1992_Stiner y Kuhn_American Anthropology Pararo

8/7/2019 1992_Stiner y Kuhn_American Anthropology Pararo

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Subsistence, Technology, and Adaptive Variation in Middle Paleolithic Italy

Author(s): Mary C. Stiner and Steven L. KuhnSource: American Anthropologist, New Series, Vol. 94, No. 2 (Jun., 1992), pp. 306-339Published by: Blackwell Publishing on behalf of the American Anthropological AssociationStable URL: http://www.jstor.org/stable/680462

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MARYC. STINERSTEVEN L. KUHNUniversityofNew Mexico

Subsistence, Technology, and Adaptive Variationin Middle Paleolithic Italy

Thiscollaborativestudyaddressesquestionsaboutvariationin behaviorwithintheMiddlePa-leolithicperiodofwest-centralItaly. Thefindingsaredirectlyrelevantto theevolutionary'fate"ofNeandertalsandtheappearanceofanatomicallymodernhumansinEurope.Analysesfocusonvariationina numberofdimensionsofsubsistenceandtechnology,includingpatternsofungulateprocurement,foodtransport,andtacticsofstonetoolmanufactureanduse.All ofthesedimensionsexhibitmarked,possibly"vectored"changein thestudyareabetween110,000and35,000yearsago.Interpretationsof thearcheologicaldataaresupportedby independentstudieslinkingfoodsearch,procurement,andtransportbehaviorstoecologicalrulesaffectingall largeterrestrialpred-ators.In thisway,variationobservedin thefaunaland lithic data sets can be shownto reflectstartlingdiversityin theresponsesof Mousterianhominidstotheworldaroundthem.The dataprovideimportantinsightsintothe natureof adaptive"rawmaterial"alreadyinplace duringtheMiddlePaleolithic,theforemostconclusionof thisstudy.Thatthisvariationin resourceex-ploitationandlandusealsoappearsdirectionalis moredifficulttoevaluate.It couldbeexplainedbyhominids'adjustmentsto localchangesin coastalhabitatas sea levelregressedor,alterna-tively, an evolutionaryshift in hominidcapabilities. Thefirst interpretationis preferred,on thebasisof availableevidence,but cannotbeadvancedas a certainconclusion.This questionprovidesmuchstimulusfor continuedresearchin thestudyarea and otherregions,using themethodspre-sented.

SOME OF THE MOSTCOMPELLINGresearch priorities in Paleolithic archeology concernthe nature of the Middle-Upper Paleolithic "transition" and the origins of anatomi-cally modern human populations. Viewpoints on this controversial period in human evo-lutionary history tend to be polarized into two camps. One group of researchers supportsa model of sudden, large-scale population replacement (e.g., Bar-Yosefet al. 1986; Mel-lars 1989; Stringer and Andrews 1988), while the other argues for continuity or in situevolution across a broad front (e.g., Clark and Lindly 1989a, 1989b; Wolpoff 1989). Forthe most part, proponents of these two positions do not cite different categories of factsor different sets of observations. Rather, the conflict arises from differing interpretationsof essentially the same bodies of data.Discussions of contrasts and similarities between the Middle and Upper Paleolithichave focused primarily on three phenomena: techniques of lithic blank production andtool forms, the ranges of prey species exploited, and frequencies of decorative or symbolicart (e.g., Chase and Dibble 1987; Clark 1987; Clark and Lindly 1989a, 1989b; Harrold1989; Lindly and Clark 1990; Mellars 1973, 1989; Orquera 1984; Straus 1983; White1982). Because it was once thought that they illustrated major discontinuities betweenthe periods, these three topics continue to dominate even the most recent syntheses. Po-MARYC. STINERis AdjunctAssistantProfessor,DepartmentofAnthropology,UniversityofNew Mexico,Albuquerque,NM87131.STEVENL.KUHNisAdjunctAssistantProfessor,DepartmentofAnthropology,UniversityofNewMexico,Albuquerque,NM 87131.

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lemical notions about the Middle and Upper Paleolithic are healthy and often produc-tive, but it is essential to recognize that most of the arguments about differences (or sim-ilarities) use information originally formulated post hoc to account for a limited range ofknown archeological facts. Researchers did not choose to study flake and blade frequen-cies or the number of mammalian species present in archeofaunas because of a theoreticalpredictionthat flake technologies or "unspecialized" hunting should be the ancestral con-dition. These were simply the only sets of facts that had been documented in comparableways for both periods. More recent observations show that Middle and Upper Paleolithicassemblages may contain quite similar ranges of species (e.g., Clark and Lindly1989b:644; Simek and Snyder 1988; Stiner 1992a) and/or comparable techniques ofblank production (Clark and Lindly 1989b:639; Straus and Heller 1988). While showingthe old generalizations to be inadequate, they provide no guarantee of the evolutionarysignificance of the facts themselves.

The major transitions in human evolution undoubtedly involved changes in technol-ogy, foraging behavior, and symbolic expression, but these are complex phenomena, andthere are many different ways to investigate them archeologically. Except for the histor-ical fact that substantial comparative data bases already exist, there is no overwhelmingreason for continuing to place so much weight on the usual triad of observations to tracethe evolution of Homosapiens. In investigating long-term evolutionary processes, the adap-tive significance of every archeological variable selected for study needs to be demon-strated rather than assumed.The problem can be illustrated most easily with a nonarcheological example: tigershave stripes and claws, leopards have spots and claws, and zebras have stripes andhooves. Looking only at these visually striking traits, we might conclude that zebras areabout as similar to tigers as are leopards. However, biologists would agree on the basisof real-life observations that the claws of a carnivore and the hooves of an ungulate aremuch more important than pelage for establishing the behavioral, physiological, andphylogenetic similarities among species. Regardless of how obvious and easily measuredstripes and spots may be, patterns of pigmentation are not all that significant at this levelof comparison.Today, most questions surrounding the "transition" and the relationship between ar-chaic and anatomically modern Homo sapiens concern the nature and rates of adaptivechange. Given that not all archeological facts are equally relevant to understanding adap-tive change or stasis, it makes sense to structure studies around evidence of prehistoric

behaviors or tendencies whose significance in this regard can be established indepen-dently of the archeological problem being studied. If evolution at the species and sub-species levels occurs as a result of selection upon variation, understanding any evolution-ary transition will also require documenting rangesof variationin adaptively linked behav-iors on either side of that transition.

It is worthwhile as well to consider the magnitude of differences that might reasonablybe expected to distinguish different subspecies of Homo sapiens. Behavioral adaptationsvary only subtly between modern subspecies of nonhuman taxa (sensuO'Brien and Mayr1991), and we should not expect huge discontinuities among late Pleistocene hominids inthe basic 4onstituents of behavior. Although more robust and apparently stronger thananatomically modern humans (e.g., Trinkaus 1983a, 1983b, 1986), Neandertals wouldhave been subject to similar physiological limitations. They would, for example, havefaced generally similar problems in getting food and "designing" artifacts to gain a phys-ical advantage, and they would have experienced similar needs for key nutrients (such asprotein and fat) in their diets. Differences between anatomically modern populations andtheir immediate predecessors are unlikely to have involved the addition or loss of any ofthe most basic components of foraging and technology. Instead, evolutionary changes aremost likely to have been manifest in how these components were integrated or organizedas problem-solving strategies for living.' Binford's (1987, 1990; see also Gamble 1986)work concerning the theoretical issues of how human adaptations vary, particularly in

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terms of economic structure and the rules of strategy combination, has been particularlystimulating in this regard.Our researchorientation does not necessarily share, however,all of the more specific characterizationsof behavioral differences between modern andarchaicHomosapiens.The research discussed in this article addresses the extent and causes of variation intechnology and subsistence on the Mousterian "side" of the transition in coastal west-central Italy. The study uses information about the relationships between food choicesand foraging strategies among modern nonhuman predatorsand hunter-gatherersas in-dependent referentsfor interpreting patterns in the Mousterian faunal data. If hominidpredatoryadaptations (admittedly only one part of the subsistence repertoire) changedsignificantlyduring the Upper Pleistocene, we should expect differencesof the magnitudethat normally distinguish other large predators in modern animal communities, regard-less of whether Neandertals and "moderns" prove to have been wholly, quasi-, or non-contemporaneoushominid populations.Trophically linked species, such as coexisting predators, may share an interest in thesame food species but exploit them differently(MacArthur 1968;MacArthurand Levins1967;Root 1975;Wiens 1977). Modern nonhuman predatorsrespond to the local abun-dances of prey within the size range dictated by their physiological adaptations (e.g.,Bertram 1979; Ewer 1973; Kruuk 1972; Schaller 1967, 1972), and species consumedwithin this range are more often than not poor indicators of adaptational differencesamong them (see, for example, Kitchener 1991:98-105). Of far greater interest are theways predators-in this case, variants of Homosapiens-might have obtained and used thesame food species, as manifest in ranging patterns, the prey age groups most commonlytargeted, processing and transport strategies, schedules of resource use, and the thresh-olds for switching among resources.Technologies can be seen as adaptations to both external factors and to the require-ments and limitations imposed by the subsistence system. What hominids did with toolsand the forms that tools took were heavily constrained by both the physical propertiesofthe human organism and the mechanics of stone fracture.Major evolutionary changes inhuman technologiesare more likely to have involved the role of technology within a largersystem than simple alterations in tool form and function. This is particularly true forMousterian and earlier technologies, since specialized stone tools for food procurementand processing were rare (Kuhn 1989a; Holdaway 1989; but cf. Shea 1989). Thus, therelationshipbetweentechnology and subsistence is here considered central to understand-ing adaptive variation in toolmaking behavior during the Upper Pleistocene. The lithicanalyses emphasize those variables expected to have been heavily influenced by foodsearchand procurement strategies. Patternsof association between lithic and faunal evi-dence are then used to assess the possible causes of technological variation.When examined from this novel perspective, the faunal and lithic data reveal consid-erablevariability in Mousterian tactics of tool manufacture and game use. While the ar-rays of tool types and mammalian prey species consumed stayed much the same, otheraspectsof technologyvaried in concert with finer details of animal exploitation. Together,the data sets document significant behavioral variation within the Mousterian as tradi-tionally defined and, perhaps, "vectored" change within one region of southern Europe.

Background and Study SampleThe animal bone and stone assemblages discussed in this article come from a series ofcave sites in Latium, a province on the Tyrrhenian (west-central) coast of Italy (Figure1). This area is characterizedby north-south trending mountains separated from the seain most places by flat, marshy coastal plains and basins. The caves yielding the Mous-terian assemblages lie very close to the modern shoreline and were never more than 10km from the sea during the period under discussion, depending on oscillations in globaltemperature.


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Figure 1Geographical locations of the four Mousterian cave sites: (B) Grotta Breuil, (G) GrottaGuattari, (M) Grotta dei Moscerini, and (S) Grotta di Sant'Agostino.Interest in the Middle Paleolithic of Latium began early in this century. The regiongained international attention in 1939 when vineyard workersaccidentally exposed theentranceof Grotta Guattari (Blanc 1939;Blanc and Segre 1953;Piperno 1976-77). Insidethe cave, a well-preservedNeandertal cranium, and later a mandible, were found exposedamid a jumble of rocks and mammal bones (see Stiner 1991a, White and Toth 1991 forrecent reanalysesof the Guattari finds). The discovery spurredan ambitious programofsurvey and excavation of coastal and inland caves throughout the 1940s and early 1950sby A. C. Blanc, A. G. Segre, L. Cardini, and other members of the Istituto Italiano diPaleontologia Umana (IIPU).The assemblages to be discussed are collections excavated by the IIPU researchgroup,along with more recently excavated material, from four caves (Table 1). Grotta Breuil(Taschini 1970; Bietti et al. 1988) and Grotta Guattari (Blanc and Segre 1953; Piperno1976-77; Taschini 1979) are located on Monte Circeo, and Grotta dei Moscerini (Vitag-liano 1984) and Grotta di Sant'Agostino (Laj-Pannocchia 1950;Tozzi 1970) are situatedin coastal cliffs near Gaeta, roughly 50 km to the south. All of the caves are relativelyshallow solution features in limestone bedrock and occur at similar elevations.The assemblages from the four caves collectively span a period beginning just before110,000years ago and ending around 35,000 years ago (Figure 2). Chronological assess-ments (Table 2) are based on electron-spin-resonance (ESR) dating of ungulate teeth,Uranium-series (U-series) dating of calcite flowstone layers (Schwarcz et al. 1990-91,

1991), and geochronological studies (e.g., Segre 1982, 1984; Blanc and Segre 1953). Atpresent, the radiometric dates are more informative about the ages of the assemblagesrelative to one another than they are for determining the absolute age of each. The studysample probably spans the portion of the classic deep-sea-core isotopic chronology(Shackleton and Opdyke 1973) between stages 5e-5d and stage 3. If correct, this timeframerepresentsa very general trend toward colder conditions, but incorporatesa num-ber of warm-cold oscillations.Because many of the assemblages are from excavations conducted 40 or 50 years ago,they are defined in terms of geological strata or artificial layers. It was necessary in thecase of Grotta dei Moscerini to combine materials from series of adjacent strata to pro-

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Table 1Origin and definition of Mousterian assemblages from the Italian cave sites.

Site Assemblage Origin/DefinitionGuattari G2 stratum 2G4 stratum 4G5 stratum 5Moscerinia M2 external strata 11-20M3 external strata 21-36M4 external strata 37-47M6 internal strata 3-4Breuil B3 stratum 3Br strata 11-15 (derived material)Sant'Agostino SO surfaceS1 arbitrarylevel 1S2 arbitrarylevel 2S3 arbitrarylevel 3

"Mosceriniproveniences M2, M3, M4, and M6 represent series of combined strata grouped onthe basis of uniform content and sediment composition. M2-M4 in fact contain over 30 culturallenses, which may or may not represent discrete occupational events.

duce assemblages of adequate size; groupings follow shifts in faunal contents and changesin sediment composition (see Table 1). In these cases, radiometric dates from multiplelevels are averaged to produce an estimated date for each strata group. Samples frommore recent (piece-plotted) excavations at Grotta Breuil are lumped into stratigraphi-cally defined groupings roughly equivalent to those derived from earlier excavations toensure comparability (see Kuhn 1990a, Stiner 1990a). None of the assemblages representsingle occupational "events"; instead, they are palimpsests of many such events. Whilethe level of resolution may not be ideal for detailed reconstruction of individual activities(Rigaud and Simek 1987), these long-term accumulations are entirely appropriate forinvestigating variation in behavioral tendencies at an evolutionary time scale.The subject lithic assemblages belong to a regional facies of the Mousterian called thePontinian, after the Pontine Plain, where first identified (Blanc 1937; Taschini 1972). ThePontinian is relatively invariant from the perspective of Bordes's (1961) Middle Paleo-lithic stone tool typology, exhibiting a strong predominance ofsidescrapers and generallylow Levallois indexes (Bietti 1980, 1982; Taschini 1972, 1979). As for the toolmakers,recent dating has shown that the Guattari I cranium was deposited no later than 51,000+ 3,000 years ago (Schwarcz et al. 1991), suggesting that all chipped stone depositedbefore this date is attributable to Neandertals. Human fossil associations for later Mous-terian assemblages are ambiguous, however. Ongoing excavations at Grotta Breuil haveyielded human remains likely to date after 50,000 B.P., but none is clearly diagnostic ofvariants of Homo sapiens (Bietti et al. 1988; Manzi and Passarello 1988).

MethodsThe data combined in this presentation were obtained from two freestanding studies,each of which employed a distinct set of analytical methods. Only the general approachesand the most important variables are described below. Details of the faunal and tech-nological analyses, data bases, sampling issues, and taphonomic considerations (in thecase of faunal remains), are presented elsewhere (see Kuhn 1990a, 1990b, 1991; Stiner1990a, 1990b, 1991a, 1991b).

[94,1992310


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ZU

30- U40-50-60-cn

< 70 -80-Y 80-90 -

100 -

110 -: U120 - I IM G S B

CAVE SITESFigure 2Chronological spans represented by the four cave strata sequences: (M) Moscerini, (G)Guattari,(S)Sant'Agostino,and (B)Breuil. Solid bars indicatespansdatedby ESRmethod,U indicates flowstone layer dated by U-series method (Schwarczet al. 1990-91, 1991),dashed bars indicate estimatedbut currentlyundatedsegments.

Methods andInterpretiveBasesfor theFaunal AnalysesThe faunal presentation concentrates on variables known to reflect two kinds of for-

aging "choices": anatomical-part transport as evident from patterns of skeletal represen-tation, and prey age selection evident from mortality patterns. The comparisons here areconfined to red and fallow deer (Cervuselaphus; Dama dama), the most common prey ex-ploited by Upper Pleistocene hominids in west-central Italy by any measure.2The faunal assemblages were deposited inside shelters, and thus represent cumulativeoutcomes of transport, rather than items discarded where food was obtained. The animalremains therefore are subsets drawn from larger resource pools, the general compositionof which can be modeled from modern analogs. In the case of body-part transportchoices, the pool is simply the ungulate body; foragers took certain parts from the car-casses of prey to new locations for processing and consumption, leaving other parts be-hind. Because scavenging could have been involved, carcass completeness at the pro-curement site is not taken for granted. Instead, the analyses assume only that the numberof parts taken away must have equaled or been less than what was present on carcasseswhen encountered by hominids.Prey age selection occurs in the context of much larger resource pools, the age struc-tures of living animal populations if hunted, or any subset thereof, if scavenged. In the

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Table 2Summary of radiometric dates for the Mousterian cave sites.

Site and Datingvertical provenience methoda Material AgebGrottadeiMoscerinicStratum 25 (M3) ESR red deer tooth 79,000 ?Stratum 26 (M3) ESR red deer tooth 67,000 -Stratum 26 (M3) ESR red deer tooth 81,000 +Stratum 33 (M3) ESR red deer tooth 89,000 -Stratum33 (M3) ESR red deer tooth 123,000 ?Stratum 35 (M3) ESR red deer tooth 66,000 ? -Stratum 38 (M4) ESR red deer tooth 96,000 ?Stratum38 (M4) ESR red deer tooth 106,000 -Stratum 39 (M4) ESR red deer tooth 96,000Stratum 39 (M4) ESR hippopotamus tooth 97,000 -GrottaGuattariGO(surface) U/Thd stalagmite 51,000 ? 3,000G4 ESR aurochs tooth 71,100 ? 2,760 (avr)G5 ESR aurochs tooth 77,500 + 9,500 (avr)GrottadiSant'AgostinoS1 ESR aurochs tooth 43,000 ? 9,000 (avr)S2 ESR aurochs tooth 53,000 ? 7,000 (avr)S3 ESR aurochs tooth 55,000 ? 11,000 (avr)Upper flowstone U/Thd stalagmite 112,000 ? 14,000Lower flowstone U/Thd stalagmite 120,000 + 15,000GrottaBreuilCapping flowstone U/Thd stalagmite 26,000 ? 12,000Stratum3 ESR aurochs and horse teeth 36,600 ? 2,700 (avr)

Source:Schwarcz et al. 1990-91, 1991.aESR = Electron-spin-resonance technique, errors calculated on the basis of multiple slices ofeach sample, using a Linear Uptake Model. U/Th = Uranium-series technique.b(avr) = Averaged value based on more than one sample.CStrataare those identified by Segre in 1949 for the exterior sequence of Grotta dei Moscerini.Designation in parentheses refers to the level group names used in this study. The cave entrancetodayis completely blockedby rubble fromhighway construction uphill of the site. For this reason,backgrounddose rates for the ESR analyses had to be estimated using soil cemented to the toothspecimens, and no errorranges are available.dNoncultural stratum.

case of scavenging, processes affecting carcass destruction and predators' abilities to lo-cate carcasses can be as important in determining age structure as the agencies of death(e.g., Blumenschine 1991; Stiner 1990b, 1991c, 1991d). More than one generation of fac-tors can influence "choice" in modern scavenging situations, greatly complicating effortsto interpret archeological cases (Stiner 1991c:9-10).Both transport patterns (anatomical representation) and prey age selection (mortalitypatterns) are closely tied to search and procurement strategies in nonhuman predators(Stiner 1990b, 1991b). From this perspective, hunting and scavenging are most interest-ing for their correspondences to fundamentally different ways of searching for food, eventhough each foraging mode can involve exactly the same prey species. Ungulate huntinginvolves capturing large, whole prey that often move about in groups while alive. In con-trast, scavenging in its more passive forms requires finding animals already dead, per-haps nutritionally depleted or ravaged, and often more dispersed on a landscape (e.g.,Houston 1979). Picking up items that are scattered and immobile makes passive scaveng-

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ing more like gathering than hunting, both in the spatial characteristics of these foodsourcesand the potential average returnsgained from them.Efforts to distinguish the faunal products of scavenging and hunting are potentiallysubjectto several sources of ambiguity, however. This point merits additional discussion,since a greatdeal hinges on how the faunas in this study areinterpreted.Generally speak-ing, both kinds of behavior can take more than one form, depending on the nature ofopportunity and details of the subsistence system in use. Moreover, some patterns thathunting and scavenging behaviors impose on faunal assemblages overlap (compare, forexample, results obtained by Blumenschine 1986, 1991;Bunn, Bartram,and Kroll 1988;Haynes 1980; O'Connell, Hawkes, and Blurton Jones 1988a, 1988b; Stiner 1990b,1991d).The general phenomenon known as "scavenging" embodies diverse alternatives, oneor more of which are practiced by nearly every carnivorousorganism. Scavenging of nat-ural deaths or predator kills can be a relatively passive activity nested within other for-aging concerns, or one involving aggressive displacement of carnivore(s) by humans wellbefore the carcass would have been discarded voluntarily. The Hadza, for example, fre-quently engage in aggressive scavenging (e.g., O'Connell, Hawkes, and BlurtonJones1988a), and tense moments between lions and people have been noted. It is important toappreciate that many or most of these contests occur during forays that anticipated(amongother things) hunting activity, and some men equipped with weapons were pres-ent. Less ambitious scavenging schemes also exist in nature, usually involving unpre-dictable access to food and lower returnsper find, yet they are profitableunder the rightconditions. The limits and strengths of each faunal variable used for interpreting Mous-terianforagingbehaviors are discussed below in light of these issues.Measures of anatomical representationare based on minimum number of skeletal ele-ment (MNE) counts for bone.Animal bones in archeological assemblages are usuallysmashed to bits for reasons beginning more or less with the processing habits of people,followed by any of a wide arrayof other destructive agencies. In using MNE, zooarcheol-ogists try to reconstruct the full, minimum array of whole animal bones on the basis ofmany fragments.This is possible because most kinds of whole bones have distinctive fea-tures ("portions" or "landmarks") that inform about what skeletal element the frag-ments came from, as well as how many whole bones the fragments represent. MNE isdeterminedfrom the most common portion or landmarkof each skeletal element and, inthis study, represents the sum of right and left sides for elements that naturally occur inpairs. The counts for limb bones use unique landmarks on or near the articular ends,3and any of a variety of distinctive bony features of the cranium and mandible (e.g., pe-trous,occipital condyle; incisive, mandibularcondyle;or bony architectureof the maxillaor mandiblebehind the cheek tooth rowifrelativelycomplete;see Stiner 1991b:460-462).Teeth are not used for the MNE determinations, because they can distort head countsrelativetopostcrania wherever preservation conditions are less than ideal. This point isvery important for understanding the anatomical comparisons, where comparability inthe face of differentialpreservationis ajust concern.Mortality patterns, in contrast, are based on teeth; specifically, tooth eruption andwear data for the lower fourth premolar sequence. The aging criteria for occlusal wear indeer are adapted from Payne's (1973) system for sheep and goats, and Lowe's (1967)system for deer. Only two adult age categories are considered here, prime adults and oldadults (see Stiner 1990b:311-313).Findings on species consumed by Middle Paleolithic hominids, and seasonality for thesame arrayof faunas, are summarized from other sources (Stiner 1990a, 1992a) as per-tinent to the discussion that follows. While not covered in this presentation, taphonomicanalyses were essential for establishing the origin of each faunal assemblage (Stiner1990a, 1991a). Cases attributable entirely or largely to the actions of nonhuman carni-vores also occur in some of the subject caves-a common situation in the Paleolithic-and are not included in the sample.4 The state of preservationfor the cave faunas is very



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good on the whole, because they were deposited in calcium-rich limestone formations.While some postdepositional bone destruction seems inevitable, particularlyfor skeletalelementscomposed mainly of trabecular (spongy) tissue (e.g., Lyman 1985), in situ boneattrition does not explain most of the variation in anatomical part representationamongthe assemblages (Stiner 1990a, 1991b). Potential biases due to differential bone decom-position are further circumvented by considering only the more durable partsof the bonyskeleton-the head and limb elements.To simplifythe presentation, patterns of variation in the archeofaunas are summarizedin terms of only three derived variables. Each is an index suitable for the interpretationof series of assemblages, but probably not for isolated archeological cases. The first vari-able is the ratio tMNE/MNI, which serves as a rough index of anatomical completenessfordeer. It is obtained by dividing the totalnumber of skeletal elements (tMNE) by theminimum number of individual animals (MNI). The ratio represents the average quan-tity of bony parts transported to shelters per carcass source. Variation in this indexloosely corresponds to differences in transported returns from hunting (high values) asopposed to scavenging (low values) in carnivore and human control cases (Stiner1990a:518-576).The second anatomical variable, called head MNE/limb MNE, is calculated by divid-ing the total minimum number of head parts by the total minimum number of limbs forred and fallow deer in each assemblage. Limb MNE includes all substantial leg elementsexcept phalanges, carpals, and tarsals other than the calcaneum and astragalus. HeadMNE includes antler as well as cranium and mandible elements, although antler is rarein the hominid-collected faunas (Stiner 1991a:112). Crania and mandibles occur inroughly equal proportions,a fact whose significance will become apparent in discussion.In effect,head MNE/limb MNE concerns relativelylarge bones, those which in principlecould representattractivelysized transportableparcels.The head MNE/limb MNE ratioaccountsfor much of the variation in anatomical representationin the Mousterian shelterfaunas (Stiner 1991b:463-465). In controlled comparisonsof other predators, the ratio isa strong predictor of the general way that prey are most commonly obtained: huntingresults in a more or less balanced anatomical ratio relative to the complete living anat-omy, while scavenging tends to lead to head- and/or horn-dominatedassemblages (Stiner1991b, 1992b).The interpretation that head-dominated faunas represent mainly scavenged sourcesmay seem counterintuitive, initially, because it is based on evidence from modern non-human predators rather than contemporary humans (Stiner 1990a, 1991b, 1991d,1992b). In fact, no modern human-generated analogs could be found for the stronglyhead-dominantpatternobserved in the Italian Mousterian. Many aspects of Middle Pa-leolithic recordsperplex archeologistsfamiliar with recent human practices, however, sofinding enigmatic faunas in the Italian Mousterian cannot be greeted with complete sur-prise or unqualified suspicion. Systematic consideration of the potential impacts of dif-ferential preservation and MNE counting methods on the study sample leave far toomuch unexplained. Likewise, taphonomic analyses firmly exclude nonhuman agencies,such as wolves and spotted hyenas, as significant bone collectors in all of the cases (Stiner1990a),except GuattariG2 and possibly G4-5 (Stiner 1991a). In contrast to the apparentlackof analogous situations for "head-collecting"among modern-dayhumans, threespe-cies of scavengingsocial carnivore (striped, brown, and spotted hyenas) frequentlycreatehead- and/or horn-dominated anatomical biases for medium-sized ungulate prey in mod-ern dens. The head-dominated pattern produced by nonhuman predators in the contextof scavenging is best explained by propertiesof ungulate hard- and soft-tissue anatomy,a point developed in a later section.The third ratio, called PRIME/OLD, refers to mortality patterns in deer, specificallythe relative frequencies of prime and old adults. These adult age categories are definedon the basis of life-history characteristics and associated physiological changes in deerand other ungulates (Stiner 1990b:308-313). The ratio registers two potential directions

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that age biases often take in predator-collectedfaunas. Comparative research indicatesthat this ratio is very responsive to the relative emphasis on ambush hunting, whichamong modern humans frequentlyresults in a prime-dominant mortality pattern, as op-posed to scavenging, which among carnivorescan result in elevated numbers of old-agedanimals (Stiner 1990b, 1991d).Not all foragingtactics leave strong or mutually exclusive signatureson mortalitypat-terns in prey death assemblages. This study is fortunate because the old-biased age pat-tern observed in some Italian Mousterian cases matches a relatively outstanding "tag"of scavenging in modern ecosystems, probably passive scavenging at that (Stiner1990b:328-329). The old-biased pattern may owe its distinctive form to at least two gen-erations of processes that commonly occur in succession: the prevailingcause(s) of deathand what happens to carcassesduring the interlude between death and when foragersarenormally able to locate them. Whether originating from predator kills or nonviolentcauses of death, the carcass pool appears to have begun with a situation of U-shapedmortality, the most common family of death patterns in nature (reviewed in Stiner1990b:321-329). U-shaped mortality patterns consist mainly ofjuvenile and old individ-uals, meanwhile being very poor in prime adults relative to their expected abundance innaturalliving populations. Such mortalitypatterns frequentlyundergofurtherdistortion,however, by losing much of the juvenile component. Small, immature bodies are moreprone to rapid decay, more complete consumption by primary feeders (e.g., Blumen-schine 1986, 1991;Stiner, unpublished study of mountain lion kills and scats, 1985), andthey are more difficult to locate (Schaller 1967, 1972). For all of these reasons,juvenilestend to drop out of the potential array of scavengeable carcasses: in the situation of U-shaped mortality, this leaves an inordinate proportion of old-aged adults (cf. Klein1989:377).The phenomenon generally known as "hunting" is no less diverse than scavenging,nor should it be any easier to demonstrate from faunal evidence in principle. Mortalityanalyses of Paleolithic faunas are a case in point. In this study, attributions of deer mor-tality patterns to hominid hunting behaviors are greatly aided by the fact that the pat-ternsoften take a relativelydistinct, prime-dominantform. This cannot be explained bydifferentialpreservation favoring adult teeth over those of juveniles, as prey mortalitypatternsassociated with hyena occupations in some of the very same caves are stronglybiased towardjuvenile prey (Stiner 1990a, 1990b).While not the only age pattern that ever resulted fromhunting, prime-dominant mor-tality is regularlyassociated with modern human hunting practices (Stiner 1990b, 199ld;contra Klein 1982a, 1982b). The pattern varies from a slight to great overabundance ofprime-agedanimals in relation to their live abundance. On the whole, prime-dominantprey age selection represents a long-established tendency in humans, generally settingthem apart from other large predators (Stiner 1990b:329-341). While targeting the mostproductiveage groups in prey populations, the relationship is potentially stable in situ-ations involvingcervidsand bovids, unless or until human subsistence no longerdependsin any way on the targeted species (Stiner 1991Id:180-184).Even with the benefit of controlled associations between transport patterns, mortalitypatterns, and foraging behaviors, it would not be appropriate to assume that each andevery whole animal procured by Mousterian hominids had to have been hunted, norevery head from an old animal scavenged. The interpretations are probabilistic assess-ments about what people usually did when presentedwith a general set of circumstances.The study seeks information about foraging practices at the level of interactions betweenpredatorand prey populations, and only the cumulative patterns representing repeatedbehaviorsare meaningful.MethodsandInterpretiveBasesfor theLithicAnalyses

The analyses of the Mousterian industries address some of the potential links betweentechnology and subsistence, particularly connections involving the shared time and en-



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316 AMERICANANTHROPOLOGIST [94,1992

ergy budget. Among modern hunter-gatherers, the timing of subsistence activities andmovement from one place to another determine the scheduling of both the needs for toolsand the opportunities to make them. Any number of technological strategies may be em-ployed to manipulate the utilityof artifacts or raw materials, thereby bridging the spatialand temporal discontinuities between need and opportunity (e.g., Binford 1977, 1979;Bleed 1986; Kuhn 1989b; Nelson 1991; Shott 1986; Torrence 1983, 1989). Tactics forproducingflakes and blanks from cores, the resharpeningor renewal of stone tools, andthe transportof tools and raw material all play a role in determining how long tools andraw materials will last. Studies of these variables should therefore provide informationabout the contingencies faced by hominids in making tools available when and wherethey were needed.Coastal Latium presents an especially interesting natural laboratory for the study ofMousterian technology. The only workable stone found within at least 40-50 km of thecoastal caves consists of small flint pebbles, normally less than 10 cm in maximum di-mension. The pebbles occur in scattered, active and fossil beach deposits on the coastalplains. Such small and dispersed raw materials would have placed a special premium ontactics for stretching raw materials and maximizing tool size.The lithic analyses here focus on two dimensions of the technological record: (1) theintensity or duration of tool use and renewal; and (2) core reduction techniques, the tac-tics for manufacturing flakes and tool blanks. Very different analytical approaches arerequiredto monitor these disparate phenomena. The intensities of tool use and reductionare measured using two variables. One is the frequency of retouched tools relative tounmodified but potentially usable flakes (>2.0 cm in length), a measure that has beenemployed in other studies (e.g., Rolland 1981; Rolland and Dibble 1990). The other isan experimentallyderived index of scraperreduction (Kuhn 1990b). This index monitorshow much of a tool blank has been removed by previous retouch, based on the angle ofthe retouched edge and the extension of retouch scars relative to the medial thickness ofthe blank.The analyses of core reduction techniques use a differentapproach and requiregreaterexplanation. Two techniques for making flakes and tool blanks, here termed centripetalandplatformcore reduction (Figure 3), were very important in the Pontinian. There are,of course, numerous variations on the two basic ways of producing flakes from pebblecores, and they may sometimes overlap (e.g., Bietti, Rossetti, and Zanzi 1989; Kuhn1990a;Laj-Pannocchia1950). Nonetheless, there is ample reason to believe that these aredistinct ways of exploiting raw material, as discussed below.The centripetal technique, schematized in Figure 3(a), is typical of the Mousterian ingeneral (Bordes 1961:26-27; Tixier, Inizian, and Roche 1980:43), although there aremany variants (e.g., Boeda 1986, 1988;Crew 1975; Geneste 1985). Centripetal core re-duction in west-central Italy involved striking flakes from around the perimeter of a flatround pebble, gradually spiraling toward the center by rotating the core with each newblow. For the purposeof this study, evidence of Levallois technique with centripetal corepreparation (representedlargely by "atypical" products) is combined with more gener-alized centripetal reduction.

Platform core reduction techniques (schematized in Figure 3[b,c]) differed from cen-tripetal techniques in that flakes were removed parallel to the long axis of a core from astrikingplatform,or from two opposing platforms, located at the end(s) of the piece. Oc-casionally, one or two flakes were also struck from the side of a core as it neared the endof its useful life (Figure3[c]). These platformcore techniques are similar but not identicalto classic prismatic blade core techniques of the Upper Paleolithic. However, the Mous-terianversions are decidedly less complex, beginning only with platform preparation atone or both ends of a pebble. The "crested blade" technique, in which a ridge is createdalong the length of the core to guide the initial removals, was not employed in the Pon-tinian Mousterian. Moreover, flakes detached from Pontinian platform cores rarely can


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a b cCentripetal Platform

Figure 3Schematized (a) centripetal and (b, c) platform core reduction techniques.

be classified as blades, primarily because small pebbles make it very difficult to manu-facture flakes that are twice as long as they are wide.Because cores preserve the traces of last removals, and earlier scars are preserved onthe backs of flakes detached subsequently, it is possible to monitor the use of differentcore reduction techniques by examining the orientations of flake scars both on cores and-on the dorsal faces of many flakes and tools. Flakes produced from centripetal cores tendto exhibit dorsal scars originating from several directions, often orthogonal to one another(Figure 3[a]). In contrast, flakes detached from platform cores generally have dorsalscars running parallel to one another along the long axis of the flake (Figure 3[b,c]): whenpebbles are used, flakes often retain remnant cortex along one edge.5The two basic core reduction techniques, platform and centripetal, have somewhatdifferent economic properties. Platform core reduction appears to maximize the numberofflakes produced per pebble core, while centripetal techniques maximize the sizes of flakesand tool blanks. To illustrate this difference, Figure 4 compares the number of large flakesand tools attributable to each technique relative to the number of cores in the total Mous-terian sample. The ratio of flakes and tools to cores is much higher for the platform re-duction technique; even when all of the unattributed pieces (specimens that cannot beassigned to any mode of manufacture) are added to the identifiable products of centripe-tal reduction, the ratio of flakes and tools per core is still much lower than that for plat-form cores. It is possible that the proportions of total products that can be reliably at-tributed to either basic technique varies between them, and the actual contrasts in pro-ductivity may have been somewhat less marked (Kuhn 1990a:232). On the other hand,the flakes produced via centripetal core reduction techniques tend to be larger in thestudy sample, with more sharp edge per piece than those made from platform cores. Theflakes and blanks attributed to centripetal reduction are approximately 25% longer inmaximum dimension and roughly 1.5 to 2 times larger in plan view area than those at-tributed to platform cores (Kuhn 1990a:226).

StinerandKuhn] 317


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AAMERICANANTHROPOLOGIST

flakes+ toolscores

2- ...... .........

1-

. L i .,Centripetal Platform

.... . unattributable pieces

Figure4Comparisonof the relative efficiencies (flakes + tools/cores) of centripetaland platformreductiontechniques in Mousterianlithic assemblages.Variation in the Faunas

In the following discussion, 12 hominid-generatedfaunal assemblages are arrangedinchronologicalorder to illustrate patterns of change over time. Assemblages at the early(M4, M3, G2, and G5-G4) and late (S1-3, B3) ends of the sequence are dated by ESRand U-series methods, whereas fourassemblages in the middle (M2, M6, SO,and Br) areranked on the basis of stratigraphiccriteriaonly (see Tables 1 and 2). Because we wantto consider both radiometricallydated and stratigraphicallyordered assemblages, andbecause the ESR dating technique remains somewhat experimental (Griin 1988), thechronologicalsequence is treated as a simple rank ordering. For ease of discussion, theassemblages are often referred to as dating before or after 55,000 B.P., the boundary ofwhich is markedon the bar graphs.Figure 5 shows a trend in the average amount of transported food returns per red orfallow deer carcass, using the bone-based ratio of anatomical completeness (tMNE/MNI). In the earlierhalf of the chronological scale, the quantity of bony parts carried tosheltersby hominids per procurementevent is relatively low. The values increase signif-icantly toward the recent end of the scale. The changes appear gradual overall, althougha sharp increase may be indicated around the 55,000-year boundary.The second trend, shown in Figure 6, traces changes in the proportion of deer headparts to limb parts transported to the caves by hominids, using the head MNE/limbMNE ratio based on bone. The expected "balanced" value for the complete deer skeleton(0.30) is indicated on they-axis. Head parts are extraordinarily abundant relative tolimbs in assemblages dating to before55,000 years ago. The ratio declines gradually earlyon and then stabilizes around the expected value for relatively complete prey.A third trend (Figure 7) is found in the mortality patterns for deer, based on tootheruption and wear. The expected value shown on they-axis (2.14) refers to the ratio ofprime-aged to old-aged adults in an idealized, stable live-prey population (from Stiner1990b;see also Lyman 1987). The expected value on the graph would represent "non-selective"harvesting by hominids, regardlessofjust how the animals wereobtained. Deermortality patterns are significantly biased toward old-aged prey in all cases dating tobefore55,000 B.P., and the bias lies well outside the range of normal variation displayedby live populations (Stiner 1990b, 1991d). The cases dating to just after 55,000 showessentiallynonselectiveprocurement,while values forthe most recent Mousterian assem-

318 [94, 1992


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StinerandKuhn] ADAPTIIZEVARIATIONINMIDDLEPALEOLITHICITALY 3l9

35

tMNE 25-MNI

' S - ,,1,|,l1llllM4 M3 G45 M2 M6 G2 Br S3 S2 S1 S0 B3110 KY 55 KY 35 KY

ASSEMBLAGEFigure 5Anatomicalcompleteness index values (tMNE/MNI)for redandfallow deer in Mousterianfaunalassemblages, arrangedin chronologicalorder (see Table 1 for assemblage designa-tions).

blages(ca.40,000-35,000B.P.) arestronglybiasedtowardprimeadultprey.Theprime-dominantcasesexceedtherangeencompassedbynormalvariationin naturallivingpop-ulations,evenwhenphasesofgrowthanddeclinearetakenintoaccount.Rankordercorrelations(Spearman'srS)ofthe threeratiosagainstchronologicalrank-ings evaluate the statisticalsignificanceof the trends.Two alternativechronologicalrankingsareused.Table3(a) listscorrelationcoeflicientsfor all datedand undatedas-semblages.Variationin eachof the threefaunalratiosis highlycorrelatedwith assem-blage age at the .01 level of probability.Not all assemblagescould be datedby radio-metrictechniques,however,andproperchronologicalorderingis veryimportantto theconclusions.In orderto check thisproblem,Table3(b) lists correlationcoeflicientsforradiometricallydatedassemblagesonly, thereforeexcludingM2, M6, S0, andBr.Cor-relationsremainsignificantfor thissubset,althoughthevalue fortMNE/MNIdrops tothe.05level ofprobability.The relationship between two of the faunal ratios, head MNE/limb MNE andPRIME/OLD,is evaluatedanotherway in Figure8. A log-logregressionof the ratiosdocumentsa strongnegativerelationshipbetweenthe occurrenceof head-biasesandprime-agedprey (r = -. 733,r2 _ .54, .02< P < .01,N = 10). Becausethefirstvariableis basedexclusivelyon bonematerial,whereasthesecondis basedexclusivelyon moreresistanttoothmaterial,thecomparisonservesas a partialcheckagainstpossiblebiasescreatedby differentialpreservationamongskeletaltissueclasses.Bones and teeth tellsimilarstoriesabout the contextin which foodwasobtained.The B3 assemblagefromGrottaBreuilis an outlierto thegraphand,while not includedin theregressioncalcu-lation,falls at theappropriateextremeof thedifferencesas described.The casefromG2of GrottaGuattariis excludedfromconsiderationbecauseof insufEcientage data andbecauseits taphonomicoriginis moreambiguousthan all of theothercases. The assem-blagesalign in a mannerconsistentwiththe generalchronologicalorderdefinedprevi-ously,althoughcasesdatingafter 55,000B.P. display less variationon the anatomicalaxis (headMNE/limbMNE).


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320

SO B3M4 M3 G*5 M2 M6110 KY 55 KY 35 KY

ASSEMBLAGEFigure 6Ratios of head to limb parts (head MNE/limb MNE) for red and fallow deer in Mousterian

faunal assemblages, arranged in chronological order.

PRIMEOLD

M4 M3 G45 M2 M6 G2 Br S3 S2 S1 SO B335 KY110 KY 55 KY

ASSEM BLAGEFigure 7Ratios of prime-aged to old-aged adult (PRIME/OLD)red and fallow deer in Mousterianfaunal assemblages, arranged in chronological order.

head MNEIimb MNE

l-W-F-G2 Br S3 S2 S1


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Faunal variable N rSa Pa. All assemblagestMNE/MNI 12 -0 744 < .01Head MNE/limb MNE 12 0.818 < .01PRIME/OLD 1l -0.773 < .01b. Datedassemblagesonly (excludesM2, M6, SO,andBr)tMNE/MNI 8 -0.714 = .05Head MNE/limb MNE 8 0.881 < .01PRIME/OLD 7 -0.905 < .01

aValues of rSindicate correlation with increasing age.

Stinerand Kuhn] ADAPTIVEVARIATIONIN MIDDLEPALEOLITHIKITALY321

Table3Spearmanrankordercorrelation(rS)offaunalvariablesagainstincreasingassemblageage.

*>55K Years* < 55 K Years

PRIMEOLD

head MNElimb MNE

Figure 8Log-log regression of head MNE/limb MNE based on bone data, and PRIME/OLDmortal-ity based on tooth data for red and fallow deer in Mousterian faunal assemblages.

In sum, low returnsfor transportedfood (low tMNE/MNI values) and head-domi-natedanatomicalpatterns(highheadMNE/limbMNE values)in the earlierMousterianfaunasfromGrottadei MosceriniandGrottaGuattariindicatea heavy relianceon scav-engingof deer carcasses.Conversely,highertransportedreturnsand higher (balanced)frequenciesof limb to head elementsin the later Mousterianfaunas from Grotta diSant'Agostinoand GrottaBreuilindicatea strongemphasison huntingof the same preyspecies.Variationin the PRIME/OLD mortalityratio likewisepoints to differingreli-ance on scavenging(old-biasedpatterns)and hunting (prime-dominantpatterns).Some additionalfeaturesof the Mousterianfaunasare of interestwith regardto thetrendsillustratedhere, as well-as to the discussionsthat follow.Data on juvenile tooth


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322 A MERICANANTHROPOLOCIST[94, 1992

eruptionand wearand the absenceof antlersuggestthat Mousterianoccupationsdatingbefore55,000yearsago mayhave centeredon springand/orearlysummer,althoughtheyprobablywerenot restrictedto theseseasons(Stinerl990a:660-721).Laterassemblagesfrom Sant'Agostinoand Breuil more clearly representfall throughlate-winteroccupa-tions. The seasonalitydata are subject to considerableerror,for reasonsof sampling,latitude,and environment.These resultsnonethelessintroducethe possibilitythat sea-sonal factors(as opposed to diachronicchanges) caused differencesamong the faunas,althoughit mustalso be recognizedthat each site representsthousandsof yearsof debrisaccumulation.In markedcontrastto other propertiesof the Mousterianfaunas,the arraysand rela-tive frequenciesof mammalian species consumedby hominidsdid not differsignificantlyamongsites or within the overallsequence(Stiner l990a:341-353, 1992a).Moreover,avarietyof statisticalanalysesrevealno significantdifferencesin the relativedietaryem-phaseson mammalianspeciesconsumedat caves by Middle Paleolithichominids,spot-ted hyenas,and (later)Upper Paleolithichumansin this regionof Italy. Such uniformitywithin the Mousteriansample, and betweenit and cave faunas createdby other largesocial predatorsin the same area, suggests that natural abundanceexerted an over-whelminginfluenceon prey species choicefor all predatorsin coastal Italy. Small-scaleexploitationof marineshellfishand aquatictortoisesby hominidsis evidencedin the pre-55,000B.P. assemblagesof Grottadei Moscerini,but not at the otherthreecaves. Shell-fish and tortoiseremainsoccur in generalassociationwith (inferred)scavengingof un-gulates,but frequenciesof shellsand mammalbonesdo not predictone anotherverywellwithinthe long stratasequence(30 fine layers)of this cave (Stinerl990a:281-283).

Variation in TechnologyThe strongestpatternsof technologicalvariationamongthe Mousterianlithic assem-blagesinvolvethe use of differentreductiontechniques.Figure9(a) comparesthe ratiosof platformcoresto centripetalcoresin the subjectassemblages,and Figure9(b) showsthe ratiosof the tools and flakesattributableto these two differentreductiontactics. It isclearthat the use of platformcore techniquesincreasedover time in the Pontiniansitesat the expenseof centripetalcorereduction.It is less certainwhetherthe trendrepresentsa gradualor suddenincreasein the use of platformcore techniquesafter55,000 B.P.Figure9 also shows that core reductiontechniquesvariedfromsite to site, as well as

throughtime.Sincethe two basictechniqueswouldbe moreconvenientlypracticedusingdifferentpebbleshapes (see Figure3), it is conceivablethat the observedvariationmightsimplyrepresenta highly expedientresponseby hominidsto the shapesof pebblesavail-able in the immediatevicinityof each shelter.Alternatively,becauseplatformcores ap-pearto yield moreflakesper core, it is possiblethat this techniquecould have been usedpreferentiallyat sites wherepebbleswere relativelyscarce.Raw materialqualitiescertainlyinfluencedmany of the generalcharacteristicsof thePontinianassemblages,particularlytool sizes (Bordes1968:l l 9; Taschini 1970:70-74) .The proximityof rawmaterialsourcesalso seemsto havedeterminedthe extentto whichcoreswereusedup, regardlessof whichtechniquewas morecommon(Kuhn 1991).How-ever, studiesof interassemblagevariationin the frequenciesof differentkinds of flakesand coresindicatethat local variationin the shapesand/or abundancesof raw materialsdoesnotaccount for variationin the frequenciesof the alternativecore-reductiontech-niquesindependentlyofthe time element(Kuhn l990a:436-439).Changesover timein tacticsof flakeand tool manufacturein coastalwest-centralItalywereaccompaniedby declinesin both the overallintensityof raw materialexploitationand the extent to which individual tools were used up. There is a markeddecreasethroughtime in the frequenciesof retouchon flakesand blankslargerthan 2 cm in max-imumdimension(Figure10). The medianreductionindex valuesfor single-edgedtoolsalso decreasewith time (Figure l l ), indicatingthat tools were resharpenedless often in


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Stinerand Kuhnl ADAPTI1ZEVARIATIONINMIDDLEPALEOLITHICITALY 323

CORES: 'Platform

Centri peta I

PRODUCTS:Plat form

Centripetal

a

b

M4 M3 G5 G4 M2 M6 G2 Br S3 S2 S1 B355 KY110KY 35 KY

ASSEM BLAGE

Figure 9Changing emphases on centripetal and platform reduction techniques in the Mousterianlithic assemblages, as measured by (a) core ratios and (b) ratios of core products. Assem-blages are arranged in chronological order.

the laterassemblages.Since sedimentswere screenedeven in the firstmajorexcavationsundertakenin this region,variationin the frequenciesof unretouchedflakes (all largerthan 2 cm) is not due to differentialrecovery.Nor can this patternbe attributedto dif-ferencesin the availabilityof lithic raw materialsfromone cave to the next;assemblagesfromthe site located closest to a sourceof pebbles (GrottaGuattari)exhibit relativelyhigh frequenciesof retouchand high levels of scraperreduction.Rankordercorrelations(Spearman'srS)betweeneach of the technologicalvariablesand assemblageage confirmthe statisticalsignificanceof trendsin the lithic data. Table4(a) lists correlationcoefEcientsfor dated and undatedlithic assemblages,showingthatfourof the variablesare highly correlatedwith time at the .01 level of probability.Thecoefficientfor a fifth variable(% withoutpebble cortex)is weaker,but still significant.Table4(b) lists correlationcoefficientsforthe datedassemblagesonly (excludesM2, M6,


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A AFERICANANTHROPOLOCIST[94,

324

80

PERCEN TW ITH

RETOUCH{> 2.0 cm}

60

4020- pw

M4 M3 G5 G4 M2 M6 G2 Br S3 S2 S1 B3110 KY 55 KY 35 KY

ASSEMBLAGEFigure 10Frequenciesof retouch on flakes and blanks largerthan 2.0 cm in Mousterianlithic assem-blages, arrangedin chronological order.

.

* . :* . * v

* * v: v.

: @

* @

MED IANREDUCTION 0.6-

INDEXVALUE

0.5-

0.4-

rs o2 | | l l l l l l l l l l

M4 M3 G5 G4 M2 M6 G2 Br S3 S2 S1 B355 KY110 KY 35 KY

ASSEMBLAGEFigure 11Median reduction index values for single-edged tools in Mousterian lithic assemblages,arrangedin chronological order.

SO,and Br). The relationships of the five variables to time remain significant in spite ofthe smaller arrayof assemblages considered, although the probabilities drop to the .05level for most.Another, less clearly defined pattern in the lithic data deserves mention. The use ofderived (and thereforescattered) pebble raw materials in the subject assemblages makes


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Lithievariable N rSa Pa. All assemblagesPlatform/centripetalcores 12 -0.769 < .01Platform/centripetalproducts 11 -0.882 < .01%retouchedpieces 11 0.800 < .01% withoutpebblecortex 12 0.634 < .05Medianreductionindex 12 0.786 < .01b. Datedassemblagesonly (excludesM2, M6, SO,andBr)Platform/centripetalcores 9 -0.733 < .05Platform/centripetalproducts 8 -0.786 < .05% retouchedpieces 8 0.833 < .01%withoutpebblecortex 9 0.650 < .05Medianreductionindex 9 0.778 < .05

aValuesofrjindicatecorrelationwith increasingage.

it difficultto obtain directmeasuresof the distancesartifactswere moved aroundthelandscape.However,alternativeanalysesof variationinartifactsizes, rawmaterialqual-ities, andthe relativefrequenciesof flakes,tools, and coressuggest that the frequenciesofartifactstransportedfromsourcesof largerraw materials(eitheroutsideor withinthecoastalzone) decreaseover time(Kuhn l990a:325-330;1991). Largesharp-edgedtoolsmadeon blanksfromcentripetalor Levalloiscores appearto have been the itemsmostfrequentlymovedaroundthe "Pontinian"area, as has beennoted for someMousteriancaseselsewherein Europe(e.g.,Geneste 1989;Otte 1991).In sum, changesthroughtime in the use of centripetalversusplatformcorereductiontechniquescorrespondto differentartifact"life histories"in the PontinianMousterian.Anemphasison the productionof relativelylargeflakesthroughcentripetalcorereduc-tioninthe earlierassemblagesis associatedwith frequentretouchand fairlyextensiveuseandrenewalof tools. The emphasison large flakeproductionprobablyassociateswithfairlyhigh frequenciesof transportedartifactsas well. The increasinguse of more"eco-nomical"platformcore techniqueswith time in west-centralItaly is actuallyaccom-paniedby less intensiveexploitationof tools and flakes,andby decliningfrequenciesoftransportedor "exotic"artifacts(Kuhn l 990a:325-330).The Connections between Subsistence and Technology

The waysthat Mousterianhominidsmadeand usedstonetoolsat the fourcavescovar-ied withdifferencesin animalexploitation,and significanttrends are documentedforbothdata sets. Shifts in animal procurementand use aresomewhatmore strikingandappearmore continuousthan the technologicalshifts. The convergenttrendsand theirbehavioralimplicationsare bestsummarizedby comparingassemblagesdepositedbeforeand afterthe 55,000-yearjuncture.Theoldergroupof assemblagessuggestsprocurementofungulateslargelybyscaveng-ing,basedon old-biasedmortalitypatternsand selectivetransportof head partsto shel-ters. All head bone cavitieshad been opened while fresh to extracttheir contents.Un-gulateprocurementactivitiesappearto have centeredonspring,althoughtheyprobablywerenotentirelyconfinedto thatseason. "Head-collecting"is generallyassociatedwithlow-levelexploitationof marineshellfishand aquatictortoisesat Grottadei Moscerini.Relativelylarge tools and flakesproducedvia centripetalcore reductiontechniquesare

StinerandKuhnl ADAPTIVEVARIATIONINMIDDLEPALEOLITHICBITALY 325

Table4Spearmanrankordercorrelation(rS)of lithic technologyvariablesagainstincreasingas-semblageage.


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326 AMERIC}ANANTHROPOLOCIST[94,

1992

morecommonin theassociatedindustries,andlithicartifactsexhibitevidenceof moreintense,prolongeduseandgreaterfrequenciesof transport.Thefaunasdatingafter55,000yearsagoareverydifferentfromthe olderones. U-n-gulatemortalitypatternsresembleeitherthestructureoflivingpreypopulations

or,morecommonly,arebiasedin favorofprime-agedadults.Substantialquantitiesof foodweretransportedto sheltersfromeachcarcassobtained,andtheremainsrepresent"meaty"ornearlycompletearraysofbodyparts.Allofthelimbboneswerethoroughlyprocessedformarrow,andsofttissuesencasedin theheadboneswereextractedaswell.The char-acteristicsof thesefaunasareentirelyconsistentwiththeexpectationsforprocurementbyhunting,as determinedfromthe controlstudies(Stinerl990a, l991b, l991d). Har-vestingbeganinfallandcontinuedthroughwinter.Theassociatedlithicassemblagesaredominatedby platformcorereductiontechniquesthatyieldedrelativelylargenumbersofsmallflakespercore.The toolsproducedin thesecontextswerenot modifiedor re-sharpenedintensively,and thereis comparatively

little evidencethat they weretrans-portedoverlongdistances.Therearenodirectorobviousfunctionallinksbetweentheformalcomponentsof tech-nologyandsubsistencein theseMousteriandata.Throughouttheperiodinquestion,thevastmajorityof toolsaresidescrapers(primarilysimpleand transversevarieties),un-likelycandidatesforfoodprocurementimplements.Thereareno substantialchangesinthefrequenciesof putative"points,"in spite of the apparentlydifferentemphasesonhuntingand scavengingor gatheringamongthe assemblages.Instead,we findstrongcorrespondencesbetweenforagingstrategiesand the "life histories"of artifacts.Wewouldarguethatthepatternsofcovariationexpresstheinfluenceofdifferingpatternsofmobilityandlanduse, revolvingaroundhowhominidssearchedforresourcesin space.Shiftsin thefrequencyof residentialmovementanddurationof particularoccupationswouldhaveresultedin changingconstraintsoverwhereandwhenlithicrawmaterialscouldbe gatheredand wheretoolscouldbe madeand maintained.The advantagesofalternativetacticsof toolmanufactureandrenewalwouldvaryaccordingly.Passivescavenging,evidentin the earlierMousterianfaunas,is rwormallya strategythattargetsrelativelydispersedresourcesandtendsto yieldlowertransportablereturnsontheaverage.It is morelikegatheringthanhunting,bothin energeticreturnsandtherangingpatternsthatmayberequired.Passivescavengingimpliesmorefrequentmove-mentandperhapswide-rangingsearchpatternson the partof hominids,regardlessofwhetherscavengingopportunitieswereactivelysoughtor (morelikely)merelytakenad-vantageofwhenencountered.It seemsno coincidencethatpassivescavenging

of ungu-latepartsoccursalongsidecollectionofsmall,moreorlesssessileanimalsatonecoastalMousteriancave.An interestin scavengingprobablydid notdrivehominidsto expandtheirforagingradii,butbecausetheywerealreadyrangingwidelyforsomeotherener-geticreason(s),thiskindofscavengingwasbothfeasibleandrewarding.Centripetalcorereductionwouldhavebeenadvantageousin thecontextof relativelyextensiveforaging,becauseit produceslargerflakesand tool blankssuitableforprolongeduse in a raw-material-poorenvironment,as evidenceof toolreductionandtransportindeedsuggests.Prolongingthepotentialuselifeof toolsthroughpersistentreductionorresharpeningis,inturn,a wayof copingwithuncertaintiesaboutjust whenthe toolswill be neededasopposedto whenandwherethenextchanceto make

newtoolswillarise.Evidenceforhuntingin the laterMousterianassemblagesincludesfarrichertrans-portedreturnsperprocurementeventandmayimplytargetingrelativelyconcentratedfoodpatches.Afterall, deerand mostotherungulatespeciestend to congregatewhilealive,especiallyin fallandearlywinter,whereascarcassesgenerallydo notdosooftheirownaccord,at leastnotcontinuously.Moreover,thenearlycompletearraysofanatom-icalelementsindicateminimallevelsof selectivediscardof less economicalbodypartspriorto reachingthe shelters.Morerestrictedrangingforfoodandlongeroccupationsprovlslonedwithhuntedmeatwouldhavemadeit lessimportanttoproducethelargestmostdurableflakesfrompebblecores.Hominidsinsteadmadegreaternumbersofrela-


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tivelysmalltoolsdestinedfor lightuse andlittle transport.Proeessingof huntedgame,whiehcertainlytookplace at thelaterMousterianshelters,mayalsohave set a premiumonunmodifiedorlightlyretouehededgessuitableforcuttinganimaltissues.Thus, in thePontinianMousterian,bigger tools and tool blankswere seleetedonly when peopleneededthings to last, suehas when theyhad to carrytoolsaroundforextendedperiodsandtheaetualmomentsofneedwereunpredictable.AccumulationRatesofBonesversusStones

Informationon thevaryingratesat whichbonesandstonesaccumulatedin strataofthefourcaveslendsadditionalperspectiveon therelationshipbetweenanimalexploita-tion and technology.We assumethat the totalamountsof bothkinds of materialsin theculturaldepositsarelargelya consequenceof twoconditions:( 1) thefrequencyorinten-sityofhominiduseofthe cavesites;and(2) thevolumeofsedimentexcavated.Therelativeproportionsofbonesandstonesamongstratashould,ontheotherhand,relateinsomewaytothe economicrelationshipbetweenthemindependentlyofsedimentationrates.Recallthat the confoundinginfluencesof bonepreservationwereconsideredin advanceandwere not foundto be suflicientlygreat to prohibitthis kind of comparison(see Stiner1990a, 1991a, alsodiscussionsin 199 1b:456-467) .If themaindifferencesbetweenaccumulationratesof bonesandstonesbeforeandafter55,000B.P. were tied to the relativeincidenceof huntingversusscavenging,we wouldexpecttheirrelativefrequenciesto showpositivecorrelationsbutdifferentslopes, in ac-cordancewithtimeperiod.This is becausescavengingby itself (i.e., withoutplantand/orsmallanimalsupplements)shouldnotoperateat a loss,butwouldtend to offerfewertransportablepartsthanhuntedsources(Stinerl 991b, l 991d) .Figure 12 plots the total frequenciesof Mousterianstone tools, flakes,and cores(tLITHICS)againstthetotalnumberofbones(tMNE) forallungulatespecies(bovids,equids, and cervids)6in each assemblage.Trianglesrepresentthe earlierMousterian

1w>55KYears f* < 55 KYears

800- /t LITHICS v /

100 200tM NE, all ungulate taxaFigure 12Scatterplotandregressions for combinedungulatetMNE(cervids,bovids, and equids)andthe frequency of lithic artifacts (tLITHICS = tools + flakes + cores) for Mousterianas-semblages before (dashedline) and after (solid line) 55,000 yearsago.


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cases from Moscerini(level groupsM2, M3, M4, and M6) and Guattari(G2, G4, andG5), dating between 110,000and 55,000 years ago. Squaresrepresentlater cases fromBreuil (Br and B3) and Sant'Agostino(S1, S2, and S3), dating between 55,000 and35,000years ago. It is immediatelyclear from the graph that the relationshipbetweenfaunaland lithic abundancesis not the same for the two groupsof assemblages.The cor-relationbetweentMNE and tLITHICSforthe earliergroupis not statisticallysignificant(r2= 0.084,P = .577,N = 7), indicatingthatthe quantitiesof stonesand ungulatebonesvarymoreor less independentlyof one another.Hominids'discardof stoneat thesecavesis not directlyexplainedor predictedby their use of ungulateresources.To the extentthat thereis anyrelationship,however,it appearsto be negative(dashedregressionline),and hints at the existenceof some componentof subsistencenot directlymonitoredbythe data for largemammalremains.In contrastto the earliergroup, bones and stonesoccurin roughlyequal proportionsin the youngerMousterianassemblages.These cases display a strongpositiverelation-ship (solid regressionline) betweentMNE and tLITHICS (r2= .986, P-.007, N= 5).The S1assemblageis much largerthan the others, hence representingan outlier to thegraph,but it is consistentwith the generalrelationshipas described.The strongpositiverelationshipfor this later group of assemblagessuggests that lithic material cycledthroughthe system more or less in tandem with the numberof ungulate body partsbroughtto the sheltersby hominids.The frequenciesof faunal and lithic debris suggest that, prior to 55,000 B.P., theamountof toolmakingthat went on in the Italiancaves didnotdependon successin ob-tainingscavengedungulateparts.Whileconspicuousfroman archeologicalperspective,scavengingmay have in fact been ancillaryto subsistenceactivities at MosceriniandGuattari.It morelikelywas integratedwithin a wide-rangingforagingpatternthat ac-tually focusedon a variety of dispersedsessile resources,includingsmall quantitiesofmarineshellfish,tortoises,and probablyvegetablefoods.In contrast,the amountoftech-nologicalactivityrepresentedin the laterassemblageswasresponsiveto the quantitiesofungulatepartsprocuredand subsequentlycarriedto the shelters.Huntedgame may ac-tually have been centralto subsistenceduringoccupationsat Grottadi Sant'Agostinoand GrottaBreuil.HeadParts,Scavenging,andFat

Mousterians'exceptionalinterestin head parts when apparentlyscavengingis unex-pectedin the sense that we are not able to find compellingevidenceof this foragingpat-ternamongmodernhunter-gatherers.Isolatedinstancesof specialinterestin head partsmay happennow and again in modernsituations,but not such that the practicedomi-natesentirefaunalsequencesin deeplystratifiedshelterdepositsformedover thousandsof years.To understandwhat "head-collecting"might have been about, it is necessaryto turnto two othersourcesof information,the habitsof nonhumanpredatorsand detailsof the ungulateanatomy.Among the carnivores,a "prefercnce"for head parts (of medium-sizedungulates)oftenemergesin the contextof scavenging(Stiner l991b). It is partlyexplainedby therelativepersistenceof craniaat find sites (e.g., Haynes 1980;Blumenschine1986:35-38)and the ease with whichheads (includingthe mandible)can be removedfromcarcasses,regardlessof theircondition(Stiner l 990a:397-399).Thereare energeticincentivesas well. Headsare complexbony containersand requireconsiderableprocessingby humansand carnivoresalike. So, movinghead parts to a se-cureplace wherethey can be workedon in peace can be worthwhile.Equallyimportantanywherefood energysourcesare periodicallyor perenniallyscarceis the uniquelyhighfat contentof soft tissuesin the head, much of which persistseven in malnourishedprey(Stiner l991b:471-474, 1992b). The fat/proteinratio in head tissues, particularlythebrain,is both high and stablethroughoutthe year, because the fat-richmyelin sheathsenclosingthe cranialnervescannot be metabolizedunderconditionsof food stress. No


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part of the postcranialprey anatomyis like this; instead,fat/proteinratios cycle in ac-cordancewith seasonalshiftsin food supply.Also significantis the fact that yellow mar-row reservesin the adult ungulatemandibleare amongthe last to be drainedmetaboli-callyin timesof food scarcity.Human beings need to consumesubstantialamountsof fat and/or carbohydratesinorderto utilize dietaryproteinseffectively(Speth and Spielmann1983). Becauseplantproductionnaturallywaxesand waneswith the seasons,animalsourcesoffat can becomevery valuableto humans,and Neandertalswould have been no exception.Under con-ditionsof scarcity,then, the nutritionalstate of the animal at the time of death shouldstronglyinfluencehuman(and carnivore)decisionsaboutwhatpartsareworthyof trans-portand extensiveprocessingat anotherplace, such as at a rock-shelteror cave.Nutritionalcontingencieswould not be confinedto scavengingsituations,however.People,whetherhuntingor scavenging,should alwaysbe interestedin heads if fats andcarbohydratesare scarce,and ethnoarcheologicalaccountsshow that they are (e.g., Bin-ford 1978;O'Connell,Hawkes,and BlurtonJones1988a;Yellen 1977). Peoplemay be-come less interestedin heads if other rich, morefavoredparts requiringless processingefTortare available,a fact that undoubtedlyaccountsformuchvariationin transportandprocessingdecisionsfromone monthto the next, dependingon the amplitudeof the sea-sons. Binford's(1978:60-61,77-90) descriptionsof ungulatebutchering,transport,andprocessingby NunamiutEskimorevealconsiderablevariationby season;the Nunamiutwere much more interestedin head parts in spring than they were in the fall. Hunter-gatherersliving in tropicalregionsdisplaymoreconsistentinterestin everypossiblefat-yieldingpartof prey (e.g., Yellen 1977).While peoplecould conceivablychoose to take only heads away fromstarvedprey ob-tainedby hunting,the effortto procurethem can quicklyoutweighthe potentialreturns(sensu Metcalfe 1989) because,rich or not, the head is a relativelysmall package.Thisreasoningassumesan importantstrategicdifferencebetweenpassivescavenging,on theone hand,and huntingand aggressivescavenging,on the other;the value of eitherclassof foragingstrategydependson the nutritionalstate and bulkmass of prey as most typ-ically encounteredin a given foragingsystem. Thus, there is a much lower probabilitythathead-dominatedfaunasrepresenthuntingby hominidson energeticgrounds,in ad-ditionto empiricalsupportabout what scavengingsocial carnivoresreallydo in currentsettings. But, if carnivoresand Neandertalscould produce strongly head-dominatedfaunasat some sites, why is it that modernpeopledo not?The answermay hinge upon the rangeofprocessingoptionsavailableto humans,manyof whichhave been aroundat least since the Upper Paleolithic.Here we referto objectsassociatedwith grinding,pounding,and/or boilinganimaland plant foods. Most of theevidencein the Paleolithicforfood processingtechniquesis indirect,such as fire-crackedcobblesthatmighthave beenusedforstoneboiling.In timesoffat scarcity,bonetrabecu-lae of the spinalcolumnof adult prey also representpotentiallyrich sourcesof fats (e.g.,Speth 1991),and could have been processedby grindingthem up and boiling them, asthey are in certainmodernhumansituations(e.g., Binford1978;Yellen 1977). For rea-sonsthat arenot fullyunderstood,researchon the Mousterianof Europehas yieldedverylimited evidenceof grindingtechnologiesor stone boiling. Heads, in contrast,can beroastedand pickedapart;processingrequirestime and firebut no additionaltechnologyfor extractingnutrients.The absenceof containertechnologyand grindingor poundingimplementsin the Mousteriancould have made heads even more attractiveto foragersin need of fat.Thus, in the Italian Mousterian,strict choices to take awayjust about only heads totwo of the caves may have been due to an advancedstate of starvationin deer, makingthemvery poorin fat at a time when this nutrientwas neededto supplementthe humandiet (Stinerl991b, 1992b).That the opportunitiesto scavengeweregeneratedprimarilyby starvationratherthan by huntingcarnivoresis supportedby the moreor less equiv-alent frequenciesof craniato mandibles(see Stiner l991b:462,471; comparewith Blu-


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mensehine1986on predator-generatedopportunities).Huntingeapabilitymay have hadlittle or nothingto do with Mousterians'deeisionsto seavenge.Rather,foragingeondi-tions made seavengingmore worthwhilegiven the rangingpatternsand teehnologiealsupportin existeneeat that time. Seavengingis reallya familyof taetieswhosediversityand eeonomierules are not well-understood.Clearly,teehnologiealeonsiderationseangreatlyaugmentany effortto investigatethis elass of foragingbehaviorsamonganeientand modernhominids.WhatDoHuntersNeedin OrdertoSucceed?

The rangeofformaltool types (as opposedto reduetionteehniques)in the Mousteriansamplestays about the same aerossthe 55,000-yearboundary,while subsisteneestrate-gies appearto ehangeradically.Why isn't therean equallydrasticchangein the kindsoftoolsbeingprodueed,eitherin the weaponsfordispatehinganimalsor in the implementsused to makeweapons?We are not in a positionto eompletelyanswerthis question,but recentstudies showthat inereasingsophistieationof formalweapons used by modernhuman huntersand"huntingsuecess"(as measuredby prey nutritionalreturns,age elassestaken,etc.) varysomewhatindependentlyof one another.Thereis morethanone solutionformaintainingregularaeeess to high-qualitylive prey. In surveysof ethnohistoriealand areheologiealeases,Stiner(199Ob,l991d) documentsanalogouspatternsof ungulatepreyage seleetionby anatomicallymodernhumansaerossa diversearrayof teehnologiealsystemsand en-vironments.Preyage seleetionis an importantdiagnostieof predator-preyrelationships,implieating"huntingsuecess"in broadecologicalterms(Stinerl991c). Alvardand Kap-lan ( 1991) comparepreyage and sex selectionof terrestrialand arborealspeciesby mod-ern native huntersin a Peruvianrain forest, finding that greaterlevels of cooperationamongbow-hunterscaneffectivelysubstitutefor the advantagesconferredby long-rangeweapons,such as shotguns.Of course,neitherbows nor shotgunswould have existedinMiddlePaleolithictimes.But, if cooperationcan in any way substituteforthe advantagesthat sophisticatedmodernweaponsoffer,then we are obligedto allow for the possibilitythat eooperationamongMousterianforagers(who ereated"modern"-lookingprey mor-talitypatternsin some sites) workedin basicallythe same way.Humanhuntersare not entirelyweapon-dependent;they are only partlyso. The ram-ificationsof these two views are quite different,and failureto appreeiatethe differeneecould presentseriousobstaclesto understandingthe relationshipsbetweenvariationinhumantechnologyand foragingstrategies.There is no questionthat weaponsean faeil-itate the food quest. It is beeomingincreasinglyclear, however,that prey seleetionandgettingenoughto eat are produetsof a complexinterplaybetweenthe teehnologiealsys-tem in use, demography-mediatedpossibilitiesfor human eooperationduringprocure-ment and subsequenttasks, and speeies-and seasonal-speeifiebehaviorsof the animalspursued(Binford1978;Frison 1991;Hudson 1991;Lyman 1991;Stiner l990b, l991c,l991d).In sum, findingevideneeof increasingemphaseson hunted prey in the Mousterian,includingprime-adultharvesting,need not demand a correspondingincrease in thequantityofstoneweaponheads producednor an increasingsophisticationin the types ofweaponsused.It couldhappen,but it does not haveto happen,becausemanyadvantagesof"formal"technologyoverlapwith thoseof beingan intenselysoeialspecies.The ItalianMousteriandata seem to illuminatea substantialblind spot in areheology,lyingjust be-tweenthe specialtiesof zooarcheologyand teehnologyresearehas eurrentlypracticed.

ConclusionCurrentresearchprioritiesfor the Mousterianand debates about the Middle-UpperPaleolithictransitioncenter on questions about adaptive variation and evolutionarychange.In contrast,mostextantdata werecollectedwithverydifferentaimsin mindand,


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in manycases,are of questionablerelevanceto theseissues.The researchdescribedhereapproachesvariationin the Italian Mousterianby combininga unique set of variablesthat are demonstrablyrelevantto how hominidsused resources.The faunal variableswerechosen based on cross-speciessurveysof modern(humanand nonhuman)preda-tors,and the technologicalvariableswereselectedbasedon expectationsabouthow tech-nologyshouldrelateto humanforagingpractices.This researchorientationrevealscon-siderablevariationin the manufactureand use of stone tools in "Pontinian"Mousterianassemblagesthat closelyparalleledshiftsin game use, especiallypatternsof preyage se-lection and body-parttransport.Such variationexists within an archeologicalrecordthat,fromthe classicalperspectivesof tool typologiesand speciesrepresentation,is quitemonotonous.It is clearfromthis researchthat some facts about the Middle Paleolithiccan only beunderstoodthroughsimultaneousconsiderationof zooarcheologicaland technologicalevidenceat the intersitelevel. Collaborationrevealsseven importantthings about therelationbetweenMousteriantechnologyand subsistencepracticesin west-centralItaly:1. Strongcovariationexists betweenartifact(coreand tool) "lifehistories"and hom-nlc . toraglng strategles.2. In situationsthat apparentlyinvolvedscavenging,Mousterianhominidspreferredto transportheads (but not many other ungulateparts) back to shelters.This relativelystrictchoicewas responsiveto the uniquenutritionalassetsofferedby cranialsoft tissuesin times of fat or carbohydratescarcityand, perhaps,the limitationsset by extant pro-cessingtechnologyduringthat culturalperiod.3. Thereare no completeor necessaryconnectionsbetweenmanyof the most obvious(i.e., visually striking)aspects of weapons technologyand the relativedependenceonhuntedlargeprey, eitherin the Mousterianor in modernhumansituations.This obser-vation is especiallycriticalfor researchon the Middle Paleolithic,whereanalystsoftendeal with questionsof presenceor absenceof stone-tippedprojectiles.4. Patternsin lithic technologyand animal exploitationconvergemost clearlyat thelevel of mobilityand land use. More significantthan any findingabout "hunters"and"scavengers,"per se, is the spatialelementimpliedby thesedata. Covariationobservedbetweenthe lithic and faunaldata sets appearsto reflecthow foragersused territory,thedistributionof opportunitiesto collect or hunt variousclasses of resources,and ways ofofEsettingneedsfor key but sometimesscarceresources,such as protein,animalfat, andworkablestone.5. It is also clear that the trendsobservedin the Middle Paleolithicof Latiumrepre-sent shifts in the frequenciesof variousaspects of technologyand foraging,not the ap-pearanceof entirelynew componentsof behavior.For example,evidenceof both centri-petal core reductionand platformcore reductionare presentthroughoutthe Pontiniansequence;platformcore reductionsimply overtakescentripetaltechniqueswith time.This is arguedto be an adjustmentof tool manufactureto the requirementsand limita-tions of foragingand land use, an adaptivefrequencyshift withinan extant rangeof al-ternatives.The same must be said for scavengingversus hunting. Mousterianscavengingprac-tices appearodd in light of what modernhumansdo, but thereis no reasonto assumethat Mousteriansnever scavengedafter 55,000 years ago, or never hunted beforethistime. Virtuallyeveryknownpredator,includingmodernpeople, engagesin both;pred-atorspeciesdifferin theiremphaseson huntingand scavengingin what is best describedas a continuum(Stiner l991b, 1992b).Juxtaposinghuntingand scavengingas if theyrepresenteddistinct hominidpredatoryniches or adaptations,ratherthan componentsof any one, is counterproductive.In otherwords,we shouldexpect to see some combina-tion of these two generalstrategyclassesin hominidadaptationsthroughoutthe UpperPleistoceneand perhapsearlier.Variationin how hunting and scavengingtactics aremanifestin foragingadaptationsis a problemof a finer order, as researchon modern


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predatorsattests. Certainly,scavengingshould not, at the level of populationrelation-ships,be modeledas the last resortof an incompetentspecies.6. If thereis anythingdistinctiveor anomalousabout the Mousterianof west-centralItaly comparedto anatom-icallymodernhuman lifeways,it is in how alternativetacticsof gameprocurementand tool manufacturewereimplementedas componentsin a largeradaptivesystem. For example,while scavengingis conspicuousin the earlierpart of theItalian sequence,it probablywas not the only option for these foragersor their singlemainstay.The Mousterianscavengingpatternstandsapartfromknownmodernhumancasesbecauseit was so clearlyseparated,spatiallyand temporally,fromhunting.It pre-sentsa strikingcontrastto ethnoarcheologicalcases wherethe productsof both strategyclassesare pooledat the same residentialplaces (e.g., Binford1978;O'Connell,Hawkes,and BlurtonJones 1988a, 1988b;Bunn, Bartram,and Kroll 1988). Because ungulateheadpartsrepresenta consistentlyrich sourceof fats, and hominids'basic physiologicalneed for dietaryfat probablyhas not changed,the Mousteriandata suggest that therewas somethingvery differentabout the ways ungulateexploitationoffsetthe use of theother resources(Stiner l991b, 1992b). Independent(possiblymildly negative)rates ofaccumulationfor bones and stones in the caves also suggestthat ungulateswere not thecoreof subsistencein sites wherepassivescavengingis evidenced.Deer head parts mayinsteadhave merelysupplementedshellfishand otherarcheologicallyinvisiblefoodsde-

, . . ,>hcoent1ntats.As concernsthe contextof passive scavengingin the Italian Mousterian,seasonalitydata areof limitedhelp. Death by starvationin ungulatepopulationspeaksaroundearlyspringin coolerenvironments(see reviewin Stiner1990a:595-600),and fat consequentlybecomesrarerfor omnivorouspredatorsbeforeplant productionis in full swing eachyear.The head-dominatedMousterianfaunasindeedappearto center(witha wide rangeof variation)on spring.However,abstractingthe seasonalityresultsto explainintersitevariationin hominidforagingactivitieswould be much too simplistic:the intersitedif-ferencesin resourceuse lasted thousandsof years,probablyspanningseveralminorcli-maticoscillations.7. Within the geographicallimits imposedby the study sample, the observedtrendsin foragingand technologydo appearsomewhat"progressive."Scavengingis "replaced"by hunting,and classic Mousteriancentripetalreductiontechniquesare supersededbysingle-and double-platformapproachesreminiscentof Upper Paleolithicblade core re-ductiontechniques.The fact that observedchangesappearto point to a stereotypically"modern"pattern does not constitute an explanation, however. The technologicalchangesmay indeedrepresentresponsesto alterationsin foragingp

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1992_Stiner y Kuhn_American Anthropology Pararo