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* Research Laboratory for Archaeology and the History of Art,
University of Oxford, 6 Keble Road, Oxford OX1 3QJ, United Kingdom;
Contributed by Erik Trinkaus, April 19, 2000
Archeological analysis of faunal remains and of lithic and bone
tools has suggested that hunting of medium to large mammals was a major
element of Neanderthal subsistence. Plant foods are almost invisible in
the archeological record, and it is impossible to estimate accurately
their dietary importance. However, stable isotope ( paleodiet | Croatia | Europe | Reconstructions of European
Neanderthal subsistence strategies have overwhelmingly focused on the
specialized hunting and scavenging of herbivores as the predominant
method of obtaining food (1-6). These reconstructions are based
principally on the analysis of the abundantly preserved faunal remains,
supplemented by artifactual evidence of lithic and wood hunting
apparatuses, as well as on the relative importance of the faunal
biomass in the environments that European Neanderthals occupied during
later oxygen isotope stage 5 and especially oxygen isotope stages 4 and
3 of the Late Pleistocene. Understanding Neanderthal diet has
implications for understanding Neanderthal land use, social organization, and behavioral complexity. Yet despite the abundant evidence for successful hunting techniques across Neanderthal Eurasia,
faunal remains can indicate only hunting or scavenging episodes; they cannot tell us about the predominant foods in
the diet over the long term.
By contrast, the measurement of the ratios of the stable isotopes of
carbon and nitrogen in mammal bone collagen provides an indication of
aspects of diet over the last few years of life (7-9). This stable
isotope evidence can therefore provide us with direct
information on Neanderthal diet. This method has been applied to
Neanderthal remains from the sites of Marillac, France (10), and
Scladina Cave, Belgium (11). These studies, focusing particularly on
their high Vindija Neanderthal and Faunal Specimens.
Recently, the Vi-207 and Vi-208 Neanderthal specimens, as well as
various other archeological materials from level
G1 of Vindija Cave, Croatia, were submitted for
accelerator mass spectrometer radiocarbon dating at the Oxford
Radiocarbon Accelerator Unit, University of Oxford (13). The two
Neanderthal specimens were dated to 29,080 ± 400 years before
present (B.P.) (OxA-8296, Vi-207) and 28,020 ± 360 years B.P.
(OxA-8295, Vi-208), making them the youngest directly dated Neanderthal
specimens in Europe (13). Because the radiocarbon sample preparation
process includes assessments of stable isotopes, in part to control for
potential contamination, this analysis also yielded stable isotope
profiles for these late archaic humans. Combined with similar data
obtained from faunal remains from level G1 and
the older level G3 of Vindija Cave, this provides
a means of assessing the dietary profiles of these Neanderthals.
Stable Isotope Analyses.
Mammal bone collagen Vindija Neanderthal and Faunal Isotope Values.
Collagen was extracted from the two Neanderthal specimens from level
G1 of Vindija Cave and from various faunal
remains from level G1 and the older level
G3 according to standard collagen extraction
procedures; the Neanderthal specimens were extracted according to the
methods outlined in Law and Hedges (19), and the faunal specimens were
extracted according to the procedure outlined in Richards and Hedges
(16). The collagen extracts varied in quality, and only those samples
that had acceptable collagen attributes were used. These attributes are
based on values determined by DeNiro (20) and Ambrose (21) and used by
the majority of stable isotope researchers and radiocarbon dating labs.
The acceptable values are a C:N ratio between 2.9 and 3.6, "percent
collagen" >1%, and %C and %N in the extracted collagen of >13%
for carbon and >5% for nitrogen. These collagen attributes allow us
to identify and exclude collagen that is heavily degraded or
contaminated. This is in contrast to stable isotope measurements of
bioapatite in bone mineral and enamel, where no such criteria exist.
The stable isotope values and various collagen attributes are given in
Table 1; based on these, we are confident
that the collagen
From the Cover
Anthropology
Neanderthal diet at Vindija and Neanderthal predation: The
evidence from stable isotopes
,
,
,
, and
Department of Archaeology, Simon Fraser University,
Burnaby, BC V5A 1S6, Canada; Keble College, Oxford OX1
3PG, United Kingdom; § Department of Anthropology, Campus
Box 1114, Washington University, St. Louis MO 63130;
¶ Unité Mixte de Recherche 5809 du Centre National de
la Recherche Scientifique, Laboratoire d'Anthropologie,
Université de Bordeaux I, 33405 Talence, France; ** Department of
Anthropology, Northern Illinois University, DeKalb, IL 60115;
Zavod za paleontologiju i geologiju kvartara, Hrvatske
akademije znanosti i umjetnosti, Ulica A. Kova
ia 5/II,
HR-10000 Zagreb, Croatia; and Arheolo
ki zavod
Filozofskog fakulteta Sveuilita u Zagrebu, I. Luia 3, HR-10000 Zagreb, Croatia
Abstract
Top
Abstract
Introduction
Summary and Conclusions
References
13C
and 15N) analysis of mammal bone collagen provides a
direct measure of diet and has been applied to two Neanderthals and
various faunal species from Vindija Cave, Croatia. The isotope evidence
overwhelmingly points to the Neanderthals behaving as top-level
carnivores, obtaining almost all of their dietary protein from animal
sources. Earlier Neanderthals in France and Belgium have yielded
similar results, and a pattern of European Neanderthal adaptation as
carnivores is emerging. These data reinforce current taphonomic
assessments of associated faunal elements and make it unlikely that the
Neanderthals were acquiring animal protein principally through
scavenging. Instead, these findings portray them as effective predators.
13C | 15N
Introduction
Top
Abstract
Introduction
Summary and Conclusions
References
15N values, indicated that the
Neanderthals measured occupied the top trophic level, obtaining nearly
all of their dietary protein from animal sources. In the context of
this finding, we undertook stable isotope analyses of the two late
Neanderthal specimens from Vindija Cave, in the Hrvatsko Zagorje
of northern Croatia [Vi-207 and Vi-208 (12)], and of the fauna
with which they were stratigraphically associated.
13C and
15N values reflect the
13C and 15N values of
dietary protein (14). They furnish a long-term record of diet, giving
the average 13C and
15N values of all of the protein consumed over
the last years of the measured individual's life.
13C values can be used to discriminate between
terrestrial and marine dietary protein in humans and other mammals (15,
16). In addition, because of the canopy effect, species that live in
forest environments can have 13C values that
are more negative than species that live in open environments (17).
15N values are, on average, 2-4
higher
than the average 15N value of the protein
consumed (18). Therefore, 15N values can be
used to determine the trophic level of the protein consumed. By
measuring the 13C and
15N values of various fauna in a
paleo-ecosystem, it is possible to reconstruct the trophic level
relationships within that ecosystem. Therefore, by comparing the
13C and 15N values of
omnivores such as hominids with the values of herbivores and carnivores
from the same ecosystem, it is possible to determine whether those
omnivores were obtaining dietary protein from plant or animal sources.
13C and
15N values reported here are robust and
reflect the organisms' original collagen 13C
and 15N values. The Neanderthal samples were
measured at the Oxford Radiocarbon Accelerator Unit, and the faunal
samples were measured at the Stable Isotope Laboratory, Research
Laboratory for Archaeology and the History of Art, University of
Oxford.
Table 1.
Bone collagen 13C and 15N
values of Neanderthals and associated fauna from Vindija Cave,
Croatia
23,000-26,000 B.P.) sites of Dolní V
stonice II
and Milovice in the Czech Republic (22). In addition, we have
contributed a single herbivore sample from the site of
Brno-Francouzská, which dates within this time range (23).
There are fluctuations in faunal 15N values
through time that are correlated with climate changes (24, 25). For
example, Richards et al. (26) observed faunal
15N values dated to 12,000 years B.P. from
Gough's Cave, U.K. that were 2 lower than the
15N values of similar species from the
Holocene. Therefore, comparing isotope values between sites, especially
sites of different ages, could be problematic. However, by employing
fauna that are as geographically and temporally as close to our samples
as possible, we should be providing an appropriate comparative
framework for the Vindija Neanderthal samples. Moreover, the relative
distribution, especially of 15N values, for
the species included in this pooled sample is similar to the
distributions derived for various faunal species from single sites (10,
11).
Fauna.
The Bos/Bison and cervid samples from
Vindija (Table 1) have herbivore 13C and
15N values that are within the ranges observed
for European Holocene specimens (25, 27). The
13C values are more indicative of open-ranging
species (20
), rather than forest-dwelling species
(22), but ranges of variation in Late Pleistocene
Bos/Bison 13C
values (24) as well as the hilly terrain in the vicinity of Vindija
Cave make it difficult to assess which of these bovine genera is most
likely represented. The cave bear samples are interesting from a
paleobiological, rather than an anthropological, perspective as they
have very low 15N values. Similarly low
Ursus spelaeus 15N values have been
observed for samples from Slovenia (28), France (29), and Belgium (30).
The low U. spelaeus values probably reflect a high degree of
herbivory (31); they may also be a result of their unusual metabolism
related to hibernation (32), although the hibernation model has been
disputed (30).
Neanderthals.
The Neanderthal samples from Vindija have high
15N values, which indicate that the
overwhelming majority of their dietary protein was from animal, rather
than plant, sources (Table 1, Fig. 1). The associated 13C values indicate the
exploitation of more open-ranging herbivores, despite the hilly terrain
of the Hrvatsko Zagorje. The Neanderthal values are close to the later
carnivore isotope values from Dolní Vstonice II and
Milovice (22), as well as those of earlier carnivores from Marillac and
Scladina (10, 11), indicating that these Neanderthals had diets similar
to nonhuman carnivores.
|
15N values from
Milovice are intriguing, as they are higher than the other herbivores.
This pattern of higher mammoth values has been observed previously (30,
33, 34) and may relate to mammoths targeting specific plant species,
whereas other herbivores consume a wider range of species. The higher
mammoth 15N values may be of relevance here,
as the Neanderthal 15N values could make sense
if their main dietary protein source was mammoths rather than the other
faunal species. However, archeological evidence for Neanderthal
exploitation of proboscideans is extremely rare, and a broader series
of fauna needs to be analyzed before the spectrum of predated
herbivores can be evaluated through stable isotope analysis.
Our findings concerning the diet of the Vindija Neanderthals are
remarkably similar to those observed by Bocherens and colleagues for
other European Neanderthals (10, 11). They obtained similar 13C and 15N values
for two Neanderthals from the site of Marillac dated to
40,000-45,000 years B.P. and for a Neanderthal specimen from Scladina Cave, Belgium, which is earlier, dated to between 80,000 and
130,000 years B.P. (Table 2). Moreover,
the high 15N for the Marillac Neanderthal
remains are most closely approached by the values for Canis
lupus and Crocuta crocuta from that site (10), whereas
the earlier Neanderthal 15N value from
Scladina is most closely approached in that site's faunal assemblage
by Panthera spelaea and secondarily by slightly lower values
for Crocuta crocuta and Canis lupus (11). For
these five Neanderthal specimens, therefore, we have stable isotope data indicating that geographically and chronologically dispersed Neanderthals consistently behaved as top-level carnivores.
|
Neanderthals as Predators. Neanderthal subsistence strategies were varied in space and time, with carcass utilization patterns varying on intersite and interspecies levels (4, 35). The role of hunting versus scavenging in meat acquisition by Middle Paleolithic humans has been debated particularly over the last two decades (3, 36, 37), and from this discussion it has become clear that the Neanderthals were capable of, and frequently engaged in, predation on mammals.
In particular, taphonomic analyses of a number of Middle Paleolithic, Neanderthal-associated mammalian faunal assemblages in recent years have concluded that focused and selective hunting strategies resulting in high meat utility acquisition were carried out by these late archaic humans in areas of Europe and the Near East as dispersed as France (Bau de l'Aubesier, La Borde, Canalettes, Coudoulous, Mauran, Le Portel), Germany (Salzgitter Lebenstedt, Wallertheim), Italy (Grotta Breuil), Croatia (Krapina), Iran (Kobeh), Israel (Kebara), and Russia (Il'skaja) (1-3, 6, 35, 38-44). These interpretations are based principally on mortality profiles and/or distributions of skeletal part frequencies of the prey species being processed, combined with direct evidence of human carcass processing with lithic tools. In the former, prime age-dominated assemblages are usually taken to indicate selective and active predation by these hominids. In the latter, a proximal limb element-dominated assemblage or a preserved skeletal distribution representative of anatomical frequencies, as opposed to a head and foot-dominated assemblage, are generally taken to indicate primary carcass access and hence active predation. However, not only do a significant number of these assemblages not meet both criteria for active predation on the part of the Neanderthals, it is increasingly apparent that a variety of factors can contribute to the mortality and skeletal element distributions documented in archeological faunal assemblages. These factors include prey population demographic dynamics, nonhuman predator prey selection patterns, carcass consumption patterns by both humans and other carnivores, human carcass element transport variation, and postdepositional processes acting differentially on skeletal elements. Moreover, it remains unclear how representative of overall Neanderthal diet such episodes are. Consequently, current taphonomic analyses of these and other archeological faunal assemblages do not always permit assessment of the degrees to which the assemblages were accumulated through active predation versus scavenging. Neanderthal predation has also been supported by the evidence for spears (stone-tipped and wooden) among both the Neanderthals and their Middle Pleistocene European predecessors (45-49), combined with rare examples of such weapons in the remains of apparent prey animals [e.g., the wooden spear in the ribs of an Elephas skeleton at Lehringen, Germany, and the Levallois point embedded an Equus cervical vertebra from Umm el Tlel in Syria (45, 50)]. In addition, indirect measures of Neanderthal subsistence such as the Levallois point to core frequencies have been used to suggest that the Neanderthals were highly predatory in the Near East (ref. 51; but see refs. 52 and 53), despite the absence of evidence for the kind of projectile weaponry seen in the Upper Paleolithic that would increase the mechanical efficiency and safety of hunting or for the patterned variance in extractive technologies widely seen in Upper Paleolithic and more recent hunter-gatherer toolkits (54). This inference of active predation on the part of the Neanderthals is further supported by their anatomical distribution of trauma, which suggests proximate encounters with large animals (55) of the kind necessitated by their predominantly heavy available weaponry (45, 47, 48, 56). Yet, their pattern of trauma does not permit distinctions between injuries sustained during hunting versus those suffered in competition with other carnivores for carcasses or space. Consequently, although several lines of evidence support active mammalian predation by the Neanderthals and contradict the previous models of the Neanderthals acquiring their animal protein principally through scavenging, the archeological data nonetheless remain frequently ambiguous as to the extent to which these late archaic humans were the primary predators of the mammals whose remains they processed. The consistent stable isotope data indicating their position as top-level carnivores provides insight into this issue. There are no true mammalian scavengers, as all are omnivores (ursids and canids) and/or actively hunt (hyenas) (57). This is because the search time for scavenging relative to the return is too expensive for terrestrial homeothermic vertebrates, and most predators actively defend their kills, thereby increasing risk to any potential terrestrial scavenger (57). If the Neanderthals were obtaining their animal protein principally through scavenging, they would have had to obtain most of their food from plants, as a reliable food source, and only supplemented this with scavenged animal products. Even though the isotope data cannot distinguish the species or even the sizes of the animals consumed, they clearly show that animal products were the overwhelming source of protein in European Neanderthal diets and that protein from plants was insignificant. It is therefore likely that scavenging, although undoubtedly practiced on an opportunistic basis by these European Neanderthals, must have been distinctly secondary to predation.| |
Summary and Conclusions |
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Top
Abstract
Introduction
Summary and Conclusions
References
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Isotope analyses of two Neanderthals and associated fauna from Vindija Cave, Croatia, have indicated that the bulk of their dietary protein came from animal sources. Comparison with faunal remains from this and other sites of similar age indicates that the Vindija Neanderthal isotope values were similar to those of other carnivores. These results are very close to the results for earlier Late Pleistocene Neanderthals from France and Belgium.
Therefore, the emerging picture of the European Neanderthal diet indicates that although physiologically they were presumably omnivores, they behaved as carnivores, with animal protein being the main source of dietary protein. This finding is in agreement with the indirect archeological evidence and strongly points to the Neanderthals having been active predators.
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Acknowledgements |
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We thank H. Bocherens, F. B. Marshall, and M. C. Stiner for helpful comments. The collection and analysis of the Vindija hominid samples was supported by the L. S. B. Leakey Foundation, by the Wenner-Gren Foundation, and by the Prehistoric Society of the United Kingdom. The analysis of the Vindija faunal samples was supported the Natural Environment Research Council (U.K.) and the Social Sciences and Humanities Research Council of Canada.
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Footnotes |
|---|
To whom reprint requests should be
addressed. E-mail: trinkaus{at}artsci.wustl.edu.
Article published online before print: Proc. Natl. Acad. Sci. USA, 10.1073/pnas.120178997.
Article and publication date are at www.pnas.org/cgi/doi/10.1073/pnas.120178997
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References |
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Abstract
Introduction
Summary and Conclusions
References
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 J. Dickson, J. H. Dickson, M. P. Richards, R. J. Hebda, P. J. Mudie, O. Beattie, S. Ramsay, N. J. Turner, B. J. Leighton, J. M. Webster, et al. Kwaday Dan Ts'inchi, the first ancient body of a man from a North American glacier: reconstructing his last days by intestinal and biomolecular analyses The Holocene, May 1, 2004; 14(4): 481 - 486. [Abstract] [PDF]
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 R. P. Heaney Constructive Interactions among Nutrients and Bone-Active Pharmacologic Agents with Principal Emphasis on Calcium, Phosphorus, Vitamin D and Protein J. Am. Coll. Nutr., October 1, 2001; 20(90005): 403S - 409. [Abstract] [Full Text]
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B. L. Hardy, M. Kay, A. E. Marks, and K. Monigal Stone tool function at the paleolithic sites of Starosele and Buran Kaya III, Crimea: Behavioral implications PNAS, September 4, 2001; (2001) 191384498. [Abstract] [Full Text] [PDF]
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M. P. Richards, P. B. Pettitt, M. C. Stiner, and E. Trinkaus Stable isotope evidence for increasing dietary breadth in the European mid-Upper Paleolithic PNAS, May 22, 2001; 98(11): 6528 - 6532. [Abstract] [Full Text] [PDF]
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R. P Heaney Reply to JE Kerstetter et al Am. J. Clinical Nutrition, May 1, 2001; 73(5): 991 - 992. [Full Text]
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R. P Heaney Protein intake and bone health: the influence of belief systems on the conduct of nutritional science Am. J. Clinical Nutrition, January 1, 2001; 73(1): 5 - 6. [Full Text] [PDF]
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B. L. Hardy, M. Kay, A. E. Marks, and K. Monigal Stone tool function at the paleolithic sites of Starosele and Buran Kaya III, Crimea: Behavioral implications PNAS, September 11, 2001; 98(19): 10972 - 10977. [Abstract] [Full Text] [PDF]
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