Tracing the footprints of the Atlas Brown Bear: a metric analysis of Holocene and Late Pleistocene remains from Northern Morocco
Author
Iken, Shaymae
Instituto Universitario de Xeoloxía (IUX), Universidade da Coruña, Edificio de Servicios Centrales de Investigación (ESCI), Campus de Elviña s / n, 15071 A Coruña (Spain) & National Institute of Archaeological Sciences and Heritage (INSAP), Madinat Al Irfane, Hay Riad, BP 6828 Rabat Institutes (Morocco)
Author
García-Vázquez, Ana
Instituto Universitario de Xeoloxía (IUX), Universidade da Coruña, Edificio de Servicios Centrales de Investigación (ESCI), Campus de Elviña s / n, 15071 A Coruña (Spain) & Archaeoscience Platform (AsP), Research Institute of the University of Bucharest (ICUB), University of Bucharest, 90 - 92 Sos. Panduri, 5 th District, 50663 Bucharest (Romania)
Author
González-Fortes, Gloria
Instituto Universitario de Xeoloxía (IUX), Universidade da Coruña, Edificio de Servicios Centrales de Investigación (ESCI), Campus de Elviña s / n, 15071 A Coruña (Spain)
Author
Bouzouggar, Abdeljalil
National Institute of Archaeological Sciences and Heritage (INSAP), Madinat Al Irfane, Hay Riad, BP 6828 Rabat Institutes (Morocco) & Max Planck Institute for Evolutionary Anthropology, Deutscher Platz 6, 04103 Leipzig (Germany)
Author
Grandal-D’Anglade, Aurora
Instituto Universitario de Xeoloxía (IUX), Universidade da Coruña, Edificio de Servicios Centrales de Investigación (ESCI), Campus de Elviña s / n, 15071 A Coruña (Spain)
aurora.grandal@udc.es
text
Comptes Rendus Palevol
2026
2026-03-11
25
4
57
76
https://doi.org/10.5852/cr-palevol2026v25a4
journal article
314543
10.5852/cr-palevol2026v25a4
0efbbe6b-2c87-4b27-9335-6ca8c35b0b01
1777-571X
19005282
urn:lsid:zoobank.org:pub:59E82E04-C7EB-4586-BF61-D9646DE136C6
APPENDIX 4. — Main measurements (in mm) of brown bear phalanges from the sites of Keft-el-Hammar (KEH) and Hattab II (HAT II). Abbreviations: APW prox, proximal anteroposterior width; H prox, proximal height; n/n, no number; TL, total length; TW dia, transversal width of diaphysis; TW dist, distal transversal width; TW prox, proximal transversal width. The asterisk (*) indicates elements that were published previously (Ouchaou 2000).| First phalanx |
|---|
| Site | Level | Number | TL | TW prox | APW prox | TW dia | TW dist | Source |
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| HAT II | 7 | 558 | 30.7 | 16.3 | 15.6 | 11.4 | 12.2 | 1 |
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| HAT II | 3 | 115 | 33.8 | 17.8 | 14.8 | 11.1 | 13.0 | 1 |
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| HAT II | 3 | 245 | 34.1 | 18.3 | 13.8 | 10.6 | 13.3 | 1 |
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| HAT II | 6 | 557 | 34.7 | 18.3 | 13.3 | 10.8 | 11.8 | 1 |
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| HAT II | 6 | 360 | 37.2 | 17.5 | 14.8 | 11.8 | 13.8 | 1 |
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| HAT II | 6 | 372 | 38.7 | 18.3 | 14.5 | 12.0 | 13.7 | 1 |
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| HAT II | 3 | 259 | 38.8 | 17.6 | 15.4 | 11.7 | 13.8 | 1 |
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| KEH | 1 | 9 | 37.3 | 16.4 | 15.0 | 9.5 | 14.1 | 2 |
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| KEH | 1 | F7/27 | 37.6 | 21.5 | 12.3 | 10.5 | 12.3 | 2 |
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| KEH | 1 | HST8 | 37.8 | 21.0 | 16.3 | 13.2 | 13.4 | 2 |
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| KEH | 1 | I9/4 | 38.0 | 17.0 | 16.4 | 17.3 | 14.0 | 2 |
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| KEH | 1 | J29 | 46.0 | 18.5 | 15.0 | 11.5 | 14.0 | 2 |
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| KEH | 2 | 27/A | 32.0 | 18.0 | 12.0 | 11.0 | 13.0 | 2 |
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| KEH | 2 | 89 | 32.0 | 17.0 | 12.0 | 11.0 | 14.0 | 2 |
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| KEH | 2 | 58 | 34.0 | 17.0 | 14.0 | 11.0 | 13.0 | 2 |
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| KEH | 2 | n/n | 36.0 | 17.0 | 16.0 | 12.0 | 14.0 | 2 |
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| KEH | 2 | n/n | 36.0 | 18.0 | 15.0 | 12.0 | 14.0 | 2 |
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| KEH | 2 | n/n | 37.2 | 17. 8 | – | 12.3 | 13.4 | 2 |
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| KEH | 2 | n/n | 38.0 | 17.0 | 15.0 | 12.0 | 14.0 | 2 |
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| KEH | 2 | n/n | 41.4 | 17.2 | 13.0 | 12.0 | 12.3 | 2 |
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| KEH | 4 | – | 36.9 | 17.5 | 14.8 | 11.4 | 13.8 | 3 |
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| KTG | C | – | 35.0 | 17.0 | 12.5 | 9.8 | 12.8 | 3 |
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| KTG | D | – | 34.0 | 17.0 | 13.4 | 10.3 | 12.4 | 3 |
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| KTG | F | – | 35.0 | 17.0 | 13.0 | 9.5 | 12.5 | 3 |
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| KTG | F | – | 37.0 | 18.3 | 14.9 | 11.2 | 13.3 | 3 |
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| KTG | F | – | 40.0 | 20.0 | 15.8 | 13.0 | 14.9 | 3 |
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| KTG | G | – | 39.0 | 20.0 | 14.0 | 12.0 | – | 3 |
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| KBS | 3c | – | 37.8 | 20.5 | 15.5 | 12.0 | 13.5 | 3 |
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| GKH | IIIb | – | 40.0 | 20.0 | 16.0 | 13.5 | 13.9 | 3 |
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| KEB | salle | – | 43.0 | 21.0 | 17.4 | 14.6 | 15.7 | 3 |
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| IOS | – | 05/21 | 51.0 | 23.0 | – | 13.2 | – | 4 |
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| IOS | – | 05/36 | 43.0 | 16.9 | 14.5 | 9.2 | – | 4 |
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| IOS | – | 05/37 | – | 19.4 | 14.5 | 11.3 | – | 4 |
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| IOS | – | 06/I6/21 | 43.0 | 23.9 | 18.3 | 14.9 | 16.4 | 4 |
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| IOS | – | 06/I8/2 | 38.0 | 17.5 | 14.4 | 11.4 | 13.9 | 4 |
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| IOS | – | 06/I10/8 | 46.0 | 17.6 | 15.0 | 9.6 | 14.0 | 4 |
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| IOS | – | 06/I10/19 | 36.3 | 17.2 | 15.0 | 11.5 | 13.8 | 4 |
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| IOS | – | 06/I10/20 | 37.5 | 19.1 | 15.0 | 11.9 | 14.0 | 4 |
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| IOS | – | 06/I10/28 | 31.0 | 16.0 | 13.0 | 9.3 | 12.1 | 4 |
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| IOS | – | 06/I10/29 | 33.0 | 16.0 | 13.0 | 9.3 | 12.1 | 4 |
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| IOS | – | 06/I11/17 | 31.0 | 15.9 | 13.3 | 9.4 | 12.2 | 4 |
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| IOS | – | 06/I11/18 | 31.5 | 15.8 | 13.5 | 9.9 | 12.1 | 4 |
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| IOS | – | 06/I11/21 | 33.5 | 18.2 | 14.4 | 10.4 | 12.0 | 4 |
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| IOS | – | 06/I11/24 | 31.0 | 16.0 | 13.3 | 10.1 | 12.2 | 4 |
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| IOS | – | 06/I11/25 | 33.4 | 17.3 | 14.1 | 10.4 | 12.8 | 4 |
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| IOS | – | 06/I11/27 | 33.3 | 18.2 | 14.3 | 10.5 | 12.2 | 4 |
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| IOS | – | 06/I11/29 | 34.0 | 17.3 | 14.4 | 10.9 | 12.9 | 4 |
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| IOS | – | 06/J4/1 | 39.2 | 24.0 | 17.7 | 13.4 | 15.2 | 4 |
|---|
| Second phalanx |
|---|
| Site | Level | Number | TL | TW prox | APW prox | TW dia | TW dist | Source |
|---|
| HAT II | 6 | 430 | 21.8 | 14.4 | 13.3 | 11.2 | 12.3 | 1 |
|---|
| KEH | 2 | n/n | 27.0 | 16.0 | 15.0 | 11.0 | 14.0 | 2 |
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| KEH | 2 | n/n | 27.0 | 16.0 | 13.0 | 11.0 | 14.0 | 2 |
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| KEH | 2 | n/n | 29.0 | 17.0 | 16.2 | 14.0 | 15.0 | 2 |
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| KEH | 2 | n/n | 30.0 | 19.5 | 15.3 | 13.0 | 14.0 | 2 |
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| KEH | 2 | n/n | 30.0 | 18.0 | 17.0 | 13.8 | 16.0 | 2 |
|---|
| KEH | 4 | 283 | 27.5 | 17.0 | 15.3 | 13.0 | 14.1 | 2 |
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| KTG | E | – | 28.8 | 15.9 | 13.4 | 11.1 | 14.0 | 3 |
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| KTG | F | – | 26.2 | – | – | 10.5 | 13.0 | 3 |
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| KTG | F | – | 27.5 | 15.7 | 13.1 | 10.2 | 14.1 | 3 |
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| KTG | F | – | 22.6 | 17.3 | 13.9 | 12.3 | 14.9 | 3 |
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| GKH | V | – | 19.9 | 16.0 | 12.9 | 11.1 | 13.0 | 3 |
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| IOS | – | 05/9 | 28.8 | 15.9 | 13.4 | 11.1 | 14.0 | 4 |
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| IOS | – | 05/38 | 26.2 | – | – | 10.5 | 13.0 | 4 |
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| IOS | – | 06/I10/6 | 27.5 | 15.7 | 13.1 | 10.2 | 14.1 | 4 |
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| IOS | – | 06/I5/2 | 22.6 | 17.3 | 13.9 | 12.3 | 14.9 | 4 |
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| IOS | – | 06/I6/17 | 19.9 | 16.0 | 12.9 | 11.1 | 13.0 | 4 |
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| IOS | – | 06/I10/21 | 20.0 | 14.9 | 11.2 | 9.8 | 12.9 | 4 |
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| IOS | – | 06/I10/31 | 18.9 | 14.1 | 12.1 | 10.7 | 12.1 | 4 |
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| IOS | – | 06/I11/10 | 19.4 | 14.2 | 11.5 | 9.8 | 12.2 | 4 |
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| IOS | – | 06/I11/16 | 18.5 | 14.1 | 12.1 | 10.5 | 12.3 | 4 |
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| IOS | – | 06/I11/20 | 17.9 | 14.9 | 11.3 | 10.4 | 12.3 | 4 |
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| IOS | – | 06/J4/10 | 21.5 | 18.3 | 15.6 | 14.3 | 15.8 | 4 |
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| Third phalanx |
|---|
| Site | Level | Number | TL | H prox | TW prox | | | Source |
|---|
| HAT II | 6 | 1041 | 44.5 | 23.8 | 14.1 | | | 1 |
|---|
| HAT II | 6 | 674 | 35.7 | 25.0 | 13.9 | | | 1 |
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| HAT II | 6 | 879 | 31.0 | 26.0 | 11.3 | | | 1 |
|---|
| KEH | surface | n/n | – | – | 11.0 | | | 2 |
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| KEH | surface | HST7 | – | 26.0 | 13.0 | | | 2 |
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| KEH | 1 | 1 | 30.0 | 21.0 | 11.2 | | | 2 |
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| KEH | 2 | 52 | 46.0 | 23.0 | 14.0 | | | 2 |
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| KEH | 2 | n/n | 36.4 | 26.0 | 15.5 | | | 2 |
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| KEH | 2 | n/n | 30.0 | 21.0 | 11.0 | | | 2 |
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| KEH | 3 | n/n | 30.0 | 26.0 | 11.0 | | | 2 |
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| KEH | 3 | n/n | 41.0 | 24.0 | 14.0 | | | 2 |
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| KEH | 3 | n/n | 49.0 | – | 13.7 | | | 3 |
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| KEH | 4 | n/n | 56.3 | 28.0 | 15.5 | | | 3 |
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| KEH | 7 | n/n | – | 24.0 | 12.3 | | | 2 |
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| KTG | D | – | 50.5 | 33.0 | 16.7 | | | 3 |
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| KTG | E | – | 56.0 | 30.0 | 14.5 | | | 3 |
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| KTG | F | – | 54.0 | 28.5 | 14.4 | | | 3 |
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| KTG | F | – | 50.0 | 29.0 | 14.2 | | | 3 |
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| KTG | F | – | 51.0 | 29.8 | 15.0 | | | 3 |
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| KTG | F | – | 45.0 | – | 14.0 | | | 3 |
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| KBS | 3c | – | 50.5 | 26.0 | 14.7 | | | 3 |
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| KEB | salle | – | 36.0 | 26.0 | 15.0 | | | 3 |
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| IOS | – | 05/13 | 45.0 | 27.5 | 14.0 | | | 4 |
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| IOS | – | 06/I10/9 | 50.0 | 28.2 | 15.0 | | | 4 |
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| IOS | – | 06/I10/11 | 33.0 | 23.0 | 12.2 | | | 4 |
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| IOS | – | 06/I10/30 | 28.0 | 22.0 | 12.4 | | | 4 |
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| IOS | – | 06/I10/32 | 31.4 | 23.2 | 12.2 | | | 4 |
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| IOS | – | 06/I11/9 | 33.1 | 23.4 | 12.7 | | | 4 |
|---|
| IOS | – | 06/I11/15 | 30.0 | 21.0 | 13.0 | | | 4 |
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| IOS | – | 06/I12/1 | 32.2 | 23.0 | 14.2 | | | 4 |
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| IOS | – | 06/I13/5 | 34.0 | 21.5 | 14.2 | | | 4 |
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| IOS | – | 06/I10/10 | 32.0 | 25.0 | 14.2 | | | 4 |
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| IOS | – | 06/J5/6 | 31.8 | 23.7 | 13.4 | | | 4 |
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