Subsistence, Technology, and Adaptive Variation in Middle Paleolithic Italy

Author(s): Mary C. Stiner and Steven L. Kuhn

Source: American Anthropologist, New Series, Vol. 94, No. 2 (Jun., 1992), pp. 306-339
Published by: Blackwell Publishing on behalf of the American Anthropological Association
Stable URL: http://www.jstor.org/stable/680462
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MARY C. STINER
STEVEN L. KUHN
of New Mexico
University

Subsistence, Technology, and Adaptive Variation

in Middle Paleolithic Italy

Thiscollaborative studyaddressesquestionsaboutvariationin behaviorwithintheMiddlePa-

leolithicperiodof west-central Italy. Thefindingsaredirectlyrelevantto theevolutionary 'fate"
of Neandertals andtheappearance of anatomically modernhumansin Europe.Analysesfocuson
variationin a numberof dimensions of subsistenceandtechnology, includingpatternsof ungulate
procurement,food transport, andtacticsof stonetoolmanufacture anduse.All of thesedimensions
exhibitmarked,possibly"vectored" changein thestudyareabetween110,000and35,000years
ago. Interpretations of thearcheological data aresupported by independent studieslinkingfood
search,procurement, andtransport behaviors toecologicalrulesaffectingall largeterrestrialpred-
ators.In this way, variationobservedin thefaunaland lithicdatasets can be shownto reflect
startlingdiversityin theresponses of Mousterianhominidsto theworldaroundthem.Thedata
provideimportantinsights into the natureof adaptive"rawmaterial"alreadyin place during
theMiddlePaleolithic,theforemost conclusionof thisstudy.Thatthis variationin resource ex-
ploitationandlandusealsoappearsdirectional is moredifficulttoevaluate.It couldbeexplained
by hominids'adjustments to localchangesin coastalhabitatas sea levelregressed or, alterna-
tively, an evolutionaryshift in hominid capabilities. Thefirst interpretationis preferred,on the
basis of available evidence,but cannotbe advancedas a certainconclusion.This questionprovides
muchstimulusfor continuedresearchin the study area and otherregions, using the methodspre-
sented.

SOME OF THE MOSTCOMPELLINGresearch priorities in Paleolithic archeology concern

the 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 supports
a 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 situ
evolution across a broad front (e.g., Clark and Lindly 1989a, 1989b; Wolpoff 1989). For
the most part, proponents of these two positions do not cite different categories of facts
or different sets of observations. Rather, the conflict arises from differing interpretations
of essentially the same bodies of data.
Discussions of contrasts and similarities between the Middle and Upper Paleolithic
have focused primarily on three phenomena: techniques of lithic blank production and
tool forms, the ranges of prey species exploited, and frequencies of decorative or symbolic
art (e.g., Chase and Dibble 1987; Clark 1987; Clark and Lindly 1989a, 1989b; Harrold
1989; Lindly and Clark 1990; Mellars 1973, 1989; Orquera 1984; Straus 1983; White
1982). Because it was once thought that they illustrated major discontinuities between
the periods, these three topics continue to dominate even the most recent syntheses. Po-

MARYC. STINER
is AdjunctAssistantProfessor,Department Universityof New Mexico,Albuquerque,
of Anthropology, NM
87131.STEVENL.
KUHNis AdjunctAssistantProfessor,DepartmentofAnthropology,
University ofNew Mexico,Albuquerque,
NM 87131.

306
Stiner and Kuhn] ADAPTIVE VARIATIONIN MIDDLE PALEOLITHIC ITALY 307

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 of
known 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 theoretical
predictionthat flake technologies or "unspecialized" hunting should be the ancestral con-
dition. These were simply the only sets of facts that had been documented in comparable
ways for both periods. More recent observations show that Middle and Upper Paleolithic
assemblages may contain quite similar ranges of species (e.g., Clark and Lindly
1989b:644; Simek and Snyder 1988; Stiner 1992a) and/or comparable techniques of
blank production (Clark and Lindly 1989b:639; Straus and Heller 1988). While showing
the old generalizations to be inadequate, they provide no guarantee of the evolutionary
significance 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, and
there are many different ways to investigate them archeologically. Except for the histor-
ical fact that substantial comparative data bases already exist, there is no overwhelming
reason for continuing to place so much weight on the usual triad of observations to trace
the evolution of Homo sapiens. 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: tigers
have stripes and claws, leopards have spots and claws, and zebras have stripes and
hooves. Looking only at these visually striking traits, we might conclude that zebras are
about as similar to tigers as are leopards. However, biologists would agree on the basis
of real-life observations that the claws of a carnivore and the hooves of an ungulate are
much more important than pelage for establishing the behavioral, physiological, and
phylogenetic similarities among species. Regardless of how obvious and easily measured
stripes and spots may be, patterns of pigmentation are not all that significant at this level
of comparison.
Today, most questions surrounding the "transition" and the relationship between ar-
chaic and anatomically modern Homo sapiens concern the nature and rates of adaptive
change. 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 reasonably
be expected to distinguish different subspecies of Homo sapiens. Behavioral adaptations
vary only subtly between modern subspecies of nonhuman taxa (sensu O'Brien and Mayr
1991), and we should not expect huge discontinuities among late Pleistocene hominids in
the basic 4onstituents of behavior. Although more robust and apparently stronger than
anatomically modern humans (e.g., Trinkaus 1983a, 1983b, 1986), Neandertals would
have been subject to similar physiological limitations. They would, for example, have
faced 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 as
protein and fat) in their diets. Differences between anatomically modern populations and
their immediate predecessors are unlikely to have involved the addition or loss of any of
the most basic components of foraging and technology. Instead, evolutionary changes are
most likely to have been manifest in how these components were integrated or organized
as 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
308 AMERICAN
ANTHROPOLOGIST [94,1992

terms of economic structure and the rules of strategy combination, has been particularly
stimulating in this regard. Our research orientation does not necessarily share, however,
all of the more specific characterizations of behavioral differences between modern and
archaic Homosapiens.
The research discussed in this article addresses the extent and causes of variation in
technology and subsistence on the Mousterian "side" of the transition in coastal west-
central Italy. The study uses information about the relationships between food choices
and foraging strategies among modern nonhuman predators and hunter-gatherers as in-
dependent referents for interpreting patterns in the Mousterian faunal data. If hominid
predatory adaptations (admittedly only one part of the subsistence repertoire) changed
significantly during the Upper Pleistocene, we should expect differences of the magnitude
that normally distinguish other large predators in modern animal communities, regard-
less of whether Neandertals and "moderns" prove to have been wholly, quasi-, or non-
contemporaneous hominid populations.
Trophically linked species, such as coexisting predators, may share an interest in the
same food species but exploit them differently (MacArthur 1968; MacArthur and Levins
1967; Root 1975; Wiens 1977). Modern nonhuman predators respond 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 consumed
within this range are more often than not poor indicators of adaptational differences
among them (see, for example, Kitchener 1991:98-105). Of far greater interest are the
ways predators-in this case, variants of Homosapiens-might have obtained and used the
same food species, as manifest in ranging patterns, the prey age groups most commonly
targeted, 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 tools
and the forms that tools took were heavily constrained by both the physical properties of
the human organism and the mechanics of stone fracture. Major evolutionary changes in
human technologies are more likely to have involved the role of technology within a larger
system than simple alterations in tool form and function. This is particularly true for
Mousterian and earlier technologies, since specialized stone tools for food procurement
and processing were rare (Kuhn 1989a; Holdaway 1989; but cf. Shea 1989). Thus, the
relationship betweentechnology and subsistence is here considered central to understand-
ing adaptive variation in toolmaking behavior during the Upper Pleistocene. The lithic
analyses emphasize those variables expected to have been heavily influenced by food
search and procurement strategies. Patterns of 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-
erable variability 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, other
aspects of technology varied 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 Sample

The animal bone and stone assemblages discussed in this article come from a series of
cave sites in Latium, a province on the Tyrrhenian (west-central) coast of Italy (Figure
1). This area is characterized by north-south trending mountains separated from the sea
in 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 10
km from the sea during the period under discussion, depending on oscillations in global
temperature.
Stiner and Kuhn] ADAPTIVE VARIATIONIN MIDDLE PALEOLITHIC ITALY 309

Figure 1
Geographical locations of the four Mousterian cave sites: (B) Grotta Breuil, (G) Grotta
Guattari, (M) Grotta dei Moscerini, and (S) Grotta di Sant'Agostino.

Interest in the Middle Paleolithic of Latium began early in this century. The region
gained international attention in 1939 when vineyard workers accidentally exposed the
entrance of Grotta Guattari (Blanc 1939; Blanc and Segre 1953; Piperno 1976-77). Inside
the cave, a well-preserved Neandertal cranium, and later a mandible, were found exposed
amid a jumble of rocks and mammal bones (see Stiner 1991a, White and Toth 1991 for
recent reanalyses of the Guattari finds). The discovery spurred an ambitious program of
survey and excavation of coastal and inland caves throughout the 1940s and early 1950s
by A. C. Blanc, A. G. Segre, L. Cardini, and other members of the Istituto Italiano di
Paleontologia Umana (IIPU).
The assemblages to be discussed are collections excavated by the IIPU research group,
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; Piperno
1976-77; 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 situated
in coastal cliffs near Gaeta, roughly 50 km to the south. All of the caves are relatively
shallow solution features in limestone bedrock and occur at similar elevations.
The assemblages from the four caves collectively span a period beginning just before
110,000 years 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). At
present, the radiometric dates are more informative about the ages of the assemblages
relative to one another than they are for determining the absolute age of each. The study
sample 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 time
frame represents a very general trend toward colder conditions, but incorporates a 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 the
case of Grotta dei Moscerini to combine materials from series of adjacent strata to pro-
310 A AERICAN ANTHROPOLOGIST [94, 1992

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

Site Assemblage Origin/Definition

Guattari G2 stratum 2
G4 stratum 4
G5 stratum 5
Moscerinia M2 external strata 11-20
M3 external strata 21-36
M4 external strata 37-47
M6 internal strata 3-4
Breuil B3 stratum 3
Br strata 11-15 (derived material)
Sant'Agostino SO surface
S1 arbitrary level 1
S2 arbitrary level 2
S3 arbitrary level 3

"Moscerini proveniences M2, M3, M4, and M6 represent series of combined strata grouped on
the basis of uniform content and sediment composition. M2-M4 in fact contain over 30 cultural
lenses, which may or may not represent discrete occupational events.

duce assemblages of adequate size; groupings follow shifts in faunal contents and changes
in sediment composition (see Table 1). In these cases, radiometric dates from multiple
levels are averaged to produce an estimated date for each strata group. Samples from
more recent (piece-plotted) excavations at Grotta Breuil are lumped into stratigraphi-
cally defined groupings roughly equivalent to those derived from earlier excavations to
ensure comparability (see Kuhn 1990a, Stiner 1990a). None of the assemblages represent
single occupational "events"; instead, they are palimpsests of many such events. While
the level of resolution may not be ideal for detailed reconstruction of individual activities
(Rigaud and Simek 1987), these long-term accumulations are entirely appropriate for
investigating variation in behavioral tendencies at an evolutionary time scale.
The subject lithic assemblages belong to a regional facies of the Mousterian called the
Pontinian, after the Pontine Plain, where first identified (Blanc 1937; Taschini 1972). The
Pontinian is relatively invariant from the perspective of Bordes's (1961) Middle Paleo-
lithic stone tool typology, exhibiting a strong predominance ofsidescrapers and generally
low 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 deposited
before this date is attributable to Neandertals. Human fossil associations for later Mous-
terian assemblages are ambiguous, however. Ongoing excavations at Grotta Breuil have
yielded human remains likely to date after 50,000 B.P., but none is clearly diagnostic of
variants of Homo sapiens (Bietti et al. 1988; Manzi and Passarello 1988).

Methods

The data combined in this presentation were obtained from two freestanding studies,
each of which employed a distinct set of analytical methods. Only the general approaches
and the most important variables are described below. Details of the faunal and tech-
nological analyses, data bases, sampling issues, and taphonomic considerations (in the
case of faunal remains), are presented elsewhere (see Kuhn 1990a, 1990b, 1991; Stiner
1990a, 1990b, 1991a, 1991b).
Stiner and Kuhn] ADAPTIVEVARIA
TIONIN MIDDLEPALEOLITHIC
ITALY 311

ZU

30- U

40- I

50-

60-
cn
< 70 -
80-
Y 80-

90 -

100 -

110 -

-
: U
120 I I
M G S B
CAVE SITES

Figure 2
Chronological spans represented by the four cave strata sequences: (M) Moscerini, (G)
Guattari,(S) Sant'Agostino,and (B) Breuil. Solid bars indicate spans dated by ESR method,
U indicates flowstone layer dated by U-series method (Schwarcz et al. 1990-91, 1991),
dashed bars indicate estimated but currently undated segments.

Methodsand InterpretiveBasesfor the Faunal Analyses

The 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 are
confined to red and fallow deer (Cervus elaphus; Dama dama), the most common prey ex-
ploited by Upper Pleistocene hominids in west-central Italy by any measure.2
The faunal assemblages were deposited inside shelters, and thus represent cumulative
outcomes of transport, rather than items discarded where food was obtained. The animal
remains therefore are subsets drawn from larger resource pools, the general composition
of which can be modeled from modern analogs. In the case of body-part transport
choices, 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 number
of parts taken away must have equaled or been less than what was present on carcasses
when 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
312 AMERICAN
ANTHROPOLOGIST [94, 1992

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

Site and Dating

vertical provenience methoda Material Ageb
Grottadei Moscerinic
Stratum 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 -
Stratum 33 (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 ?
Stratum 38 (M4) ESR red deer tooth 106,000 -
Stratum 39 (M4) ESR red deer tooth 96,000
Stratum 39 (M4) ESR hippopotamus tooth 97,000 -
GrottaGuattari
GO(surface) U/Thd stalagmite 51,000 ? 3,000
G4 ESR aurochs tooth 71,100 ? 2,760 (avr)
G5 ESR aurochs tooth 77,500 + 9,500 (avr)
Grottadi Sant'Agostino
S1 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 ? 14,000
Lower flowstone U/Thd stalagmite 120,000 + 15,000
GrottaBreuil
Capping flowstone U/Thd stalagmite 26,000 ? 12,000
Stratum 3 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 of
each 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 entrance
today is completely blocked by rubble from highway construction uphill of the site. For this reason,
background dose rates for the ESR analyses had to be estimated using soil cemented to the tooth
specimens, and no error ranges 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 efforts
to interpret archeological cases (Stiner 1991c:9-10).
Both transport patterns (anatomical representation) and prey age selection (mortality
patterns) 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, even
though each foraging mode can involve exactly the same prey species. Ungulate hunting
involves 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-
Stiner and Kuhn] ADAPTIVE VARIATION IN MIDDLE PALEOLITHIC ITALY 313

ing more like gathering than hunting, both in the spatial characteristics of these food
sources and the potential average returns gained from them.
Efforts to distinguish the faunal products of scavenging and hunting are potentially
subject to several sources of ambiguity, however. This point merits additional discussion,
since a great deal hinges on how the faunas in this study are interpreted. Generally speak-
ing, both kinds of behavior can take more than one form, depending on the nature of
opportunity and details of the subsistence system in use. Moreover, some patterns that
hunting and scavenging behaviors impose on faunal assemblages overlap (compare, for
example, 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, one
or more of which are practiced by nearly every carnivorous organism. 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 well
before the carcass would have been discarded voluntarily. The Hadza, for example, fre-
quently engage in aggressive scavenging (e.g., O'Connell, Hawkes, and Blurton Jones
1988a), and tense moments between lions and people have been noted. It is important to
appreciate that many or most of these contests occur during forays that anticipated
(among other 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 returns per find, yet they are profitable under the right
conditions. The limits and strengths of each faunal variable used for interpreting Mous-
terian foraging behaviors are discussed below in light of these issues.
Measures of anatomical representation are based on minimum number of skeletal ele-
ment (MNE) counts for bone.Animal bones in archeological assemblages are usually
smashed to bits for reasons beginning more or less with the processing habits of people,
followed by any of a wide array of other destructive agencies. In using MNE, zooarcheol-
ogists try to reconstruct the full, minimum array of whole animal bones on the basis of
many 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 is
determined from the most common portion or landmark of each skeletal element and, in
this study, represents the sum of right and left sides for elements that naturally occur in
pairs. The counts for limb bones use unique landmarks on or near the articular ends,3
and any of a variety of distinctive bony features of the cranium and mandible (e.g., pe-
trous, occipital condyle; incisive, mandibular condyle; or bony architecture of the maxilla
or mandible behind the cheek tooth row if relatively complete; see Stiner 1991b:460-462).
Teeth are not used for the MNE determinations, because they can distort head counts
relativeto postcrania wherever preservation conditions are less than ideal. This point is
very important for understanding the anatomical comparisons, where comparability in
the face of differential preservation is a just concern.
Mortality patterns, in contrast, are based on teeth; specifically, tooth eruption and
wear data for the lower fourth premolar sequence. The aging criteria for occlusal wear in
deer 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 old
adults (see Stiner 1990b:311-313).
Findings on species consumed by Middle Paleolithic hominids, and seasonality for the
same array of faunas, are summarized from other sources (Stiner 1990a, 1992a) as per-
tinent to the discussion that follows. While not covered in this presentation, taphonomic
analyses were essential for establishing the origin of each faunal assemblage (Stiner
1990a, 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 preservation for the cave faunas is very
314 ANTHROPOLOGIST
AM1ERICAN [94, 1992

good on the whole, because they were deposited in calcium-rich limestone formations.
While some postdepositional bone destruction seems inevitable, particularly for skeletal
elements composed mainly of trabecular (spongy) tissue (e.g., Lyman 1985), in situ bone
attrition does not explain most of the variation in anatomical part representation among
the assemblages (Stiner 1990a, 1991b). Potential biases due to differential bone decom-
position are further circumvented by considering only the more durable parts of the bony
skeleton-the head and limb elements.
To simplify the presentation, patterns of variation in the archeofaunas are summarized
in terms of only three derived variables. Each is an index suitable for the interpretation
of 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 completeness
for deer. It is obtained by dividing the totalnumber of skeletal elements (tMNE) by the
minimum number of individual animals (MNI). The ratio represents the average quan-
tity of bony parts transported to shelters per carcass source. Variation in this index
loosely corresponds to differences in transported returns from hunting (high values) as
opposed to scavenging (low values) in carnivore and human control cases (Stiner
1990a: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 for
red and fallow deer in each assemblage. Limb MNE includes all substantial leg elements
except phalanges, carpals, and tarsals other than the calcaneum and astragalus. Head
MNE includes antler as well as cranium and mandible elements, although antler is rare
in the hominid-collected faunas (Stiner 1991a:112). Crania and mandibles occur in
roughly equal proportions, a fact whose significance will become apparent in discussion.
In effect, head MNE/limb MNE concerns relatively large bones, those which in principle
could represent attractively sized transportable parcels. The head MNE/limb MNE ratio
accounts for much of the variation in anatomical representation in the Mousterian shelter
faunas (Stiner 1991b:463-465). In controlled comparisons of other predators, the ratio is
a strong predictor of the general way that prey are most commonly obtained: hunting
results 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-dominated assemblages (Stiner
1991b, 1992b).
The interpretation that head-dominated faunas represent mainly scavenged sources
may 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 strongly
head-dominant pattern observed in the Italian Mousterian. Many aspects of Middle Pa-
leolithic records perplex archeologists familiar with recent human practices, however, so
finding 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 too
much unexplained. Likewise, taphonomic analyses firmly exclude nonhuman agencies,
such as wolves and spotted hyenas, as significant bone collectors in all of the cases (Stiner
1990a), except Guattari G2 and possibly G4-5 (Stiner 1991a). In contrast to the apparent
lack of analogous situations for "head-collecting" among modern-day humans, three spe-
cies of scavenging social carnivore (striped, brown, and spotted hyenas) frequently create
head- 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 context
of scavenging is best explained by properties of 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, specifically
the relative frequencies of prime and old adults. These adult age categories are defined
on the basis of life-history characteristics and associated physiological changes in deer
and other ungulates (Stiner 1990b:308-313). The ratio registers two potential directions
Stiner and Kuhn] ADAPTIVE V4RIA TION IN MIDDLE PALEOLITHIC ITALY 315

that age biases often take in predator-collected faunas. Comparative research indicates
that this ratio is very responsive to the relative emphasis on ambush hunting, which
among modern humans frequently results in a prime-dominant mortality pattern, as op-
posed to scavenging, which among carnivores can result in elevated numbers of old-aged
animals (Stiner 1990b, 1991d).
Not all foraging tactics leave strong or mutually exclusive signatures on mortality pat-
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 (Stiner
1990b: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 prevailing cause(s) of death
and what happens to carcasses during the interlude between death and when foragers are
normally able to locate them. Whether originating from predator kills or nonviolent
causes of death, the carcass pool appears to have begun with a situation of U-shaped
mortality, the most common family of death patterns in nature (reviewed in Stiner
1990b: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 in
natural living populations. Such mortality patterns frequently undergo further distortion,
however, by losing much of the juvenile component. Small, immature bodies are more
prone 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), and
they are more difficult to locate (Schaller 1967, 1972). For all of these reasons, juveniles
tend 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. Klein
1989:377).
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. Mortality
analyses 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-
terns often take a relatively distinct, prime-dominant form. This cannot be explained by
differential preservation favoring adult teeth over those of juveniles, as prey mortality
patterns associated with hyena occupations in some of the very same caves are strongly
biased towardjuvenile prey (Stiner 1990a, 1990b).
While not the only age pattern that ever resulted from hunting, prime-dominant mor-
tality is regularly associated with modern human hunting practices (Stiner 1990b, 199ld;
contra Klein 1982a, 1982b). The pattern varies from a slight to great overabundance of
prime-aged animals in relation to their live abundance. On the whole, prime-dominant
prey age selection represents a long-established tendency in humans, generally setting
them apart from other large predators (Stiner 1990b:329-341). While targeting the most
productive age groups in prey populations, the relationship is potentially stable in situ-
ations involving cervids and bovids, unless or until human subsistence no longer depends
in any way on the targeted species (Stiner 1991Id: 180-184).
Even with the benefit of controlled associations between transport patterns, mortality
patterns, and foraging behaviors, it would not be appropriate to assume that each and
every whole animal procured by Mousterian hominids had to have been hunted, nor
every head from an old animal scavenged. The interpretations are probabilistic assess-
ments about what people usually did when presented with a general set of circumstances.
The study seeks information about foraging practices at the level of interactions between
predator and prey populations, and only the cumulative patterns representing repeated
behaviors are meaningful.
MethodsandInterpretive
Basesfor theLithicAnalyses
The analyses of the Mousterian industries address some of the potential links between
technology and subsistence, particularly connections involving the shared time and en-
316 AMERICAN
ANTHROPOLOGIST [94,1992

ergy budget. Among modern hunter-gatherers, the timing of subsistence activities and
movement from one place to another determine the scheduling of both the needs for tools
and 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 spatial
and temporal discontinuities between need and opportunity (e.g., Binford 1977, 1979;
Bleed 1986; Kuhn 1989b; Nelson 1991; Shott 1986; Torrence 1983, 1989). Tactics for
producing flakes and blanks from cores, the resharpening or renewal of stone tools, and
the transport of tools and raw material all play a role in determining how long tools and
raw materials will last. Studies of these variables should therefore provide information
about the contingencies faced by hominids in making tools available when and where
they were needed.
Coastal Latium presents an especially interesting natural laboratory for the study of
Mousterian technology. The only workable stone found within at least 40-50 km of the
coastal 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 coastal
plains. Such small and dispersed raw materials would have placed a special premium on
tactics for stretching raw materials and maximizing tool size.
The lithic analyses here focus on two dimensions of the technological record: (1) the
intensity 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 are
required to monitor these disparate phenomena. The intensities of tool use and reduction
are measured using two variables. One is the frequency of retouched tools relative to
unmodified but potentially usable flakes (>2.0 cm in length), a measure that has been
employed in other studies (e.g., Rolland 1981; Rolland and Dibble 1990). The other is
an experimentally derived index of scraper reduction (Kuhn 1990b). This index monitors
how much of a tool blank has been removed by previous retouch, based on the angle of
the retouched edge and the extension of retouch scars relative to the medial thickness of
the blank.
The analyses of core reduction techniques use a different approach and require greater
explanation. Two techniques for making flakes and tool blanks, here termed centripetal
and platformcore reduction (Figure 3), were very important in the Pontinian. There are,
of course, numerous variations on the two basic ways of producing flakes from pebble
cores, and they may sometimes overlap (e.g., Bietti, Rossetti, and Zanzi 1989; Kuhn
1990a;Laj-Pannocchia 1950). Nonetheless, there is ample reason to believe that these are
distinct ways of exploiting raw material, as discussed below.
The centripetal technique, schematized in Figure 3(a), is typical of the Mousterian in
general (Bordes 1961:26-27; Tixier, Inizian, and Roche 1980:43), although there are
many 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 flat
round pebble, gradually spiraling toward the center by rotating the core with each new
blow. For the purpose of this study, evidence of Levallois technique with centripetal core
preparation (represented largely 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 a
striking platform, 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 end
of its useful life (Figure 3[c]). These platform core techniques are similar but not identical
to classic prismatic blade core techniques of the Upper Paleolithic. However, the Mous-
terian versions are decidedly less complex, beginning only with platform preparation at
one or both ends of a pebble. The "crested blade" technique, in which a ridge is created
along 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
Stinerand Kuhn] ADAPTIVE VARIATIONIN MIDDLE PALEOLITHIC ITALY 317

a b c

Centripetal Platform

Figure 3
Schematized (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 on
the backs of flakes detached subsequently, it is possible to monitor the use of different
core 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 tend
to exhibit dorsal scars originating from several directions, often orthogonal to one another
(Figure 3[a]). In contrast, flakes detached from platform cores generally have dorsal
scars running parallel to one another along the long axis of the flake (Figure 3[b,c]): when
pebbles are used, flakes often retain remnant cortex along one edge.5
The two basic core reduction techniques, platform and centripetal, have somewhat
different economic properties. Platform core reduction appears to maximize the numberof
flakes produced per pebble core, while centripetal techniques maximize the sizes of flakes
and tool blanks. To illustrate this difference, Figure 4 compares the number of large flakes
and 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 be
assigned 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 the
study sample, with more sharp edge per piece than those made from platform cores. The
flakes and blanks attributed to centripetal reduction are approximately 25% longer in
maximum dimension and roughly 1.5 to 2 times larger in plan view area than those at-
tributed to platform cores (Kuhn 1990a:226).
318 ANTHROPOLOGIST
AAMERICAN [94, 1992

flakes+ tools
cores

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

1-

.Li .,
Centripetal Platform

.... . unattributable pieces

Figure 4
Comparisonof the relative efficiencies (flakes + tools/cores) of centripetal and platform
reduction techniques in Mousterianlithic assemblages.

Variation in the Faunas

In the following discussion, 12 hominid-generated faunal assemblages are arranged in
chronological order 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 ESR
and U-series methods, whereas four assemblages in the middle (M2, M6, SO,and Br) are
ranked on the basis of stratigraphic criteria only (see Tables 1 and 2). Because we want
to consider both radiometrically dated and stratigraphically ordered assemblages, and
because the ESR dating technique remains somewhat experimental (Griin 1988), the
chronological sequence is treated as a simple rank ordering. For ease of discussion, the
assemblages are often referred to as dating before or after 55,000 B.P., the boundary of
which is marked on the bar graphs.
Figure 5 shows a trend in the average amount of transported food returns per red or
fallow deer carcass, using the bone-based ratio of anatomical completeness (tMNE/
MNI). In the earlier half of the chronological scale, the quantity of bony parts carried to
shelters by hominids per procurement event is relatively low. The values increase signif-
icantly toward the recent end of the scale. The changes appear gradual overall, although
a 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 head
parts to limb parts transported to the caves by hominids, using the head MNE/limb
MNE 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 to
limbs in assemblages dating to before 55,000 years ago. The ratio declines gradually early
on 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 tooth
eruption and wear. The expected value shown on they-axis (2.14) refers to the ratio of
prime-aged to old-aged adults in an idealized, stable live-prey population (from Stiner
1990b; see also Lyman 1987). The expected value on the graph would represent "non-
selective" harvesting by hominids, regardless ofjust how the animals were obtained. Deer
mortality patterns are significantly biased toward old-aged prey in all cases dating to
before 55,000 B.P., and the bias lies well outside the range of normal variation displayed
by live populations (Stiner 1990b, 1991d). The cases dating to just after 55,000 show
essentially nonselective procurement, while values for the most recent Mousterian assem-
Stinerand Kuhn] ADAPTI
IZE
VARIA
TION
INMIDDLE
PALEOLITHIC
ITALY 3l9

35

tMNE 25-
MNI

'S- ,,1,|,l1llll
M4 M3 G45 M2 M6 G2 Br S3 S2 S1 S0 B3

110 KY 55 KY 35 KY

ASSEM BLAGE

Figure 5
Anatomicalcompleteness index values (tMNE/MNI)for red and fallow deer in Mousterian
faunal assemblages, arranged in chronological order (see Table 1 for assemblage designa-
tions).

blages(ca. 40,000-35,000B.P.) are stronglybiasedtowardprimeadultprey.The prime-

dominantcasesexceedthe rangeencompassedby normalvariationin naturallivingpop-
ulations,even whenphasesof growthand declineare takeninto account.
Rankordercorrelations(Spearman'srS)of the threeratiosagainstchronologicalrank-
ings evaluate the statisticalsignificanceof the trends. Two alternativechronological
rankingsare used. Table 3(a) lists correlationcoeflicientsfor all dated and undatedas-
semblages.Variationin each of the threefaunalratiosis highly correlatedwith assem-
blage age at the .01 level of probability.Not all assemblagescould be dated by radio-
metrictechniques,however,and properchronologicalorderingis very importantto the
conclusions.In orderto check this problem,Table 3(b) lists correlationcoeflicientsfor
radiometrically dated assemblagesonly, thereforeexcludingM2, M6, S0, and Br. Cor-
relationsremainsignificantfor this subset,althoughthe value for tMNE/MNI dropsto
the .05 level of probability.
The relationship between two of the faunal ratios, head MNE/limb MNE and
PRIME/OLD,is evaluatedanotherway in Figure8. A log-log regressionof the ratios
documentsa strong negative relationshipbetween the occurrenceof head-biasesand
prime-agedprey(r = -. 733,r2 _ .54, .02 < P < .01,N = 10). Becausethe firstvariable
is basedexclusivelyon bone material,whereasthe secondis based exclusivelyon more
resistanttoothmaterial,the comparisonservesas a partialcheckagainstpossiblebiases
createdby differentialpreservationamong skeletaltissue classes. Bones and teeth tell
similarstoriesabout the contextin which food was obtained.The B3 assemblagefrom
GrottaBreuilis an outlierto the graphand, while not includedin the regressioncalcu-
lation,falls at the appropriateextremeof the differencesas described.The case fromG2
of GrottaGuattariis excludedfrom considerationbecauseof insufEcientage data and
becauseits taphonomicoriginis moreambiguousthanall of the othercases.The assem-
blagesalign in a mannerconsistentwith the generalchronologicalorderdefinedprevi-
ously, althoughcases dating after55,000 B.P. display less variationon the anatomical
axis (headMNE/limbMNE).
 AMERICANANTlfROPOLO(;IST
320

[94, 1992

head MNE
Iimb MNE

M4 M3 G*5 M2 M6
l-W-F-
G2 Br S3 S2 S1 SO B3

110 KY 55 KY 35 KY

ASSEMBLAGE
Figure 6
Ratios of head to limb parts (head MNE/limb MNE) for red and fallow deer in Mousterian
faunal assemblages, arranged in chronological order.

PRIME
OLD

M4 M3 G45 M2 M6 G2 Br S3 S2 S1 SO B3

110 KY 55 KY 35 KY

ASSEM BLAGE

Figure 7
Ratios of prime-aged to old-aged adult (PRIME/OLD)red and fallow deer in Mousterian
faunal assemblages, arranged in chronological order.
Stinerand Kuhn] ADAPTIVEVARIA
TIONIN MIDDLEPALEOLITHIK
ITALY

321
Table3
Spearmanrankordercorrelation(rS)of faunalvariablesagainstincreasingassemblageage.

Faunal variable N rSa P

a. All assemblages
tMNE/MNI 12 -0 744 < .01
Head MNE/limb MNE 12 0.818 < .01
PRIME/OLD 1l -0.773 < .01
b. Datedassemblages
only(excludesM2, M6, SO,andBr)
tMNE/MNI 8 -0.714 = .05
Head MNE/limb MNE 8 0.881 < .01
PRIME/OLD 7 -0.905 < .01

aValues of rSindicate correlation with increasing age.

*>55K Years
* < 55 K Years

PRIME
OLD

head MNE
limb MNE

Figure 8
Log-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 Mousterianfaunal assemblages.

In sum, low returnsfor transportedfood (low tMNE/MNI values) and head-domi-

natedanatomicalpatterns(highheadMNE/limbMNEvalues)in the earlierMousterian
faunasfromGrottadei Mosceriniand GrottaGuattariindicatea heavyrelianceon scav-
engingof deer carcasses.Conversely,highertransportedreturnsand higher (balanced)
frequenciesof limb to head elements in the later Mousterianfaunas from Grotta di
Sant'Agostinoand GrottaBreuilindicatea strongemphasison huntingof the sameprey
species.Variationin the PRIME/OLD mortalityratio likewisepoints to differingreli-
anceon scavenging(old-biasedpatterns)and hunting(prime-dominantpatterns).
Some additionalfeaturesof the Mousterianfaunasare of interestwith regardto the
trendsillustratedhere, as well-asto the discussionsthat follow.Data on juvenile tooth
322 A MERICANANTHROPOLOCIST

[94, 1992
eruptionand wearand the absenceof antlersuggestthat Mousterianoccupationsdating
before55,000yearsago mayhavecenteredon springand/orearlysummer,althoughthey
probablywerenot restrictedto theseseasons(Stinerl990a:660-721).Laterassemblages
fromSant'Agostinoand Breuil more clearlyrepresentfall throughlate-winteroccupa-
tions. The seasonalitydata are subjectto considerableerror,for reasonsof sampling,
latitude,and environment.These resultsnonethelessintroducethe possibilitythat sea-
sonal factors(as opposedto diachronicchanges)causeddifferencesamong the faunas,
althoughit mustalso be recognizedthat each site representsthousandsof yearsof debris
accumulation.
In markedcontrastto otherpropertiesof the Mousterianfaunas,the arraysand rela-
tive frequenciesof mammalian species consumedby hominidsdid not differsignificantly
amongsites or withinthe overallsequence(Stiner l990a:341-353, 1992a).Moreover,a
varietyof statisticalanalysesrevealno significantdifferencesin the relativedietaryem-
phaseson mammalianspeciesconsumedat caves by MiddlePaleolithichominids,spot-
ted hyenas,and (later)UpperPaleolithichumansin this regionof Italy. Suchuniformity
within the Mousteriansample,and betweenit and cave faunascreatedby other large
social predatorsin the same area, suggests that natural abundanceexerted an over-
whelminginfluenceon prey specieschoicefor all predatorsin coastalItaly. Small-scale
exploitationof 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,butfrequenciesof shellsand mammalbonesdo not predictone anotherverywell
withinthe long stratasequence(30 fine layers)of this cave (Stinerl990a:281-283).

Variation in Technology
The strongestpatternsof technologicalvariationamongthe Mousterianlithic assem-
blagesinvolvethe use of differentreductiontechniques.Figure9(a) comparesthe ratios
of platformcoresto centripetalcoresin the subjectassemblages,and Figure9(b) shows
the ratiosof the toolsand flakesattributableto thesetwo differentreductiontactics.It is
clearthat the use of platformcore techniquesincreasedover time in the Pontiniansites
at the expenseof centripetalcorereduction.It is less certainwhetherthe trendrepresents
a gradualor suddenincreasein the use of platformcore techniquesafter55,000B.P.
Figure9 also shows that core reductiontechniquesvariedfromsite to site, as well as
throughtime.Sincethe two basictechniqueswouldbe moreconvenientlypracticedusing
differentpebbleshapes(see Figure3), it is conceivablethat the observedvariationmight
simplyrepresenta highlyexpedientresponseby hominidsto the shapesof pebblesavail-
able in the immediatevicinityof each shelter.Alternatively,becauseplatformcoresap-
pearto yield moreflakesper core,it is possiblethat this techniquecouldhave been used
preferentiallyat sites wherepebbleswererelativelyscarce.
Raw materialqualitiescertainlyinfluencedmany of the generalcharacteristicsof the
Pontinianassemblages,particularlytool sizes (Bordes1968:l l 9; Taschini 1970:70-74).
The proximityof rawmaterialsourcesalso seemsto havedeterminedthe extentto which
coreswereusedup, regardlessof whichtechniquewas morecommon(Kuhn 1991).How-
ever, studiesof interassemblagevariationin the frequenciesof differentkinds of flakes
and coresindicatethat local variationin the shapesand/orabundancesof rawmaterials
doesnot accountfor variationin the frequenciesof the alternativecore-reductiontech-
niquesindependentlyofthe time element(Kuhn l990a:436-439).
Changesovertimein tacticsof flakeand tool manufacturein coastalwest-centralItaly
wereaccompaniedby declinesin both the overallintensityof raw materialexploitation
and the extent to which individualtools were used up. There is a markeddecrease
throughtime in the frequenciesof retouchon flakesand blankslargerthan2 cm in max-
imumdimension(Figure10). The medianreductionindexvaluesfor single-edgedtools
also decreasewith time (Figurel l ), indicatingthat tools wereresharpenedless oftenin
Stinerand Kuhnl ADAPTI
1ZE
VARIA
TION
INMIDDLE
PALEOLITHIC
ITALY 323

CORES: '
Platform
Centri peta I

PRODUCTS:
Plat form
Centripetal

M4 M3 G5 G4 M2 M6 G2 Br S3 S2 S1 B3

55 KY 35 KY
110KY

ASSEM BLAGE

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

the laterassemblages.Sincesedimentswerescreenedeven in the firstmajorexcavations

undertakenin this region,variationin the frequenciesof unretouchedflakes (all larger
than 2 cm) is not due to differentialrecovery.Nor can this patternbe attributedto dif-
ferencesin the availabilityof lithic rawmaterialsfromone cave to the next;assemblages
fromthe site locatedclosest to a sourceof pebbles (GrottaGuattari)exhibit relatively
highfrequenciesof retouchand high levelsof scraperreduction.
Rankordercorrelations(Spearman'srS)betweeneach of the technologicalvariables
and assemblageage confirmthe statisticalsignificanceof trendsin the lithicdata. Table
4(a) lists correlationcoefEcientsfor dated and undatedlithic assemblages,showingthat
fourof the variablesare highly correlatedwith time at the .01 level of probability.The
coefficientfor a fifthvariable(% withoutpebblecortex)is weaker,but still significant.
Table4(b) listscorrelationcoefficientsforthe datedassemblagesonly (excludesM2, M6,
 1992

324 CANANTHROPOLOCIST
A AFERI

[94,

80

PERCEN T
W ITH
60
RETOUCH
{> 2.0 cm}

40

20- pw

M4 M3 G5 G4 M2 M6 G2 Br S3 S2 S1 B3

110 KY 55 KY 35 KY

ASSEMBLAGE

Figure 10
Frequenciesof retouch on flakes and blanks larger than 2.0 cm in Mousterianlithic assem-
blages, arrangedin chronological order.

MED IAN * . :

REDUCTION 0.6- * . * v
* * v
INDEX
: v
VALUE .

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 B3

110 KY 55 KY
35 KY

ASSEMBLAGE

Figure 11
Median 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 of
the smaller array of assemblages considered, although the probabilities drop to the .05
level for most.
Another, less clearly defined pattern in the lithic data deserves mention. The use of
derived (and therefore scattered) pebble raw materials in the subject assemblages makes
Stinerand Kuhnl ADAPTIVE
VARIA
TION
INMIDDLE
PALEOLITHICB
ITALY 325

Table4
Spearmanrankordercorrelation(rS)of lithic technologyvariablesagainstincreasingas-
semblageage.

Lithievariable N rSa P
a. All assemblages
Platform/centripetal
cores 12 -0.769 < .01
Platform/centripetal
products 11 -0.882 < .01
% retouchedpieces 11 0.800 < .01
% withoutpebblecortex 12 0.634 < .05
Medianreductionindex 12 0.786 < .01
b. Datedassemblages
only(excludesM2, M6, SO,andBr)
Platform/centripetal
cores 9 -0.733 < .05
Platform/centripetal
products 8 -0.786 < .05
% retouchedpieces 8 0.833 < .01
% withoutpebblecortex 9 0.650 < .05
Medianreductionindex 9 0.778 < .05

aValuesof rj indicatecorrelationwith increasingage.

it difficultto obtain direct measuresof the distancesartifactswere moved aroundthe

landscape.However,alternativeanalysesof variationin artifactsizes,rawmaterialqual-
ities, and the relativefrequenciesof flakes,tools, and coressuggestthat the frequencies
of artifactstransportedfromsourcesof largerrawmaterials(eitheroutsideor withinthe
coastalzone) decreaseover time (Kuhn l990a:325-330;1991).Largesharp-edgedtools
madeon blanksfromcentripetalor Levalloiscoresappearto have been the items most
frequentlymovedaroundthe "Pontinian"area,as has been notedfor some Mousterian
caseselsewherein Europe(e.g., Geneste 1989;Otte 1991).
In sum, changesthroughtime in the use of centripetalversusplatformcore reduction
techniquescorrespondto differentartifact"lifehistories"in the PontinianMousterian.
An emphasison the productionof relativelylargeflakesthroughcentripetalcore reduc-
tionin the earlierassemblagesis associatedwithfrequentretouchandfairlyextensiveuse
and renewalof tools. The emphasison large flakeproductionprobablyassociateswith
fairlyhigh frequenciesof transportedartifactsas well. The increasinguse of more "eco-
nomical"platformcore techniqueswith time in west-centralItaly is actually accom-
paniedby less intensiveexploitationof tools and flakes,and by decliningfrequenciesof
transportedor "exotic"artifacts(Kuhn l 990a:325-330).

The Connections between Subsistence and Technology

The waysthatMousterianhominidsmadeand usedstonetoolsat the fourcavescovar-
ied with differencesin animal exploitation,and significanttrendsare documentedfor
both data sets. Shiftsin animal procurementand use are somewhatmore strikingand
appearmorecontinuousthan the technologicalshifts.The convergenttrendsand their
behavioralimplicationsarebestsummarizedby comparingassemblagesdepositedbefore
and afterthe 55,000-yearjuncture.
The oldergroupof assemblagessuggestsprocurementof ungulateslargelyby scaveng-
ing, basedon old-biasedmortalitypatternsand selectivetransportof head partsto shel-
ters.All head bone cavitieshad been openedwhile freshto extracttheir contents.Un-
gulateprocurementactivitiesappearto havecenteredon spring,althoughthey probably
werenot entirelyconfinedto that season."Head-collecting"is generallyassociatedwith
low-levelexploitationof marineshellfishand aquatictortoisesat Grottadei Moscerini.
Relativelylargetools and flakesproducedvia centripetalcore reductiontechniquesare
 ANTHROPOLOCIST
AMERIC}AN
326 1992

[94,
lithic artifactsexhibitevidenceof more
commonin the associatedindustries,and
more
prolongeduse and greaterfrequenciesof transport.
intense, are very differentfromthe older ones. U-n-
Thefaunasdating after55,000 years ago of livingpreypopulationsor, more
mortalitypatternsresembleeitherthe structure
gulate Substantialquantitiesof foodwere
commonly,are biasedin favorof prime-agedadults. the remainsrepresent"meaty"
to sheltersfromeach carcassobtained,andboneswerethoroughlyprocessed
transported
nearly
or completearraysof bodyparts.All of the limbwereextractedas well. The char-
marrow,
for and soft tissuesencasedin the head bones expectationsfor procurement
the
of thesefaunasare entirelyconsistentwith
acteristics l990a, l991b, l991d). Har-
studies (Stiner
hunting,as determinedfromthe control
by The associatedlithicassemblagesare
beganin falland continuedthroughwinter.
vesting yieldedrelativelylarge numbers
dominatedby platformcore reductiontechniquesthatcontextswere not modifiedor re-
in these
smallflakesper core. The tools produced
of little evidencethat they were trans-
sharpenedintensively, and there is comparatively
overlong distances.
ported betweentheformalcomponentsof tech-
Thereare no director obviousfunctionallinks Throughoutthe periodin question,the
and subsistencein theseMousteriandata. simple and transversevarieties),un-
nology
majorityof tools are sidescrapers(primarily Thereare no substantialchangesin
vast
candidatesfor foodprocurementimplements.
likely of the apparentlydifferentemphaseson
frequenciesof putative "points,"in spitethe assemblages.Instead, we find strong
the
huntingand scavengingor gatheringamong histories"of artifacts.We
between foragingstrategiesand the "life
correspondences the influenceof differingpatternsof
wouldarguethat the patternsof covariationexpress searchedfor resourcesin space.
mobilityand land use, revolvingaroundhow hominids
and durationof particularoccupations
in the frequencyof residentialmovement whereand when lithic raw materials
Shifts
would have resultedin changingconstraintsover and maintained.The advantagesof
couldbe gatheredand where tools could be made wouldvary accordingly.
alternativetacticsof tool manufactureand renewal a strategy
Mousterianfaunas,is rwormally
Passivescavenging,evident in the earlier tendsto yield lowertransportablereturns
targetsrelativelydispersedresourcesand
that hunting,both in energeticreturnsand the
onthe average.It is morelike gatheringthan scavengingimpliesmorefrequentmove-
patternsthat may be required.Passive
ranging on the part of hominids,regardlessof
mentand perhapswide-rangingsearchpatterns or (morelikely)merelytakenad-
whether scavengingopportunitieswereactivelysought that passivescavengingof ungu-
vantage of when encountered.It seemsno coincidence
moreor less sessileanimalsat one coastal
latepartsoccursalongsidecollectionof small, did not drivehominidsto expand
Mousterian cave. An interestin scavengingprobably
alreadyrangingwidelyfor some other ener-
theirforagingradii,but becausethey werebothfeasibleand rewarding.Centripetalcore
geticreason(s),thiskindof scavengingwasin the contextof relativelyextensiveforaging,
reductionwouldhave been advantageous blankssuitablefor prolongeduse in a raw-
becauseit produceslargerflakesand tooltool reductionand transportindeedsuggests.
material-poor environment,as evidenceof
persistentreductionor resharpeningis,
Prolongingthe potentialuse life of toolsthrough
aboutjust when the tools will be needed
as
in turn,a way of copingwith uncertainties to makenew toolswill arise.
opposedto whenand wherethe next chance assemblagesincludesfar richertrans-
Evidencefor huntingin the later Mousterian
may imply targetingrelativelyconcentrated
portedreturnsper procurementevent and ungulatespecies tend to congregatewhile
foodpatches.Afterall, deer and most other carcassesgenerallydo not do so of their
alive,especiallyin fall and earlywinter,whereas the nearlycompletearraysof anatom-
own accord,at least not continuously.Moreover, discardof less economicalbody parts
ical elementsindicateminimallevels of selective rangingfor food and longeroccupations
priorto reachingthe shelters.More restricted
madeit less importantto producethe largest
provlslonedwith huntedmeatwouldhave insteadmadegreaternumbersof rela-
mostdurableflakesfrompebblecores.Hominids
Stinerand Kuhn] ADAPTIVEVARIA
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THICI TALY 327

tivelysmall tools destinedfor light use and little transport.Proeessingof huntedgame,

whiehcertainlytookplaceat the later Mousterianshelters,may also haveset a premium
on unmodifiedor lightlyretouehededgessuitableforcuttinganimaltissues.Thus, in the
PontinianMousterian,bigger tools and tool blanks were seleeted only when people
neededthingsto last, sueh as when they had to carrytools aroundfor extendedperiods
and the aetualmomentsof need wereunpredictable.
Accumulation RatesofBonesversusStones
Informationon the varyingratesat which bones and stones accumulatedin strataof
the fourcaveslends additionalperspectiveon the relationshipbetweenanimalexploita-
tion and technology.We assumethat the totalamounts of both kinds of materialsin the
culturaldepositsare largelya consequenceof two conditions:( 1) the frequencyor inten-
sityof hominiduseofthe cavesites;and (2) thevolumeof sedimentexcavated.The relative
proportionsof bonesand stonesamongstratashould,on the otherhand,relatein someway
to the economicrelationshipbetweenthemindependentlyof sedimentationrates.Recall
that the confoundinginfluencesof bone preservationwere consideredin advance and
were not found to be suflicientlygreat to prohibitthis kind of comparison(see Stiner
1990a, 1991a, also discussionsin 1991b:456-467).
If the maindifferencesbetweenaccumulationratesof bonesandstonesbeforeandafter
55,000B.P. were tied to the relativeincidenceof huntingversusscavenging,we would
expecttheirrelativefrequenciesto show positivecorrelationsbut differentslopes,in ac-
cordancewith time period.This is becausescavengingby itself (i.e., withoutplant and/
or smallanimalsupplements)shouldnot operateat a loss, but wouldtend to offerfewer
transportablepartsthan huntedsources(Stinerl 991b, l 991d).
Figure 12 plots the total frequenciesof Mousterianstone tools, flakes, and cores
(tLITHICS)againstthe total numberof bones (tMNE) forall ungulatespecies(bovids,
equids, and cervids)6in each assemblage.Triangles representthe earlierMousterian

w>55KYears f
* < 55 KYears

800- /

t LITHICS v /

100 200
tM NE, all ungulate taxa

Figure 12
Scatterplotand regressions for combined ungulate tMNE (cervids, bovids, and equids) and
the frequency of lithic artifacts (tLITHICS = tools + flakes + cores) for Mousterian as-
semblages before (dashed line) and after (solid line) 55,000 years ago.
328 A3IFRI(ANANTHROPOLO(;IST [94, 1992

casesfrom Moscerini(level groupsM2, M3, M4, and M6) and Guattari(G2, G4, and
G5), dating between 110,000and 55,000 years ago. Squaresrepresentlater cases from
Breuil (Br and B3) and Sant'Agostino(S1, S2, and S3), dating between 55,000 and
35,000years ago. It is immediatelyclearfrom the graphthat the relationshipbetween
faunaland lithicabundancesis not the sameforthe two groupsof assemblages.The cor-
relationbetweentMNEand tLITHICSforthe earliergroupis not statisticallysignificant
(r2= 0.084,P = .577,N = 7), indicatingthatthe quantitiesof stonesandungulatebones
varymoreor less independentlyof one another.Hominids'discardof stoneat thesecaves
is not directlyexplainedor predictedby their use of ungulateresources.To the extent
thatthereis anyrelationship,however,it appearsto be negative(dashedregressionline),
and hints at the existenceof some componentof subsistencenot directlymonitoredby
the data forlargemammalremains.
In contrastto the earliergroup,bonesand stonesoccurin roughlyequal proportions
in the youngerMousterianassemblages.These cases displaya strongpositiverelation-
ship (solid regressionline) betweentMNE and tLITHICS (r2= .986, P-.007, N= 5).
The S1assemblageis much largerthan the others,hence representingan outlierto the
graph,but it is consistentwith the generalrelationshipas described.The strongpositive
relationshipfor this later group of assemblagessuggests that lithic material cycled
throughthe system more or less in tandem with the numberof ungulatebody parts
broughtto the sheltersby hominids.
The frequenciesof faunal and lithic debris suggest that, prior to 55,000 B.P., the
amountof toolmakingthat went on in the Italiancaves didnotdependon successin ob-
tainingscavengedungulateparts.Whileconspicuousfroman archeologicalperspective,
scavengingmay have in fact been ancillaryto subsistenceactivitiesat Mosceriniand
Guattari.It morelikelywas integratedwithin a wide-rangingforagingpatternthat ac-
tually focusedon a varietyof dispersedsessile resources,includingsmall quantitiesof
marineshellfish,tortoises,and probablyvegetablefoods.In contrast,the amountoftech-
nologicalactivityrepresentedin the laterassemblageswasresponsiveto the quantitiesof
ungulatepartsprocuredand subsequentlycarriedto the shelters.Huntedgamemay ac-
tually have been centralto subsistenceduringoccupationsat Grottadi Sant'Agostino
and GrottaBreuil.
HeadParts,Scavenging,
andFat
Mousterians'exceptionalinterestin head partswhen apparentlyscavengingis unex-
pectedin the sense that we are not able to find compellingevidenceof this foragingpat-
ternamongmodernhunter-gatherers. Isolatedinstancesof specialinterestin head parts
may happennow and again in modernsituations,but not such that the practicedomi-
natesentirefaunalsequencesin deeplystratifiedshelterdepositsformedover thousands
of years.To understandwhat "head-collecting"might have been about, it is necessary
to turnto twoothersourcesof information,the habitsof nonhumanpredatorsand details
of the ungulateanatomy.
Among the carnivores,a "prefercnce"for head parts (of medium-sizedungulates)
oftenemergesin the contextof scavenging(Stiner l991b). It is partlyexplainedby the
relativepersistenceof craniaat findsites (e.g., Haynes 1980;Blumenschine1986:35-38)
and the easewith whichheads (includingthe mandible)can be removedfromcarcasses,
regardlessof theircondition(Stinerl 990a:397-399).
Thereareenergeticincentivesas well. Headsare complexbonycontainersand require
considerableprocessingby humansand carnivoresalike.So, movinghead partsto a se-
cureplacewherethey can be workedon in peace can be worthwhile.Equallyimportant
anywherefoodenergysourcesare periodicallyor perenniallyscarceis the uniquelyhigh
fat contentof soft tissuesin the head, muchof whichpersistseven in malnourishedprey
(Stiner l991b:471-474, 1992b). The fat/proteinratio in head tissues, particularlythe
brain,is both high and stablethroughoutthe year, becausethe fat-richmyelin sheaths
enclosingthe cranialnervescannot be metabolizedunderconditionsof food stress. No
Stinerand Kuhn] ADAPTIVE
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ITALY 329

part of the postcranialprey anatomyis like this; instead,fat/proteinratioscycle in ac-

cordancewith seasonalshiftsin foodsupply.Also significantis the fact that yellowmar-
rowreservesin the adult ungulatemandibleare amongthe last to be drainedmetaboli-
callyin timesof foodscarcity.
Humanbeings need to consumesubstantialamountsof fat and/or carbohydratesin
orderto utilizedietaryproteinseffectively(Spethand Spielmann1983). Becauseplant
productionnaturallywaxesandwaneswiththe seasons,animalsourcesoffat can become
very valuableto humans,and Neandertalswould have been no exception.Under con-
ditionsof scarcity,then, the nutritionalstate of the animalat the time of death should
stronglyinfluencehuman(andcarnivore)decisionsaboutwhatpartsareworthyof trans-
portand extensiveprocessingat anotherplace, such as at a rock-shelteror cave.
Nutritionalcontingencieswould not be confinedto scavengingsituations,however.
People,whetherhuntingor scavenging,shouldalwaysbe interestedin headsif fats and
carbohydrates are scarce,and ethnoarcheological accountsshow that they are (e.g., Bin-
ford 1978;O'Connell,Hawkes,and BlurtonJones1988a;Yellen 1977). Peoplemay be-
comeless interestedin heads if otherrich, morefavoredpartsrequiringless processing
efTort areavailable,a factthat undoubtedlyaccountsformuchvariationin transportand
processingdecisionsfromone monthto the next, dependingon the amplitudeof the sea-
sons. Binford's(1978:60-61,77-90) descriptionsof ungulatebutchering,transport,and
processingby NunamiutEskimorevealconsiderablevariationby season;the Nunamiut
weremuch moreinterestedin head parts in springthan they were in the fall. Hunter-
gathererslivingin tropicalregionsdisplaymoreconsistentinterestin everypossiblefat-
yieldingpartof prey (e.g., Yellen 1977).
Whilepeoplecould conceivablychooseto takeonly heads away fromstarved prey ob-
tainedby hunting,the effortto procurethem can quicklyoutweighthe potentialreturns
(sensu Metcalfe1989) because,rich or not, the head is a relativelysmall package.This
reasoningassumesan importantstrategicdifferencebetweenpassivescavenging,on the
one hand,and huntingand aggressivescavenging,on the other;the valueof eitherclass
of foragingstrategydependson the nutritionalstate and bulkmass of preyas most typ-
icallyencounteredin a given foragingsystem.Thus, thereis a much lower probability
thathead-dominatedfaunasrepresenthuntingby hominidson energeticgrounds,in ad-
ditionto empiricalsupportaboutwhat scavengingsocial carnivoresreallydo in current
settings. But, if carnivoresand Neandertalscould produce strongly head-dominated
faunasat some sites, why is it that modernpeopledo not?
The answermay hingeuponthe rangeofprocessing optionsavailableto humans,many
of whichhave been aroundat least since the Upper Paleolithic.Here we referto objects
associatedwith grinding,pounding,and/or boilinganimaland plant foods. Most of the
evidencein the Paleolithicforfoodprocessingtechniquesis indirect,suchas fire-cracked
cobblesthatmighthavebeenusedforstoneboiling.In timesoffat scarcity,bonetrabecu-
lae of the spinalcolumnof adultpreyalso representpotentiallyrichsourcesof fats (e.g.,
Speth 1991),and could have been processedby grindingthem up and boilingthem, as
they are in certainmodernhumansituations(e.g., Binford1978;Yellen 1977).For rea-
sonsthatarenotfullyunderstood,researchon the Mousterianof Europehas yieldedvery
limitedevidenceof grindingtechnologiesor stone boiling. Heads, in contrast,can be
roastedand pickedapart;processingrequirestimeand firebut no additionaltechnology
forextractingnutrients.The absenceof containertechnologyand grindingor pounding
implementsin the Mousteriancould have made heads even moreattractiveto foragers
in needof fat.
Thus, in the Italian Mousterian,strictchoicesto take awayjust about only heads to
two of the caves may have been due to an advancedstate of starvationin deer, making
themverypoorin fat at a time when this nutrientwas neededto supplementthe human
diet (Stinerl991b, 1992b).That the opportunitiesto scavengeweregeneratedprimarily
by starvationratherthan by huntingcarnivoresis supportedby the moreor less equiv-
alentfrequenciesof craniato mandibles(see Stiner l991b:462,471; comparewith Blu-
330 AMERICAN
ANTWROPOLOGIST

[94, 1992
mensehine1986on predator-generated opportunities).Huntingeapabilitymayhavehad
little or nothingto do with Mousterians'deeisionsto seavenge.Rather,foragingeondi-
tions made seavengingmore worthwhilegiven the rangingpatternsand teehnologieal
supportin existeneeat that time. Seavengingis reallya familyof taetieswhosediversity
and eeonomierules are not well-understood.Clearly,teehnologiealeonsiderationsean
greatlyaugmentany effortto investigatethis elass of foragingbehaviorsamonganeient
and modernhominids.
What
DoHunters
NeedinOrder
toSucceed?
The rangeofformaltool types (as opposedto reduetionteehniques)in the Mousterian
samplestays about the same aerossthe 55,000-yearboundary,while subsisteneestrate-
gies appearto ehangeradically.Why isn't therean equallydrasticchangein the kindsof
toolsbeingprodueed,eitherin the weaponsfordispatehinganimalsor in the implements
usedto makeweapons?
We are not in a positionto eompletelyanswerthis question,but recentstudiesshow
that inereasingsophistieationof formalweaponsused by modernhuman huntersand
"huntingsuecess"(as measuredby preynutritionalreturns,age elassestaken,etc.) vary
somewhatindependentlyof one another.Thereis morethanone solutionformaintaining
regularaeeess to high-qualitylive prey. In surveysof ethnohistoriealand areheologieal
eases,Stiner(199Ob,l991d) documentsanalogouspatternsof ungulatepreyage seleetion
by 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,findingthat greaterlevels of cooperation
amongbow-hunterscaneffectivelysubstitutefor the advantagesconferredby long-range
weapons,such as shotguns.Of course,neitherbows nor shotgunswouldhave existedin
MiddlePaleolithictimes.But, if cooperationcan in anywaysubstituteforthe advantages
thatsophisticatedmodernweaponsoffer,then we are obligedto allowfor the possibility
thateooperationamongMousterianforagers(whoereated"modern"-looking preymor-
talitypatternsin somesites) workedin basicallythe same way.
Humanhuntersare not entirelyweapon-dependent; they are only partlyso. The ram-
ificationsof these two views are quite different,and failureto appreeiatethe differenee
couldpresentseriousobstaclesto understandingthe relationshipsbetweenvariationin
humantechnologyand foragingstrategies.There is no questionthat weaponsean faeil-
itate the food quest. It is beeomingincreasinglyclear,however,that prey seleetionand
gettingenoughto eat are produetsof a complexinterplaybetweenthe teehnologiealsys-
tem in use, demography-mediated possibilitiesfor humaneooperationduringprocure-
mentand subsequenttasks,and speeies-and seasonal-speeifiebehaviorsof the animals
pursued(Binford1978;Frison 1991;Hudson 1991;Lyman 1991;Stiner l990b, l991c,
l991d).
In sum, findingevideneeof increasingemphaseson huntedprey in the Mousterian,
includingprime-adultharvesting,need not demand a correspondingincrease in the
quantityof stone weaponheadsproducednor an increasingsophisticationin the typesof
weaponsused.It couldhappen,but it doesnot haveto happen,becausemanyadvantages
of"formal"technologyoverlapwiththoseof beingan intenselysoeialspecies.The Italian
Mousteriandata seem to illuminatea substantialblindspot in areheology,lyingjust be-
tweenthe specialtiesof zooarcheologyand teehnologyresearehas eurrentlypracticed.

Conclusion
Currentresearchprioritiesfor the Mousterianand debatesabout the Middle-Upper
Paleolithictransitioncenter on questions about adaptive variationand evolutionary
change.In contrast,mostextantdatawerecollectedwithverydifferentaimsin mindand,
. . . .

Stinerand Kuhn] ADAPTI

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INMIDDLE
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THIC
I TALY 331

in manycases,are of questionablerelevanceto theseissues.The researchdescribedhere

approachesvariationin the Italian Mousterianby combininga uniqueset of variables
that are demonstrablyrelevantto how hominidsused resources.The faunalvariables
werechosenbasedon cross-speciessurveysof modern(humanand nonhuman)preda-
tors,and the technologicalvariableswereselectedbasedon expectationsabouthow tech-
nologyshouldrelateto humanforagingpractices.This researchorientationrevealscon-
siderablevariationin the manufactureand use of stonetoolsin "Pontinian"Mousterian
assemblagesthat closelyparalleledshiftsin game use, especiallypatternsof preyage se-
lection and body-parttransport.Such variationexists within an archeologicalrecord
that,fromthe classicalperspectivesof tool typologiesand speciesrepresentation,is quite
monotonous.
It is clearfromthis researchthat somefacts about the MiddlePaleolithiccan only be
understoodthroughsimultaneousconsiderationof zooarcheologicaland technological
evidenceat the intersitelevel. Collaborationrevealsseven importantthings about the
relationbetweenMousteriantechnologyand subsistencepracticesin west-centralItaly:
1. Strongcovariationexists betweenartifact(coreand tool) "lifehistories"and hom-
nlc . toraglng strategles.
2. In situationsthat apparentlyinvolvedscavenging,Mousterianhominidspreferred
to transportheads (but not manyotherungulateparts)back to shelters.This relatively
strictchoicewas responsiveto the uniquenutritionalassetsofferedby cranialsoft tissues
in timesof fat or carbohydratescarcityand, perhaps,the limitationsset by extantpro-
cessingtechnologyduringthat culturalperiod.
3. Thereare no completeor necessaryconnectionsbetweenmanyof the mostobvious
(i.e., visually striking)aspects of weaponstechnologyand the relativedependenceon
huntedlargeprey,eitherin the Mousterianor in modernhumansituations.This obser-
vationis especiallycriticalfor researchon the Middle Paleolithic,whereanalystsoften
deal with questionsof presenceor absenceof stone-tippedprojectiles.
4. Patternsin lithic technologyand animalexploitationconvergemost clearlyat the
level of mobilityand land use. More significantthan any findingabout "hunters"and
"scavengers,"per se, is the spatialelementimpliedby thesedata. Covariationobserved
betweenthe lithicand faunaldata sets appearsto reflecthow foragersused territory,the
distributionof opportunitiesto collector hunt variousclassesof resources,and ways of
ofEsetting needsfor key but sometimesscarceresources,such as protein,animalfat, and
workablestone.
5. It is also clearthat the trendsobservedin the MiddlePaleolithicof Latiumrepre-
sent shiftsin the frequenciesof variousaspectsof technologyand foraging,not the ap-
pearanceof entirelynew componentsof behavior.For example,evidenceof both centri-
petal core reductionand platformcore reductionare presentthroughoutthe Pontinian
sequence;platformcore reductionsimply overtakescentripetaltechniqueswith time.
This is arguedto be an adjustmentof tool manufactureto the requirementsand limita-
tionsof foragingand land use, an adaptivefrequencyshift withinan extant rangeof al-
ternatives.
The same must be said for scavengingversushunting.Mousterianscavengingprac-
tices appearodd in light of what modernhumansdo, but thereis no reasonto assume
that Mousteriansnever scavengedafter 55,000 years ago, or never hunted beforethis
time. Virtuallyeveryknownpredator,includingmodernpeople,engagesin both;pred-
atorspeciesdifferin theiremphaseson huntingand scavengingin what is best described
as a continuum(Stiner l991b, 1992b).Juxtaposinghuntingand scavengingas if they
representeddistincthominidpredatorynichesor adaptations,ratherthan components
of any one, is counterproductive. In otherwords,we shouldexpect to see some combina-
tion of these two generalstrategyclassesin hominidadaptationsthroughoutthe Upper
Pleistoceneand perhapsearlier.Variationin how hunting and scavengingtactics are
manifestin foragingadaptationsis a problemof a finerorder,as researchon modern
332 AAIERICAN
ANTHROPOLOGIST

[94, 1992

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-central
Italy comparedto anatom-ically modernhumanlifeways,it is in how alternativetactics
of gameprocurementand tool manufacturewereimplementedas componentsin a larger
adaptivesystem. For example,while scavengingis conspicuous in the earlierpart of the
Italiansequence,it probablywas not the only option for these foragersor their single
mainstay.The Mousterianscavengingpatternstandsapartfromknownmodernhuman
casesbecauseit was so clearlyseparated,spatiallyand temporally,fromhunting.It pre-
sentsa strikingcontrastto ethnoarcheological caseswherethe productsof both strategy
classesarepooledat the sameresidentialplaces(e.g., Binford1978;O'Connell,Hawkes,
and BlurtonJones 1988a, 1988b;Bunn, Bartram,and Kroll 1988). Becauseungulate
headpartsrepresenta consistentlyrichsourceof fats, and hominids'basicphysiological
need for dietaryfat probablyhas not changed,the Mousteriandata suggestthat there
was somethingvery differentabout the ways ungulateexploitationoffsetthe use of the
otherresources(Stiner l991b, 1992b). Independent(possiblymildly negative)rates of
accumulationfor bonesand stonesin the caves also suggestthat ungulateswerenot the
coreof subsistencein sites wherepassivescavengingis evidenced.Deer head partsmay
insteadhave merelysupplementedshellfishand otherarcheologicallyinvisiblefoodsde-
hcoent1ntats.
, . . ,>

As concernsthe contextof passivescavengingin the Italian Mousterian,seasonality

dataareof limitedhelp. Death by starvationin ungulatepopulationspeaksaroundearly
springin coolerenvironments(seereviewin Stiner1990a:595-600),andfat consequently
becomesrarerfor omnivorouspredatorsbeforeplant productionis in full swing each
year.The head-dominatedMousterianfaunasindeedappearto center(witha widerange
of variation)on spring.However,abstractingthe seasonalityresultsto explainintersite
variationin hominidforagingactivitieswould be much too simplistic:the intersitedif-
ferencesin resourceuse lasted thousandsof years,probablyspanningseveralminorcli-
maticoscillations.
7. Withinthe geographicallimits imposedby the study sample,the observedtrends
in foragingand technologydo appearsomewhat"progressive."Scavengingis "replaced"
by hunting,and classic Mousteriancentripetalreductiontechniquesare supersededby
single-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 technological
changesmay indeedrepresentresponsesto alterationsin foragingpatterns,but the ulti-
mate causes of long-termvariationin subsistenceremain unclear.The trends in the
Mousteriancouldrepresentsimplechangesin how theseparticularcave sites wereused
or, alternatively,the trendscould reflecta permanentevolutionaryshift in hominidfor-
agingnichewithinthe Mousterian.Oscillationsin sea level between110,000and 35,000
yearsago certainlyalteredthe settingof each shelterand probablyits utilityto hominid
foragers.Duringwarmclimatephases, all of the caves would have been situatedquite
closeto the beachjust as they are now, whereasin colderperiodsthey wouldhave been
separatedfromthe sea by ribbonsof dunes, coastalplain, and/orsmallmarshylagoons.
The trendswe see couldsimplyreflecthominids'adjustmentsto localchangesin habitat,
causedby gradual,if irregular,loweringof the MediterraneanSea.
Becausethis studydealsonly withone smallregion,it wouldbe a mistaketo generalize
the trendsto the Mousterianas a whole.The Middle-UpperPaleolithic"transition"by
definitiontookplaceovera widearea.The FrenchMousterianshowsneithera consistent
declineover time in the frequenciesof retouchedtools (Rolland 1988:170)nor a clear
increasein the productionof blade-likeflakesand tool blanks(cf. Harrold1989:691;Pe-
legrin 1990). Our recent researchon Mousterianassemblagesfrom Riparo Mochi, a
deeplystratifiedcoastalshelterin the BalziRossinearthe French-Italianborder,reveals
a technologicaltrendcontraryto that observedin the Pontiniancases, namelya decrease
StinerandKuhn] ADAPTIVEVARIA
TIONIN MIDDLEPALEOLITHICL
I TALY 333

throughtime in the use of bladesor blade-likeflakes.The Mochi lithic assemblagesare

as yet undated,however,so theirehronologicalrelationshipto the Latiumeavesremains
unelear.
Of the two possibleexplanationsfor why the trendsexist adjustments by hominidsto
changesin local resoureeopportunitiesversusevolutionarychange we preferthe first,but
we arenot in a positionto advanceit as a eertaineonelusion.The methodsand basesfor
interpretationpresentedin this study help providemeans for addressingthe two hy-
potheses,but rigoroustestingwill requireexpansionof the geographicalscaleconsidered
or, at least, examininga largerarrayof long stratigraphicsequencesin the studyareaas
heredefined.This is an importantstimulusfor continuedresearchin west-centralItaly
(e.g., the thickbut as yet unexcavatedportionsof GrottaBreuil),as well as in other re-
gionsof the Mediterranean.
Regardlessof whetherthe trendsobservedin west-centralItaly prove to represent
broaderevolutionarychangesin Mousterianbehavioror merelyroutineadjustmentsto
localizedchangesin foragingconditions,the data clearlydemonstrate- that the Middle
Paleolithicwas neithermonotonousnor immutable.When examinedin termsof vari-
ables that can be expectedto show evidenceof adaptiveresponse,Mousterianforaging
practices,use of places, and technologywere quite variable.The findingsof this study
provideinsightsinto the kinds of adaptive"rawmaterial"alreadyin place duringthe
MiddlePaleolithic.

Notes
Acknowledgments. We thankLewisBinford,Diane Gifford-Gonzalez, LawrenceStraus,and Erik
Trinkausfor theircriticalinputthroughoutthe courseof our research.Commentsby threeanon-
ymousAA reviewershave made this a betterpaper.We are deeplyindebtedto our Europeancol-
leagues,Aldo G. Segre,EugeniaSegre-Naldini,AmilcareBietti, PieroCassoli,DaniellaCocchi,
GiorgioManzi, MarcelloPiperno,AntonioRadmilli,and CarloTozzi, withoutwhoseassistance
the researchsimplywouldneverhave happened.The studiesweresupportedby grantsfromthe
L. S. B. LeakeyFoundationand the Institutefor InternationalEducation(FulbrightFellowships)
to Kuhn and to Stiner,the AmericanAssociationof UniversityWomen(to Stiner),and the Na-
tionalScienceFoundation(dissertationimprovementgrantto Stiner,BNS-8618410).
lThe term "strategy"is employedin an abstractsense in this study, as used in evolutionary
biologyand strategicmodeling(e.g., Toobyand DeVore 1987).Behavioris analyzedas if it were a
goal-orientedeconomicstrategy.The actualmotivationor proximatecauseof the behavioris not
assumedto be ideationalor even intentional.
2Reddeer are much morecommonthan fallowdeer in all of the faunalassemblages.The two
taxa are combinedinto one prey categorybecausemanyskeletalelementfragmentsof these two
deerspeciescannotbe distinguishedto the specieslevel.
3Theanalyst'schoiceof landmarksfordetermininglimbMNEcan affectthe numbersperceived.
In recognitionof this potentialproblem,controlledcomparisonsof the relativefrequenciesof epi-
physeallandmarksversusmidshaftforaminaewereconductedforfaunasexcavatedfromthreecave
sitesin the studyarea,each involvingcompleterecoveryof bone.These are recentexcavationsat
GrottaBreuil(MiddlePaleolithic),RiparoSalvini(late Upper Paleolithic),and Bucadella Iena
(a seriesof Upper Pleistocenecarnivoreden occupations,primarilyspotted hyenas). The sites
sharesimilarsedimentarysettings.In all cases, the epiphysis-basedmethodyieldedequivalentor
superiorMNE countsfor limbs relativeto counts basedon midshaftlandmarks(primarilyfora-
minae). The resultsof this controlstudyjustifythe use of the epiphysis-basedcountingmethodfor
the oldercollections,which wererecoveredusing systematic,but moreselective,criteria(Stiner
1991b:460-462).
4ThefaunalassemblagefromG2 of GrottaGuattarirepresentssome mixtureof hominidand
spottedhyenacomponents(Stiner l991a), and thus is not an ideal case for this comparison.G2
exhibitsa numberof qualities(manyof whichare not presentedhere) that morecloselyresemble
hominid-associated patternsoffaunal exploitationin neighboringsites than patternstypicallyas-
sociatedwithspottedhyenas.Forthesereasons,it has beenincludedin a sampleof otherwisemore
definitelyhominid-associated cases. Acceptanceor deletionof this case does not underminethe
overallresultsof the study.
334 A AfERlC'ANANTHROPOLOCIST

[94, 1992
5Thetwocorereductionschemesrepresentidealizedmodels,basictacticsfortakingaparta piece
of stone.In reality,a singlecorecan changeformin the courseof its usefullife, althoughthe small
raw materiaIsavailableto hominidsin the study area probablylimitedthe extentof these trans-
formations.It is clearin somecasesthatplatformcoresweremodifiedintosomethinglikecentripe-
tal coresnearthe end of the reductionsequence(Bietti,Rossetti,and Zanzi 1989),as observedin
Mousterianindustriesfromotherregions(e.g., Baumler1988).FIowever,by examiningvariation
in the frequenciesof bothcoresand the flakesor toolsmadefromthem,it is possibleto controlfor
the kindsof flakeproductiontechniquesused before coresreacheda state of exhaustionand were
discarded.
6Thisarticlefocuseson red and fallowdeerremainsbecausetheyare verycommonin all of the
faunalassemblages.The generalpatternsof exploitationby hominidsnonethelessapply to the
otherungulatespeciesrepresentedin theseassemblages.

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