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Caprine dairy exploitation on the Iranian Plateau from the seventh millennium BC

Nature Human Behaviour (2026)Cite this article

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Abstract

Animal domestication and development of pastoralism in southwest Asia revolutionized human subsistence strategies. Various centres of ruminant domestication and diffusion routes of agropastoralism have been identified. The area between the northern and central Zagros Mountains on the Iranian Plateau is a cradle for goat domestication and eastward spread of agropastoralism. However, the early exploitation of ruminant milk by pastoral communities in the Zagros remains insufficiently studied. Here we show residues of caprine dairy products that were detected from the analysis of lipid residues in pottery vessels and protein residues in human dental calculus. These results, combined with the faunal spectra and radiocarbon analyses directly on the dairy residues, show that sheep and goat dairy products were widely exploited in the Zagros from the seventh millennium BC. This pattern parallels the contemporaneous exploitation of cattle milk in Anatolia. Neolithic communities in both regions reveal similarly complex dynamics of early ruminant milk use, marking the emergence of independent yet synchronous trajectories in the diffusion of agropastoral lifeways.

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Fig. 1: Localities of the studied sites plotted on a map with relative abundance of faunal remains and lipid residue sources.
Fig. 2: Degraded animal fats dominate the lipid profiles in the pottery vessels from the Iranian Plateau.
Fig. 3: Ruminant dairy and adipose products were exploited regardless of settlement type and age.
Fig. 4: Non-supervised clustering performed to generate principal component analyses and protein and peptide heat maps using Perseus73.
Fig. 5: The chronological model identifies the beginning of ruminant dairy exploitation in the seventh millennium BC.

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Data availability

Reusable processed data (blank corrected, calibrated and so on) are accessible in the main manuscript, Extended Data and Supplementary Information files. Proteomic raw and results files are deposited in Massive (https://massive.ucsd.edu/ProteoSAFe/private-dataset.jsp?task=cf70550391664926ae7b8068854ea6b0). For lipids, raw files are available on request from the corresponding author.

Code availability

The modelling code used in OxCal is fully accessible in Extended Data Table 10. There are no restrictions for reuse, other than citation of the current paper.

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Acknowledgements

The work was carried out under a Marie Skłodowska-Curie individual fellowship, project Vargah (grant no. MSCA-IF-2020, 101024503), awarded to E.C. We thank the ATM (Transversal Actions of the MNHN) of the Museum national d’Histoire naturelle in Paris for providing additional funds for analyses through the projects SIALK (grant no. ATM 2022 HE-7209) and MAHI (grant no. ATM 2023 HE-7209) awarded to E.C. and co-funding with the UMR7209 a Master’s project for A.B.-S. The National Museum of Iran, M. Hesari and R. Shirazi from the Iranian Centre for Archaeological Research, B. Omrani and M. Amiri from Research Institute of Cultural Heritage and Tourism of Iran, Y. Miyake from the University of Tsukuba and H. McDonald from the University of Chicago are warmly thanked for the authorization and access to the collection of materials studied. We thank the team SAPOA of the UMR7209 for supporting the acquisition of laboratory material. The SSMIM facility, M. Balasse, E. Dufour, L. Rémusat, N. Khodrova and M. Lecasble are thanked for the access to gas chromatography–flame ionization detector and gas chromatography–combustion–isotope ratio mass spectrometry analysis instruments. GCMS and ECHoMICADAS instruments of the PANOPLY platform at the LSCE were used for this project. NanoLC-MS/MS equipment of the SMBP facility was subsidized by the Conseil Régional d’Île-de-France (Sesame no. 10022268).

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Authors and Affiliations

  1. Bioarchéologies Interactions Sociétés Environnements, UMR7209 CNRS, Muséum national d’Histoire naturelle, Paris, France

    Emmanuelle Casanova, Antoine Zazzo, Audrey Boco-Simon & Marjan Mashkour

  2. Laboratoire des Sciences du Climat et de l’Environnement, UMR8212 CEA CNRS UVSQ, Université Paris-Saclay, Gif-sur-Yvette, France

    Emmanuelle Casanova, Jérémy Jacob, Christine Hatté, François Thil & Caroline Gauthier

  3. Bioarchaeology Laboratory, Central Laboratory, University of Tehran, Tehran, Iran

    Hossein Davoudi, Homa Fathi & Marjan Mashkour

  4. National Museum of Iran, Tehran, Iran

    Hossein Davoudi, Jebrael Nokandeh, Fereidoun Biglari & Omolbanin Ghafoori

  5. Molécules de Communication et Adaptation des Micro-organismes, UMR7245CNRS, Muséum national d’Histoire naturelle, Paris, France

    Séverine Zirah

  6. Silesian University of Technology, Gliwice, Poland

    Christine Hatté

  7. Research Institute of Cultural Heritage and Tourism Organization of Iran, Tehran, Iran

    Jebrael Nokandeh

  8. Institute for the Study of Ancient Cultures, The University of Chicago, Chicago, IL, USA

    Abbas Alizadeh

  9. Institute for Near Eastern Archaeology, Free University of Berlin, Berlin, Germany

    Reinhard Bernbeck & Susan Pollock

  10. Faculty of Humanities and Social Sciences, University of Tsukuba, Tsukuba, Japan

    Akira Tsuneki

  11. Departement of Anthropology, Yale University, New Haven, CT, USA

    Frank Hole

  12. Biological Mass Spectrometry and Proteomics SMBP, ESPCI Paris, LPC CNRS UAR2051, PSL University, Paris, France

    Emmanuelle Demey-Thomas & Joëlle Vinh

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  1. Emmanuelle Casanova
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Contributions

E.C. and M.M. designed the study and wrote the paper. Analyses and data processing were performed by M.M. and H.D. for the faunal analyses; E.C., A.B.-.S., C.G. and J.J. undertook the lipid analyses, E.C., S.Z., E.D.-T. and J.V. the proteomic analyses and E.C., A.Z., C.H. and F.T. the radiocarbon analyses. H.D., M.M., J.N., A.A., F.B., R.B., S.P., A.T., O.G., H.F. and F.H. directed excavations and/or curated and selected materials for analyses. All authors read and approved the manuscript.

Corresponding authors

Correspondence to Emmanuelle Casanova or Marjan Mashkour.

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Extended data

Extended Data Table 1 Summary of studied sites and material for protein residues analysis in human dental calculus and lipid residue analysis in pottery vessels
Full size table
Extended Data Table 2 Stable carbon isotopes on individual fatty acid measured on lipids extracted from modern references of wild caprine bones and dairy products from the Iranian
Full size table
Extended Data Table 3 List of studied pottery which revealed animal fats residues, their total lipids concentrations and source assignment for the permanent settlements 7-6th Millennium BC (full details in Supplementary Information 2.1, Tables S2.3, S2.4 and S2.7)
Full size table
Extended Data Table 4 List of studied pottery which revealed animal fats residues, their total lipids concentrations and source assignment for the permanent settlements 6-4th Millennium BC, including already published data from Qazvin plain42 (full details in Supplementary Information 2.1, Tables S2.1, S2.2 and S2.8)
Full size table
Extended Data Table 5 List of studied pottery which revealed animal fats residues, their total lipids concentrations and source assignment for the open-air campsites 7-6th Millennium BC. (Full details in Supplementary Information 2.1, Tables S2.6 and S2.8)
Full size table
Extended Data Table 6 List of studied pottery which revealed animal fats residues, their total lipids concentrations and source assignment for the cave campsites 6-4th Millennium BC (Full details in Supplementary Information 2.1, Tables S2.5 and S2.10)
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Extended Data Table 7 List and description of studied human teeth from Abdul Hosein5 and Sang-e Chakhmaq53 for dental calculus analysis
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Extended Data Table 8 List of studied human teeth from Abdul Hosein (AHO) and Sang-e Chakhmaq (TSC) for dental calculus analysis, proteins, peptide chain, and taxonomy
Full size table
Extended Data Table 9 Component 1 and component 2 major factors of ACP presented in Fig. 4 (A and B)
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Extended Data Table 10 Chronological modelling code performed in OxCal V4.4.4
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Supplementary information

Supplementary Information

Supplementary Information 1. Sites, chronology, faunal remains and material description. Supplementary Information 2. Analyses parameters and data acquired. Supplementary Information 3. Supplementary References.

Reporting Summary

Supplementary Table 1

PCA analysis results.

Supplementary Table 2

Blast analysis results.

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Casanova, E., Davoudi, H., Zazzo, A. et al. Caprine dairy exploitation on the Iranian Plateau from the seventh millennium BC. Nat Hum Behav (2026). https://doi.org/10.1038/s41562-025-02396-y

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