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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 (δ13C and δ15N) 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.

Reconstructions of European Neanderthal subsistence strategies have overwhelmingly focused on the specialized hunting and scavenging of herbivores as the predominant method of obtaining food (16). 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 (79). 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 δ15N 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.
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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.
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Stable Isotope Analyses.

Mammal bone collagen δ13C and δ15N values reflect the δ13C and δ15N values of dietary protein (14). They furnish a long-term record of diet, giving the average δ13C and δ15N values of all of the protein consumed over the last years of the measured individual's life. δ13C 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 δ13C values that are more negative than species that live in open environments (17). δ15N values are, on average, 24

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