Susana Carvalho


Primate Models for Behavioural Evolution Lab
University of Oxford
64 Banbury Road
United Kingdom


Centre for Functional Ecology
University of Coimbra



Research Interests

Susana Carvalho’s current funded projects include tackling the environmental andbehavioural contexts of the earliest tool users in the Koobi Fora formation, Kenya (funded by N.S.F., with D.R. Braun); investigating human origins and adaptations in complex environments with the Paleo-Primate Project Gorongosa, Mozambique (funded by the John Fell Fund, National Geographic Society, and Gorongosa Restoration Project); and testing hypotheses for how technological-related behaviours emerged in hominin evolution via using wild chimpanzees as models, e.g. the project ‘Stones on the move: the real life of a chimpanzee tool’, funded by the National Geographic Society. Researchers in Susana’s new lab ‘Primate Models for Behavioural Evolution’ conduct a broad variety of exciting studies on the evolution of human behaviourusing extant primates as models (e.g. cumulative culture in non-human primates; primate behavioural responses to shifts in predation pressures; automated approaches to find fossil sites, etc.).

Short Bio

Carvalho is a primatologist and palaeoanthropologist. She is an Associate Professor of Paleoanthropology and Fellow at St. Hugh's College. She has been one of main founders of the field of Primate Archaeology. She has been studying stone tool use by wild chimpanzees in Bossou, Guinea, West Africa, since 2006, and carries archaeological research in the Koobi Fora area, Kenya, East Africa since 2008, with a current focus on the archaeology of the Pliocene. She is the director of the Paleo-Primate Project Gorongosa since 2015, where an international team of 20 senior researchers is carrying an unprecedented interdisciplinary approach to understanding hominin origins and adaptations.

Key Publications

• Carvalho, S. and Beardmore-herd, M. (submitted). Technological origins: Primate perspectives and Early hominin tool use. To Encyclopaedia of African History.

• Habermann, J., Alemseged, Z., Aldeias, V., Archer, W., Bamford, M., Biro, D., Bobe, R., Braun, D.R., Capelli, C., Cunha, E., Ferreira da Silva, M.J., Luedecke, T., Madiquida, H., Martinez, F., Negash, E., Paulo, L.M., Pinto, M., Stalmans, M., Tatá, F., Wynn, J. and Carvalho, S. (submitted). Gorongosa by the sea: Neogene coastal-plain estuarine record in the Southern termination of the Rift Valley. Paleoenvironmental and paleoecological implications for the ancient East African margin and its primate land use potential. To Nature Communications.

• Bobe, R. and Carvalho, S. (in review). High hominin diversity and environmental variability in the Okote Member, Koobi Fora Formation, Kenya. Journal of Human Evolution.

• Biro, D., Carvalho, S., Gwyther, A. and Matsuzawa, T. (in review). Stone-tool “recycling” increases efficiency in wild juvenile chimpanzees. Proceedings of the Royal Society B.

• Mendonça, R., Dahl, C., Carvalho, S., Matsuzawa, T. and Adachi, I. (in review). Clear prosocial choice by chimpanzees (Pan troglodytes) and its social determinants.  Scientific Reports.

• Tibble, L. and Carvalho, S. (in review). Rethinking the evolution of property and possession: A review and methodological proposition.  Evolutionary Anthropology.

• Thompson, J., Carvalho, S., Marean, C. and Alemseged, Z. (in press). The origins of the human predatory pattern: The transition to large animal exploitation by early hominins. (Current Anthropology, special article with series of comments).

• Campbell, R. and Carvalho, S. (2017). Tool use and manufacture in the last common ancestor of Pan and Homo. In M. Muller, R. Wrangham and D. Pilbeam (eds.), Chimpanzees and Human Evolution, pp. 602-644. Harvard: Harvard University Press. ISBN 9780674967953

• Pereira, T., Paixão, E., Carvalho, V., Carvalho, S. and Gomes, T. (2017). Raw material diversity, distribution and acquisition in the river Lis basin, central Portugal. In T. Pereira, X. Terradas and N. Bicho (eds.), Raw materials exploitation in Prehistory: sourcing, processing and distribution, pp. 16-29. Cambridge: Cambridge Scholars Publishing. ISBN (10): 1-4438-9597-0 / ISBN (13): 978-1-4438-9597-2

• Carvalho, S. (2015). Standing on the shoulders of giants: The contribution of Claudia Sousa for the foundation of primate archaeology. In M. Fernandes, A. Frazão-Moreira, K.J. Hockings and F. Alves Cardoso (eds.), Special issue: Apes, People and Nature. The legacy of Cláudia Sousa. Etnográfica, 20(3): 648-652.

• Hockings, J.K., Bryson-Morrison, N., Carvalho, S., Fujisawa, M., Humle, T., McGrew, W.C., Nakamura, M., Ohashi, G., Yamanashi, Y., Yamakoshi, G. and Matsuzawa, T.  (2015). Tools to tipple: Ethanol ingestion by wild chimpanzees using leaf-sponges. Royal Society Open Scie, 2: 150150.  Link

• Benito-Calvo, A., Carvalho, S., Arroyo, A., Matsuzawa, T. and de la Torre, I. (2015). First GIS Analysis of Modern Stone Tools Used by Wild Chimpanzees (Pan troglodytes verus) in Bossou, Guinea, West Africa. PLoS ONE, 10(3): e0121613. doi:10.1371. [I.F.: 3.534]  Link

• Hockings, K.J., McLennan, M.R., Carvalho, S., Ancrenaz, M., Bobe, R., Byrne, R., Dunbar, R.I.M., Matsuzawa, T., McGrew, W.C., Wood, B., Wrangham, R. and Hill, C.M. Apes in the Anthropocene: Adaptation & Survival. Trends in Ecology and Evolution, 30(4): 215-222. [I.F.: 15.353]  Link

• Assis, S., Campanacho, V., Carvalho, V., Cruz, C., Curate, F., Gonçalves, D., Marques, R., Matos, V., Pereira, T. and Carvalho, S. (2014). Looking back at a decade of science communication in the field of human evolution. Notes in Human Evolution/Cadernos do GEEvH, 4(1):8-24.

• Correia, M.A., Cunha, E. and Carvalho, S. (2014). Finding our way: A review on the human evolution of spatial cognition. Notes in Human Evolution/Cadernos do GEEvH, 3(1): 7-37.

• Caruana, M., Carvalho, S., Braun, D.R., Presnyakov,a D., Haslam, M., Archer, W., Bobe, R. and Harris, J.W.K. (2014). Quantifying traces of tool use: A novel morphometric analysis of damage patterns on percussive tools. PLoS ONE, 9(11): e113856. [I.F.: 3.534]  Link

• Haslam, M., Gumert, M., Biro, D., Carvalho, S. and Malaivijitnond, S. (2013). Use-Wear patterns on wild macaque stone tools reveal their behavioural history. PLoS ONE, 8(8): e72872. doi:10.1371/journal.pone.0072872. [I.F.: 3.534]  Link

• Carvalho, S., Matsuzawa, T. and McGrew, W.C. (2013). From pounding to knapping: How living apes can help us model hominin lithics. In C. Sanz, J. Call and C. Boesch (eds.), Tool Use in Animals: Cognition and Ecology, pp. 225-241. Cambridge: Cambridge University Press.

• Carvalho, S., Biro, D., Cunha, E., Hockings, K., McGrew, W.C., Richmond, B. and Matsuzawa, T. (2012). Chimpanzee carrying behavior and the origins of human bipedality. Current Biology, 22(6): R180-181. [I.F.: 9.916]  Link

• Carvalho, S. and McGrew, W.C. (2012). The origins of the Oldowan: Why chimpanzees (Pan troglodytes) still are good models for technological evolution in Africa. In M. Domínguez-Rodrigo (ed.), Stone Tools and Fossil Bones: Debates in the Archaeology of Human Origins, pp. 222-244. Cambridge: Cambridge University Press.

• Assis, S. and Carvalho, S. (2012). Evolutionism and the teaching of science: How Portugal has been “Playing with the big tree of evolution”. Evolution: Education and Outreach, 5(3): 445-452.  Link

• Hockings, K., Humle, T., Carvalho, S. and Matsuzawa T. (2012). Chimpanzee interactions with other species in an anthropogenic habitat.  Behaviour, 149(3/4): 299-324. [I.F.: 1.401]

• Carvalho, S. (2011). Extensive surveys of chimpanzee stone tools: From the telescope to the magnifying glass. In T. Matsuzawa, T. Humle and Y. Sugiyama (eds.), The Chimpanzees of Bossou and Nimba: A Cultural Primatology, pp. 145-156. Tokyo: Springer.

• Carvalho, S. (2011) Diecké Forest, Guinea: Delving into chimpanzee behavior using stone tool surveys. In T. Matsuzawa, T. Humle and Y. Sugiyama (eds.), The Chimpanzees of Bossou and Nimba: A Cultural Primatology, pp. 301-312. Tokyo: Springer.

• Biro, D., Carvalho, S. and Matsuzawa, T. (2010). Tools, traditions and technologies: Interdisciplinary approaches to chimpanzee nut-cracking. In E.V. Lonsdorf, S.R. Ross and T. Matsuzawa (eds.), The Mind of the Chimpanzee: Ecological and Experimental Perspectives, pp. 141-155. Chicago: University of Chicago Press.

• Carvalho, S., Yamanashi, Y., Yamakoshi, G. and Matsuzawa, T. (2010). Bird in the hand: Bossou chimpanzees capture West African wood-owls (Ciccaba woodfordi) but not to eat. Pan African News, 17(1): 6-9.  Link

• Carvalho, S., Biro, D., McGrew, W.C. and Matsuzawa, T. (2009). Tool-composite reuse in wild chimpanzees (Pan troglodytes): Archaeologically invisible steps in the technological evolution of early hominins? Animal Cognition, 12: 103-114. [I.F.: 2.631]  Link

• Haslam, M., Hernandez-Aguílar, A., Ling, V., Carvalho, S., de la Torre, I., De Stefano, A., Du, A., Hardy, B., Harris, J.W.K., Marchant, L., Matsuzawa, T., McGrew, W., Mercader, J., Mora, R., Petraglia, M., Roche, H., Stout, D., Visalberghi, E. and Warren, R. (2009). Primate archaeology. Nature 460: 339-444. [I.F.: 42.351]  Link

• Carvalho, S., Matsuzawa, T., Sousa, C. and Cunha, E. (2009). Hammers, anvils & nuts: Chimpanzee technology? Applying the concept of chaîne opératoire to nut-cracking. In E. Cunha and Group of Studies in Human Evolution (eds.), Humans, evolution and environment, British Archaeological Reports (B.A.R) International Series, S2026, 22: 137-142.

• Ling, V., Hernandez-Aguilar, A., Haslam, M. and Carvalho, S. (2009). The origins of percussive technology: A smashing time in Cambridge. Evolutionary Anthropology, 18: 48-49. [I.F.: 4.886]

• Carvalho, S., Cunha, E., Sousa, C. and Matsuzawa, T. (2008). Chaînes Opératoires and resource exploitation strategies in chimpanzee nut-cracking (Pan troglodytes). Journal of Human Evolution, 55(1): 148-163. [I.F.: 3.867]  Link

• Carvalho, S., Sousa, C. and Matsuzawa, T. (2007). New nut-cracking sites in Diecké forest, Guinea: An overview of the etho-archaeological surveys. Pan African News, 14(1): 11-13.


Other scholarly publications
• Carvalho, S. and Hockings, K.J. (2015). Apes under pressure show their ingenuity – and hint at our own evolutionary past. The Conversation, US edition, online.  Link

• Mendonça, R. and Carvalho, S. (2015). Obituary: [In memory of Cláudia Sousa, the pioneer of Cognitive Primatology in Portugal]. Notes in Human Evolution/Cadernos do GEEvH, 3(2).

• Carvalho, S. (2011). Book review: Beautiful Minds, the Parallel Lives of Great Apes and Dolphins, by Stanford C., and Bearzi M. Antropologia Portuguesa, 26: 443-459.

• Carvalho, S., Carvalho, A., Angelucci, D.E. and Tavares, J. (2009). [Industrial Archaeology - The Paper Mill], pp. 32-49. Leiria: Câmara Municipal de Leiria. [in Portuguese]

• Carvalho, S. and Tavares, J. (2005). [The flint source and Martinela knapping site]. In S. Carvalho (ed.), [Inhabitants and Habitats – Pre and Proto history in the Lis River Basin], pp. 26-33. Leiria: Câmara Municipal de Leiria.

• Zambujo, G. and Carvalho, S. (2005). [Quinta do Bispo – Parceiros: The first Mesolithic site in the Lis River basin]. In S. Carvalho (ed.), [Inhabitants and Habitats – Pre and Proto history in the Lis River Basin], pp. 84-103. Leiria: Câmara Municipal de Leiria.

• Carvalho, S. (2003). [Preventive test pits in the Castle of the Outeiro or Carrilhas], Arqueologia Medieval, 8: 83-97.

• Carvalho, S. (2003). [Emergency excavation in the Old Cloister of the Monastery of S. João de Deus – Montemor-o-Novo], Almansor, 2: 111-144.

• Carvalho, S. (2002). [The Castle of Outeiro or Carrilhas (Lavre)], Almansor, 1: 7-32.


• 2016-2019: National Science Foundation: ‘Technological Origins: Environmental and Behavioral Context of the Earliest Tool Users”. (Senior Personnel, P.I. D.R. Braun). $320,000. Awarded.
• 2016-2018: John Fell Fund: ‘Human origin and adaptation in complex environments: Gorongosa, the unexplored African Rift Valley’. (PI) £43,908. Awarded.
• 2016-2017: National Geographic Society’s Global Exploration Fund: The unexplored Rift Valley: human origins research at Gorongosa National Park’ (Co-P.I. with R. Bobe). €19,759. Awarded.
• 2006: International Team for Bossou-Nimba Research, Kyoto University, Japan: ‘Seeking the evolutionary origins of technology’. Long-term collaboration with Japan and Guinea. Current funding bodies: MEXT grant #20002001 and #24000001, JSPS-U04-PWS to T. Matsuzawa