or during play. When encounters are not antici-pated, foragers may not be carrying tools to exploittridacnids efficiently resulting in a substantial increasein the distance at which field processing the clamswould be profitable. In 54 follows during whichforagers collected Hippopus and/or Tridacna, shell wastransported to residences nine times. Lack of efficienttechnology carried by children or men accounts forseven of the nine follows not matching the predictions.

To account for other follows which are unantici-pated by the model we might investigate variability in

the value of the shell. Our analysis assumes that theshells of molluscs have no value and that field process-ing decisions would not be affected by the benefitsaccrued by transporting the clams in-shell. Thisassumption is occasionally violated. Hippopus andTridacna spp. shells are occasionally used to define

land boundaries, collect water in gardens, or make adecorative border around cultivated plants. In the past,tridacnid shells were a source of vital tool material,rain water containers, and cooking utensils. It may alsobe the case that because women do most of the homeprocessing regardless of who acquired the resource,childrens and mens field processing decisions areaffected by the fact that they may not be required toprocess the resources after returning home. Given thatprocessing costs associated with these clams are rela-tively small, this seems improbable. Furthermore, thisfact would not change the predicted z values becausethe model assumes no processing costs to the forager

upon return to the residence.

Lambis lambis.Figure 6 presents the relationshipbetween the maximum terminal foraging distanceduring 52 Lambis follows the distance at which fieldprocessing will increase foraging efficiency (zdf) and theproportion of transported Lambis that consisted ofwaste material. On eight follows the MTFD did notexceed the predicted z value. During these episodes, asexpected, no field processing occurred. However on 22of the 44 follows in which the distance walked exceededthe z value, at least some of this prey type wastransported home whole. In total, 30 of the 52 follows(58%) match the predictions that no shell will be

transported if the MTFD extended beyond z or that noprocessing will occur during follows not exceeded by z.If we include all 66 follows, using mean z values forchildren (4034 m) and adults (2008 m) on follows forwhich we were unable to calculate z, 59% (39 of 66follows) of the variability is accounted for by theprediction.

Regardless of the increase in variability for Lambisfollows relative to the tridacnids, the hypothesis thatincreases in the MTFD will be associated with anincrease in the likelihood that field processing willoccur is strongly supported by the data. Figure 7 showsthe negative correlation between the MTFD and the

Lambis waste weight transported (r

2

=0197, F-test,df=67, P


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was significantly longer than the mean associated withLambis loads of only whole specimens (N=30,mean=5853 m, S.D.=4334) (two-tailed t-test,l df=33,P


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distance and proportion of transported Lambis loadsconsisting of waste material. Because women are moreeffective at cracking Lambis shells than are children,the point at which field processing will increase awomans overall rate of edible flesh yield whileforaging (195 m) is substantially reduced from the

combined mean zdfof 2787 m (total mean versus adultfemale mean, two-tailed t-test, df=25, P=0087). Assuch, about 35% of the variability in womens followsremains unaccounted for by the model. Seventeen ofthe 26 follows (65%) with z values during which womencollected Lambis fit our predictions, compared to thetotal of 5859% inclusive of all follows.

Two important points about collecting Lambis helpto illustrate how reef flat collecting might violate someof the models assumptions. First, especially amongwomen, loads of Lambis transported back to theresidence occasionally contain both fully processedflesh and whole animals. Of the 13 loads that women

transported to a central locale that contained at leastsome shell material, four (31%) were mixed with fullyprocessed and whole specimens. On all of these fol-lows, foragers collected without field processing onlyon the return trip, as the tide flooded the reef. Thisdemonstrates that travel and collection are not alwaysmutually exclusive and provides good support for ahypothesis that the decision to field process while onthe reef flat does not remain constant during mostcollecting bouts.

The second point about Lambis collecting dealsmore specifically with those follows exceeding the zvalue during which no field processing occurred(N=30). Foragers often commented during such fol-

lows that they wanted to transport whole Lambis sothat they could roast them in the shell. This suggeststhat other assumptions of the model may not alwayshold true for reef flat collecting: Lambis shells may notbe completely without value. The shell can serve as itsown cooking container and Lambis roasted in shellseem to be much easier to crack than uncookedanimals. Such may be the payoffof roasting Lambis inthe shell, that foragers reported occasions in the pastwhen sand and wood were hauled out on the reef whilethis and other reef flat resources were collected andcooked at the site of procurement. Here, foragerstransported the central place to the site of procure-

ment rather than transporting the resources to acentral place.Dinner-time camps might also be a solution to the

trade-off between overall efficiency while foraging andhome processing benefits. Resources are thus fullyprocessed prior to travel back to the residence, but thebenefits of roasting the shellfish whole are not lost. Wemight expect dinner-time camps to occur on episodesduring which collection occurs relatively far from theresidence, and where gathering is a collective endeavor.Lambis were processed at dinner-time camps on fourfocal follows. These follows all occurred during picnictrips in which groups of four or five foragers traveled

relatively far from the village, collected reef flat re-sources (processing the tridacnids on encounter asusual), roasted, field processed, and ate the cookedLambis, and later, transported the tridacnids back totheir residences. Although we do not have enough ofthese episodes for statistical analysis, compared to the

average episode during which Lambis were collected,picnic trips with dinnertime camps involved largergroups and took place at substantially greater dis-tances from home (average for all follows=21 foragersper episode, and 338 m from residence to collectionstarting location; average for picnic trips= 45 foragersper episode, and 19 km from residence to collectionstarting location).

Rocky shore harvesting

Rocky shore harvesting matches the models predic-tions better than do the results of the Lambis analysis.

However, rocky shore harvesting is uncommon relativeto reef flat collecting. Thus our conclusions concerningAsaphis violascens and Nerita undata harvesting arebased on a much smaller data set: 10 foraging followsin this patch were fully recorded. Neither of thesespecies was passed over while harvesting in this patch.No significant travel or transport occurred within therocky shore patch.

Asaphis violascens.The point at which field processingbecomes profitable for this prey type is significantlygreater than the z value reported for reef flat resources(mean=2419 m, S.D.=1584). This is primarily a resultof increased field processing times that foragers will

accrue if Asaphis is culled of waste material beforetransport. Field processing would require the flesh ofeach animal to be removed from the shell and stomachcontents scraped clean.

Only one follow during which Asaphis wereharvested was associated with an MTFD greater thanits associated z value. In this case, no field processingoccurred when the amount of flesh returned to theresidence could have been greater if the Asaphis hadbeen field processed. For all Asaphis follows, the modelpredicts 86% of the variability (six of seven). Figure 8shows this relationship and further supports a moregeneral hypothesis that the Asaphis waste weight trans-

ported home decreases with decreases in the diff

erencebetween the MTFD and the z value. Follows withincreasingly large amounts of waste weight transportedwere consistently associated with walking distancesthat fell increasingly short of the point at which fieldprocessing would be profitable (r2=09, F-test, df=6,P


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consumed before returning home. Eger lies 2700 and3100 walking m from the foragers respective resi-

dences. Both episodes would have incorporated walk-ing distances that exceeded the average predicted valueat which field processing should occur. Foragers oftentold us that if they target Asaphis on apek (literallyother side) trips they will process and consume loadsbefore returning home.

Nerita undata.The average distance at which we wouldexpect Nerita field processing to occur is over 5 km(53557 m, S.D.=31338), and is never less than 3 km, adistance far greater than any other shellfish species.The six follows during which Nerita undata wereharvested never exceeded 750 walking m from central

place to rocky shore patch and never came close toexceeding their associated z values. As expected, no

Nerita were ever field processed. All loads of thisspecies had similar proportions of waste material trans-ported (roughly 60% waste). Figure 9 presents therelationship between MTFD minus zd1 and the wasteweight transported. During three follows less than05 kg waste weight was returned home. On Mer,Dauar, and Waier it is not possible to walk anywherethat lies beyond about 35 km from a starting point. Assuch, in most cases there is no point at which theMeriam should field process Nerita undata. In fact,when we asked foragers if they ever thought aboutdiscarding these shells before returning home, theyalways said that it would take too much time.

1000

6

05000

MTFD minus zd1 (m)

kgwaste

1000

2

5

4

3

04000 3000 2000

1

Outlier

Figure 8. Relationship between actual Asaphis violenscens waste transported to a central locale and the difference between the MTFD (in m)and the predicted point (in one way m) at which field processing will increase the weight of edible flesh harvested and transported to a central

locale (zd1). Explanation as in Figure 4 (seven follows, one outlier). y=0001x+1046; r2

=090; df=6; F-test, P


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Summary

In general, the Meriam cull shell material prior totransport in a manner that maximizes the rate at whichthey deliver edible flesh to a central locale. Prey typeswhich are relatively difficult to field process, providerelatively little increase in proportion of edible flesh iffield processed and were gathered near the foragersresidence or temporary camp were transported whole.Species which require relatively little time to fieldprocess, provide relatively large increases in the pro-portion of edible flesh if field processed and wereexploited by foragers who walked far from a centrallocale were routinely culled of waste. The tridacnidswere almost always field processed, because theyrequire so little time to field process, and the quality ofload of clams can be greatly improved by field process-ing. The conch Lambis lambis is usually processedbefore transport as expected by the model. However, itappears that processing decisions based on encounters

at variable distances from the central place and un-measured benefits of transporting whole specimensoccasionally cause foragers to transport unprocessedLambis in a manner unanticipated by our predictions.Although our samples of Asaphis violascens andNerita undata harvesting are relatively small, rockyshore resources were almost always transported aspredicted by the model. These resources were neverfield processed. Only one follow exceeded the point atwhich field processing would increase the overall rateat which flesh was delivered to a central place.

Archaeological Implications

Under the circumstances specified above, archaeolo-gists should expect substantial variability in shellfishassemblage composition as a result of differentialtransport of shell material. Our analysis suggests thatdelineating these effects in archaeological situationsshould be possible with estimates of the models essen-tial parameters. Meriam intertidal gathering and thehypothesis analysed above have important implica-tions for (1) understanding transport decisions andrelationships between resource processing costs,foraging range, and patterns of discard; (2) archaeo-logical reconstructions of settlement patterns and site

function; and (3) current thought about the importanceof littoral resources in prehistoric diets.

Transport decisions

Archaeologists have long recognized the importance ofdifferential transport as a potential agent of variabilityin bone assemblages (e.g. White, 1952, 1953; Perkins &Daly, 1986). Recently, especially as a result of ethno-graphic work among modern hunters (Binford, 1978;Bunn et al ., 1988; OConnell et al ., 1988, 1990;OConnell & Marshall, 1989; Bartram, 1993; Emerson,1993), researchers have begun to pay increasing atten-

tion to the benefits of field processing and foragingdistance as factors that determine carcass handling andtransport decisions. These studies have shown thatwhen constrained by load size, hunters often selectanatomical parts for transport with respect to theirvalue (i.e. parts with higher associated nutritional value

are less likely to be discarded before returning toresidential camps). OConnell et al . (1988, 1990),guided by theoretical constructs, have argued explicitlythat the net benefit of any particular anatomical partwill be influenced by the costs of field processing andtransport distance. Our results concur with thesestudies and demonstrate that the same variables affecttransport decisions in an entirely different context.Moreover, our research provides the first test of amodel that analyses the relationship between process-ing costs and foraging range and its effect on process-ing location. Insofar as our analysis demonstrates amatch between the models predictions and the

Meriam data, we have shown that variability in discardpatterns is shaped by the trade-off between fieldprocessing and transport. Although researchers haveyet to develop simple models (like the one tested here)appropriate for analysing the transport of more struc-turally complex resources (but see Metcalfe & Barlow,1992: 350351), decisions about processing and trans-porting resources like shellfish that have few parts canbe readily explained with estimates of variabilityin expected field processing costs and procurementlocation.

The implications of this for interpreting assemblagesof subsistence remains are well illustrated in BettyMeehans (1982) landmark study of intertidal gather-

ing among the Gidjingali of coastal Arnhem Land,Australia. The Gidjingali focus their shellfish gatheringeffort toward the collection of only a small fraction ofthe wide array of species available for exploitation. Ofthe 29 species of shellfish observed to have beencollected, 95% of the total weight was contributed byonly six species, with 61% of the total made up by onespecies, Tapes hiantina. Meehan notes that observedpatterns in Gidjingali foraging behavior seem to be insharp contrast to the prehistoric evidence from thesame region. For example, in samples taken from threeseparate prehistoric middens in Gidjingali territory,Tapes constituted merely 14% and 3% of two of the

assemblages by count, while the third sample had noTapes. Meehan argues that this indicates a shift in preychoice over time resulting from environmental changes(1982: 168). This hypothesis has yet to be tested butwould require details about the local paleoenvironmentand its relationship to intertidal gathering strategies.

Another testable explanation of the contrastbetween the ethnographic and archaeological evidenceis that the shellfish waste material surviving in archaeo-logical deposits may have been differentially trans-ported to the sites in question. Meehan observes thatoften the Gidjingali process Tapes hiantina at dinner-time camps near the site of procurement prior to



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returning to the residential camp, even when residentialcamps are within 1 km of the shell beds. As such,variability in the taxonomic representations of theshellfish remains from the tested prehistoric sites mayreflect differences in the extent of processing near thesite of procurement rather than relative importance of

the shellfish types in the prehistoric diet. An instructivetest of similar hypotheses could also be investigated insituations where researchers have noted that tridacnidsare conspicuously missing or rare in sites where reefenvironments or ethnographic observations suggestthat they were available to foragers (e.g. Janetski, 1976;Swadling & Chowning, 1981; Beaton, 1985; Swadling,1986; Spennemann, 1987; Sullivan & Sassoon, 1987).

In many cases, the predictions of central placeforaging models and their implications for estimatingthe degree to which faunal assemblages might bebiased by differential transport could be investigatedwith actualistic data on the time required to collect and

field process resources and distances of the sites fromthe littoral [see Barlow & Metcalfe (1995) for anexample dealing with plant transport]. Where sitesfarther from intertidal zones consistently lack par-ticular species that are represented in sites closer toshell beds, archaeologists could begin to tease out theeffects of differential transport (if any) on shell middenvariability. The predictions of central place foragingmodels may be especially illuminating where thenumber of prey types exploited from similar environsremains relatively constant in various assemblageswhile others fluctuate. We expect that species whichdisappear from assemblages as distance from the coastincreases would be associated with z values less than

the estimated travel distances to the littoral areas. Atest of this hypothesis would require sampling anumber of middens in the same area where foragerswould have been harvesting shellfish at known dis-tances from procurement areas. This underscores theimportance of regional approaches to archaeologicalproblems. Such a project is feasible in areas likenorthern Arnhem Land and Cape York Peninsula ofAustralia where various middens from the same areaare abundant. Alternatively, situations where the goalof collecting shellfish was not directly related tomaximizing the rate of flesh delivery (for instancewhere shell may have had value as tool or ornamenta-

tion material) should be very clear exceptions to themodels predictions.

Site function and settlement patterns

Many archaeological studies have proposed that vari-ability in the relative types and amounts of shellfish cancontribute toward understanding site function andprehistoric settlement patterns (e.g. Shawcross, 1967;Bailey, 1975; Bowdler, 1977; Barber, 1982; Buchananet al., 1984; Beaton, 1985). Our results show that therole of a site in a settlement system varies as a functionof resource processing characteristics and foraging

range. Processing sites, dinner-time camps, and resi-dential locales become distinct in the archaeologicalrecord as differences between these variables increase.In turn, where the benefits of field processing are greatenough, these sites are likely to be associated withpatterned biases in the relative representation of shell

and intrasite variability is unlikely to provide reliableevidence of residential patterns. These results againemphasize the importance of regional approaches forreconstructing prehistoric settlement systems.

Central place foraging models might also be a sourcefor interpretive hypotheses of site function and settle-ment patterns. One might predict that sites near shell-fish habitats with faunal assemblages containing anabundance of homogeneous shellfish remains associ-ated with low relative z values to indicate processingsites or dinner-time camps, not residential sites. Sitesnear intertidal or riverine environs containing shellfishassociated with both relatively low and high z values

might represent residential bases. Sites relatively farfrom procurement areas with assemblages consistingprimarily of shellfish with high z values are also morelikely to be residential loci. These hypotheses couldthen be tested against independently derived data thatmight also indicate residential or processing sites (e.g.presence or absence of extensive secondary deposition,the relative abundance of artifactual material, indica-tions of the presence or lack of structures, habitations,and storage).

The role of intertidal resources in the diet

The Meriam data are relevant for many studies of

the importance of shellfish in prehistoric diets (e.g.Parmalee & Kippel, 1974; Bailey, 1975; Osborn, 1977;Perlman, 1980; Yesner, 1981; Claassen, 1985, 1986;Glassow & Wilcoxon, 1988; Erlandson, 1988, 1991;Raab, 1992) insofar as they demonstrate circumstanceswhere shellfish remains are likely to be closely tied tothe littoral. In cases where the benefits of homeprocessing do not offset the benefits of culling shellwaste, the model predicts that on walking trips beyondabout 5 km from a central locale, foragers should fieldprocess most shellfish resources before transport. Wesuggest that incidents where archaeologists find signifi-cant amounts of shell remains in sites at distances

which are significantly farther from the intertidalwould indicate clear violations of the model and givereason for us to suspect that something other than therate of flesh delivery may be determining processingand transport decisions. The predictions of the modelwould likely be violated where shellfish are consistentlytransported whole in order to keep them fresh for laterconsumption or trade, or where the demand forshell itself is great. An important implication for anarchaeological test of this model would lie in itsability to identify unanticipated patterns in shellremains, allowing for researchers to delineate aspectsof variability that might be related to prehistoric



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opportunities and constraints other than immediatenutrition (e.g. storage, trade and technology).

Our analysis demonstrates that as a result of circum-stances related to the costs and benefits of shellfishgathering, midden assemblage composition may varyindependently of prey choice: all other factors being

equal, among foragers interested in maximizing theirrate of resource delivery to a central place, rela-tively high ranked shellfish resources (resources that onencounter have high energy yield per unit time process-ing; in the Meriam case, Hippopus and Tridacna spp.)are likely to be under-represented in residentialdeposits while low ranked shellfish (rocky shoreresources) are likely to be over-represented, irrespectiveof their importance in the diet. These relativedifferences will become more distinct in the archaeo-logical record with increasing distances between theprocurement location and central base.

Conclusions

A common objection to the application of ethno-graphic observations to archaeological problems isthat, because behavior is bound to its historical andcultural context, knowledge of modern humanforaging is ultimately limited in its ability to informresearch on the past. This critique has been leveledagainst many ethnoarchaeological studies, includingBinfords (1976, 1978) influential study of Nunamiuttransport (see comments in Hall, 1976). We agree withthese objections: if we treat Meriam shellfish gatheringas a model for interpreting archaeological remains, the

ethnographic observations are relevant to prehistoricstudies only in cautionary terms, regardless of howsimilar or dissimilar we feel modern and ancientcontexts might be.

However, our research among the Meriam isdesigned to test explanations of factors that determinevariability in field processing and transport behavior.The problem we address is not one of how appropriatemodern Meriam intertidal subsistence is as an analogyfor the past; rather, we attempt to understand thefactors that influence variation in subsistence behavior.Here lies the real value of formal models from behav-ioral ecology: these models nominate fitness related

factors that are likely to influence behavior in anycontext and provide precise predictions amenable totesting. While the complex and flexible reality of inter-tidal gathering behavior is not always reflected inthe central place foraging model we have tested, theprecision and generality of the simplest expression ofthis model are instructive. Our analysis shows thatmuch of the variability in Meriam field processing andtransport behavior can be explained in terms of therelationship between foraging range and processingefficiency. While these factors will take on differentvalues in different settings, they should be operativeand measurable in most contexts where individuals are

(or were) interested in efficiently delivering resources toa central place. We would suspect that in most cases,individuals who did so would be at a distinct advantageover individuals who did not. We do not attempt tonominate the Meriam as an analogue for understand-ing prehistoric huntergatherers; rather, we attempt to

evaluate with a specific ethnographic case the utility ofa hypothesis which could have application for a broadset of anthropological and archaeological problems notspecific to the Meriam or shellfish gathering.

The Meriam data suggest that researchers interestedin making inferences about shellfish gathering behaviorbased on archaeological patterning should considerthat under certain circumstances relative to processingcosts and foraging range, differential field processingand transport will be an important source of shellmidden variability. More importantly, the results indi-cate that variability in contemporary Meriam shelldeposition is patterned in predictable ways as a result

of subsistence decisions that are anticipated by centralplace foraging models. These subsistence decisions are,in principle, likely to be patterned in many settings andpredictable with certain known parameters inherent tothe resources (distribution and profitability). Wherepredictions and observations do not match, tests offormal models can isolate unanticipated variability andbe a source for alternative hypotheses. Most import-ant, the Meriam data contribute toward a growingunderstanding of how general theory can guide expla-nations of human decision making and its archaeo-logical consequences (Beaton, 1973; Winterhalder &Smith, 1982; Simms, 1992; Grayson, 1993; OConnell,1995). More studies, both ethnographic and actualistic,

investigating the range of costs and benefits involved inshellfish gathering will enhance our understanding ofhow differences in foraging goals, decisions, and con-straints are likely to affect variability in shell middencomposition.

Acknowledgements

This study was supported by a dissertation improve-ment grant from National Science Foundation, as wellas funds from the Leakey Foundation, the WennerGren Foundation, and the Australian Institute forAboriginal and Torres Strait Islander Studies. We owe

a great debt of gratitude to the Meriam, especially thePassi, Kaddy, Cowley, Lui, Mabo, Day, Rice, andKabere families, and the Mer Island CommunityCouncil for their support and immeasurable hospital-ity. We particularly thank our research assistant EdnaKabere for all of her dedicated work. The authors arealso grateful to James OConnell, Renee Barlow,Kristen Hawkes, Robert Bettinger, and DuncanMetcalfe for discussion and comments which greatlyimproved this paper. Jon Erlandson and David Rhodealso read an early draft and made many useful sugges-tions. Finally, we thank John Beaton for his unfailingassistance throughout this project.



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Appendix

Definition of variables in Tables AE:Follow= 001099 (children, age 15 and younger), 100199 (adult males, 200299 (adult females).tc=time (min) spent collecting.MTFD=Maximum Terminal Foraging Distance.# Harv=number harvested.# Gr tr= number of gross (whole) specimens trans-ported to a central locale.kg fl=weight of edible flesh (observed).kg gr=weight of gross shellfish (observed).# FP=number of specimens field processed.kg FP=weight of field processed edible flesh.Time FP=time (min) spent field processing.kg waste=weight of waste material transported.Prop waste=proportion of prey type load consisting ofwaste material.tfp per kgfp=time (min) field processing per kg edibleflesh.tfp per #fp=time (min) field processing per specimen.L=total load weight actually transported (kg) (all preytypes).w0=weight (kg) without field processing (calculated).w1=weight (kg) with field processing (calculated).0=proportion of prey type load consisting of edibleflesh without field processing.1=proportion of prey type load consisting of edibleflesh with field processing.

x0=costs (min) without field processing.x1=costs (min) with field processing.y0=value (kg) of unprocessed load.y1=value (kg) of processed load.zt=round trip travel time at which field processing willincrease the amount of edible flesh returned to acentral locale.zd1=one way travel distance at which field processingwill increase the amount of edible flesh returned to acentral locale.



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Table A. Hippopus hippopus: observational and calculated data

Follow tc MTFD#

Harv# Gr.

tr kg fl kg gr # FP kg FP Time FP kg wastePropwaste tfp per kgfp

004 3775 785 3 056 3 056 355 0 0 634005 4462 795 2 2 246 2219 0902 007 38 1810 1 0229 1 0229 10 0 0 4367011 85 3100 2 04 2 04 5 0 0 125015 58 200 3 3 12 1082 0902 018 505 1 0066 1 0066 162 0 0 2455019 1820 1 005 1 005 2 0 0 40020 64 3105 1 015 1 015 2 0 0 1333027 16 200 1 1 118 1064 0902 028 4263 810 2 037 2 037 237 0 0 641033 65 3115 3 3 0455 48 3 0455 4 0 0 879134 901 510 1 1 03 311 1 03 19 0 0 633136 61 2000 1 1 014 14 1 014 1 0 0 714142 1 0145 157 1083158 2977 695 4 24 4 24 1048 0 0 437161 9967 1600 1 03 1 03 125 0 0 417162 2500 26 82 26 82 0 0 163 71 1645 7 18 7 18 112 0 0 622164 100 300 3 3 6 5412 0902 165 7817 1750 6 087 6 087 493 0 0 567173 1867 1100 2 11 2 11 667 0 0 606180 820 1 1 01 09 1 01 06 0 0 6181 5681 700 5 13 5 13 517 0 0 398237 504 350 2 04 2 04 16 0 0 4238 8612 1650 3 027 3 027 346 0 0 1281243 9075 450 3 1 02 07 2 02 2 0632 0702 10245 4458 705 4 08 4 08 43 0 0 538246 457 495 5 04 5 04 367 0 0 918247 8848 1695 16 238 16 238 128 0 0 538248 8752 1700 12 13 12 13 1125 0 0 865251 4133 820 7 2555 7 2555 6 0 0 235252 521 305 12 45 12 45 189 0 0 42253 32 260 5 067 5 067 6 0 0 896254 12356 3210 6 06 6 06 545 0 0 908256 12262 1690 28 4055 27 4055 2302 0 0 568257 3749 1310 1 008 1 008 058 0 0 725260 1450 1 1 01 102 1 01 0 0 266 627 710 11 12 11 12 44 0 0 367

267 8185 1655 5 04 5 04 351 0 0 877268 4455 2510 10 2 10 2 12 0 0 6269 5691 1710 5 0525 5 0525 287 0 0 547270 4038 1715 12 216 12 216 987 0 0 457271 9693 1750 35 402 35 402 2915 0 0 725272 16325 1705 27 216 27 216 1048 0 0 485274 5573 660 16 336 16 336 1227 0 0 365275 7578 1610 9 1841 9 1841 59 0 0 32276 6372 650 2 02 2 02 128 0 0 64277 40 240 1 1 11 0992 0902 279 5635 950 8 22 8 22 865 0 0 393Total 2297 323Mean 651 12879 67S.D. 299 8392 79



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Table A. Continued

Follow L w0 w1 0 1 x0 x1 y0 y1 zt zd1

004 2077 5714 056 0098 1 1372 15318 0204 2077 143 3577005 27 246 0241 0098 1 4897 55244 0265 27 571 14267007 0252 2337 0229 0098 1 41 5282 0025 0252 12 2989011 1801 4082 04 0098 1 375 40523 0176 1801 245 6114015 287 12 0118 0098 1 13872 146637 0282 287 607 15185018 0466 0673 0066 019 0151 051 005 020 1207 1531 015 0098 1 5046 53108 0118 1207 175 437027 393 118 0116 0098 1 5329 61864 0386 393 835 20869028 562 3776 037 0098 1 6345 68351 0551 562 391 9776033 0455 48 0455 0095 1 616 69 0043 0455 042 1047134 10193 3109 03 0096 1 2954 312585 0984 10193 689 17236136 58 14 014 01 1 25271 256857 058 58 46 11508142 473 158 495 2449 24 0098 1 602 8302 0485 495 235 5871161 395 3061 03 0098 1 12862 132878 0387 395 179 4471162 134 83673 82 163 35 18367 18 0098 1 1353 15983 0343 35 237 5915164 8562 6 0588 0098 1 1427 150942 0839 8562 579 14477165 23 8878 087 0098 1 2025 21969 0225 23 142 354

173 42 11224 11 0098 1 699 9675 0412 42 2

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