PALAEO.ZOOARCH
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PALAEO.ZOOARCH investigates how Pre-Neanderthals, Neanderthals and Anatomically Modern Humans adapted their subsistence strategies to environmental and socio-cultural change in Atlantic Iberia. Focusing on faunal assemblages dating from Marine Isotope Stage 11c to MIS 2, approximately 424,000–12,000 years ago, the project examines changes in animal exploitation across the Early, Middle and Upper Palaeolithic. By integrating zooarchaeology, taphonomy, biometrics, isotopic evidence and experimental archaeology, it seeks to move beyond traditional models of Pleistocene subsistence centred primarily on large-game hunting.
Project Overview
Research on Palaeolithic diets has often portrayed earlier human populations, particularly Neanderthals, as specialised hunters dependent on large terrestrial mammals. However, evidence from Portuguese archaeological sites demonstrates that Neanderthals also exploited tortoises, birds, shellfish, crabs and other small or aquatic resources. These finds reveal flexible decision-making, detailed ecological knowledge and a capacity to respond creatively to locally available resources.
PALAEO.ZOOARCH develops this research through a diachronic comparison of human-animal interactions during two major transitional periods: the Early-to-Middle and Middle-to-Upper Palaeolithic. Its geographical focus is central Portugal, particularly the Almonda karst system and the Nabão valley, supplemented by comparative evidence from sites such as Gruta da Figueira Brava and Lapa dos Furos. Principal case studies include Gruta da Oliveira, Gruta do Caldeirão and Abrigo da Senhora das Lapas.
The project will analyse faunal remains to reconstruct prey selection, carcass transport, butchery, cooking, consumption and discard. Taxonomic and anatomical identification, mortality profiles, skeletal representation, biometrics and bone-surface analysis will be used to distinguish human activity from carnivore action and other natural processes. These results will be integrated with palaeoenvironmental and isotopic evidence to investigate animal mobility, habitat use, seasonality and human landscape exploitation.
Experimental archaeology constitutes a central component of the project. Controlled experiments involving birds, turtles and crabs will replicate different methods of skinning, butchery and cooking. The resulting fragmentation patterns and surface modifications will provide comparative criteria for recognising small-prey processing in archaeological assemblages, extending previous experimental work.
PALAEO.ZOOARCH develops this research through a diachronic comparison of human-animal interactions during two major transitional periods: the Early-to-Middle and Middle-to-Upper Palaeolithic. Its geographical focus is central Portugal, particularly the Almonda karst system and the Nabão valley, supplemented by comparative evidence from sites such as Gruta da Figueira Brava and Lapa dos Furos. Principal case studies include Gruta da Oliveira, Gruta do Caldeirão and Abrigo da Senhora das Lapas.
The project will analyse faunal remains to reconstruct prey selection, carcass transport, butchery, cooking, consumption and discard. Taxonomic and anatomical identification, mortality profiles, skeletal representation, biometrics and bone-surface analysis will be used to distinguish human activity from carnivore action and other natural processes. These results will be integrated with palaeoenvironmental and isotopic evidence to investigate animal mobility, habitat use, seasonality and human landscape exploitation.
Experimental archaeology constitutes a central component of the project. Controlled experiments involving birds, turtles and crabs will replicate different methods of skinning, butchery and cooking. The resulting fragmentation patterns and surface modifications will provide comparative criteria for recognising small-prey processing in archaeological assemblages, extending previous experimental work.
Aims and Objectives
The principal aim is to establish a new diachronic model of Pleistocene subsistence in Atlantic Iberia, explaining how human populations adapted their use of animal resources during periods of climatic, ecological and cultural transformation.
The project will:
The project will:
- Compare subsistence strategies among Pre-Neanderthals, Neanderthals and Anatomically Modern Humans across the Early, Middle and Upper Palaeolithic.
- Assess the influence of environmental and socio-cultural factors on prey choice, food provisioning and resource diversification during major transitional periods.
- Reconstruct animal acquisition and processing, including hunting, transport, butchery, cooking, consumption and discard practices.
- Determine the contribution of small prey and aquatic resources to Pleistocene diets and evaluate whether their exploitation reflects necessity, local availability, technological choices or cultural preferences.
- Develop improved taphonomic and biometric criteria for identifying human modifications on bones and shells and distinguishing them from natural processes.
Funding Body and Collaborations
Supported by Portuguese funds through FCT - Fundação para a Ciência e a Tecnologia in the framework of the project 2024.08182.CEECIND
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Selected Bibliography
Angelucci, D., Nabais, M., Zilhão, J., 2023. Formation processes, fire use, and patterns of human occupation across the Middle Palaeolithic (MIS 5a-5b) of Gruta da Oliveira (Almonda karst system, Torres Novas, Portugal) PLoS ONE, 18(10): e0292075 https://doi.org/10.1371/journal.pone.0292075
Aubry, T., Dimuccio, L., Almeida, M., Neves, M., Angelucci, D., Cunha, L., 2011. Palaeoenvironmental forcing during the Middle-Upper Palaeolithic transition in central-western Portugal. Quaternary research, 75, 66-79. https://doi.org/10.1016/j.yqres.2010.11.002
Daura, J., Sanz, M., Arsuaga, JL., Hoffmann, D., Quam, R., Cruz Ortega, M., Santos, E., Gómez, S., Rubio, A., Villaescusa, L., Souto, P., Maurício, J., Rodrigues, F., Ferreira, A., Godinho, P., Trinkaus, E., Zilhão, J., 2017. New Middle Pleistocene hominin cranium from Gruta da Aroeira (Portugal). Proceedings of the National Academy of Sciences, 114: 3397-3402. https://doi.org/10.1073/pnas.1619040114
Deschamps, M., Zilhão, J., 2018. Assessing site formation and assemblage integrity through stone tool refitting at Gruta da Oliveira (Almonda karst system, Torres Novas, Portugal): A Middle Paleolithic case study. PLoS ONE, 13: e0192423. https://doi.org/10.1371/journal.pone.0192423
Deschamps, M., Martín-Lerma, I., Linares-Matás, G., Zilhão, J., 2022. Organization of residential space, site function variability, and seasonality of activities among MIS 5 Iberian Neandertals. Scientific Reports, 12: 2022 https://www.nature.com/articles/s41598-022-24430-z
Linscott, B., Pike, A., Angelucci, D.E, Zilhão, J., 2023. Reconstructing Middle and Upper Paleolithic human mobility in Portuguese Estremadura through laser ablation strontium isotope analysis. PNAS 120 (20): e2204501120. https://doi.org/10.1073/pnas.2204501120
López-García, J.M., Póvoas, L., Zilhão, J., 2020. Note on the taxonomy of the Microtus (Iberomys) (Arvicolinae, Rodentia) from the Late Pleistocene of Gruta do Caldeirão (Tomar, Portugal) and paleoclimatic interpretation of the rodent assemblage. Estudios Geológicos, 76: e128. https://doi.org/10.3989/egeol.43622.542
López-García, JM, Fagoaga, A., Nabais, M., Póvoas, L., Zilhão, J., 2022. Late Quaternary (MIS 5a-5b) climate and environments of western Iberia inferred from the small-mammal assemblage of Gruta da Oliveira, Torres Novas, Portugal. Palaeogeography, Palaeoclimatology, Palaeoecology, 603: 111194. http://dx.doi.org/10.1016/j.palaeo.2022.111194
Nabais, M., Zilhão, J., 2019. The consumption of tortoise among Last Interglacial Iberian Neanderthals. Quaternary Science Reviews, 217: 225-246. https://doi.org/10.1016/j.quascirev.2019.03.024
Nabais, M., Zilhão, J., 2025. Ungulate exploitation and procurement strategies in the late MIS-5 Neanderthal occupation sequence of Gruta da Figueira Brava (Portugal). Frontiers in Environmental Archaeology, 4. https://doi.org/10.3389/fearc.2025.1564495
Nabais, M., Blasco, R., Boneta, I., Gonçalves, D., Igreja, M., Lubrano, V., Rufà, A., 2026. Raw vs Roasted: Experimental Approaches to Turtle Butchery and Their Implications for Early Human Cognition and Behaviour. Scientific Reports, 16: 1913. https://doi.org/10.1038/s41598-025-31738-z
Nabais, M., Dupont, C., Zilhão, J., 2023. The exploitation of crabs by Last Interglacial Iberian Neanderthals: the evidence from Gruta da Figueira Brava (Portugal). Frontiers in Environmental Archaeology, 2: 1097815. https://doi.org/10.3389/fearc.2023.1097815
Nabais, M., Pimenta, C., Zilhão, J., 2023. Human-bird interaction in last Interglacial Iberia: A combined approach using skeletal part analysis, bone surface modification, bird ethology and ethnography. Journal of Archaeological Science: Reports 49, 104023. https://doi.org/10.1016/j.jasrep.2023.104023
Nabais, M., Rufà, A., Igreja, M., 2024. Experimental replication of early human behaviour in bird preparation: a pilot-study focusing on bone surface modification and breakage patterns. Frontiers in Environmental Archaeology 3: 1411853. https://doi.org/10.3389/fearc.2024.1411853
Sanz, M., Daura, J., Cabanes, D., Éguez, N., Carrancho, A., Badal, E., Souto, P., Rodrigues, F., Zilhão, J., 2020. Early evidence of fire in south-western Europe: the Acheulean site of Gruta da Aroeira (Torres Novas, Portugal). Scientific Reports, 10: 12053. https://doi.org/10.1038/s41598-020-68839-w
Sanz, M., Sala, N., Daura, J., Pantoja-Pérez, A., Santos, E., Zilhão, J., 2018. Taphonomic inferences about Middle Pleistocene hominins: The human cranium of Gruta da Aroeira (Portugal). American Journal of Biological Anthropology, 167: 615-627 https://doi.org/10.1002/ajpa.23689
Zilhão, J., Angelucci, D.E., Arnold, L.J., Demuro, M., Hoffmann, D.L., Pike, A.W.G., 2021. A revised, Last Interglacial chronology for the Middle Palaeolithic sequence of Gruta da Oliveira (Almonda karst system, Torres Novas, Portugal). Quaternary Science Reviews, 258: 106885. https://doi.org/10.1016/j.quascirev.2021.106885
Zilhão, J., Angelucci, D., Araújo-igreja, M., Arnold, L., Badal, E., Callapez, P., Cardoso, J., D’Errico, F., Daura, J., Demuro, M., Deschamps, M., Dupont, C., Gabriel, S., Hoffmann, D., Legoinha, P., Matias, H., Monge soares, A., Nabais, M., Rodrigues, A., Souto, P., Wainer, K., 2020. Last Interglacial Iberian Neandertals as Fisher-Hunter-Gatherers. Science, 367: eaaz7943. http://dx.doi.org/10.1126/science.aaz7943
Zilhão, J., Angelucci, D., Arnold, L., D’Errico, F., Dayet, L., Demuro, M., Deschamps, M., Fewlass, H., Gomes, L., Linscott, B., Matias, H., Pike, A., Steier, P., Talamo, S., Wild, E., 2021. Revisiting the Middle and Upper Palaeolithic archaeology of Gruta do Caldeirão (Tomar, Portugal). PLoS One, 16: e0259089. https://doi.org/10.1371/journal.pone.0259089
Aubry, T., Dimuccio, L., Almeida, M., Neves, M., Angelucci, D., Cunha, L., 2011. Palaeoenvironmental forcing during the Middle-Upper Palaeolithic transition in central-western Portugal. Quaternary research, 75, 66-79. https://doi.org/10.1016/j.yqres.2010.11.002
Daura, J., Sanz, M., Arsuaga, JL., Hoffmann, D., Quam, R., Cruz Ortega, M., Santos, E., Gómez, S., Rubio, A., Villaescusa, L., Souto, P., Maurício, J., Rodrigues, F., Ferreira, A., Godinho, P., Trinkaus, E., Zilhão, J., 2017. New Middle Pleistocene hominin cranium from Gruta da Aroeira (Portugal). Proceedings of the National Academy of Sciences, 114: 3397-3402. https://doi.org/10.1073/pnas.1619040114
Deschamps, M., Zilhão, J., 2018. Assessing site formation and assemblage integrity through stone tool refitting at Gruta da Oliveira (Almonda karst system, Torres Novas, Portugal): A Middle Paleolithic case study. PLoS ONE, 13: e0192423. https://doi.org/10.1371/journal.pone.0192423
Deschamps, M., Martín-Lerma, I., Linares-Matás, G., Zilhão, J., 2022. Organization of residential space, site function variability, and seasonality of activities among MIS 5 Iberian Neandertals. Scientific Reports, 12: 2022 https://www.nature.com/articles/s41598-022-24430-z
Linscott, B., Pike, A., Angelucci, D.E, Zilhão, J., 2023. Reconstructing Middle and Upper Paleolithic human mobility in Portuguese Estremadura through laser ablation strontium isotope analysis. PNAS 120 (20): e2204501120. https://doi.org/10.1073/pnas.2204501120
López-García, J.M., Póvoas, L., Zilhão, J., 2020. Note on the taxonomy of the Microtus (Iberomys) (Arvicolinae, Rodentia) from the Late Pleistocene of Gruta do Caldeirão (Tomar, Portugal) and paleoclimatic interpretation of the rodent assemblage. Estudios Geológicos, 76: e128. https://doi.org/10.3989/egeol.43622.542
López-García, JM, Fagoaga, A., Nabais, M., Póvoas, L., Zilhão, J., 2022. Late Quaternary (MIS 5a-5b) climate and environments of western Iberia inferred from the small-mammal assemblage of Gruta da Oliveira, Torres Novas, Portugal. Palaeogeography, Palaeoclimatology, Palaeoecology, 603: 111194. http://dx.doi.org/10.1016/j.palaeo.2022.111194
Nabais, M., Zilhão, J., 2019. The consumption of tortoise among Last Interglacial Iberian Neanderthals. Quaternary Science Reviews, 217: 225-246. https://doi.org/10.1016/j.quascirev.2019.03.024
Nabais, M., Zilhão, J., 2025. Ungulate exploitation and procurement strategies in the late MIS-5 Neanderthal occupation sequence of Gruta da Figueira Brava (Portugal). Frontiers in Environmental Archaeology, 4. https://doi.org/10.3389/fearc.2025.1564495
Nabais, M., Blasco, R., Boneta, I., Gonçalves, D., Igreja, M., Lubrano, V., Rufà, A., 2026. Raw vs Roasted: Experimental Approaches to Turtle Butchery and Their Implications for Early Human Cognition and Behaviour. Scientific Reports, 16: 1913. https://doi.org/10.1038/s41598-025-31738-z
Nabais, M., Dupont, C., Zilhão, J., 2023. The exploitation of crabs by Last Interglacial Iberian Neanderthals: the evidence from Gruta da Figueira Brava (Portugal). Frontiers in Environmental Archaeology, 2: 1097815. https://doi.org/10.3389/fearc.2023.1097815
Nabais, M., Pimenta, C., Zilhão, J., 2023. Human-bird interaction in last Interglacial Iberia: A combined approach using skeletal part analysis, bone surface modification, bird ethology and ethnography. Journal of Archaeological Science: Reports 49, 104023. https://doi.org/10.1016/j.jasrep.2023.104023
Nabais, M., Rufà, A., Igreja, M., 2024. Experimental replication of early human behaviour in bird preparation: a pilot-study focusing on bone surface modification and breakage patterns. Frontiers in Environmental Archaeology 3: 1411853. https://doi.org/10.3389/fearc.2024.1411853
Sanz, M., Daura, J., Cabanes, D., Éguez, N., Carrancho, A., Badal, E., Souto, P., Rodrigues, F., Zilhão, J., 2020. Early evidence of fire in south-western Europe: the Acheulean site of Gruta da Aroeira (Torres Novas, Portugal). Scientific Reports, 10: 12053. https://doi.org/10.1038/s41598-020-68839-w
Sanz, M., Sala, N., Daura, J., Pantoja-Pérez, A., Santos, E., Zilhão, J., 2018. Taphonomic inferences about Middle Pleistocene hominins: The human cranium of Gruta da Aroeira (Portugal). American Journal of Biological Anthropology, 167: 615-627 https://doi.org/10.1002/ajpa.23689
Zilhão, J., Angelucci, D.E., Arnold, L.J., Demuro, M., Hoffmann, D.L., Pike, A.W.G., 2021. A revised, Last Interglacial chronology for the Middle Palaeolithic sequence of Gruta da Oliveira (Almonda karst system, Torres Novas, Portugal). Quaternary Science Reviews, 258: 106885. https://doi.org/10.1016/j.quascirev.2021.106885
Zilhão, J., Angelucci, D., Araújo-igreja, M., Arnold, L., Badal, E., Callapez, P., Cardoso, J., D’Errico, F., Daura, J., Demuro, M., Deschamps, M., Dupont, C., Gabriel, S., Hoffmann, D., Legoinha, P., Matias, H., Monge soares, A., Nabais, M., Rodrigues, A., Souto, P., Wainer, K., 2020. Last Interglacial Iberian Neandertals as Fisher-Hunter-Gatherers. Science, 367: eaaz7943. http://dx.doi.org/10.1126/science.aaz7943
Zilhão, J., Angelucci, D., Arnold, L., D’Errico, F., Dayet, L., Demuro, M., Deschamps, M., Fewlass, H., Gomes, L., Linscott, B., Matias, H., Pike, A., Steier, P., Talamo, S., Wild, E., 2021. Revisiting the Middle and Upper Palaeolithic archaeology of Gruta do Caldeirão (Tomar, Portugal). PLoS One, 16: e0259089. https://doi.org/10.1371/journal.pone.0259089
