<table><caption>Tableau 17. Inventaire complet des vestiges humains mis au jour, découverts ou redécouverts par l'équipe scientifique</caption><thead><tr><th>N°</th><th>Nature du matériel</th><th>Date de la mise au jour</th><th>Carré</th><th>Sous-carré</th><th>Faciés / Couche</th><th>N° fouille</th><th>Fouilleur</th><th>Inventeur</th><th>Date de l'invention</th><th>Remontages</th></tr></thead><tbody><tr><td>1</td><td>Diaphyse fémur droit immature</td><td>24.10.1967</td><td>G12</td><td>A déterminer</td><td>Couche Rouge</td><td>3</td><td>? Battail</td><td>S. Costamagno</td><td>Début 2002</td><td></td></tr><tr><td>2</td><td>Incisive permanente supérieure latérale droite</td><td>19.08.1970</td><td>F12</td><td>A déterminer</td><td></td><td>54</td><td>Mario Chech</td><td>S. Costamagno</td><td>Début 2002</td><td></td></tr><tr><td>3</td><td>Fragment de voûte crânienne, mature</td><td>1971</td><td>F10</td><td>A déterminer</td><td></td><td>10</td><td>Non mentionné</td><td>B. Maureille</td><td>24.07.2003</td><td></td></tr><tr><td>4</td><td>Incisive déciduale latérale inférieure droite</td><td>14.08.1972</td><td>E09</td><td>A déterminer</td><td></td><td>10</td><td>? Normand</td><td>S. Costamagno</td><td>Début 2002</td><td></td></tr><tr><td>5</td><td>Incisive permanente centrale supérieure (droite ?)</td><td>10.08.1972</td><td>E11</td><td>A déterminer</td><td></td><td>10</td><td>336</td><td>S. Costamagno</td><td>Début 2002</td><td></td></tr><tr><td>6</td><td>Première prémolaire supérieure gauche</td><td>15.08.1972</td><td>I10</td><td>A déterminer</td><td></td><td>11</td><td>22</td><td>B. Vandermeersch</td><td>Idem mise au jour<br>Isabelle Crévecoeur &<br>Naomi Amar</td><td></td></tr><tr><td>7</td><td>Fragment de diaphyse de radius droit, mature</td><td>07.08.1980</td><td>D10</td><td>A déterminer</td><td>9b</td><td>54</td><td>Non mentionné</td><td>B. Maureille</td><td>02.08.2003</td><td></td></tr><tr><td>8</td><td>Partie antérieure, branche montante droite, mandibule mature</td><td>18.08.1980</td><td>D11</td><td>A déterminer</td><td></td><td>10</td><td>Patrick Péridy</td><td>S. Costamagno</td><td>Début 2002</td><td></td></tr><tr><td>9</td><td>Fragment diaphyse fibula gauche (taille mature)</td><td>18.08.1980</td><td>D11</td><td>A déterminer</td><td></td><td>10</td><td>Patrick Péridy</td><td>S. Costamagno</td><td>Début 2002</td><td></td></tr><tr><td>10</td><td>Première (?) molaire inférieure gauche, usée mais intacte, adulte</td><td>11.08.2001</td><td>E10</td><td>SE</td><td>Remanié</td><td>Rem. H01</td><td>B. Maureille</td><td>B. Maureille</td><td>Idem mise au jour</td><td></td></tr><tr><td>11</td><td>Deuxième prémolaire inférieure d., peu usée, adolescent ou adulte</td><td>16.08.2001</td><td>D13</td><td>NE</td><td>Fa 3</td><td>1</td><td>Hélène David</td><td>H. David</td><td>Idem mise au jour</td><td></td></tr><tr><td>12</td><td>Temporal droit, fragmentaire, mature</td><td>19.08.2001</td><td>D08</td><td>SW</td><td>Remanié</td><td>Rem. H02</td><td>Hélène Rougier</td><td>H. Rougier</td><td>Idem mise au jour</td><td></td></tr><tr><td>13</td><td>Maxillaire droit, région alvéolaire avec M3 à C', adulte</td><td>22.08.2001</td><td>D11</td><td>NW & NE</td><td>Remanié mais presque en place</td><td>Rem. H03</td><td>Guillaume Seguin</td><td>G. Seguin</td><td>Idem mise au jour</td><td></td></tr><tr><td>14</td><td>Fragment gauche d'occipital, mature (?)</td><td>23.08.2001</td><td>C11</td><td>NW</td><td>Remanié mais presque en place</td><td>Rem. H04</td><td>G. Seguin</td><td>G. Seguin & B. Maureille</td><td>Idem mise au jour</td><td></td></tr><tr><td>15</td><td>Fragment gauche d'occipital, immature</td><td>26.08.2001</td><td>E12</td><td>SE</td><td>Remanié</td><td>Rem. H05</td><td>B. Maureille</td><td>B. Maureille</td><td>Idem mise au jour</td><td></td></tr><tr><td>16</td><td>Fragment gauche d'occipital, mature</td><td>27.08.2001</td><td>E11</td><td>SW</td><td>Remanié</td><td>Rem. H06</td><td>B. Maureille</td><td>B. Maureille</td><td>Idem mise au jour</td><td></td></tr><tr><td>17</td><td>Fragment droit d'occipital, mature (?)</td><td>07.08.2001</td><td>D09</td><td>Indéterminé</td><td>Remanié</td><td>Rem. H07</td><td>Indéterminé</td><td>H. Rougier</td><td>20 Décembre 2004</td><td></td></tr><tr><td>18</td><td>Fragment de bord alvéolaire (face externe, recolle avec la prémolaire)</td><td>07.08.2002</td><td>D13</td><td>NE</td><td>Fa 3</td><td>2</td><td>B. Maureille</td><td>B. Maureille</td><td>Idem mise au jour</td><td></td></tr><tr><td>19</td><td>Fragment d'écaille temporal gauche, immature (?)</td><td>07.08.2002</td><td>E12</td><td>SE</td><td>Fa 2a (?) - Rem</td><td>1</td><td>Mélanie Frelat</td><td>H. Rougier</td><td>20 Décembre 2004</td><td></td></tr><tr><td>20</td><td>Fragment de voûte crânienne, immature (?)</td><td>07.08.2002</td><td>E12</td><td>SE</td><td>Fa 2a (?) - Rem</td><td>2</td><td>M. Frelat</td><td>B. Maureille</td><td>21 Décembre 2004</td><td></td></tr><tr><td>21</td><td>Fragment de voûte crânienne, immature (?)</td><td>07.08.2002</td><td>E12</td><td>SE</td><td>Fa 2a (?) - Rem</td><td>5</td><td>M. Frelat</td><td>B. Maureille</td><td>21 Décembre 2004</td><td></td></tr><tr><td>22</td><td>Divers fragments d'un même occipital</td><td>26.05.2003</td><td>D11</td><td>NE</td><td>Fa 2a</td><td>96, 97, 99, 100 à 104<br>108 à 112, 114 à 119<br>120, 121, 122, 125</td><td>B. Maureille<br>Linda Bouchneb</td><td>B. Maureille<br>L. Bouchneb</td><td>29 Mai 2003<br>et<br>Idem mise au jour</td><td></td></tr><tr><td>23</td><td>Deuxième molaire déciduale supérieure gauche</td><td>06.08.2003</td><td>D10</td><td>SW</td><td>2a</td><td>287</td><td>Dorothée Archambault</td><td>D. Archambault</td><td>Idem mise au jour</td><td></td></tr><tr><td>24</td><td>Patella droite, mature</td><td>15.08.2003</td><td>D15</td><td>SE</td><td>2b</td><td>Surf./001</td><td>Yann Heuzé</td><td>Y. Heuzé & D. Archambault</td><td>Idem mise au jour</td><td></td></tr><tr><td>25</td><td>Fragment postéro-latéral d'un parietal g., immature</td><td>03.08.2004</td><td>D12</td><td>SE</td><td>Fa 4a</td><td>113</td><td>L. Bouchneb</td><td>L. Bouchneb</td><td>Idem mise au jour</td><td></td></tr><tr><td>26</td><td>Fragment d'un occipital (?) g. immature (?)</td><td>08.08.2004</td><td>D12</td><td>SW</td><td>Fa 4a</td><td>314</td><td>D. Archambault</td><td>D. Archambault</td><td>Idem mise au jour</td><td></td></tr><tr><td>27</td><td>Fragment supéro-antérieur d'un occipital g., immature (?)</td><td>09.08.2004</td><td>D12</td><td>SE</td><td>Fa 4a</td><td>352</td><td>D. Archambault</td><td>D. Archambault</td><td>Idem mise au jour</td><td></td></tr><tr><td>28</td><td>Première molaire déciduale inférieure droite</td><td>09.08.2004</td><td>D12</td><td>SW</td><td>Fa 4a</td><td>411</td><td>D. Archambault</td><td>D. Archambault</td><td>Idem mise au jour</td><td></td></tr><tr><td>29</td><td>Fragment postéro-latéral d'un parietal droit, mature</td><td>13.08.2004</td><td>D12</td><td>SE</td><td>Fa 2b</td><td>718</td><td>D. Archambault</td><td>D. Archambault</td><td>Idem mise au jour</td><td></td></tr><tr><td>30</td><td>Fragment postéro-latéral d'un parietal droit, mature</td><td>13.08.2004</td><td>D12</td><td>SE</td><td>Fa 2b</td><td>719</td><td>D. Archambault</td><td>D. Archambault</td><td>Idem mise au jour</td><td></td></tr><tr><td>31</td><td>Fragment postéro-latéral d'un occipital g., immature (?)</td><td>15.08.2004</td><td>D12</td><td>SE</td><td>Fa 2b</td><td>751</td><td>D. Archambault</td><td>D. Archambault</td><td>Idem mise au jour</td><td></td></tr><tr><td>32</td><td>Fragment crânien avec zone suturaire</td><td>18.08.2004</td><td>D12</td><td>NE</td><td>Fa 2b</td><td>863</td><td>G. Seguin</td><td>G. Seguin</td><td>Idem mise au jour</td><td></td></tr><tr><td>33</td><td>Fragment de diaphyse tibiale droite, mature (?)</td><td>16.08.2004</td><td>D11</td><td>NE</td><td>Fa 2a</td><td>263</td><td>A. Clark</td><td>C. Beauval & B. Maureille</td><td>02 Décembre 2004</td><td></td></tr><tr><td>34</td><td>Deuxième prémolaire supérieure droite, mature</td><td>12.08.2004</td><td>D11</td><td>NE</td><td>Fa 4a</td><td>163</td><td>A. Clark</td><td>B. Maureille</td><td>02 Décembre 2004</td><td></td></tr><tr><td>35</td><td>Fragment de diaphyse tibiale droite, mature (?)</td><td>13.08.2004</td><td>D11</td><td>NE</td><td>Fa 4a</td><td>191</td><td>A. Clark</td><td>C. Beauval & B. Maureille</td><td>02 Décembre 2004</td><td></td></tr><tr><td>36</td><td>Fragment de diaphyse tibiale droite, mature (?)</td><td>15.08.2004</td><td>D11</td><td>NE</td><td>Fa 4a</td><td>217</td><td>A. Clark</td><td>C. Beauval</td><td>02 Décembre 2004</td><td></td></tr><tr><td>37</td><td>Cunéiforme intermédiaire gauche</td><td>15 Juillet</td><td>E12</td><td>S1/S2</td><td>Rem.</td><td>Rem H08</td><td>M. Arbuthnot</td><td>B. Maureille</td><td>16 Juillet</td><td></td></tr><tr><td>38</td><td>Fragment de voûte crânienne</td><td>15 Juillet</td><td>E12</td><td>S1/S2</td><td>Rem.</td><td>Rem H09</td><td>M. Arbuthnot</td><td>B. Maureille</td><td>16 Juillet</td><td></td></tr><tr><td>39</td><td>Fragment de frontal, région sus-orbitaire à gauche</td><td>15 Juillet</td><td>E12</td><td>S1/S2</td><td>Rem.</td><td>Rem H10</td><td>M. Arbuthnot</td><td>B. Maureille</td><td>1er Septembre</td><td></td></tr><tr><td>40</td><td>Fragment de parietal</td><td>27 Juillet</td><td>D12</td><td>S3</td><td>Fa. 2a</td><td>1053</td><td>M. Arbuthnot</td><td>M. Arbuthnot</td><td>Idem mise à jour</td><td></td></tr><tr><td>41</td><td>Fragment de voûte crânienne</td><td>27 Juillet</td><td>D12</td><td>S3</td><td>Fa. 2a</td><td>1059</td><td>M. Arbuthnot</td><td>M. Arbuthnot</td><td>Idem mise à jour</td><td></td></tr><tr><td>42</td><td>Fragment de parietal droit</td><td>28 Juillet</td><td>D12</td><td>S3</td><td>Fa. 2a</td><td>1069</td><td>M. Arbuthnot</td><td>M. Arbuthnot</td><td>Idem mise à jour</td><td></td></tr><tr><td>43</td><td>Fragment crâne (maxillaire ?)</td><td>31 Juillet</td><td>D09</td><td>-</td><td>Rem.</td><td>Rem H11</td><td>B. Maureille</td><td>B. Maureille</td><td>Idem mise à jour</td><td></td></tr><tr><td>44</td><td>Phalange moyenne de la main (RIII ou RIV)</td><td>05 Août</td><td>D12</td><td>S3</td><td>Fa. 2a</td><td>1170</td><td>J. Nassar</td><td>J. Nassar</td><td>Idem mise à jour</td><td></td></tr><tr><td>45</td><td>Germe de troisième molaire inférieure droite</td><td>12 Août</td><td>D12</td><td>S3</td><td>Fa. 2a</td><td>1307</td><td>J. Nassar</td><td>J. Nassar</td><td>Idem mise à jour</td><td></td></tr></tbody></table>

Dans les colonnes remontages : cases noircies dans une même colonne = remontage ; case grisée = appartenance possible au même ossement, au même sujet
**Annexes :**
figures
<figure><img src="image_12.png" /></figure>

<figure><img src="image_13.png" /><figcaption>Figure 1. Localisation du gisement, extrait de la carte IGN n° 1831 Ouest et du plan cadastral section B, feuille 1, n° 515.</figcaption></figure>
<figure><img src="image_14.png" /></figure>

<figure><img src="image_15.png" /><figcaption>Figure 2. Extrait de la carte géologique de la France à 1/50.000, feuille La Rochefoucauld, (Bourgueil *et al.*, 1982)</figcaption></figure>
<figure><img src="image_16.png" /></figure>

<figure><img src="image_17.png" /><figcaption>Figure 3. Contexte géomorphologique du site des Pradelles.</figcaption></figure>
<figure><img src="image_18.png" /></figure>

<figure><img src="image_19.png" /><figcaption>Figure 4. Plan topographique général et orientation du carroyage.</figcaption></figure>
<figure><img src="image_20.png" /><figcaption>Figure 5. Relevé en plan de la dépression, de l'abri sous-roche et des principales galeries (et localisation du carroyage).</figcaption></figure>
<figure><img src="image_21.png" /><figcaption>Figure 6. Relevé en plan du témoin Est et position des fouilles clandestines.</figcaption></figure>
<figure><img src="image_22.png" /><figcaption>Fig. 7. Feuille de relevé : grille pour le dessin.</figcaption></figure>
<figure><img src="image_23.png" /><figcaption>Figure 9. Localisation, description et coordonnées des points de référence EDM dans
le carroyage Vandermeersch.</figcaption></figure>
<figure><img src="image_24.png" /><figcaption>Figure 10. Localisation des secteurs fouillés entre 2002 et 2005.</figcaption></figure>
<figure><img src="image_25.png" /><figcaption>Figure 11. Les Pradelles, témoins Ouest et Est, fouilles 2002-2005.
A : localisation en plan des objets archéologiques coordonnés avec mise en exergue (couleur grise) des pièces exhumées en 2005.</figcaption></figure>

B : objets archéologiques par type de vestiges (blanc = restes osseux, dentaires, bois de cervidés et coprolithes, bleu = restes lithiques).
<figure><img src="image_26.png" /></figure>

Figure 12. Les Pradelles, témoins Est, Ouest et galerie aux poules, fouilles 2002-2005. Transects sagittaux et localisation des objets archéologiques coordonnés (vues sagittales depuis le Nord du carroyage).

A : par campagne archéologique (2005 = couleur grise)
B : par type de vestiges (rouge = restes osseux, dentaires, bois de cervidés et coprolithes, bleu = restes lithiques).
<figure><img src="image_27.png" /><figcaption>Figure 13. Les Pradelles 2002-2005, témoins Est, Ouest et galerie aux poules.</figcaption></figure>

Transects frontaux (vue frontale depuis l'Ouest du carroyage).

A : localisation en plan des objets archéologiques coordonnés avec mise en exergue (couleur grise) des pièces exhumées en 2005.

B : objets archéologiques par type de vestiges (rouge = restes osseux, dentaires, bois de cervidés et coprolithes, bleu = restes lithiques).
<figure><img src="image_28.png" /></figure>

Figure 14. Transects frontaux et localisation des objets archéologiques coordonnés avec mise en exergue des pièces exclues du corpus d'étude (en gris foncé). Vue frontale depuis l'Ouest du carroyage.
<figure><img src="image_29.png" /></figure>

<figure><img src="image_30.png" /><figcaption>Figure 15. A : la patte de *Bovinae* et le maxillaire de Cheval (carrés S4 & S5) mis au jour dans la galerie aux poules en Août 2005.</figcaption></figure>

<caption>B : Le frontal de bison avec ses chevilles osseuses mis au jour en 1967 à la base de la stratigraphie probablement en E-F/10 à 12.</caption>
<figure><img src="image_31.png" /><figcaption>Figure 16. Vue des dépôts mis au jour dans le carré D6 (soulignés par un trait) et interprétés comme une coulée de débris sub-actuelle.</figcaption></figure>
<figure><img src="image_32.png" /></figure>

Figure 17. Coupes stratigraphiques observées sur les parois sud, ouest et nord du sondage réalisé à l'entrée d'une galerie karstique s'ouvrant dans la partie sud-ouest du site. 0,1, 2, 3 : unités stratigraphiques. L'étoile marque l'emplacement d'un crâne d'Equidé.
<figure><img src="image_33.png" /><figcaption>Figure 18. Vue de la paroi sud et d'une partie de la paroi ouest du sondage. Au premier plan, le crâne d'Equidé associé à l'unité 2.</figcaption></figure>
<figure><img src="image_34.png" /></figure>

Légende. 1 : calcaire. 2 : F - Horizon A. 3 : F - Horizon Sca. 4 : F - Horizon Cca. 5 : cailloutis stratifié. 6 : éboulis ouvert à semi-ouvert. 7 : diamicton à support clastique. 8 : lentilles jaune rougeâtre à support clastique colmaté.

9 : éboulis semi-ouvert à matrice brun foncée. 10 : éboulis colmaté à matrice brun foncé. 11 : argiles limoneuses.

12 : lentille jaune rougeâtre à fraction grossière hétérométrique. 13 : zones masquées. 14 : galerie et vides.

<caption>Figure 19. Litho-stratigraphie du témoin principal.</caption>

A, B, C, D, E, F, G : unités lithostratigraphiques. Les chiffres indiqués entre parenthèses renvoient au lithofaciès. Les lettres affichées entre parenthèses à côté de "F" désignent les horizons du sol de surface.
<figure><img src="image_35.png" /></figure>

Figure 20. Vue de la partie supérieure de la série stratigraphique montrant l'orientation des blocs de la partie supérieure de l'unité D (flèches), les éboulis stratifiés et le calcosol développé sur l'unité F.
<figure><img src="image_36.png" /></figure>

Figure 21. Certains des vestiges osseux ramassés en surface dans la galerie aux poules et ayant motivé notre décision de réaliser enfin ce sondage. On note de haut en bas et de la gauche vers la droite : 2 rochers digérés de chevaux, 1 esquille d'os long d'ongulé de grande taille très corrodée et un métacarpien d'Auroch.
<figure><img src="image_37.png" /></figure>

<figure><img src="image_38.png" /></figure>

<figure><img src="image_39.png" /><figcaption>Figure 22. Projections des objets coordonnés dans la « galerie aux poules » et raccords de remontage.
Couleur rouge : intermédiaire ou remanié (unité 0) ; rose : remanié intermédiaire ou remanié inférieur (unité interface 0 / 2, 1 / 2) ; jaune : jaune (unité 2)</figcaption></figure>
<figure><img src="image_40.png" /><figcaption>Figure 23. Projection sur un plan frontal des vestiges coordonnés du témoin Est.
\ : marmotte, × : cheval, o : bovin ; + : autres taxons
Bleu ciel : faciès 5 ; jaune : faciès 4b ; rouge : faciès 4a ; rose : faciès 2b ; violet : faciès 2a ; noir : autres faciès</figcaption></figure>
<figure><img src="image_41.png" /><figcaption>Figure 24. Projection sur plan sagittal des vestiges coordonnés du témoin Est</figcaption></figure>
<figure><img src="image_42.png" /><figcaption>Figure 25. Os d'Oiseaux provenant de la Galerie aux Poules.</figcaption></figure>

<figure><img src="image_43.png" /><figcaption>Figure 26. Représentation différentielle des principaux ossements crâniens de *Gallus gallus domesticus* exprimée en NME.</figcaption></figure>

<figure><img src="image_44.png" /><figcaption>et état général de fragmentation du squelette céphalique de Poule et de Pintade (en haut à droite)</figcaption></figure>
<figure><img src="image_45.png" /></figure>

Figure 27. Stries de découpe sur la face inférieure d'un crâne de *Gallus gallus domesticus* (à gauche), agrandissement (à droite).

<figure><img src="image_46.png" /></figure>

Figure 28. Stries de découpe sur crânes de *Gallus gallus domesticus* en vue inférieure. Deux coups ont été portés sur chacun des crânes.

<figure><img src="image_47.png" /></figure>

Figure 29. Surface de tranchage verticale sur un crâne de *Gallus gallus domesticus* en vue inférieure.
<figure><img src="image_48.png" /><figcaption>Figure 30. Coupures dans le prolongement de surfaces de tranchage verticale sur des crânes de Gallus gallus domesticus, en vue inférieure.</figcaption></figure>

<figure><img src="image_49.png" /><figcaption>Figure 31. Coupure et surface de tranchage verticale sur des crânes de *Gallus gallus domesticus*, en vue inférieure</figcaption></figure>

<figure><img src="image_50.png" /><figcaption>Figure 32. Surfaces horizontales de tranchage sur des crânes de *Gallus gallus domesticus* en vue inférieure (à gauche), agrandissements (à droite).</figcaption></figure>
<figure><img src="image_51.png" /></figure>

Figure 33. A gauche, agrandissement de la figure 8 montrant des surfaces horizontales de tranchage et des écailles d'os en bas à droite; à droite, agrandissement des stries de tranchage orientées grosso modo d'avant en arrière du crâne.
<figure><img src="image_52.png" /></figure>

Figure 34. Fragments mandibulaires en vue inférieure montrant des surfaces sub-horizontales de tranchage au niveau de l'articulaire.

<figure><img src="image_53.png" /></figure>

Figure 35. Fragment de mandibule en vue inférieure montrant des stries et des écailles d'os.

<figure><img src="image_54.png" /></figure>

Figure 36. Humérus et tarsométatarse de *Gallus gallus domesticus* montrant des traces de manducation (punctures, pits) laissées par un petit carnivore.
<figure><img src="image_55.png" /><figcaption>Figure 37. Neurocrâne de *Gallus gallus domesticus* provenant d'un terrier de petits carnivores et portant des traces de manducation à l'arrière du crâne.</figcaption></figure>

<figure><img src="image_56.png" /><figcaption>Figure 38. "Saigneur de volaille" et bouchon servant à protéger la pointe.</figcaption></figure>
<figure><img src="image_57.png" /><figcaption>Figure 39. Pourcentage de représentation (PR) de chaque éléments anatomiques.</figcaption></figure>

Fragment de tibio fibula S46.21 (Bufo ?) présentant une importante dégradation de la corticale compatible avec l'action de sucs digestifs de carnivores.

<figure><img src="image_58.png" /></figure>

Tibio fibula R4.9 (Bufo juvénile) présentant une légère altération de la corticale.

<figure><img src="image_59.png" /></figure>

Scapula droite S46.26 (Bufo juvénile) présentant d'importantes pertes osseuses.

<figure><img src="image_60.png" /></figure>

Humérus droit Q1.1 (Bufo calamita femelle) présentant une perforation de l'acétabulum.

<figure><img src="image_61.png" /><figcaption>Figure 40. Traces de prédation sur Bufo.</figcaption></figure>
<figure><img src="image_62.png" /></figure>

Figure 41. Pourcentages des restes présentant des traces de prédation par catégorie d'éléments anatomiques.

<figure><img src="image_63.png" /></figure>

Figure 42. Comparaison de l'ilion droit chez *Bufo*.

L'ilion de *Bufo calamita* se caractérise par sa tubérosité dorsale en pointe. Outre un plus grand format et une plus grande robustesse, l'ilion de *Bufo bufo* porte une tubérosité plus mousse parfois polylobée.

L'ilion de *Bufo bufo* présente une patine de l'os cortical caractéristique de l'action de sucs digestifs.

Les disparitions d'os spongieux dans la partie postérieure de l'ilion de *Bufo calamita* sont récentes et ont certainement étaient occasionnées lors de la fouille ou durant le tamisage. Ce type de dégradation n'a pas été enregistré comme lié à une action de prédation.
<figure><img src="image_64.png" /></figure>

Bufo calamita
Faciès 5

<figure><img src="image_65.png" /></figure>

Bufo viridis
Faciès 4b & 5

<figure><img src="image_66.png" /></figure>

<figure><img src="image_67.png" /></figure>

Bufo bufo juvéniles
Faciès 5 et 3

Figure 43. Comparaison du sphénethmoïde chez plusieurs espèces de *Bufo*.

*Bufo calamita* : léger épaississement du plancher de chacune des deux alvéoles antérieures (*sella amplificans*).

*Bufo viridis* : *sella amplificans* très développés.

*Bufo bufo* : sphénethmoïde plus large que long. Absence de *sella amplificans*.
### Alytes obstetricans

S44.2 Faciès 5.

Pas de crête dorsale, apophyses transverses courtes et larges

<figure><img src="image_68.png" /></figure>

1mm

<figure><img src="image_69.png" /></figure>

### Bufo calamita

S24.1. Faciès 5.

crête dorsale développée, absence d'apophyse transverse, cotyle d'articulation avec la vertèbre sacrée double et partiellement soudé médialement.

<figure><img src="image_70.png" /><figcaption>Figure 44. Identification d'Alytes obstetricans.</figcaption></figure>

### Rana temporaria

-Humérus droit Rana temporaria actuelle, Haute Pyrénées.

De gauche à droite :

- R4.2 Faciès 5

- S34.1 Faciès 4b

- R8.5 Faciès 5

### Rana dalmatina

-Humérus gauche Rana dalmatina actuelle, Gironde.

De gauche à droite :

- S21.1 Faciès 4a

- S47.6 Faciès 5

- R8.15 Faciès 5

### Rana juvéniles indéterminés

De gauche à droite :

- S4.1 Faciès 2a

- R20 Faciès 5

- R12.7 Faciès 5

- S52.8 Faciès 5

- S53.12 Faciès 5

<caption>Figure 45. Comparaison des humérus de Rana.</caption>
<figure><img src="image_71.png" /><figcaption>Cunéiforme intermédiaire gauche – LP05 E12 Rem. H08</figcaption></figure>

<figure><img src="image_72.png" /></figure>

<figure><img src="image_73.png" /><figcaption>Fragment de frontal à droite – LP05 E12 Rem H10</figcaption></figure>

<figure><img src="image_74.png" /><figcaption>Figure 46. les quatre pièces humaines trouvées en position remaniée.</figcaption></figure>
A

<figure><img src="image_75.png" /></figure>

B

<figure><img src="image_76.png" /></figure>

C

<figure><img src="image_77.png" /><figcaption>Figure 47. Fragment de pariétal droit LP05 D12 S3 Fa 2a 1053.</figcaption></figure>

A : vestige en place.

B : face exo-crânienne à gauche et endo-crânienne à droite

C : détails des stries de découpe
<figure><img src="image_78.png" /><figcaption>Figure 48. Fragment de frontal à droite LP05 D12 S3 Fa 2a 1059.
A : vestige en place.
B : face endo-crânienne à gauche et exo-crânienne à droite</figcaption></figure>
<figure><img src="image_79.png" /><figcaption>Figure 49. Fragment de pariétal droit LP05 D12 S3 Fa 2a 1069.</figcaption></figure>

A : vestige en place.

B : face exo-crânienne à gauche et endo-crânienne à droite

C : détails des stries de découpe
<figure><img src="image_80.png" /><figcaption>Figure 50. Le fragment de voûte crânienne reconstitué à partir du remontage des trois pièces mises au jour en D12.</figcaption></figure>
<figure><img src="image_81.png" /><figcaption>Figure 51. Phalange moyenne de la main. LP05 D12 S3 Fa 2a 1170
A : en place sur le terrain
B : vue dorsale en haut, vue palmaire en base (épiphyse distale en haut)</figcaption></figure>

<caption>Figure 52. Germe de troisième molaire inférieure droite. LP05 D12 S3 Fa 2a 1307
A : en place sur le terrain.
B : face occlusale en haut et face apicale en bas (face mésiale en haut).</caption>
<figure><img src="image_82.png" /></figure>

Figure 53. A. Témoins Est et Ouest. Localisation en plan des objets archéologiques coordonnées par type de vestiges avec mise en exergue des pièces exhumées lors de la campagne 2005. En rouge : restes osseux, dentaires, bois de Cervidés et coprolithes coordonnés en 2005. En bleu : restes lithiques coordonnés en 2005. En blanc : tous les vestiges coordonnés lors des campagnes 2002 à 2004.

B. Témoins Est et Ouest. Localisation en plan des objets archéologiques coordonnés avec mise en exergue des restes humains : 2001 (en noir), 2002 (en vert), 2003 (en bleu), 2004 (en rouge) et 2005 (en jaune). Les restes figurés par un carré ont été trouvés en contexte remanié. En blanc : reste faunique ou lithique.
<figure><img src="image_83.png" /><figcaption>Figure 54. Les remontages entre les restes humains dans les carrés D12 et D12. Résultats des fouilles 2003, 2004 et 2005</figcaption></figure>
Annexes :

Publications 2005 jointes
<figure><img src="image_84.png" /></figure>

ELSEVIER

Available online at www.sciencedirect.com

SCIENCE @ DIRECT*

Journal of Human Evolution 49 (2005) 71–87

<figure><img src="image_85.png" /></figure>

# Isotopic evidence for diet and subsistence pattern of the Saint-Césaire I Neanderthal: review and use of a multi-source mixing model

Hervé Bocherens<sup>a,b,*</sup>, Dorothée G. Drucker<sup>c</sup>, Daniel Billiou<sup>d</sup>, Marylène Patou-Mathis<sup>e</sup>, Bernard Vandermeersch<sup>f</sup>

Received 10 December 2004; accepted 16 March 2005

## Abstract

The carbon and nitrogen isotopic abundances of the collagen extracted from the Saint-Césaire I Neanderthal have been used to infer the dietary behaviour of this specimen. A review of previously published Neanderthal collagen isotopic signatures with the addition of 3 new collagen isotopic signatures from specimens from Les Pradelles allows us to compare the dietary habits of 5 Neanderthal specimens from OIS 3 and one specimen from OIS 5c. This comparison points to a trophic position as top predator in an open environment, with little variation through time and space. In addition, a comparison of the Saint-Césaire I Neanderthal with contemporaneous hyaenas has been performed using a multi-source mixing model, modified from Phillips and Gregg (2003, Oecologia 127, 171). It appears that the isotopic differences between the Neanderthal specimen and hyaenas can be accounted for by much lower amounts of reindeer and much higher amounts of woolly rhinoceros and woolly mammoth in the dietary input of the Neanderthal specimen than in that of hyaenas, with relatively similar contributions of bovinae, large deer and horse for both predators,

<footnote><sup>a</sup> Institut des Sciences de l'Evolution, UMR 5554, Université Montpellier 2, Place E. Bataillon, F-34095 Montpellier cedex 05, France</footnote>

<footnote><sup>b</sup> PNWRC, Canadian Wildlife Service, Environment Canada, 115 Perimeter Road, Saskatoon, Saskatchewan, S7N 0X4 Canada</footnote>

<footnote><sup>c</sup> PNWRC, Canadian Wildlife Service, Environment Canada, 115 Perimeter Road, Saskatoon, Saskatchewan, S7N 0X4 Canada</footnote>

<footnote><sup>d</sup> Laboratoire de Biogéochimie des Milieux Continentaux, I.N.A.P.G., EGER-INRA Grignon, URM 7618, 78026 Thiverval-Grignon, France</footnote>

<footnote><sup>e</sup> Institut de Paléontologie Humaine, 1 rue René Panhard, 75 013 Paris, France</footnote>

<footnote><sup>f</sup> C/Nuñez de Balboa 4028001 Madrid, Spain</footnote>

<footnote>* Corresponding author. Institut des Sciences de l'Evolution, UMR 5554, Université Montpellier 2, Place E. Bataillon, F-34095 Montpellier cedex 05, France. Tel.: +33 4 67 14 32 60; fax: +33 4 67 14 36 10.</footnote>

<footnote>E-mail addresses: bocheren@isem.univ-montp2.fr (H. Bocherens), dorothee.drucker@tele2.fr (D.G. Drucker), billiou@grignon.inra.fr (D. Billiou), patmath@cimrs1.mnhn.fr (M. Patou-Mathis), bvanderm@bio.ucm.es (B. Vandermeersch).</footnote>
a conclusion consistent with the zooarchaeological data. The high proportion of very large herbivores, such as woolly rhinoceros and woolly mammoth, in Neanderthal's diet compare to that of the scavenging hyaenas suggests that Neanderthals could not acquire these prey through scavenging. They probably had to hunt for proboscideans and rhinoceros. Such a prey selection could result from a long lasting dietary tradition in Europe.
© 2005 Elsevier Ltd. All rights reserved.

Keywords: Neanderthal; Saint-Césaire; Diet; Stable isotopes; Collagen; Hyaena; Hunting strategies; Western Europe

## Introduction

Palaeodietary reconstruction using stable carbon and nitrogen isotopic signatures of collagen of European Pleistocene Hominids was first evaluated with Neanderthals of Marillac Cave (Charentes, France; today referred as “Les Pradelles”) (Bocherens et al., 1991; Fizet et al., 1995). Since this pioneering research, the palaeodiet of further Neanderthal specimens has been investigated using isotopic approaches, for specimens from Belgium (Bocherens et al., 1999, 2001), Slovenia (Richards et al., 2000), and more recently the Neanderthal I specimen from Saint-Césaire (Bocherens and Drucker, 2003a; Drucker and Bocherens, 2004). An attempt to extend this approach to a Middle Eastern Neanderthal failed due to poor collagen preservation (Ambrose, 1998). All of these investigations have suggested that isotopic signatures of Neanderthal were similar to those of contemporary animal predators indicating that the diet of Neanderthals was dominated by the meat of large herbivores. Such a conclusion is consistent with the results of other archaeological approaches aimed at reconstructing Neanderthal's diet (e.g. zooarchaeology in Gaudzinski, 1996 and tooth wear patterns in Lalueza Fox and Pérez-Pérez, 1993).

However, other questions related to Neanderthal's dietary behaviour are crucial for palaeoanthropologists. For instance, it is still debated whether Neanderthals hunted actively or merely acquired ungulate meat through scavenging (e.g. Binford, 1988; Mellars, 1989; Patou, 1989; Marean, 1998), whether the hunting strategies of Neanderthals were different from those of early modern Humans (e.g. Chase, 1987; Stiner, 1994; Marean and Assefa, 1999; Pike-Tay et al., 1999; Hardy et al., 2001; Speth and Tchernov, 2001), and whether diet and health differences can

account for the transition between Middle and Upper Palaeolithic and for the extinction of Neanderthals (e.g. Skinner, 1996; Cachel, 1997). The isotopic approach is destructive, and since Neanderthal specimens are very rare and precious, bringing insights into these important questions should now be the goal of future applications of isotopic investigations. Some limitations of the isotopic approach will remain, such as the limited number of Neanderthal samples available for analyses. However, certain limitations can be overcome to provide more detailed information about Neanderthal dietary habits, hunting strategies, and the possible role of dietary changes in the process of their extinction.

The Neanderthal specimen Saint-Césaire I is very useful for the purpose of inferring the influence of dietary habits and hunting strategies in the modalities of extinction of these hominids because it has been excavated from a rich archaeological layer with numerous faunal remains. Moreover, it is located in a region rich in archaeological and palaeontological sites of similar age, as well as with older and younger sites. Finally, it is dated from the transition between Middle and Upper Palaeolithic, and thus it represents one of the youngest Neanderthal specimens in Western Europe, possibly contemporaneous with early modern Humans in Europe.

This paper aims at addressing two key questions:

(1) Is there any isotopic evidence indicating a change in dietary strategies for Neanderthals between Middle-Upper Palaeolithic transition and Middle Palaeolithic Neanderthals? If a change is documented, this could indicate a dietary shift linked to increasing competition with other predators, such as anatomically modern humans for instance.
(2) Did the Saint-Césaire Neanderthal have a similar prey selection as animal predators such as hyaena? Hyaena being a scavenger, a different prey selection for this Neanderthal could indicate a different strategy for prey acquisition, i.e. hunting rather than scavenging.

Finally, the implications for the extinction of Neanderthals of our findings on dietary habits and the hunting strategies of these hominids will be considered.

## Material and Methods

Bone and tooth material were selected from the Chatelperonian layers Ejop and Ejop sup from Saint-Césaire, which yielded the Neanderthal remains. Prior to collagen extraction, nitrogen amount in bones has been measured to screen out specimens devoid of organic matter. Only specimens with nitrogen content higher than 0.4 % have been chosen for collagen extraction (Drucker et al., 1999). In this last study, only four bones out of 28 yielded acceptable collagen. Although four additional bones from the Chatelperonian layers yielded reliable collagen, the number of faunal remains occuring with the human skeleton and useful for stable isotope analyses is still limited.

The scarcity of well preserved bones in the Chatelperonian layers of Saint-Césaire led us to look for additional faunal material from other sites from the same region of similar age. Two sites were selected: Camiac and La Berbie. These two sites are located about 120 to 150 km south-east of Saint-Césaire (Fig. 1). Both sites are natural accumulations that have yielded rich faunal assemblages dated around 35,000 years BP (Gaudelli, 1987; Madelaine, 1999). The proportion of bones and teeth still containing well preserved collagen is high, with only one specimen from La Berbie not yielding collagen. Material from the main ungulate species possibly consumed by Neanderthals and animal predators has been selected: large bovids (bovinae), i.e. *Bison priscus* and *Bos primigenius*, both species being present and difficult to distinguish from fragmentary

skeletal material; large deer, i.e. giant deer *Mega loceros giganteus* and red deer *Cervus elaphus*, whose isotopic signatures are very similar; reindeer *Rangifer tarandus*; horse *Equus caballus*; woolly rhinoceros *Ceolodonta antiquitatis*; and woolly mammoth *Mammuthus primigenius*. In order to complete the trophic reconstruction of an animal predator, we have selected bone material from cave hyaena *Crocuta spelaea*, a hunter and scavenger of ungulates (e.g. Baryshnikov, 1999).

In large mammals, bone collagen is considered to reflect several years of the lifetime of the studied individual prior to its death (e.g. Klepinger, 1984; Parfitt, 2002; Price et al., 2002). Tooth dentine from cervids and carnivores has not been sampled since this tissue has been shown to yield nitrogen isotopic signatures different from bone from the same individuals, possibly due to the influence of ¹⁵N-enriched collagen incorporated in dentine during the nursing period (Bocherens et al., 1994; Bocherens and Mariotti, 1997).

Collagen was extracted from bones with more than 0.4 % nitrogen, which corresponds to a maximum collagen loss of 90 % (Bocherens et al., in press). The protocol routinely used for Pleistocene samples has been used to extract collagen (Bocherens et al., 1997). Carbon and nitrogen were evolved from the extracted collagen as CO₂ and N₂, respectively, via flash combustion followed by gas chromatography in an Elemental Analyser (Carlo Erba NA 1500) connected to an Isotopic Ratio Mass Spectrometer (VG Optima) in the department of Earth Sciences, University P. & M. Curie (Paris, France). This allows the measurement of the percentage of carbon and nitrogen and the isotopic ratios ¹³C/¹²C and ¹⁵N/¹⁴N in the analysed collagen. Isotopic ratios are expressed as δ (delta) values, as follows: δ¹³C = (Rsample/Rstandard - 1)·1000 (‰), where X stands for C or N, E stands for 13 or 15 respectively, and R stands for the isotopic ratios ¹³C/¹²C and ¹⁵N/¹⁴N respectively. The standard is the marine carbonate VPDB for carbon and atmospheric nitrogen (AIR) for nitrogen. Analytical error is 0.1 ‰ and 0.2 ‰ for δ¹³C values and δ¹⁵N values, respectively.

A mathematical mixing model based on isotopic data from western France can be used to
<figure><img src="image_86.png" /><figcaption>Fig. 1. Map depicting the location of the sites discussed in the text.</figcaption></figure>

determine the likely composition of the diet of Neanderthals and that of an animal predator such as hyaena. Until recently, mixing models have been developed in order to cope with a situation with two isotopic pairs, such as $^{13}$C/$^{12}$C and $^{15}$N/$^{14}$N, and no more than 3 possible dietary sources (e.g., Phillips, 2001; Phillips and Gregg, 2001; Phillips and Koch, 2002). A preliminary attempt to quantitatively estimate reindeer, mammoth, and large ungulates (horse and bovinae) in the Saint-Césaire Neanderthal's diet using the model published by Phillips and Koch (2002) has been presented previously (Drucker and Bocherens, 2004). The recent publication of an upgraded mixing model that can cope with a situation with

two isotopic pairs and more than 3 dietary sources (Phillips and Gregg, 2003; Newsome et al., 2004) allows us to consider in the present study all the different possible prey available to Saint-Césaire Neanderthal.

Two modifications have been performed relative to the original approach developed by Phillips and Gregg (2003). Firstly, we subtracted the fractionation factors from the consumer signatures rather than adding it to the prey signatures. The aim of this modification was to enable the use of the isotopic signatures measured on the fossil material. Secondly, we used for these calculations a range of carbon and nitrogen isotopic fractionations between prey and predator
collagen instead of a constant value for these
fractionation factors. Previous work has shown
that the δ¹³C values of predator collagen is 0.8 to
1.3 ‰ more positive than that of its average prey,
whereas the δ¹⁵N values of predator collagen is 3
to 5 ‰ more positive than that of its δ¹⁵N of its
average prey (Bocherens and Drucker, 2003b;
Drucker and Bocherens, 2004). This calculation
yields four extreme carbon and nitrogen isotopic
signatures for the prey collagen of each predator,
which can be graphically depicted as a rectangle
that includes all the possible isotopic signatures for
the collagen of prey.

Using this approach, we obtain average carbon
and nitrogen isotopic signatures of the collagen of
six potential prey (i.e. Bovinae, large deer,
reindeer, horse, woolly rhinoceros, and woolly
mammoth). These six isotopic signatures were
compared with two rectangles of carbon and
nitrogen isotopic signatures representing the aver-
age isotopic signatures possible for the prey of
Neanderthal and for those of Hyaena (Fig. 2).
Following Phillips and Gregg (2003), the isotopic
values of the collagen of the six prey delineated
a four-sided surface delimited by the average
isotopic signature of woolly mammoth, horse,
reindeer, and woolly rhinoceros, with the isotopic
values of bovinae and large deer being enclosed
inside this surface (Fig. 2). Both rectangles of the
isotopic signatures of the collagen of the possible
prey collagen intersect this surface at two points
for Hyaena and at four points for Neanderthal
(Fig. 2). The possible percentage of each prey in
the predator's diet for these intersection points as
well as for the extreme predator isotopic signature
 rectang
( rectangle corner) that are also enclosed in surface
delimited by the average isotopic signature of
potential prey have been calculated using the
computer program IsoSource, available at http://
www.epa.gov/wed/pages/models.htm (Fig. 2). For
each chosen pair of carbon and nitrogen isotopic
signatures, this program calculates the statistical
distribution of the possible percentage of each prey
that is compatible with the measured isotopic
values. The calculation takes into account a toler-
ance value and a source increment, set at 0.05 and
2 %, respectively. For each chosen point, results
are presented as the average percent values with

<figure><img src="image_87.png" /><figcaption>Fig. 2. Summary of the average isotopic signature of prey species in Saint-Césaire, La Berbie and Camiac, compared with the possible range of prey collagen isotopic signatures for Hyaena and the Saint-Césaire Neanderthal. Fractionation factors have been subtracted from the consumer signatures. δ¹³C and δ¹⁵N values are given for the points of intersection between the surface delimited by the average isotopic signature of potential prey (see Appendix) and the rectangles of the possible prey collagen isotopic signatures, as well as for the extreme values of the rectangles of the possible prey collagen isotopic signatures enclosed into this surface. The percentage calculations have been performed for these points.</figcaption></figure>

their standard-deviation for each prey, as well as
the possible range of percentage for each prey.
Finally, we combined the results obtained for the
extreme points chosen to estimate the most likely
range of possible percentage for each prey, with
standard-deviation and extreme possible ranges.
## Results and Discussion

*Isotopic evidence for possible changes
in dietary strategy between Middle Palaeolithic
and Middle-Upper Palaeolithic
transition Neanderthals*

The first question that we address is whether isotopic evidence points to a change in dietary strategy for a Neanderthal specimen from the Middle-Upper Palaeolithic transition, Saint-Césaire I, relative to Middle Palaeolithic Neanderthals. In order to accomplish this goal, we first review the published isotopic signatures of other Neanderthal specimens, and then compare the suitable isotopic data to the isotopic signatures of Saint-Césaire I specimen.

*Review of the isotopic signatures of Neanderthal
specimens usable for palaeodietary reconstruction*

So far, twelve Neanderthal specimens were analysed for the carbon and nitrogen isotopic signature of their bone collagen. Ten adults and two children (Table 1), including three new isotopic analyses performed on specimens from Les Pradelles (formerly known as “Marillac”) are presented in this paper. However, isotopic signatures of prehistoric human collagen need to follow a number of requirements in order to be usable for palaeodietary reconstruction. The first requirement is to assess the chemical reliability of the extracted collagen. The second requirement has to do with the period of life reflected in the extracted collagen, especially with respect to nursing. The third requirement is to have a reasonably good set of isotopic data on contemporaneous herbivorous and carnivorous mammals in order to reconstruct the trophic structure of the foodweb.

The first aspect to consider is the reliability of the isotopic data obtained on Neanderthals. Reliability of the isotopic signatures of ancient collagen is addressed through its chemical composition, most notably percent carbon (%C) and nitrogen (%N), and C:N ratio (DeNiro, 1985; Ambrose, 1990). Only collagen extracts with C:N ratios ranging from 2.9 to 3.6 and %N > 5 % are considered uncontaminated and not significantly altered. Among the isotopic results obtained on

Neanderthals, two from Les Pradelles have been measured on collagen extracts that had ambiguous chemical compositions. The first one was originally published by Bocherens et al. (1991) at a time when carbon and nitrogen percent determinations on collagen were not performed routinely. This specimen had its amino acid composition determined instead, and the resulting pattern was close but not identical to that of fresh collagen (Bocherens et al., 1991). The measured isotopic signatures are probably close to the original ones, but they may have been slightly shifted due to chemical degradation. It is thus preferable to discard this result from further discussion. Two of the three new isotopic measurements (M300 and M400) exhibit chemical compositions well within the range of fresh collagen chemical compositions. In contrast, M100 exhibits carbon and nitrogen concentrations lower than those of fresh collagen, although its C:N ratio is within the range of that of fresh collagen (Table 1). Using the criteria proposed by Ambrose (1990), this extract can be considered at the lower limit of reliability, and therefore we prefer to be conservative and discard this specimen also from palaeodietary interpretations. It is noteworthy that this last specimen has been sampled from an occipital bone exhibiting cut marks (Le Mort, 1987). This specimen might then correspond to an individual who was submitted to a particular treatment. This raises questions about the possible causes of the isotopic particularity of this specimen relative to other Neanderthals. For example this individual was perhaps linked to a different state of preservation or a specific status of this individual during his lifetime reflected in his diet, or both. With the recent reactivation of excavation in Les Pradelles, additional human remains have been unearthed (Beauval et al., 2002; Maureille, 2003), and further isotopic work will most likely clarify some of these uncertainties.

The second aspect to consider is the individual age of the Neanderthal specimens, in order to take into account the possible nursing effects on δ¹⁵N values of children. Indeed, nursing mammals, including humans, are enriched in δ¹⁵N relative to their mother because milk presents δ¹⁵N values similar to those of the mother's tissues (e.g. Fogel et al., 1989; Katzenberg and Pfeiffer, 1995).
<table><caption>Table 1
Summary of the Neanderthal specimens which collagen has been analysed for δ¹³C and δ¹⁵N values</caption><thead><tr><th>No analysis</th><th>Site</th><th>Age</th><th>Piece</th><th>No excavation</th><th>Yield (mg g<sup>-1</sup>)</th><th>%C</th><th>%N</th><th>C/N</th><th>δ<sup>13</sup>C</th><th>δ<sup>15</sup>N</th><th>Reference</th></tr></thead><tbody><tr><td>27801</td><td>Les Pradelles (Marillac-le-Franc, Charente, France)</td><td>OIS3</td><td>skull fragment</td><td></td><td>7.0</td><td>n.d.</td><td>n.d.</td><td>n.d.</td><td>-20.2*</td><td>9.3*</td><td>Bocherens et al. (1991)</td></tr><tr><td>64801</td><td>Les Pradelles (Marillac-le-Franc, Charente, France)</td><td>OIS3</td><td>skull fragment</td><td></td><td>82.4</td><td>38.8</td><td>13.7</td><td>3.3</td><td>-19.1</td><td>11.6</td><td>Fizet et al. (1995)</td></tr><tr><td>M300</td><td>Les Pradelles (Marillac-le-Franc, Charente, France)</td><td>OIS3</td><td>skull fragment</td><td>M70 c10 F10-41</td><td>107.4</td><td>41.1</td><td>14.7</td><td>3.3</td><td>-19.1</td><td>11.5</td><td>this paper</td></tr><tr><td>M400</td><td>Les Pradelles (Marillac-le-Franc, Charente, France)</td><td>OIS3</td><td>mandible</td><td>H1</td><td>41.5</td><td>37.6</td><td>13.1</td><td>3.3</td><td>-19.5</td><td>11.4</td><td>this paper</td></tr><tr><td>M100</td><td>Les Pradelles (Marillac-le-Franc, Charente, France)</td><td>OIS3</td><td>skull fragment°</td><td>H2</td><td>13.4</td><td>18.9</td><td>7.0</td><td>3.2</td><td>-21.8**</td><td>8.4**</td><td>this paper</td></tr><tr><td>RPB7000</td><td>La Roche-à-Pierrot (Saint-Césaire, Charentes-Maritimes, France)</td><td>OIS3</td><td>fibula</td><td></td><td>26.0</td><td>40.8</td><td>14.2</td><td>3.3</td><td>-19.8</td><td>11.4</td><td>Bocherens and Drucker (2003a)</td></tr><tr><td>MT500</td><td>Scladina Cave (Sclayn, Belgium)</td><td>OIS5b?§</td><td>phalanx</td><td>SCLA 1B 4</td><td>n.d.</td><td>39.9</td><td>14.7</td><td>3.2</td><td>-21.2</td><td>11.8</td><td>Bocherens et al. (2001)</td></tr><tr><td>SC18800</td><td>Scladina Cave (Sclayn, Belgium)</td><td>OIS5c</td><td>skull fragment</td><td>SCLA 4A 2</td><td>62.0</td><td>38.7</td><td>14.1</td><td>3.2</td><td>-19.9</td><td>10.9</td><td>Bocherens et al. (1999)</td></tr><tr><td>MT100</td><td>Awirs Cave (Belgium)</td><td>OIS3</td><td>skull fragment</td><td>Engis 2</td><td>176.4</td><td>41.7</td><td>14.4</td><td>3.4</td><td>-19.6#</td><td>12.6#</td><td>Bocherens et al. (2001)</td></tr><tr><td>MT200</td><td>Spy (Betche-al-Roche) Cave (Belgium)</td><td>OIS3</td><td>scapula</td><td>SPY OMO 1</td><td>123.9</td><td>41.5</td><td>14.4</td><td>3.4</td><td>-19.8</td><td>11.0</td><td>Bocherens et al. (2001)</td></tr><tr><td>VI-207</td><td>Vindija Cave (Croatia)</td><td>OIS3</td><td>mandible</td><td></td><td>65</td><td>37.1</td><td>13.5</td><td>3.2</td><td>-19.5</td><td>10.1</td><td>Smith et al. (1999)</td></tr><tr><td>VI-208</td><td>Vindija Cave (Croatia)</td><td>OIS3</td><td>parietal</td><td></td><td>42</td><td>36.1</td><td>11.7</td><td>3.6</td><td>-20.5</td><td>10.8</td><td>Smith et al. (1999)</td></tr></tbody></table>

<footnote>§ this specimen has been relocated since the initial publication (Bocherens et al., 2001) and belongs to layer 3 (OIS 5b) instead of layer 1B (OIS 3)</footnote>

<footnote>* at the time of the original study, no measurement of %C and %N were performed, and amino acid analysis showed a pattern close to collagen but not identical; the isotopic reliability of this collagen cannot be proven</footnote>

<footnote>° this specimen exhibits cut-marks</footnote>

<footnote>** this specimen exhibits chemical characteristics acceptable for well preserved collagen, but not identical to fresh collagen</footnote>

<footnote># this specimen is juvenile and may present an extra-¹⁵N-enrichment due to milk consumption</footnote>
Therefore, a nursing infant behaves similar to a predator, one trophic level higher than adult individuals of his population, as far as $\delta^{15}$N values are concerned. The Engis Neanderthal child is only 5-6 years old based on the dental eruption pattern (Tillier, 1983), and a child this age may still be affected by such an effect if weaning occurred later than for most modern human populations (Bocherens et al., 2001). Therefore, due to the uncertainty regarding his nursing status, this specimen will not be discussed in palaeodietary terms. Another Neanderthal child is the specimen from layer 4A in Scladina Cave. This specimen has been aged at approximately 11 years (Otte et al., 1993). Consequently, this specimen was old enough to have erased the nursing isotopic signal and thus, it is acceptable for palaeodietary reconstruction.

A final aspect to consider is the chronological distribution of the studied Neanderthals and the availability of isotopic data on contemporary faunal material. Indeed, considering pertinent co-eval fauna is crucial before drawing palaeodietary conclusions. For instance, significant variations occurred in herbivorous large mammals from Western Europe between 33,000 and 15,000 years ago (Drucker et al., 2003), and comparing the isotopic signatures of prehistoric humans with those of faunal specimens from a different age can be misleading (e.g. Drucker and Bocherens, 2004). A prerequisite is thus to compare the isotopic signatures of ancient humans with those of herbivores and carnivores from the same site and age, or at least from sites relatively close geographically and with a reasonable chronological similarity.

The two Neanderthals from Vindija do not meet this prerequisite, since there is only one deer from the same layer ($G_1$) than the human specimens for which isotopic signatures have been measured (Richards et al., 2000). The few other members of the fauna from the same site are from an older stratum, layer $G_3$ (Richards et al., 2000). Since this layer is more than 12,000 years older than the one that yielded the Neanderthal specimens (Ahern et al., 2004), they cannot be directly compared to the Neanderthal specimens. A palaeodietary reconstruction of Vindija Neanderthals' diet is thus pending isotopic data on contemporary herbivorous and carnivorous fauna.

In Belgium, the Neanderthal from Spy was found during the 19th century and no faunal remains clearly associated with the human could be selected. However, in this case, a comparison can be performed with the isotopic data evolved from the rich contemporary mammal fauna of layer 1A from Scladina Cave, both dated around 40,000 years ago and only a few kilometres apart (Bocherens et al., 2001). The Neanderthal specimen from Scladina Cave called “SCLA-1B” was originally attributed to layer 1B and thus compared to the mammal fauna from layers 1A and 1B from Scladina Cave (Bocherens et al., 2001). However, a recent revision of Scladina Cave material reassessed this specimen to layer 3 (D. Bonjean, pers. comm. 2003). Therefore, this specimen is contemporary with Oxygen Isotopic Stage (OIS) 5b, around 90,000 years BP (Ellwood et al., 2004). This indicates that the specimen is approximately 50,000 years older than previously thought. As a result, an isotopic study of the mammal fauna of layer 3 of Scladina Cave is currently under way in order to determine the trophic position of this specimen. In sum, it is still premature to attempt a palaeodietary reconstruction for this Neanderthal. The oldest Neanderthal specimen from Scladina Cave comes from layer 4A, and is contemporary to OIS 5c, around 100,000 years BP. This specimen was compared with the isotopic data obtained on the fauna from the same layer (Bocherens et al., 1999).

In France, the three Neanderthal specimens with chemically reliable collagen from Les Pradelles can be compared with the isotopic data obtained on the fauna from the same site, dating from OIS 3 (Fizet et al., 1995). The Saint-Césaire I Neanderthal meets the prerequisite presented above since we have gathered isotopic data on herbivorous and carnivorous mammals from the same region and the same age, around 35,000 years ago (this paper).

In conclusion, among the 12 Neanderthal specimens with isotopic signatures, only 5 specimens from OIS 3 and one specimen from OIS 5c can be considered so far for palaeodietary reconstruction after considering the prerequisite for chemical, physiological and stratigraphical characteristics.
### Isotopic palaeodietary comparison of the Saint-Césaire Neanderthal with other Neanderthal specimens

The Saint-Césaire I Neanderthal will first be compared with the Neanderthals from Les Pradelles since these sites are separated by only a few dozen kilometres. Les Pradelles yielded a Mousterian lithic assemblage together with human Neanderthal osteological material during excavations directed by B. Vandermeersch (1976). A rich faunal assemblage was recovered during this excavation (Fizet et al., 1995). Isotopic results obtained on faunal specimens from layer 7 have been discarded due to a general increase in δ¹⁵N values for all the components of the food web during this period (Fizet et al., 1995) and since none of the Neanderthal specimens originate from this particular layer. The age of the Neanderthals is considered to be contemporary to Oxygen Isotopic Stage 3, between 40,000 and 45,000 years BP (Fizet et al., 1995). As discussed in the previous paragraph, only three of the five Neanderthal specimens from Les Pradelles for which isotopic measurements have been performed are considered to be totally reliable and will be discussed further. These three specimens exhibit very similar isotopic signatures (Table 1, Fig. 3). Although the diversity of the species for which isotopic data are available in Les Pradelles is not as high as for Saint-Césaire, it is possible to compare the relative position of the isotopic signatures of Les Pradelles Neanderthal specimens with the same ungulate species, i.e. reindeer, bovinae and horse, and with the predatory hyaena (Fig. 3). It appears that Les Pradelles Neanderthal specimens present δ¹³C values similar to those of animal predators, such as hyaena and wolf, in a similar way than the Saint-Césaire Neanderthal specimen presents similar δ¹³C values to hyaena. In both cases, Neanderthals exhibit δ¹⁵N values significantly more positive than those of animal predators (Fig. 3). Although the chronological and cultural contexts are different for Les Pradelles and Saint-Césaire Neanderthals, their trophic position seems to be similar in both cases.

The Neanderthal from Spy together with the rich mammal fauna from layer 1A from Scladina Cave offers the possibility of a thorough compar-

ison with the Saint-Césaire Neanderthal. Both sites present a similar pattern (Fig. 3). Among herbi-vores, woolly mammoth and woolly rhinoceros exhibit the highest δ¹⁵N values, whereas reindeer exhibit the most positive δ¹³C values. In both cases, hyaenas present the highest δ¹⁵N values measured on faunal samples which is expected for carnivores, and Neanderthals present δ¹⁵N values about 2 ‰ higher than those of hyaenas (Fig. 3). The similarity of the patterns of isotopic variations in both cases strongly suggest that Neanderthals from Saint-Césaire and Spy had a similar trophic ecology. However, both sites are several hundreds of kilometres away, and Spy Neanderthal is several thousand years older than the Chatelperronian individual from Saint-Césaire.

A final comparison can be made between the Saint-Césaire Neanderthal and the Neanderthal from layer 4A in Scladina Cave (Scladina-4A). In this case, not only are the sites well separated spatially, but also chronologically and ecological-ly, since the Belgian specimen is about 65,000 years older than the French one and the environment during OIS 5c was more temperate and forested than during OIS 3 (Cordy and Bastin, 1992). These ecological differences are clearly reflected in the δ¹³C values of herbivorous mammals. The samples from Scladina-4A have more negative δ¹³C values than those from western France, which is likely due to a more pronounced canopy effect under a forested vegetation than would be under steppe vegetation (Bocherens et al., 1999). Hence, the Neanderthal from Scladina-4A relied on prey from an open environment rather than on prey from the forest. Despite dramatic environmental differen-ces, the isotopic signature of the Neanderthal from Scladina-4A is very similar to that of the other Neanderthals including the specimen from Saint- Césaire, implying a stable trophic position through time and ecological variations in Western Europe.

The comparison of the isotopic signatures of these Western European Neanderthals points to a trophic position as top predator in an open environment, with little variation through time and space. It is worth noting that in each case, the δ¹⁵N values of Neanderthals appear more positive than those of animal predators such as hyaena or wolf. A detailed quantitative study of
<figure><img src="image_88.png" /><figcaption>Fig. 3. δ¹³C and δ¹⁵N values of Neanderthal collagen from southwestern France (Saint-Césaire, Camiac, La Berbie), Les Pradelles, Wallonia (Spy and Scladina – 1A), and Scladina – 4A, compared to those of their respective contemporary mammal fauna (average ± standard-deviation for different herbivorous and carnivorous species).</figcaption></figure>

the situation in Western France, using isotopic data from Saint-Césaire, La Berbie and Camiac provides insights into such an intriguing pattern.

*Comparison of Saint-Césaire Neanderthal with contemporaneous hyaenas using a multi-source mixing model*

The results of the multi-source mixing model are presented on Fig. 4. The range of extreme percent represents the entire range of mathematically feasible solutions, whereas the range of mean percent does give some idea of how the central tendency of the distribution of feasible results shifts between Neanderthal and hyaena. The following discussion is based on the range of mean percent since, although extreme percents are mathematically feasible, they would require fractionation

values to be at the extreme opposites of their possible end values for each predator. Thus the results presented below are indicative, but not definitive.

The obtained patterns are similar for both predators – Neanderthal and hyaena – for Bovinae, large deer and horse, but they are different for reindeer, woolly rhinoceros and woolly mammoth (Fig. 4). Both predators exhibit a similar pattern of consumption of Bovinae and large deer, with a range of most likely contribution from approximately 0 to 25 %. For both predators, horse exhibits a large interval of possible contribution, ranging from a few percent to approximately 70 to 85 %, for Neanderthal and hyaena, respectively. The patterns are very different for reindeer, with a contribution to dietary input lower than approximately 5 % for Neanderthal whereas hyaena could have consumed from approximately 5 % to 40 % reindeer. Woolly
<figure><img src="image_89.png" /><figcaption>Fig. 4. Summary of the mean and extreme possible ranges of prey percentage for Neanderthal and Hyaena in southwestern France around 36,000 years BP. The solid black bar takes the means from the distributions of solutions from IsoSource for the 4 or 5 key data points in Fig. 2 (corners of rectangles and intersections of rectangles with the mixing polygon) and shows the range of those means.</figcaption></figure>

rhinoceros and woolly mammoth are apparently
preyed upon much more by Neanderthals than by
hyaenas. Woolly rhinoceros contribution range
from a few to as much as approximately 60 % in
the diet of Neanderthals, whereas it composes less
than approximately 20 % of the diet of hyaenas.
The contrast is even greater in the case of woolly
mammoth since its contribution ranges from
approximately 15 to 70 % in Neanderthal’s diet,
whereas it is not higher than approximately 15 %
in hyaena’s diet. In conclusion, the isotopic differ-
ences between Neanderthals and hyaenas reflect
essentially much lower amounts of reindeer and
much higher amounts of woolly rhinoceros and
woolly mammoth in the dietary input of Nean-
derthal than in that of hyaena, with relatively
similar contributions of bovinae, large deer and
horse for both predators.

The dietary analysis presented above does not
take plant food into account. Collagen isotopic
signatures reflect principally the isotopic signa-
tures of the protein fraction of the diet (e.g.
Ambrose and Norr, 1993; Tieszen and Fagre,
1993). Since it is much poorer in proteins than
meat, plant food proteins would be almost in-
visible in a mixed diet containing few plants and
a lot of meat, as it was probably the case of peri-
arctic dwellers like Neanderthals during OIS 3.
Moreover, plant foods are ¹⁵N-depleted relatively

to herbivore meat. Therefore, the inclusion of
plant food in significant proportion in Neander-
thal diet would require the inclusion of even more
¹⁵N-enriched food resource to account for the
observed collagen isotopic values, meaning that
the contribution of mammoth and rhinoceros
meat would need to be even higher than calculated
by the mixing model. Neglecting plant food in this
discussion does not change the general picture of
dietary resources consumed by Neanderthal.

A comparison with the conclusions based on
the zooarchaeological remains from layer Ejop sup
in Saint-Césaire yields a similar picture. The
identified remains of large mammals from this
layer are diverse. Considering the MNI (Minimal
Number of Individuals), Bovinae (Aurochs and
Bison) are dominant relative to reindeer and horse
(Patou-Mathis, 1993). When the meat weight is
estimated for each species, Bovinae remain the
most important potential meat providers, but
mammoth and woolly rhinoceros yielded more
meat than cervids (Patou-Mathis, 1993).

Finally, the Neanderthals that led to the
deposit of layer Ejop sup in Saint-Césaire con-
sumed more meat from large herbivore species
than meat from middle size and small species. This
contrasts with hyaena, which consumed relatively
more middle size and small herbivore species than
large ones.
of the results obtained independently with both approaches indicates that a similar subsistence strategy is reflected at different demographic, spatial, and chronological scales. Such consistency reinforces the contribution of large-sized herbivore species in Chatelperronian Neanderthals' diet.

The comparison of prey selection by animal predators such as hyaenas and by Neanderthals does not support scavenging as the principal subsistence pattern of these humans. This result is in keeping with recent reviews of archaeological evidence that indicate an active hunting for Neanderthals as opposed to scavenging (e.g. Marean and Kim, 1998). Isotopic data reinforce the picture of western European Neanderthals as accomplished hunters, capable of taking down prey species as large as woolly mammoth on a regular basis. Such a hunting strategy involving effective large-mammal exploitation qualifies for modern behaviour and should be taken into account in the current debate about the origin of modern human behaviour (e.g. Henshilwood and Marean, 2003).

## Acknowledgements

We are very thankful to Jean-Luc Guadelli and Stéphane Madelaine for providing samples from Camiac and La Berbie, respectively. Jean-Jacques Cleyet-Merle, head curator of the Musée National de Préhistoire (Les Eyzies de Tayac, France), and François Lévêque (Direction Régional de l'Archéologie, Poitiers, France) have been kind enough to authorise sampling on fossil skeletal material. Keith Hobson and Dan Mazerolle (Canadian Wildlife Service, Saskatoon) are warmly thanked for insightful discussions. We are also thankful to Don Phillips for his thorough comments about the use of the multi-source mixing model The CNRS programme PEH (Paléoenvironnement, Evolution des Hominidés) provided financial support for some travelling and analytical costs. Naturalia et Biologia has partly founded the Ph.D thesis of Dorothée Drucker, in the course of which this study has been performed.

## Appendix. List of collagen chemical compositions and carbon and nitrogen isotopic signatures for skeletal material from Saint-Césaire, La Berbie and Camiac.

<table><thead><tr><th>Analysis number</th><th>Species</th><th>Sample</th><th>Site</th><th>Yield (mg g<sup>-1</sup>)</th><th>%C<sub>coll</sub></th><th>%N<sub>coll</sub></th><th>C/N<sub>coll</sub></th><th>δ<sup>13</sup>C</th><th>δ<sup>15</sup>N</th></tr></thead><tbody><tr><td>RPB8000</td><td>Bovine (Bos or Bison)</td><td>metapodium</td><td>Saint-Césaire</td><td>3.0</td><td>30.1</td><td>11.3</td><td>3.1</td><td>-20.9</td><td>6.1</td></tr><tr><td>LBR100</td><td>Bison (Bison priscus)</td><td>metatarsus</td><td>La Berbie</td><td>31.9</td><td>39.5</td><td>14.5</td><td>3.2</td><td>-20.3</td><td>6.1</td></tr><tr><td>LBR200</td><td>Bison (Bison priscus)</td><td>metatarsus</td><td>La Berbie</td><td>14.8</td><td>37.8</td><td>14.0</td><td>3.2</td><td>-20.1</td><td>5.6</td></tr><tr><td>LBR300</td><td>Bison (Bison priscus)</td><td>metatarsus</td><td>La Berbie</td><td>27.6</td><td>35.3</td><td>13.1</td><td>3.1</td><td>-20.8</td><td>4.7</td></tr><tr><td>LBR400</td><td>Bison (Bison priscus)</td><td>metatarsus</td><td>La Berbie</td><td>12.5</td><td>36.9</td><td>13.7</td><td>3.1</td><td>-20.3</td><td>5.6</td></tr><tr><td>LBR500</td><td>Bison (Bison priscus)</td><td>metatarsus</td><td>La Berbie</td><td>26.1</td><td>40.3</td><td>14.8</td><td>3.2</td><td>-20.2</td><td>6.1</td></tr><tr><td>LBR600</td><td>Bison (Bison priscus)</td><td>metatarsus</td><td>La Berbie</td><td>18.2</td><td>29.0</td><td>11.1</td><td>3.0</td><td>-20.2</td><td>6.0</td></tr><tr><td>LBR700</td><td>Bison (Bison priscus)</td><td>metatarsus</td><td>La Berbie</td><td>41.0</td><td>38.4</td><td>14.3</td><td>3.1</td><td>-20.9</td><td>4.6</td></tr><tr><td>LBR3100</td><td>Bison (Bison priscus)</td><td>radius</td><td>La Berbie</td><td>36.3</td><td>39.0</td><td>14.9</td><td>3.0</td><td>-20.5</td><td>5.3</td></tr><tr><td>CAM100</td><td>Aurochs? (cf. Bos primigenius)</td><td>femur or tibia</td><td>Camiac</td><td>66.9</td><td>40.8</td><td>14.7</td><td>3.2</td><td>-20.0</td><td>6.1</td></tr><tr><td>CAM200</td><td>Aurochs? (cf. Bos primigenius)</td><td>femur or tibia</td><td>Camiac</td><td>58.7</td><td>42.0</td><td>15.1</td><td>3.2</td><td>-20.7</td><td>6.2</td></tr><tr><td>CAM300</td><td>Aurochs? (cf. Bos primigenius)</td><td>femur or tibia</td><td>Camiac</td><td>52.3</td><td>41.6</td><td>14.9</td><td>3.3</td><td>-20.4</td><td>7.4</td></tr><tr><td>CAM400</td><td>Aurochs? (cf. Bos primigenius)</td><td>tibia</td><td>Camiac</td><td>23.4</td><td>39.5</td><td>14.3</td><td>3.2</td><td>-20.1</td><td>8.6</td></tr><tr><td>CAM500</td><td>Aurochs? (cf. Bos primigenius)</td><td>tibia</td><td>Camiac</td><td>14.4</td><td>39.1</td><td>14.0</td><td>3.3</td><td>-20.0</td><td>6.0</td></tr><tr><td>CAM600</td><td>Aurochs? (cf. Bos primigenius)</td><td>femur or tibia</td><td>Camiac</td><td>35.9</td><td>41.0</td><td>14.7</td><td>3.3</td><td>-20.4</td><td>5.3</td></tr><tr><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td>av.</td><td>-20.4</td><td>6.0</td></tr><tr><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td>s.d.</td><td>0.3</td><td>1.0</td></tr><tr><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td>n</td><td>15</td><td>15</td></tr><tr><td>RPB650</td><td>Giant deer (Megaloceros giganteus)</td><td>maxillary bone</td><td>Saint-Césaire</td><td>28.9</td><td>34.1</td><td>12.7</td><td>3.1</td><td>-20.5</td><td>5.0</td></tr><tr><td>CAM900</td><td>Giant deer (Megaloceros giganteus)</td><td>femur</td><td>Camiac</td><td>10.0</td><td>33.2</td><td>11.9</td><td>3.2</td><td>-20.0</td><td>6.4</td></tr><tr><td>CAM1000</td><td>Giant deer (Megaloceros giganteus)</td><td>long bone</td><td>Camiac</td><td>36.8</td><td>26.7</td><td>9.6</td><td>3.3</td><td>-20.1</td><td>7.7</td></tr></tbody></table>

(continued on next page)
Appendix (continued)

<table><thead><tr><th>Analysis number</th><th>Species</th><th>Sample</th><th>Site</th><th>Yield (mg g<sup>-1</sup>)</th><th>%C<sub>coll</sub></th><th>%N<sub>coll</sub></th><th>C/N<sub>coll</sub></th><th>δ<sup>13</sup>C</th><th>δ<sup>15</sup>N</th></tr></thead><tbody><tr><td>CAM1100</td><td>Red deer (Cervus elaphus)</td><td>long bone</td><td>Camiac</td><td>24.4</td><td>37.5</td><td>13.4</td><td>3.3</td><td>-20.0</td><td>6.9</td></tr><tr><td>CAM1200</td><td>Red deer (Cervus elaphus)</td><td>long bone</td><td>Camiac</td><td>8.7</td><td>35.4</td><td>13.4</td><td>3.1<br>av.<br>s.d.<br>n</td><td>-20.6<br>-20.2<br>0.3<br>5</td><td>6.2<br>6.4<br>0.9<br>5</td></tr><tr><td>RPB1200</td><td>Reindeer (Rangifer tarandus)</td><td>jawbone</td><td>Saint-Césaire</td><td>9.5</td><td>23.6</td><td>8.6</td><td>3.2</td><td>-18.8</td><td>4.9</td></tr><tr><td>RPB7200</td><td>Reindeer (Rangifer tarandus)</td><td>metapodium</td><td>Saint-Césaire</td><td>31.7</td><td>41.5</td><td>15.0</td><td>3.2</td><td>-18.3</td><td>7.3</td></tr><tr><td>LBR900</td><td>Reindeer (Rangifer tarandus)</td><td>humerus</td><td>La Berbie</td><td>18.7</td><td>40.4</td><td>14.7</td><td>3.2</td><td>-19.2</td><td>3.7</td></tr><tr><td>LBR1000</td><td>Reindeer (Rangifer tarandus)</td><td>humerus</td><td>La Berbie</td><td>20.1</td><td>38.3</td><td>14.0</td><td>3.2</td><td>-19.4</td><td>3.9</td></tr><tr><td>LBR1100</td><td>Reindeer (Rangifer tarandus)</td><td>jawbone</td><td>La Berbie</td><td>14.6</td><td>39.9</td><td>14.4</td><td>3.2</td><td>-19.1</td><td>7.6</td></tr><tr><td>LBR3400</td><td>Reindeer (Rangifer tarandus)</td><td>femur</td><td>La Berbie</td><td>16.8</td><td>39.4</td><td>13.9</td><td>3.3<br>av.<br>s.d.<br>n</td><td>-19.4<br>-19.0<br>0.4<br>6</td><td>5.8<br>5.5<br>1.5<br>6</td></tr><tr><td>RPB2300</td><td>Horse (Equus caballus)</td><td>upper 2nd premolar</td><td>Saint-Césaire</td><td>14.1</td><td>17.7</td><td>6.5</td><td>3.1</td><td>-20.5</td><td>7.3</td></tr><tr><td>RPB2400</td><td>Horse (Equus caballus)</td><td>lower tooth</td><td>Saint-Césaire</td><td>6.2</td><td>34.7</td><td>13.1</td><td>3.1</td><td>-21.7</td><td>5.7</td></tr><tr><td>LBR1600</td><td>Horse (Equus caballus)</td><td>metacarpus 2 or 4</td><td>La Berbie</td><td>22.4</td><td>40.3</td><td>14.7</td><td>3.2</td><td>-20.4</td><td>7.5</td></tr><tr><td>LBR1700</td><td>Horse (Equus caballus)</td><td>femur</td><td>La Berbie</td><td>36.5</td><td>41.9</td><td>15.3</td><td>3.2</td><td>-20.9</td><td>3.5</td></tr><tr><td>CAM1600</td><td>Horse (Equus caballus)</td><td>tibia</td><td>Camiac</td><td>56.1</td><td>41.9</td><td>14.7</td><td>3.2</td><td>-20.7</td><td>5.2</td></tr><tr><td>CAM1700</td><td>Horse (Equus caballus)</td><td>tibia</td><td>Camiac</td><td>34.6</td><td>40.6</td><td>14.6</td><td>3.2<br>av.<br>s.d.<br>n</td><td>-20.5<br>0.4<br>6</td><td>5.2<br>5.7<br>1.4<br>6</td></tr><tr><td>RPB1400</td><td>Rhinoceros (Coelodonta antiquitatis)</td><td>lower tooth</td><td>Saint-Césaire</td><td>7.8</td><td>31.4</td><td>12.6</td><td>2.9</td><td>-20.9</td><td>5.4</td></tr><tr><td>LBR2200</td><td>Rhinoceros (Coelodonta antiquitatis)</td><td>skull</td><td>La Berbie</td><td>27.6</td><td>39.1</td><td>14.4</td><td>3.2</td><td>-19.6</td><td>7.2</td></tr><tr><td>CAM1300</td><td>Rhinoceros (Coelodonta antiquitatis)</td><td>humerus</td><td>Camiac</td><td>22.6</td><td>38.1</td><td>13.7</td><td>3.2</td><td>-19.9</td><td>8.4</td></tr><tr><td>CAM1400</td><td>Rhinoceros (Coelodonta antiquitatis)</td><td>humerus</td><td>Camiac</td><td>27.0</td><td>39.3</td><td>14.3</td><td>3.2</td><td>-20.1</td><td>7.1</td></tr><tr><td>CAM1500</td><td>Rhinoceros (Coelodonta antiquitatis)</td><td>humerus</td><td>Camiac</td><td>78.1</td><td>40.3</td><td>14.7</td><td>3.2<br>av.<br>s.d.<br>n</td><td>-20.6<br>-20.2<br>0.5<br>5</td><td>7.1<br>7.0<br>1.0<br>5</td></tr><tr><td>LBR1900</td><td>Mammoth (Mammuthus primigenius)</td><td>femur</td><td>La Berbie</td><td>11.9</td><td>37.2</td><td>13.4</td><td>3.2</td><td>-21.8</td><td>8.5</td></tr><tr><td>CAM700</td><td>Mammoth (Mammuthus primigenius)</td><td>femur</td><td>Camiac</td><td>57.2</td><td>39.1</td><td>14.1</td><td>3.2</td><td>-21.2</td><td>8.7</td></tr><tr><td>CAM800</td><td>Mammoth (Mammuthus primigenius)</td><td>femur</td><td>Camiac</td><td>32.3</td><td>40.6</td><td>14.7</td><td>3.2<br>av.<br>s.d.<br>n</td><td>-21.8<br>-21.6<br>0.3<br>3</td><td>7.7<br>8.3<br>0.4<br>3</td></tr><tr><td>LBR1300</td><td>Hyaena (Crocuta crocuta)</td><td>humerus</td><td>La Berbie</td><td>41.0</td><td>40.6</td><td>14.8</td><td>3.2</td><td>-19.9</td><td>9.5</td></tr><tr><td>LBR1400</td><td>Hyaena (Crocuta crocuta)</td><td>humerus</td><td>La Berbie</td><td>26.4</td><td>39.9</td><td>14.7</td><td>3.2</td><td>-19.1</td><td>8.5</td></tr><tr><td>LBR3300</td><td>Hyaena (Crocuta crocuta)</td><td>neck vertebra</td><td>La Berbie</td><td>14.7</td><td>34.5</td><td>12.0</td><td>3.3</td><td>-19.1</td><td>9.8</td></tr><tr><td>CAM1800</td><td>Hyaena (Crocuta crocuta)</td><td>long bone</td><td>Camiac</td><td>72.3</td><td>41.6</td><td>14.9</td><td>3.3</td><td>-19.3</td><td>9.8</td></tr><tr><td>CAM1900</td><td>Hyaena (Crocuta crocuta)</td><td>long bone</td><td>Camiac</td><td>53.1</td><td>40.6</td><td>14.6</td><td>3.2<br>av.<br>s.d.<br>n</td><td>-19.0<br>-19.3<br>0.3<br>5</td><td>9.1<br>9.3<br>0.5<br>5</td></tr><tr><td>RPB7000</td><td>Neandertal (Homo neandertalensis)</td><td>fibula</td><td>Saint-Césaire</td><td>26.0</td><td>40.8</td><td>14.2</td><td>3.3</td><td>-19.8</td><td>11.4</td></tr></tbody></table>
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Tieszen, L.L., Fagre, T., 1993. Experimental evidence for the relationship of the carbon isotope ratios of whole diet and dietary protein to those of bone collagen and carbonate. In: Lambert, J., Grupe, G. (Eds.), Prehistoric Human Bone, Archaeology at the Molecular Level. Springer, Berlin, pp. 121–155.

Vandermeersch, B., 1976. Les Néandertaliens en Charente. In: La préhistoire française, Civilisations Paléolithiques et Mésolithiques. CNRS T. II, Paris. 584–587.
**Final Program**

Paleoanthropology Society

April 5, 6, 2005

Milwaukee Wisconsin

**Poster Session**

**Tuesday, April 5**
4:30 - 6:30

**Coate, J., D. Neuweger and D. Curnoe**

Taxonomic integrity above the species level in primate evolution and systematics

**Cross, A. and A. Nelson**

A test of models used for estimating total body surface area in hominins

**Delezene, L. and C. Schrein**

Is *Ouranopithecus macedoniensis* a hominin? A critical assessment of evidence from canine and P3 morphology

**Doyle, S.**

Postcranial robusticity in a carnivore: implications for understanding skeletal robusticity in hominins

**Hawks, J.**

Population size, disease and the evolution of aging

**Kilberger, M. and J. Enloe**

Maxillary dentition and hunting strategies: reindeer teeth from the Magdalenian

**Kovarovic, K., P. Andrews and L. Aiello**

A palaeoenvironmental reconstruction of Laetoli, Tanzania using a bovid community ecomorphological survey

**McCall, G.**

Multivariate approaches to the lithics of Klasies River Mouth

**Pilbrow, V. and S. Bailey**

Patterns of intraspecific variation in dental metrics in *Gorilla gorilla*, *Pan troglodytes* and *Homo sapiens*: a comparison using Wright's (1969) *Fst* statistic

**Prat, S. and the West Turkana Archaeological Project**

The Lokalalei site complex (2.3-2.4 Myr): the oldest evidence of the genus *Homo* in the Nachukui Formation, Kenya

**Sisk, M. and J. Shea**

Experimental use of Levallois points as projectile points
Smith, H.

Patterns of fracture among anatomically modern humans: implications for hunting techniques

Sylvester, A.

The decoupling hypothesis: a new idea for the origin of bipedalism

Teague, R.

Comparison of the faunal contexts of *Homo erectus* in East Africa and East Asia during the early Pleistocene

Terhune, C. and A. Deane

Shape of the temporal squama in fossil hominins: relationships to cranial shape and a
determination of character polarity

Oral Presentations

Tuesday, April 5: 9:00 – 12:30

9:00 – 9:15 Introduction

9:15 Reed, D.

Calibrating paleoenvironmental analysis using modern micromammal assemblages from Tanzania

9:30 Su, D. and T. Harrison

Large mammal differences between fossil and modern communities: implications for the reconstruction of hominin paleoenvironments

9:45 Zihlman, A.

Walking, running, climbing and jumping to conclusions: interpreting locomotion in early humans

10:00 Marchal, F., G. Berillon and J. Millet

Integrative biomechanical geometric morphometrics. A new approach to study the evolution of the hominid locomotor skeleton

(10:15 – 10:30: Coffee Break)

10:30 Turner, L., Z. Throckmorton and J. Langdon

Since when are humans different?

10:45 Sherwood, R., D. Duren, T. Dyer, S. Cole, R. Siervogel, J. Blangero and B. Towne

The search for the real missing link: finding the genes that influence craniofacial
morphology
11:00 Carlson, B.
Did/does docosahexaenoic acid constrain encephalization?

11:15 Tostevin, G.
The use and abuse of the *chaine operatoire* vs. reduction sequence debate in Paleolithic archaeology

11:30 Carlson, K., M. Whalen and F. Grine.
Robusticity and sexual dimorphism in the postcranium of modern hunter-gatherers from Australia

11:45 Walker, A., H. Dunsworth, S. Kozakowski, G. Krovitz, J. Orkin and K. Welker
Thinking outside the book: escaping traditional representations of hominin evolution through computer anamation

12:00 Shackelford, L.
Regional variation in Late Pleistocene human postcranial robusticity

12:15 Henry-Gambier, D., J. Bruzek, A. Schmitt, F. Houet and P. Murail
Probabilistic approach to age and sex assessment in human fossils – application to burials from the Upper Paleolithic in Italy and France

Tuesday, April 5: 2:00 – 4:00

2:00 Trinkaus, E., J. Zilhao, H. Rougier, J. Quiles, S. Milota, M. Gherase, R. Rodrigo, L. Sarcina, V. Codrea, I. Baltean and O. Moldovan
The Pestera cu Oase: cave bear, canids, caprids, cervids and a cranium

2:15 McCarthy, R. and D. Strait
Morphological constraints on hominin speech production

2:30 Yokley, T. and R. Franciscus
Variation in nasal passage surface-area-to-volume ratios of recent and fossil humans

2:45 Meyer, M., M. Chang and J. Lewis
Functional morphology of the Neandertal nose

3:00 Bouchneb, L., M. Bruno, P. Semal, S. Louryan and J-J Hublin
The bony labyrinth of Neandertals: additional results

3:15 Mann, A., C. Beauval, S. Costamagno, J.-G. Bordes, L. Bourguignon, I. Couchoud, Fr. Lacrampe, V. Larouladnie, J.-Cl. Marquet, L. Meignen, J.-P Texier, B. Vandermeersch and B. Maureille
Carnivore and Neandertal interactions at the Les Pradelles site (Charante, Southwest France)

3:30 Cook, J. D. Frayer and J. Radovcic
New evidence for symbolic behavior by the Krapina Neandertals

3:45 Skinner, A. H. Dibble, S. McPherron, A. Turq, D. Sandgathe, B. Blackwell and S. Martin
Electron spin resonance dating of the Neandertal and Mousterian site, Roc de Marsal, Dordogne, France

4:00 Monge, J., A.-M. Tillier and A. Mann
Perikymata number and spacing on early modern human teeth: evidence from Qafzeh Cave
(Northern Israel)

## Poster Session

4:15 – 6:30 Location to be announced

### Wednesday, April 6: 8:30 – 12:30

8:30 Shea, J.
The origins and antiquity of stone projectile points

8:45 Villa, P., S. Soriano, S. Wurz, A. Delanges, L. Wadley
Development, decline and disappearance of the Howiesons poort industry

9:00 Ogola, C., T. Pickering, K. Kuman and R. Clarke
Modified bone from Sterkfontein Cave: implications for the technological behavior of
Middle Stone Age hominids in South Africa

9:15 Tryon, C., S. McBrearty and P.-J. Texier
Levallois lithic technology from the Kapthurin Formation, Kenya: Acheulean origin and
Middle Stone Age diversity

9:30 Haradon, C.
Middle Pleistocene faunal change at the Acheulean to Middle Stone Age transition

9:45 Berger, G. A. Perez-Gonzalez, E. Carbonell, J. Arsuaga, J.-M Bermudez de Castro
Luminescence chronology for cave sediments at the Atapuerca paleoanthropological
Site, Spain

10:00 Stringer, C.
Major gaps in the early human occupation of Britain

(10:15 – 10:30 Coffee Break)

10:30 Steele, T., G. Avery, R. Klein, T. Volman
EFTN HD2: a Mid-Pleistocene artifact-and-mammal-bone occurrence at Elandsfontein,
Western Cape Province, South Africa

10:45 Gavronski, E., E. Bowers and R. Quinlan
Early *Homo erectus/ergaster*: one or more species?
11:00 Grimaud-Herve, D., H. Widianto, F. Detroit and T. Jacob
The new skull of Bukuran from Sangiran Dome (Java, Indonesia)

11:15 Arif, J., H. Baba. And Y. Kaifu
The Skull IX (Tjg-1993.05) as one of the variant examples of Asian *Homo erectus* from
Indonesia

11:30 Ambrose, S., S. Katoh, Y. Bayene, and B. Asfaw
Isotopic paleoecology of Plio-Pleistocene mammals and paleosols, Konso Formation,
Southern Ethiopia

11:45 Lepre, C. and R. Quinn
Precessional forcing of Plio-Pleistocene lake levels from Koobi Fora, Kenya

12:00 DeGusta, D.
How primitive was the postcranial skeleton of *Australopithecus africanus*?

12:15 Rak, Y.
On derived characters that *Australopithecus afarensis* shares with the robust clade

Wednesday, April 6: 2:00 – 6:15

2:00 Harlacker, L.
The biomechanics of stone toolmaking: kinematics and kinetics of the arm during
Oldowan/Mode I knapping

2:15 Braun, D., T. Plummer, J. Ferraro, L. Bishop, P. Ditchfield, R. Potts and J. Harris
Oldowan technology at Kanjera South, Kenya: the context of technological diversity

2:30 Copes, L. and R. Potts
Are hominin fossils and paleoenvironmental data precisely associated in the stratigraphic
records of Turkana and Olduvai?

2:45 Kingston, J. and T. Harrison
Ostrich eggshells as paleoenvironmental indicators in the Pliocene Laetoli Succession,
N. Tanzania

3:00 Campisano, C. and C. Feibel
Tephrostratigraphy and regional correlations of the artifact bearing Kada Hadar Member,
Hadar Formation, Ethiopia

(3:15 – 3:30 Coffee Break)

3:30 Adams, J.
Faunal and paleoecological analyses of the Gondolin GD2 Plio-Pleistocene fossil
assemblage, North West Province, South Africa
3:45 Heaton, J. and T. Pickering
Fossil collection biases at Sterkfontein cave and their impact on primate taxonomy

4:00 Kibii, J.
Comparative taxonomic, taphonomic and paleoenvironmental analysis of hominid infills, Member 4 and Jacovec Cavern, within Sterkfontein, South Africa

4:15 Pickering, T., M. Dominguez-Rodrigo, C. Egeland and C. Brain
The earliest evidence of hominid butchery in southern Africa: new zooarchaeological and taphonomic data on the use of large animal carcasses at Swartkrans Members 1 - 3

4:30 DeSilva, J., E. Shoreman, L. MacLatchy
A fossil hominoid proximal femur from Kikorongo crater, southwestern Uganda

4:45 Dunsworth, H.
Ontogeny and proportions of new *Proconsul heseloni* tarsals and metatarsals from the Kaswanga Primate Site, Rusinga Island, Kenya

5:00 Benefit, B., E. Lynch, M. McCrossin, S. Ghoshroy and S. Kersey
Preliminary analysis of dental microwear and diet in kenyapithecine hominoids from the Middle Miocene of Fort Ternan and Maboko Island, Kenya

5:15–6:15 Business Meeting
# Carnivore and Neandertal Interactions at the Les Pradelles Site, Charente, Southwest France

A. E. Mann<fnref n="1" />, C. Beauval<fnref n="2" />, S. Costamagno<fnref n="3" />, J.-G. Bordes<fnref n="2" />, L. Bourguignon<fnref n="4" />, I. Couchoud<fnref n="2" />, Fr. Lacrampe<fnref n="2" />, V. Laroulandie<fnref n="2" />, J.-Cl. Marquet<fnref n="5" />, L. Meignen<fnref n="6" />, J.-P. Texier<fnref n="2" />, B. Vandermeersch<fnref n="7" />, B. Maureille<fnref n="2" />

Excavations at the collapsed cave site of Les Pradelles, Charente, France, have uncovered a series of Late Pleistocene stratigraphic levels containing a wide variety of Würmian age fauna, including reindeer, horse, bison, marmots, wolves, cave lion and cave hyena. Also discovered in these deposits have been numerous Middle Paleolithic artifacts (Mousterian of La Quina techno-complex) and more than 38 fragmentary Neandertal fossils, including cranial, dental and postcranial elements. Analysis of the artifacts and especially the human and animal bones, have revealed that over the course of time, the site was intermittently occupied, sometimes by Neandertals and at other times by cave carnivores. The human occupations have been interpreted as most likely those of a hunting camp where animal carcasses, mainly reindeer but also horses and bovids, were brought for processing.

Examination of teeth that had been provisionally identified as deciduous bovid or cervid teeth have now been shown to be maxillary permanent incisors of Neandertals. These teeth have been markedly modified by the gastric secretions of a carnivore, probably a cave hyena, and were subsequently regurgitated. This process has given these teeth the superficial appearance of herbivore deciduous incisors.

As determined from the number of occipital fragments, the MNI of Neandertals at the site is five, with the bones of four of five of these individuals showing evidence of cut marks, and incisions, moreover, that mirror those found on the faunal bones.

This evidence suggests that Neandertals treated some of their colleagues in the same manner as hunted animals, processing the eatable portions and leaving other parts to be scavenged by carnivores. The abundance of animal remains makes it difficult to interpret this treatment of the human bones as evidence of dietarily necessary cannibalism; other possible explanations will be assessed.

<footnote n="1">¹ Dpt Anthropology, Princeton University, NJ, USA</footnote>

<footnote n="2">² UMR 5199 CNRS, PACEA, Université Bordeaux 1, Talence, France</footnote>

<footnote n="3">³ UMR 5608 CNRS, UTAH, Toulouse, France</footnote>

<footnote n="4">⁴ INRAP GGrand Sud-Ouest Pessac, France</footnote>

<footnote n="5">⁵ Musée de Préhistoire, Le Grand Pressigny, France</footnote>

<footnote n="6">⁶ UMR 6130 CNRS, Centre d'étude de la préhistoire, Valbonne, France</footnote>

<footnote n="7">⁷ C/O Garralda, Antropologia, Universidad Complutense, Madrid, Spain</footnote>
# LES PRADELLES
(Commune de Marillac-le-Franc, Charente)

<figure><img src="image_90.png" /><figcaption>Vue de la coupe stratigraphique le 20.08.2005, témoin EST
Photo : équipe de fouille des Pradelles</figcaption></figure>

La campagne de fouille s'est déroulée du 12 Juillet au 20 Août. Cela représente 33 jours sur le terrain. Les fouilleurs bénévoles étaient d'abord de jeunes américains de l'Université de Princeton (16 étudiants) qui valident ainsi une part de leur cursus universitaire. Puis, du 30 Juillet au 20 Août, un groupe d'une vingtaine de jeunes européens a remplacé le précédent. Cette année, divers aménagements ont été réalisés sur le site afin de faciliter le travail des fouilleurs (et de l'équipe scientifique) ainsi que la sécurité (cabane de chantier, abri sous roche). Mais, il devient extrêmement urgent de refaire le toit qui protège les deux tiers du gisement et de l'étendre vers l'Ouest. Les fouilles du témoins EST ont porté sur moins d'une dizaine de m2 et ont surtout concerné les faciès 2a, 2b, 3, 4a, 4b, 8. Le faciès 2a' a été également fouillé dans les travées K et L (du lion des cavernes a ainsi été mis au jour). Les résultats de ces travaux ne bouleversent pas nos connaissances sur la cette partie du gisement. De nombreux restes humains ont été mis au jour dans le carré D12. En ce qui concerne la partie supérieure de la stratigraphie, elle livre du matériel qui est plutôt la conséquence de l'activité de carnivores (loup, etc). Le spectre faunique reste le même. Le principal résultat nouveau des travaux de cette année porte sur la galerie située à l'angle S-O de la dépression que forme le gisement. Cette galerie karstique a reçu les visites de blaireaux qui ont remanié du matériel archéologique. Or, ce dernier met en évidence l'existence d'une tanière de carnivore. Nous avons donc décidé de faire un sondage pour avoir une idée de la stratigraphie encore en place. Le niveau correspondant au repère d'hyène est remanié mais il est intéressant. Il pourrait correspondre à celui remarqué par P. David en 1933. Une argile sableuse jaunâtre a livré un maxillaire d'étalon et une patte en connexion d'un boviné. Les fouilles futures dans cette cavité permettront sûrement d'aller plus loin dans nos interprétations. Le site des Pradelles reste le gisement majeur de Poitou-Charentes quant à nos connaissances sur les Hommes de Néandertal et certaines de leurs activités comme le cannibalisme, la gestion des matières premières en fonction d'activités programmées d'acquisition de ressources alimentaires carnées.

Mots-clefs : Paléolithique moyen, Moustérien charentien de type Quina, Néandertalien, *hunting camp*, aven.

Programme 03 : Les peuplements néandertaliens *lato sensu* (stades isotopiques 8 à 4 : 300.000 à 40.000 ans) ; Paléolithique moyen *lato sensu*.