News and Views Hervé Bocherens, Daniel Billiou, New isotopic evidence for dietary habits of Neandertals from Belgium Laboratoire de Biogéochimie Isotopigue, UMR 7618, CNRS-INRA-UPMC, Universite P. et M. Curie, 4 place Jussieu, André Mariotti Michel Toussaint F-75252 Paris cedex 05 (France) Direction des Fouilles du Ministére de la Région Wallonne, rue des Brigades d'Irlande 1, B5100 Jambes, Belgium Maryléne Patou-Mathis Dominique Bonjean France Belgium Marcel Otte Institut de Paléontologie Humaine, 1 rue René Panhard, 75013 Paris, Asbl Archéologie Andennaise, Grand’Place 132, B-5300 Sclayn, Centre de Recherche sur les Civilisations Paléolithiques en Europe, Université de Liége, Service de Préhistoive, 7 place du 20 Aoit, B-4000 Liége, Belgium Journal of Human Evolution (2001) 40, 497-505 “doi: 10.1006/jhev.2000.0352 ra Available online at hutp://www-ideatibrary.com on IDE Pek Carbon and nitrogen isotopic signatures in fossil collagen provide useful information for reconstructing the diets of ancient popula- tions. Indeed, the collagen synthesized in living bone records the isotopic signature of the average dietary proteic fraction. This phenomenon allows the identification of the type of plants at the basis of the trophic web (forested vs open environment in the case of arctic and temperate continental climates) by using the carbon isotopic signature. The position of an individual within the food web it belongs to can be deciphered by the nitro- gen isotopic signature, since the amount of *SN increases between the food and its con- sumer. This approach has recently indicated that Neandertals from Marillac (France) and from layer 4A in Scladina cave (Belgium), i.e. maxillary SCLA 4A-2, obtained much of their protein from open country herbivores (Bocherens et al., 1991, 1999; Fizet et al., 1995). Very similar results have been published recently by Richards et al, (2000) about Neandertals from Vindija (Croatia). Although highly valuable in the (0047-2484/01/060497 +.09835.00/0 @ debates regarding Neandertal dietary strategies (¢.g., Mellars, 1989; Patou, 1989; Webb, 1989; Lalueza et al., 1993; Stiner, 1994; Marean, 1998; Richards er al., 2000), this information is still limited due to the small number of individuals analysed so far. ‘Thus it is necessary to expand this field of research and, in this respect, the goal of this paper is to report new isotopic measure ments performed on three Neandertal specimens from Wallonia (Belgium), a region that has yielded many important Neandertal specimens since the 19th century (Toussaint, 1996). The studied specimens include one pedal phalanx I “SCLA-IB-4” from layer 1B in Scladina Cave, a skull fragment “Engis 2” of a 5-6 year old child (Tillier, 1983) from Awirs Cave, and a piece of the right scapula frag- ment “Spy” from Betche-al-Roche Cave (Table), and their palaeodietary implica- tions are discussed. The sites are located in the Maas valley, about 40 km apart (Toussaint, 1996) (Figure 1). The Spy and Engis specimens are tentatively dated © 2001 Academic Press498 H, BOCHERENS ET AL, BRUXELLES. Figure 1. Map of Belgium with the location of the three sites mentioned in this paper. around 35-40,000 years BP on the basis of their associated fauna (Cordy, 1988), and the Scladina-I1B specimen has been excavated from a layer dated about 40,000 BP (Simonet, 1992) In order to be able to reconstruct palaeo- diets from collagen isotopic signatures, it is necessary (1) to test the quality of preserva- tion of such collagen, and (2) to calibrate the ecosystem using isotopic signatures of coeval fauna (Bocherens, 1997), since isotopic variations can occur within a given species due to local nondietary environmental parameters such as aridity or soil acidity (e.g. Fizet et al, 1995; Drucker et al., 2000). After extraction following the now routine protocol used for Upper Pleistocene samples (Bocherens et al., 19972), collagen indigeneity has been checked using carbon and nitrogen amounts in the extracted col- lagen, as well as C:N ratios (DeNiro, 1985; Ambrose, 1990). Collagen isotopic signa- tures measured on herbivore and carnivore species from layer 1A in Scladina have been used to calibrate the trophic web (Bocherens et al. 1997a). Indeed this layer is slightly younger than layer 1B in Scladina Cave but presents a faunal composition quite similar to layer 1B, reflecting a similar environment (Simonet, 1992). Additional isotopic measurements have been performed on horse, reindeer, cave bear and hyena from Layer 1B in Scladina Cave, as well as on reindeer from layer 1A since isotopic signa- tures for this species were not available in Bocherens et al.’s (1997a) paper (Table 1). Isotopic compositions are expressed as 81°C and 8'°N values relative to international standards (Bocherens et al., 1997) All collagens present chemical positions within the range of acceptable values, with %C>30% and %N>10%, and with C:N ratios ranging from 3-1 to 3-4 (Table 1). Noteworthy are the high collagen yields obtained on the Neandertal speci- mens, especially for the “Engis 2” child, com-NEW ISOTOPIC EVIDENCE FOR DIETARY HABITS OF BELGIAN NEANDERTALS with 176-4 mg/g, close to the yield of fresh bone (around 210 mg/g). Dentine collagen isotopic compositions have been separated from those of bone since it is known that dentine tends to have higher 6'°N than bone in species such as reindeer, bear and hyena, probably due to a suckling isotopic signal kept in dentine and eliminated in bone (Bocherens & Mariotti, 1997). Conversely, dentine and bone collagen isotopic compo- sitions seem to be equivalent in horse and bovines, probably due to their having teeth with prolonged growth periods that record a similar chronological period to remodelling bone. They are thus considered together for palaeoenvironmental implications. The fau- nal specimens from Scladina Cave exhibit clusters of isotopic values according to the species, regardless of their stratigraphic: origin (Figure 2). This suggests that the isotopic calibration of the mammal fauna has not changed dramatically within the period considered here, and probably not within the limited geographical range. The differences observed between species are similar to those seen in other Eurasian ecosystems of Weichselian age (Oxygen Iso- topic Stage 3: Bocherens, 2001). Herbivores show a pattern with reindeer having the highest 5'°C, probably due to lichen con- sumption (Fizet et al., 1995; Drucker et al., 2001), and mammoth having low 81°C and high §!°N, as in many other sites from Eurasia (Bocherens et al., 1997a; Tacumin et al., 2000). Such high 8!°N is possibly due to consumption of grass from disturbed environments (Bocherens, 2001). Cave bear presents low 5!°C and 5'°N, most probably reflecting a vegetarian diet, as in many other European sites (Bocherens et al., 1994, 1997a). Hyena exhibits the highest 5'°N, as expected for top carnivores. All Neandertals exhibit 8'°C within the range observed for the fauna, and high 8'°N, slightly more positively than those of hyenas. The °C of the Scladina-1B specimen is less positive than those of the Spy and Engis 499 specimens, whereas the Engis specimen exhibits the heaviest 8!°N. The total range is 1-6%o for both 8°C and 8!°N Neandertal values. Such a range exceeds that observed for a supposedly homogeneous Mesolithic hunter-gatherer population (Bocherens et al., 19976), and thus might reflect dietary differences between the analysed specimens. ‘The Spy specimen isotopic composition is very similar to that of Marillac Neandertals as presented by Fizet et al. (1995) when compared with their coeval fauna (Figure 3). In both cases, the isotopic compositions seem to reflect the consumption of dietary proteins from large herbivores, especially those with rather positive 8'°C and 8!°N, such as reindeer and suckling herbivores, or even omnivorous mammals such as bears. Indeed, some Neandertal and pre- Neandertal populations have been shown to exploit the meat of such animals, as in Biache-Saint-Vaast (Auguste, 1995) and ‘Taubach (Brathing, 2001). The Scladina- 1B specimen is more '*C-depleted and more 15N-enriched than the Spy specimen, which may also reflect a diet based on large herbi- vore meat, but with more mammoth meat and possibly more freshwater fish in the diet of the former specimen, since both resources have more positive 5!°N than most terres- trial herbivores (Bocherens et al., 19974; Dufour er ai., 1999). Mammoth meat may be more likely since no evidence for fishing or fish remains have been found in Neandertal sites in Belgium, not even in Scladina cave where the sediment was systematically sieved during the excavation. However, fish remains found in Mousterian sites from Southwestern France and Spain may suggest that, in some places, some Neandertal populations could fish in freshwater (Le Gall, 1992). The Engis child exhibits a 8!°C similar to that of the Spy specimen but is more !*N-enriched. 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BOCHERENS ET AL © Horse (bid) i @ Bovine Mopac a > Megaceros(b) D> Megaceros (@) 2 A Reindeer (6) | A Reindeer (d) 10. © Rhinoceros (d) 9. © Mammoth (4 zB 8 © Cave Bear () 1 a | © Cave Bear a) © Brown Bear(d) 6 Hyena 5 Hyena (@) 4 % Neandertal (6) A EEE] _berbivores (except mammoths) , smammoths 242 RU 20 to Is 17-16 Hyenas (bone) Beal Neandertals Figure 2. Carbon and nitrogen isotopic compositions of collagen extracted from Scladina 1A and 1B fauna, and from Scladina 1B, Engis and Spy Neandertals. (b) Stands for bone isotopic values and (d) stands for dentine isotopic values. The stippled ureas include only bone collagen isotopic values. than post-weaning diet (Fogel et al., 1989). Bone growth and renewing leads to a decrease of 5!5N after weaning (Fogel et al., 1989; Bocherens et ala, 1994; Bocherens & Mariotti, 1997). Isotopic studies of archaeo- logical demographic series suggest that bone reaches equilibrium 1-2 years after weaning, and individuals older than 4 years exhibit 8°°N similar to those of adults when wean- ing occurs around 2 years old (Fogel et al., 1989; Tuross & Fogel, 1994; Katzenberg & Pfeiffer, 1995). If the high 5'°N of the Engis child relative to the adult from Spy indicates that the isotopic equilibrium is not reached, this could mean that weaning actually occurred at around 1-2 years for the child but that this individual is younger than previously thought on the basis of dental eruption degree (Tillier, 1983). This indi- vidual is possibly younger than 4 years, which may be deduced through advanced dental maturation relative to modern humans, as suggested by some authors (e.g., Thompson & Nelson, 2000). An alternative is that the child is indeed 5-6 years old, as indicated by dental eruption degree, but that weaning occurred late, when the individual was around 4 years old. Another possibility is that this child belongs to a Neandertal population with a different diet than the Spy and Scladina-1B specimens, with more ‘N-enriched proteins in addition to large herbivore meat, possibly freshwater fish, mammoth meat being too '*C-depleted toNEW ISOTOPIC EVIDENCE FOR DIETARY HABITS OF BELGIAN NEANDERTALS. 2 503 2 indie Manas Sehadina ta nye ud un 10; * 10. a 10. > 9 of % F] of | 8 3 ° 8 eon 7 7 27 rr 8 Es Zoli? 2 6 a es 5 @ sl ¢ s|° o | he, a yet oy 4 af BR, 4 \ 3 34 0 3 ft 2 a 24 ° 2. i “ 14 1 0 ° ° - 219-18 22 -31-20-19-18 36-95-2433 22-20-19 18 BBC 8c Bovine © Horse(bed) % Horse (bea) > Cenvid > Reindeer) > Fallow Deer ) A. Cave Bear Bovine &) a Cave Bear (6) Neandertal & Hyena (b) vy Brown Bear (b) © Wate) Bm Cave Lion @ Fox (b) m Hyena (b) Neandertal () a Wolf) Neandertal (b) Figure 3. Carbon and nitrogen isotopic compositions of collagen extracted from Neandertals and associated fauna in Vindija (Croatia, ca. 28-29,000 BP; Richards et ai., 2000); Marillac (Charentes, France, ca. 40-45,000 BP; Fizet eral, 1995) and layer 4A from Scladina Cave (Belgium, ca. 110-120,000 BP; Bocherens er al., 1999). (b) Stands for bone isotopic values and (4) stands for dentine isotopic values. Table 2_ Isotopic values of Neandertal bones Yield No. Piece Site (mg) %C NN eC BN MT500 Pedal phalanx 1 Scladina-IB_ SCLAIB4 nd. 399-4732 212 1B MTI100 Left parietal ——-Engis ENG 2 164 417 14d 34-196 126 MT200 Right scapula Spy SPYOMO1 1239 415 144 34-108 11-0, explain the difference. Future studies will help to decipher Neandertal diet and its possible geographical and chronological variations, At this stage, it seems that Neandertal diet is characterized by high pro- portions of open environment herbivore proteins, even during forested periods (Figure 3, Bocherens et al., 1999). Such a504 high protein diet needs balancing with energy-rich nutrients (Speth & Spielmann, 1983), It is noteworthy in this regard that herbivores hunted during middle Palaco- lithic times belong mainly to fat-tich species, such as bovids, and that the seasonality of hunting shows an avoidance of animals with Jean meat, such as males after the rut period and females after giving birth to calves. Moreover, marrow was sought by Neandertals, as documented by the high degree of bone fracturation and the fact that bone fat was extracted by boiling (Patou-Mathis, 1995). This reliance on herbivore meat from open environments might reflect a fundamental adaptation of the Neandertals to the rather cold environ- mental conditions of the European Upper Pleistocene, as already suggested by some of their anatomical features (Condemi, 1998). Acknowledgements This research has been funded by the CRNS programme “Paléoenvironnements, Evolution des Hominidés.” Thanks are due to Nick van der Merwe and one anonymous reviewer for valuable comments which helped to improve this manuscript. References Ambrose, S. H. (1990). Preparation and characteriz- ation of bone and tooth collagen for isotopic analysis. J. Archaeol, Sci, 17, 431-451 Auguste, P. (1995), Chasse ot paléolithique moyen: apport Biache-Saint-Vaast (Pas-de priist. Fr. 92, 155-167, Bocherens, H. (1997). Isotopic biogeochemistry as a marker of Neandertal’s diet. Anthrop. Ane. 55, 101-120, Bocherens, H. (2001). Isotopic biogeochemistry and the paleoecology of the mammoth steppe fauna Deinsa (in press) Bockerens, H. & Mariotti, A. (1997). Comments on! Diet, physiology and ecology of fossil mammals as infezzed from stable carbon and nitrogen isotope biochemistry: Implications for Pleistocene bears bby Bocherens et al—Reply. Palacogr., Palacoclimato., Palaeoscol. 128(1/4)362-364. charognage au du gisement de ais). Bull. Soc H, BOCHERENS ET AL, Bocherens, H., Fizet, M., Mariotti, A., Lange-Badré, B., Vandermeersch, B., Borel, J-P. & Bellon, G. (1091), Isotopic biochemistry CC, °N) of fossil vertebrate collagen: implications for the study of fossil food web including Neandertal Man. hum, Ecol, 20, 481-492, Bocherens, H., Fizet, M. & Mariowt, A. (1994). Diet, ‘physiology and ecology of fossil mammals as inferred bby stable carbon and nitrogen isotopes biogeo- chemistry: implications for Pleistocene bears. Palaeogr., Palacoclimatol, Palacoecol. 107, 213— 225, Bocherens, H., Billiou, D., Patou-Mathis, M., Bonjean, D., Otte, M. & Mariotti, A. (1997@). Is0- topie biogeochemistry (°C, "°N) of fossil mammal collagen from Scladina cave (Sclayn, Belgium). ‘Quatem. Res. 48, 370-380. Bocherens, H., Grupe, G., Mariott, A. & Turban-Just, S., (19978). Molecular and isotopic preservation of mesolithic human finds from the Ofner Cave Bavaria, Germany). Anshop. Ans. 55, 121-129. Bocherens, H., Billiou, D., Patou-Mathis, M., Otte M,, Bonjean, D., Toussaint, M. & Mariott, A. (1999). Palacoenvironmental and palacodietary ‘implications of isotopic biogeochemistry of late inter- glacial Neandertal and mammal bones in Seladina Cave (Belgium). J. Archaeal. Sc. 26, 599-607. Bratiung, B. (2001). Anthropogenic factors in the thanatocoenose of the last interglacial travetines at Taubach, Germany. In (E. Turner & S. Gaudzinski, ais) Die Rolle des Menschen in Faunenakkumulationen des Alt nnd Mitelpalaolithitumn Ewropas. Mainz. (in ress). Condemi, (1998). ‘The Neanderthals: a cold- adapted European Middle Pleistocene population? Anihropologie 36, 35-42. Gordy, J-M. (1988). Apport de Ia paléozoologie & la paléoécologic et % la chronostratigraphie en Europe ‘du novd-oceidental. In L'Homme de Neandertal, 2, Lrenvironnement, Etudes et Rercherches Archéologiquet de Université de Litge (ERA.U.L), pp. 55-64. Lidge: Université de Ligge DeNiro, M. J. (1985). Postmortem preservation and alteration of im tivo bone collagen isotope ratios in relation to palacodietary reconstruction. Naswe 317, 306-809. Drucker, D., Bocherens, H. & Mariotti, A. (2000) Contribution de la biogéochimie isotopique & Ia paléobiologie des grands mammiféres pléistocénes supérieurs: Application aux rennes et aux chevaux duu Pleistocéne supérieur du Sud-Ouest de la France Mém. Soc. prihist. Fr, 28, 13-27. Dufour, E., Bocherens, H. & Mariotti, A. (1999), Palaeodietary implications of isotopic variability in Eurasian lacustrine fish. J. Archaeol, Sci, 26, 627-627 Fizet, M., Mariotti, A., Bocherens, H., Lange-Badré, B., Vandermeersch, B., Borel, J. P. & Bellon, (1995). Effect of diet, physiology and climate on carbon and nitrogen isotopes of collagen in a late Pleistocene anthropic paleoecosystem (France, Charente, Marillac). J: Archaeol Sci. 22, 67-79.NEW ISOTOPIC EVIDENCE FOR DIETARY HABITS OF BELGIAN NEANDERTALS: Fogel, M. L. Tuross, N. & Owsley, D. W. (1988). Nitrogen Kotope aces of human lactation in ‘modem and archeological populations. Ann. Rep Dir. Geophys. Lab., Carnegie Institution 1988-1989, 111-417. Kavzenberg, M. A. & Phifer, S. (1995). Nitrogen isotope evidence for weaning age in @ nineteenth century canadian skeletal sample. In (A. L. Graver, Ed.) Body of Evidence, pp. 221-235. New York Wiley-Liss. lacumin, P., Nikolaev, V. I. & Ramigni, M. (2000). C ‘and N stable isotope measurements on Eurasian fossil mammals, 40,000 0 10,000. years BP: Herbivore physiologies and’ palaeoenvironmental reconstructions, Palazogr., Palacclimato.,Palaevecol 163, 22-47, Lalueza, C., Pérez-Pérez, A. & Turbon, D. (1993) Microscopic study of the Banyoles mandible (Girona, Spain: diet, cultural activity and toothpick Use. J. nom. Bool. 24, 281-300, Le Gall, 0, (1992). Poitsons et péches au Paléoithique (quelques données de Europe occidentale). Anthropologie (Par) 96(1)121-134 Marean, C. W. (1998). A critique of the evidence for scavenging by Neandertals and easly modem hhumans: new data from Kobeh Cave Zagros Mountains, Iran) and Die Kelders Cave 1 layer 10 (South Arca). J, um. Evol. 35, 111-136. ‘Mellars, P. (1989). Major issues ia the emergence of Modem Humans. Curr. Anihrop. 30, 349-385. Patou, M. (1985). Subsistance et approvisionnaement au Paléolithique moyen. In (L. Freeman & M. Patou, Eds) Léhomme de Neandertal, vol. 6, La subsistance, Enudes et Rercherches Archéoogiques de PUniversté de Lilge (E.R.A.U.L)), 33, pp. 11-18. Lidge: Université de Livee. Patou-Mathis, M. (1095). Stress biologiques et com- portements de subsistance au Paléoithique moyen et supérieur en Europe. In (M. Otte, Ed.) Naniow et Gulture, Etudes at Rerchercher Avchéologiques de PUniverité de Litge (E.R.A.U.L.), 68, pp. 437-453. Lidge: Université de Lidge. 505 Richards, M. Pa Pettitt, P. B., Trinkaus, E., Smith, F. H., Paunovic, M. & Karavanic, I.” (2000). Neandertal dict at Vindija and Neandertal predation: ‘The evidence from stable isotopes. Pos. Natl. Acad. Sei, 91(13)7663-7666, Simonet, P. (1992). Les associations de grands mammiféres du gisement dela grotte Scladina 4 Sclayn (Namur, Belgique). In (M. Otte, Ed.) Recherches aice grotes de Selayn, vol. 1, le contexte, Endes et Rercherches Archéoogiques de. VUniversité de Lidge (ER-AU.L), 27, pp. 127-151. Liege: Université de Lidge Speth, J.D. & Spielmann, K. A (1983). Energy source, protein metabolism, and hunter-gatherer subsistence strategies. J. Anthrop. Archaeol 2, 1-31 Stiner, M.-C. (1994). Honor Among Thieves, a Zooarchacological Study of | Neandertal Ecology. Princeton: Princeton University Press ‘Thompson, J. L. & Nelson, A. J. (2000). The place of ‘Neandertals in the evolution of hominid patterns of growth and development. J. fun. Evol. 38, 475-495. Tiller, A. M, (1983). Le crane d’enfant 'Engis 2: un exemple de distribution des caractéres juvéniles, primitifs et néandertaliens. Bull. Soc. R. belge Anthrop. Prohist. 34, 51-75. Toussaint, M. (1996). D’Engis 4 Sclayn, les Néandertaliens ‘mosans. In (D. Bonjean, Ed.) Neandertal, pp. 48-61. Andenne: ASBL Archéologie Andennaise, ‘Tuross, N. & Fogel, M. L. (1994), Stable isotope analysis and. subsistence patterns at the Sully Site, South Dakota. In (D. W. Ousley & R. Jantz, Eds) ‘Skeletal Biology of the Plains: Migration, Warfare, Health and Subsistence, pp. 283-289. Washington D.G.: Smithsonian Institution Press. ‘Webb, R. E, (1989). A reassessment of the faunal ‘evidence for neandertal diet based on some western European collections. In (L. Freeman & M. Patou, Eds) L’Homme de Neandertal, vol. 6, La subsisiance, Etudes et Rercherches Archiologiques dle UUniversité de Liige (E.R.A.U.L.), 33, pp. 155-178. Lidge: Univer- site de Lidge.