title: Chimpanzee accumulative stone throwing.
abstract: The study of the archaeological remains of fossil hominins must rely on reconstructions to elucidate the behaviour that may have resulted in particular stone tools and their accumulation. Comparatively, stone tool use among living primates has illuminated behaviours that are also amenable to archaeological examination, permitting direct observations of the behaviour leading to artefacts and their assemblages to be incorporated. Here, we describe newly discovered stone tool-use behaviour and stone accumulation sites in wild chimpanzees reminiscent of human cairns. In addition to data from 17 mid- to long-term chimpanzee research sites, we sampled a further 34 Pan troglodytes communities. We found four populations in West Africa where chimpanzees habitually bang and throw rocks against trees, or toss them into tree cavities, resulting in conspicuous stone accumulations at these sites. This represents the first record of repeated observations of individual chimpanzees exhibiting stone tool use for a purpose other than extractive foraging at what appear to be targeted trees. The ritualized behavioural display and collection of artefacts at particular locations observed in chimpanzee accumulative stone throwing may have implications for the inferences that can be drawn from archaeological stone assemblages and the origins of ritual sites.
authors.affiliation: Max Planck Institute for Evolutionary Anthropology (MPI EVAN), Deutscher Platz 6, 04103 Leipzig, Germany.|German Centre for Integrative Biodiversity Research (iDiv) Halle-Leipzig-Jena, Deutscher Platz 5e, 04103 Leipzig, Germany.
authors.forename: Hjalmar S
authors.initials: HS
authors.lastname: Kühl
authors.affiliation: Max Planck Institute for Evolutionary Anthropology (MPI EVAN), Deutscher Platz 6, 04103 Leipzig, Germany.
authors.forename: Ammie K
authors.initials: AK
authors.lastname: Kalan
authors.affiliation: Max Planck Institute for Evolutionary Anthropology (MPI EVAN), Deutscher Platz 6, 04103 Leipzig, Germany.
authors.forename: Mimi
authors.initials: M
authors.lastname: Arandjelovic
authors.affiliation: Wild Chimpanzee Foundation (WCF), Deutscher Platz 6, 04103 Leipzig, Germany.
authors.forename: Floris
authors.initials: F
authors.lastname: Aubert
authors.affiliation: Wild Chimpanzee Foundation (WCF), Deutscher Platz 6, 04103 Leipzig, Germany.
authors.forename: Lucy
authors.initials: L
authors.lastname: D'Auvergne
authors.affiliation: Chimbo Foundation, Amstel 49, 1011 PW Amsterdam, Netherlands.
authors.forename: Annemarie
authors.initials: A
authors.lastname: Goedmakers
authors.affiliation: Max Planck Institute for Evolutionary Anthropology (MPI EVAN), Deutscher Platz 6, 04103 Leipzig, Germany.
authors.forename: Sorrel
authors.initials: S
authors.lastname: Jones
authors.affiliation: Wild Chimpanzee Foundation (WCF), Deutscher Platz 6, 04103 Leipzig, Germany.
authors.forename: Laura
authors.initials: L
authors.lastname: Kehoe
authors.affiliation: Wild Chimpanzee Foundation (WCF), Deutscher Platz 6, 04103 Leipzig, Germany.
authors.forename: Sebastien
authors.initials: S
authors.lastname: Regnaut
authors.affiliation: Max Planck Institute for Evolutionary Anthropology (MPI EVAN), Deutscher Platz 6, 04103 Leipzig, Germany.
authors.forename: Alexander
authors.initials: A
authors.lastname: Tickle
authors.affiliation: Max Planck Institute for Evolutionary Anthropology (MPI EVAN), Deutscher Platz 6, 04103 Leipzig, Germany.|Chimbo Foundation, Amstel 49, 1011 PW Amsterdam, Netherlands.
authors.forename: Els
authors.initials: E
authors.lastname: Ton
authors.affiliation: Max Planck Institute for Evolutionary Anthropology (MPI EVAN), Deutscher Platz 6, 04103 Leipzig, Germany.|Chimbo Foundation, Amstel 49, 1011 PW Amsterdam, Netherlands.
authors.forename: Joost
authors.initials: J
authors.lastname: van Schijndel
authors.affiliation: Ebo Forest Research Project, BP3055, Messa, Cameroon.
authors.forename: Ekwoge E
authors.initials: EE
authors.lastname: Abwe
authors.affiliation: Max Planck Institute for Evolutionary Anthropology (MPI EVAN), Deutscher Platz 6, 04103 Leipzig, Germany.
authors.forename: Samuel
authors.initials: S
authors.lastname: Angedakin
authors.affiliation: Max Planck Institute for Evolutionary Anthropology (MPI EVAN), Deutscher Platz 6, 04103 Leipzig, Germany.
authors.forename: Anthony
authors.initials: A
authors.lastname: Agbor
authors.affiliation: Max Planck Institute for Evolutionary Anthropology (MPI EVAN), Deutscher Platz 6, 04103 Leipzig, Germany.
authors.forename: Emmanuel Ayuk
authors.initials: EA
authors.lastname: Ayimisin
authors.affiliation: Max Planck Institute for Evolutionary Anthropology (MPI EVAN), Deutscher Platz 6, 04103 Leipzig, Germany.
authors.forename: Emma
authors.initials: E
authors.lastname: Bailey
authors.affiliation: Max Planck Institute for Evolutionary Anthropology (MPI EVAN), Deutscher Platz 6, 04103 Leipzig, Germany.
authors.forename: Mattia
authors.initials: M
authors.lastname: Bessone
authors.affiliation: The Aspinall Foundation, Port Lympne Wild Animal Park, Hythe, Kent, UK.
authors.forename: Matthieu
authors.initials: M
authors.lastname: Bonnet
authors.affiliation: Max Planck Institute for Evolutionary Anthropology (MPI EVAN), Deutscher Platz 6, 04103 Leipzig, Germany.
authors.forename: Gregory
authors.initials: G
authors.lastname: Brazolla
authors.affiliation: Max Planck Institute for Evolutionary Anthropology (MPI EVAN), Deutscher Platz 6, 04103 Leipzig, Germany.
authors.forename: Valentine Ebua
authors.initials: VE
authors.lastname: Buh
authors.affiliation: West Chester University, Departments of Anthropology &Sociology and Psychology, West Chester, PA, USA.
authors.forename: Rebecca
authors.initials: R
authors.lastname: Chancellor
authors.affiliation: Wildlife Conservation Society (WCS), 2300 Southern Boulevard. Bronx, New York 10460, USA.
authors.forename: Chloe
authors.initials: C
authors.lastname: Cipoletta
authors.affiliation: Max Planck Institute for Evolutionary Anthropology (MPI EVAN), Deutscher Platz 6, 04103 Leipzig, Germany.
authors.forename: Heather
authors.initials: H
authors.lastname: Cohen
authors.affiliation: Max Planck Institute for Evolutionary Anthropology (MPI EVAN), Deutscher Platz 6, 04103 Leipzig, Germany.
authors.forename: Katherine
authors.initials: K
authors.lastname: Corogenes
authors.affiliation: Max Planck Institute for Evolutionary Anthropology (MPI EVAN), Deutscher Platz 6, 04103 Leipzig, Germany.
authors.forename: Charlotte
authors.initials: C
authors.lastname: Coupland
authors.affiliation: The Aspinall Foundation, Port Lympne Wild Animal Park, Hythe, Kent, UK.
authors.forename: Bryan
authors.initials: B
authors.lastname: Curran
authors.affiliation: Max Planck Institute for Evolutionary Anthropology (MPI EVAN), Deutscher Platz 6, 04103 Leipzig, Germany.
authors.forename: Tobias
authors.initials: T
authors.lastname: Deschner
authors.affiliation: Max Planck Institute for Evolutionary Anthropology (MPI EVAN), Deutscher Platz 6, 04103 Leipzig, Germany.
authors.forename: Karsten
authors.initials: K
authors.lastname: Dierks
authors.affiliation: Max Planck Institute for Evolutionary Anthropology (MPI EVAN), Deutscher Platz 6, 04103 Leipzig, Germany.
authors.forename: Paula
authors.initials: P
authors.lastname: Dieguez
authors.affiliation: Wildlife Conservation Society (WCS), 2300 Southern Boulevard. Bronx, New York 10460, USA.
authors.forename: Emmanuel
authors.initials: E
authors.lastname: Dilambaka
authors.affiliation: Korup Rainforest Conservation Society, c/o Korup National Park, P.O. Box 36 Mundemba, South West Region, Cameroon.
authors.forename: Orume
authors.initials: O
authors.lastname: Diotoh
authors.affiliation: Wild Chimpanzee Foundation (WCF), Deutscher Platz 6, 04103 Leipzig, Germany.
authors.forename: Dervla
authors.initials: D
authors.lastname: Dowd
authors.affiliation: Wildlife Conservation Society (WCS), 2300 Southern Boulevard. Bronx, New York 10460, USA.
authors.forename: Andrew
authors.initials: A
authors.lastname: Dunn
authors.affiliation: Max Planck Institute for Evolutionary Anthropology (MPI EVAN), Deutscher Platz 6, 04103 Leipzig, Germany.
authors.forename: Henk
authors.initials: H
authors.lastname: Eshuis
authors.affiliation: Max Planck Institute for Evolutionary Anthropology (MPI EVAN), Deutscher Platz 6, 04103 Leipzig, Germany.
authors.forename: Rumen
authors.initials: R
authors.lastname: Fernandez
authors.affiliation: Max Planck Institute for Evolutionary Anthropology (MPI EVAN), Deutscher Platz 6, 04103 Leipzig, Germany.
authors.forename: Yisa
authors.initials: Y
authors.lastname: Ginath
authors.affiliation: Lukuru Foundation, 1235 Avenue des Poids Lourds/Quartier de Kingabois, Kinshasa, DRC.
authors.forename: John
authors.initials: J
authors.lastname: Hart
authors.affiliation: The Aspinall Foundation, Port Lympne Wild Animal Park, Hythe, Kent, UK.
authors.forename: Daniela
authors.initials: D
authors.lastname: Hedwig
authors.affiliation: WWF Cameroon Country Office, BP6776; Yaoundé, Cameroon.
authors.forename: Martijn
authors.initials: M
authors.lastname: Ter Heegde
authors.affiliation: Max Planck Institute for Evolutionary Anthropology (MPI EVAN), Deutscher Platz 6, 04103 Leipzig, Germany.
authors.forename: Thurston Cleveland
authors.initials: TC
authors.lastname: Hicks
authors.affiliation: Max Planck Institute for Evolutionary Anthropology (MPI EVAN), Deutscher Platz 6, 04103 Leipzig, Germany.|Wildlife Conservation Society (WCS), 2300 Southern Boulevard. Bronx, New York 10460, USA.
authors.forename: Inaoyom
authors.initials: I
authors.lastname: Imong
authors.affiliation: Agence National des Parcs Nationaux (ANPN) Batterie 4, BP20379, Libreville, Gabon.|School of Natural Sciences, University of Stirling, Stirling, UK.|Institute de Recherche en Ecologie Tropicale, Libreville, Gabon.
authors.forename: Kathryn J
authors.initials: KJ
authors.lastname: Jeffery
authors.affiliation: Max Planck Institute for Evolutionary Anthropology (MPI EVAN), Deutscher Platz 6, 04103 Leipzig, Germany.
authors.forename: Jessica
authors.initials: J
authors.lastname: Junker
authors.affiliation: University of Cambridge, Pembroke Street, Cambridge, UK CB2 3QG.
authors.forename: Parag
authors.initials: P
authors.lastname: Kadam
authors.affiliation: Max Planck Institute for Evolutionary Anthropology (MPI EVAN), Deutscher Platz 6, 04103 Leipzig, Germany.
authors.forename: Mohamed
authors.initials: M
authors.lastname: Kambi
authors.affiliation: Max Planck Institute for Evolutionary Anthropology (MPI EVAN), Deutscher Platz 6, 04103 Leipzig, Germany.
authors.forename: Ivonne
authors.initials: I
authors.lastname: Kienast
authors.affiliation: Wildlife Conservation Society (WCS), 2300 Southern Boulevard. Bronx, New York 10460, USA.
authors.forename: Deo
authors.initials: D
authors.lastname: Kujirakwinja
authors.affiliation: Arizona State University, PO Box 872402, Tempe, AZ 85287-2402 USA.
authors.forename: Kevin
authors.initials: K
authors.lastname: Langergraber
authors.affiliation: Wild Chimpanzee Foundation (WCF), Deutscher Platz 6, 04103 Leipzig, Germany.
authors.forename: Vincent
authors.initials: V
authors.lastname: Lapeyre
authors.affiliation: Max Planck Institute for Evolutionary Anthropology (MPI EVAN), Deutscher Platz 6, 04103 Leipzig, Germany.
authors.forename: Juan
authors.initials: J
authors.lastname: Lapuente
authors.affiliation: Max Planck Institute for Evolutionary Anthropology (MPI EVAN), Deutscher Platz 6, 04103 Leipzig, Germany.
authors.forename: Kevin
authors.initials: K
authors.lastname: Lee
authors.affiliation: Wild Chimpanzee Foundation (WCF), Deutscher Platz 6, 04103 Leipzig, Germany.
authors.forename: Vera
authors.initials: V
authors.lastname: Leinert
authors.affiliation: Max Planck Institute for Evolutionary Anthropology (MPI EVAN), Deutscher Platz 6, 04103 Leipzig, Germany.
authors.forename: Amelia
authors.initials: A
authors.lastname: Meier
authors.affiliation: Max Planck Institute for Evolutionary Anthropology (MPI EVAN), Deutscher Platz 6, 04103 Leipzig, Germany.
authors.forename: Giovanna
authors.initials: G
authors.lastname: Maretti
authors.affiliation: Max Planck Institute for Evolutionary Anthropology (MPI EVAN), Deutscher Platz 6, 04103 Leipzig, Germany.
authors.forename: Sergio
authors.initials: S
authors.lastname: Marrocoli
authors.affiliation: Max Planck Institute for Evolutionary Anthropology (MPI EVAN), Deutscher Platz 6, 04103 Leipzig, Germany.
authors.forename: Tanyi Julius
authors.initials: TJ
authors.lastname: Mbi
authors.affiliation: Agence National des Parcs Nationaux (ANPN) Batterie 4, BP20379, Libreville, Gabon.
authors.forename: Vianet
authors.initials: V
authors.lastname: Mihindou
authors.affiliation: Max Planck Institute for Evolutionary Anthropology (MPI EVAN), Deutscher Platz 6, 04103 Leipzig, Germany.
authors.forename: Yasmin
authors.initials: Y
authors.lastname: Moebius
authors.affiliation: Wildlife Conservation Society (WCS), 2300 Southern Boulevard. Bronx, New York 10460, USA.|Lester E. Fisher Center for the Study and Conservation of Apes, Lincoln Park Zoo, 2001 North Clark Street, Chicago, Illinois 60614 USA.
authors.forename: David
authors.initials: D
authors.lastname: Morgan
authors.affiliation: Ebo Forest Research Project, BP3055, Messa, Cameroon.|Institute for Conservation Research, Zoological Society of San Diego, Escondido, CA 92025, USA.
authors.forename: Bethan
authors.initials: B
authors.lastname: Morgan
authors.affiliation: Wildlife Conservation Society (WCS), 2300 Southern Boulevard. Bronx, New York 10460, USA.
authors.forename: Felix
authors.initials: F
authors.lastname: Mulindahabi
authors.affiliation: Max Planck Institute for Evolutionary Anthropology (MPI EVAN), Deutscher Platz 6, 04103 Leipzig, Germany.
authors.forename: Mizuki
authors.initials: M
authors.lastname: Murai
authors.affiliation: Wildlife Conservation Society (WCS), 2300 Southern Boulevard. Bronx, New York 10460, USA.
authors.forename: Protais
authors.initials: P
authors.lastname: Niyigabae
authors.affiliation: Wild Chimpanzee Foundation (WCF), Deutscher Platz 6, 04103 Leipzig, Germany.
authors.forename: Emma
authors.initials: E
authors.lastname: Normand
authors.affiliation: Wildlife Conservation Society (WCS), 2300 Southern Boulevard. Bronx, New York 10460, USA.
authors.forename: Nicolas
authors.initials: N
authors.lastname: Ntare
authors.affiliation: Max Planck Institute for Evolutionary Anthropology (MPI EVAN), Deutscher Platz 6, 04103 Leipzig, Germany.
authors.forename: Lucy Jayne
authors.initials: LJ
authors.lastname: Ormsby
authors.affiliation: School of Natural Sciences and Psychology, Liverpool John Moores University, James Parsons Building, Rm653 Byrom Street, Liverpool L3 3AF, UK.
authors.forename: Alex
authors.initials: A
authors.lastname: Piel
authors.affiliation: Iowa State University, Department of Anthropology, 324 Curtiss Hall, Ames, IA 50011, USA.
authors.forename: Jill
authors.initials: J
authors.lastname: Pruetz
authors.affiliation: West Chester University, Department of Psychology, 700 S High St., West Chester, PA, 19382 USA.
authors.forename: Aaron
authors.initials: A
authors.lastname: Rundus
authors.affiliation: Wildlife Conservation Society (WCS), 2300 Southern Boulevard. Bronx, New York 10460, USA.|Washington University Saint Louis, Department of Anthropology, One Brookings Drive, St. Louis, MO 63130, USA.
authors.forename: Crickette
authors.initials: C
authors.lastname: Sanz
authors.affiliation: University College London, Department of Anthropology, 14 Taviton Street, London WC1H 0BW, UK.
authors.forename: Volker
authors.initials: V
authors.lastname: Sommer
authors.affiliation: University of Cambridge, Pembroke Street, Cambridge, UK CB2 3QG.
authors.forename: Fiona
authors.initials: F
authors.lastname: Stewart
authors.affiliation: KMDA, Centre for Research and Conservation, Royal Zoological Society of Antwerp, Koningin Astridplein 20-26, B-2018 Antwerp, Belgium.
authors.forename: Nikki
authors.initials: N
authors.lastname: Tagg
authors.affiliation: Wildlife Conservation Society (WCS), 2300 Southern Boulevard. Bronx, New York 10460, USA.
authors.forename: Hilde
authors.initials: H
authors.lastname: Vanleeuwe
authors.affiliation: Wild Chimpanzee Foundation (WCF), Deutscher Platz 6, 04103 Leipzig, Germany.
authors.forename: Virginie
authors.initials: V
authors.lastname: Vergnes
authors.affiliation: KMDA, Centre for Research and Conservation, Royal Zoological Society of Antwerp, Koningin Astridplein 20-26, B-2018 Antwerp, Belgium.
authors.forename: Jacob
authors.initials: J
authors.lastname: Willie
authors.affiliation: Max Planck Institute for Evolutionary Anthropology (MPI EVAN), Deutscher Platz 6, 04103 Leipzig, Germany.|Taï Chimpanzee Project, Centre Suisse de Recherches Scientifiques, BP 1301, Abidjan 01, Côte d'Ivoire.
authors.forename: Roman M
authors.initials: RM
authors.lastname: Wittig
authors.affiliation: Université de Neuchâtel, Institut de Biologie, Rue Emile-Argand 11, 2000 Neuchâtel, Switzerland.
authors.forename: Klaus
authors.initials: K
authors.lastname: Zuberbuehler
authors.affiliation: Max Planck Institute for Evolutionary Anthropology (MPI EVAN), Deutscher Platz 6, 04103 Leipzig, Germany.|Wild Chimpanzee Foundation (WCF), Deutscher Platz 6, 04103 Leipzig, Germany.
authors.forename: Christophe
authors.initials: C
authors.lastname: Boesch
mesh_terms: D000354:Africa, Western; D000818:Animals; D001522:Behavior, Animal; D005843:Geography; D002679:Pan troglodytes