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In this setting, ochre use is considered a multifaceted proxy for modern behaviour, allowing inferences about technical and symbolic aspects of Homo sapiens. Powdered ochre is reported in multiple southern African ethnographic and archaeological contexts alongside various organic materials, some of which are protein-based. Yet, the recovery of proteinaceous residues is rare in the MSA. Since proteins can offer insights into their tissue and species of origin, identifying proteins in association with ochre may help to infer its function. At Klipdrift Shelter (South Africa), the 60\u2005000-year-old Howiesons Poort layers contain a laterally continuous, ochre-rich sediment surface, possibly over 1\u2005m long, 40\u2005cm wide, and up to 1\u2005cm thick. We used palaeoproteomics to identify proteinaceous traces from this feature. Our analysis identified type I collagen confidently matching common eland (Taurotragus oryx) in two samples. Together with the feature's highly complex nature, these findings suggest the deliberate mixing of eland-derived proteins with ochre powder. Our results confirm that palaeoproteomics can be a reliable tool for reconstructing the protein component of the early human material culture.<\/jats:p>","DOI":"10.1098\/rspb.2025.3300","type":"journal-article","created":{"date-parts":[[2026,10,21]],"date-time":"2026-10-21T00:18:16Z","timestamp":1792541896000},"source":"Crossref","is-referenced-by-count":1,"title":["Palaeoproteomic investigation of archaeological sediments associated with ochre use in the Middle Stone Age of southern Africa"],"prefix":"10.1098","volume":"293","author":[{"ORCID":"https:\/\/orcid.org\/0009-0005-7679-3929","authenticated-orcid":false,"given":"Zahra","family":"Haghighi","sequence":"first","affiliation":[{"name":"SFF Centre for Early Sapiens Behaviour (SapienCE), University of Bergen 1 , 5007 Bergen, Norway"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-5104-5223","authenticated-orcid":false,"given":"Takumi","family":"Tsutaya","sequence":"additional","affiliation":[{"name":"Research Center for Integrative Evolutionary Science, The Graduate University for Advanced Studies (SOKENDAI) 2 , Hayama, Kanagawa 240-0193, Japan"},{"name":"Institute for Advanced Study, Kyushu University 3 , Fukuoka 819-0395, Japan"},{"name":"Globe Institute, University of Copenhagen 4 , 1350 Copenhagen K, Denmark"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Elizabeth C.","family":"Velliky","sequence":"additional","affiliation":[{"name":"SFF Centre for Early Sapiens Behaviour (SapienCE), University of Bergen 1 , 5007 Bergen, Norway"},{"name":"Interdisciplinary Center for Archaeology and Evolution of Human Behaviour (ICArEHB), University of Algarve 5 , 8005-139 Faro, Portugal"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Simon","family":"Armitage","sequence":"additional","affiliation":[{"name":"SFF Centre for Early Sapiens Behaviour (SapienCE), University of Bergen 1 , 5007 Bergen, Norway"},{"name":"Department of Geography, Royal Holloway, University of London 6 , Egham, Surrey TW20 0EX, UK"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Christopher","family":"Miller","sequence":"additional","affiliation":[{"name":"SFF Centre for Early Sapiens Behaviour (SapienCE), University of Bergen 1 , 5007 Bergen, Norway"},{"name":"Institute for Archaeological Sciences and Senckenberg Centre for Human Evolution and Palaeoenvironment, University of T\u00fcbingen 7 , 72074 T\u00fcbingen, Germany"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-0763-7592","authenticated-orcid":false,"given":"Meaghan","family":"Mackie","sequence":"additional","affiliation":[{"name":"Globe Institute, University of Copenhagen 4 , 1350 Copenhagen K, Denmark"},{"name":"Novo Nordisk Foundation Center for Protein Research, University of Copenhagen 8 , 2200 Copenhagen, Denmark"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-2422-3079","authenticated-orcid":false,"given":"Francesco","family":"d'Errico","sequence":"additional","affiliation":[{"name":"SFF Centre for Early Sapiens Behaviour (SapienCE), University of Bergen 1 , 5007 Bergen, Norway"},{"name":"CNRS UMR5199 PACEA, Universit\u00e9 de Bordeaux 9 , 33615 Pessac Cedex, France"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Gaudry","family":"Troche","sequence":"additional","affiliation":[{"name":"Novo Nordisk Foundation Center for Protein Research, University of Copenhagen 8 , 2200 Copenhagen, Denmark"},{"name":"CNRS UMR5199 PACEA, Universit\u00e9 de Bordeaux 9 , 33615 Pessac Cedex, France"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Jesper","family":"Olsen","sequence":"additional","affiliation":[{"name":"Novo Nordisk Foundation Center for Protein Research, University of Copenhagen 8 , 2200 Copenhagen, Denmark"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Karen","family":"van Niekerk","sequence":"additional","affiliation":[{"name":"SFF Centre for Early Sapiens Behaviour (SapienCE), University of Bergen 1 , 5007 Bergen, Norway"},{"name":"School of Geography, Archaeology and Environmental Studies, University of the Witwatersrand 10 , Johannesburg 2000, South Africa"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Christopher","family":"Henshilwood","sequence":"additional","affiliation":[{"name":"SFF Centre for Early Sapiens Behaviour (SapienCE), University of Bergen 1 , 5007 Bergen, Norway"},{"name":"Evolutionary Studies Institute, University of the Witwatersrand 11 , Johannesburg 2000, South 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PhD thesis, University of London."},{"key":"2026102020180475900_RSPB20253300C21","doi-asserted-by":"crossref","first-page":"e0131273","DOI":"10.1371\/journal.pone.0131273","article-title":"A milk and ochre paint mixture used 49,000 years ago at Sibudu, South Africa","volume":"10","author":"Villa","year":"2015","journal-title":"PLoS One"},{"key":"2026102020180475900_RSPB20253300C22","doi-asserted-by":"crossref","first-page":"392","DOI":"10.1016\/j.jhevol.2010.07.006","article-title":"The pigments from Pinnacle Point Cave 13B, Western Cape, South Africa","volume":"59","author":"Watts","year":"2010","journal-title":"J. Hum. Evol."},{"key":"2026102020180475900_RSPB20253300C23","doi-asserted-by":"crossref","first-page":"587","DOI":"10.1016\/j.jhevol.2005.06.007","article-title":"Putting ochre to the test: replication studies of adhesives that may have been used for hafting tools in the Middle Stone Age","volume":"49","author":"Wadley","year":"2005","journal-title":"J. Hum. 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Field Archaeol."},{"key":"2026102020180475900_RSPB20253300C36","doi-asserted-by":"crossref","first-page":"e0136090","DOI":"10.1371\/journal.pone.0136090","article-title":"Evaluating the photoprotective effects of ochre on human skin by in vivo SPF assessment: implications for human evolution, adaptation and dispersal","volume":"10","author":"Rifkin","year":"2015","journal-title":"PLoS One"},{"key":"2026102020180475900_RSPB20253300C37","doi-asserted-by":"crossref","first-page":"674","DOI":"10.1086\/203205","article-title":"Ochre as medicine: a suggestion for the interpretation of the archaeological record","volume":"25","author":"Velo","year":"1984","journal-title":"Curr. Anthropol."},{"key":"2026102020180475900_RSPB20253300C38","doi-asserted-by":"crossref","first-page":"2397","DOI":"10.1016\/j.jas.2010.04.012","article-title":"Cemented ash as a receptacle or work surface for ochre powder production at Sibudu, South Africa, 58,000 years ago","volume":"37","author":"Wadley","year":"2010","journal-title":"J. Archaeol. Sci."},{"key":"2026102020180475900_RSPB20253300C39","doi-asserted-by":"crossref","first-page":"38","DOI":"10.1007\/s12520-025-02376-3","article-title":"Ochre and manganese pigments from the Middle Stone Age layers at Klipdrift Shelter","volume":"18","author":"Velliky","year":"2026","journal-title":"Archaeol. Anthropol. Sci."},{"key":"2026102020180475900_RSPB20253300C40","doi-asserted-by":"crossref","first-page":"e0310741","DOI":"10.1371\/journal.pone.0310741","article-title":"Reconcilable differences: using retrospective photogrammetry to bridge the divide between analogue and digital site data collected during long-term excavation projects","volume":"19","author":"Unhammer","year":"2024","journal-title":"PLoS One"},{"key":"2026102020180475900_RSPB20253300C41","first-page":"519","article-title":"Long-distance transport of red ocher by Clovis foragers","volume":"25","author":"Zarzycka","year":"2019","journal-title":"J. Archaeol. Sci. Rep."},{"key":"2026102020180475900_RSPB20253300C42","volume-title":"Die Gei\u00dfenkl\u00f6sterle-H\u00f6hle im Achtal bei Blaubeuren I. Fundhorizontbildung und Besiedlung im Mittelpal\u00e4olithikum und im Aurignacien [The Gei\u00dfenkl\u00f6sterle Cave in the Ach Valley near Blaubeuren I: site formation and occupation in the Middle Palaeolithic and Aurignacian]","author":"Hahn","year":"1988"},{"key":"2026102020180475900_RSPB20253300C43","unstructured":"Dippon\n              G\n            \n          . 2003Die taphonomie der Aurignacienhorizonte der Gei\u00dfenkl\u00f6sterle-H\u00f6hle bei Blaubeuren [The taphonomy of the Aurignacian occupation horizons at Gei\u00dfenkl\u00f6sterle Cave, Blaubeuren]. Master's thesis, University of T\u00fcbingen. [In German.]"},{"key":"2026102020180475900_RSPB20253300C44","doi-asserted-by":"crossref","first-page":"84","DOI":"10.1016\/j.jas.2015.03.033","article-title":"Analysis of the red ochre of the El Mir\u00f3n burial (Ramales de la Victoria, Cantabria, Spain)","volume":"60","author":"Rom\u00e1n","year":"2015","journal-title":"J. Archaeol. 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Int."},{"key":"2026102020180475900_RSPB20253300C57","doi-asserted-by":"crossref","first-page":"13208","DOI":"10.1073\/pnas.1202629109","article-title":"Border Cave and the beginning of the Later Stone Age in South Africa","volume":"109","author":"Villa","year":"2012","journal-title":"Proc. Natl Acad. Sci. USA"},{"key":"2026102020180475900_RSPB20253300C58","first-page":"172","article-title":"The southern African Stone Age sequence updated (II)","volume":"77","author":"Lombard","year":"2022","journal-title":"S. Afr. Archaeol. Bull."},{"key":"2026102020180475900_RSPB20253300C59","doi-asserted-by":"crossref","first-page":"155","DOI":"10.1080\/0067270X.2015.1039236","article-title":"Those marvellous millennia: the Middle Stone Age of Southern Africa","volume":"50","author":"Wadley","year":"2015","journal-title":"Azania"},{"key":"2026102020180475900_RSPB20253300C60","doi-asserted-by":"crossref","first-page":"411","DOI":"10.1007\/s10437-016-9232-0","article-title":"Earliest evidence for the ivory trade in southern Africa: isotopic and ZooMS analysis of seventeenth-century AD ivory from KwaZulu-Natal","volume":"33","author":"Coutu","year":"2016","journal-title":"Afr. Archaeol. Rev."},{"key":"2026102020180475900_RSPB20253300C61","doi-asserted-by":"crossref","first-page":"2419","DOI":"10.1007\/s12520-018-0688-5","article-title":"Identifying the animal species used to manufacture bone arrowheads in South Africa","volume":"11","author":"Bradfield","year":"2019","journal-title":"Archaeol. Anthropol. Sci."},{"key":"2026102020180475900_RSPB20253300C62","doi-asserted-by":"crossref","first-page":"14427","DOI":"10.1038\/s41598-020-71374-3","article-title":"Palaeoproteomics gives new insight into early southern African pastoralism","volume":"10","author":"Le Meillour","year":"2020","journal-title":"Scient. Rep."},{"key":"2026102020180475900_RSPB20253300C63","doi-asserted-by":"crossref","first-page":"231002","DOI":"10.1098\/rsos.231002","article-title":"The name of the game: palaeoproteomics and radiocarbon dates further refine the presence and dispersal of caprines in eastern and southern Africa","volume":"10","author":"Le Meillour","year":"2023","journal-title":"R. Soc. Open Sci."},{"key":"2026102020180475900_RSPB20253300C64","doi-asserted-by":"crossref","first-page":"108380","DOI":"10.1016\/j.quascirev.2023.108380","article-title":"Peptide mass fingerprinting as a tool to assess micromammal biodiversity in Pleistocene South Africa: the case of Klipdrift Shelter","volume":"322","author":"Nel","year":"2023","journal-title":"Quat. Sci. Rev."},{"key":"2026102020180475900_RSPB20253300C65","doi-asserted-by":"crossref","first-page":"2","DOI":"10.1016\/j.quaint.2015.07.041","article-title":"Subsistence strategies in the southern Cape during the Howiesons Poort: taphonomic and zooarchaeological analyses of Klipdrift Shelter, South Africa","volume":"404","author":"Reynard","year":"2016","journal-title":"Quat. Int."},{"key":"2026102020180475900_RSPB20253300C66","doi-asserted-by":"crossref","first-page":"284","DOI":"10.1016\/j.jas.2014.01.033","article-title":"Klipdrift Shelter, southern Cape, South Africa: preliminary report on the Howiesons Poort layers","volume":"45","author":"Henshilwood","year":"2014","journal-title":"J. Archaeol. Sci."},{"key":"2026102020180475900_RSPB20253300C67","doi-asserted-by":"crossref","first-page":"25","DOI":"10.1007\/s10533-007-9102-6","article-title":"Role of proteins in soil carbon and nitrogen storage: controls on persistence","volume":"85","author":"Rillig","year":"2007","journal-title":"Biogeochemistry"},{"key":"2026102020180475900_RSPB20253300C68","doi-asserted-by":"crossref","first-page":"293","DOI":"10.1016\/B978-044481516-3\/50008-6","volume-title":"Humic substances in terrestrial ecosystems","author":"Nannipieri","year":"1996"},{"key":"2026102020180475900_RSPB20253300C69","doi-asserted-by":"crossref","first-page":"2156","DOI":"10.1016\/j.soilbio.2008.05.001","article-title":"Comparison of extraction methods for recovery of extracellular \u03b2-glucosidase in two different forest soils","volume":"40","author":"Masciandaro","year":"2008","journal-title":"Soil Biol. Biochem."},{"key":"2026102020180475900_RSPB20253300C70","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1007\/s003740050335","article-title":"The chemistry of soil organic nitrogen: a review","volume":"26","author":"Schulten","year":"1997","journal-title":"Biol. Fertil. Soils"},{"key":"2026102020180475900_RSPB20253300C71","doi-asserted-by":"crossref","first-page":"12841","DOI":"10.1038\/s41598-019-49183-0","article-title":"Palaeoproteomic identification of breast milk protein residues from the archaeological skeletal remains of a neonatal dog","volume":"9","author":"Tsutaya","year":"2019","journal-title":"Scient. Rep."},{"key":"2026102020180475900_RSPB20253300C72","doi-asserted-by":"crossref","first-page":"103","DOI":"10.1038\/s41586-019-1555-y","article-title":"Early Pleistocene enamel proteome from Dmanisi resolves Stephanorhinus phylogeny","volume":"574","author":"Cappellini","year":"2019","journal-title":"Nature"},{"key":"2026102020180475900_RSPB20253300C73","doi-asserted-by":"crossref","first-page":"2085","DOI":"10.1038\/s41596-024-00975-3","article-title":"Deep-time phylogenetic inference by paleoproteomic analysis of dental enamel","volume":"19","author":"Taurozzi","year":"2024","journal-title":"Nat. Protoc."},{"key":"2026102020180475900_RSPB20253300C74","doi-asserted-by":"crossref","first-page":"1896","DOI":"10.1038\/nprot.2007.261","article-title":"Protocol for micro-purification, enrichment, pre-fractionation and storage of peptides for proteomics using StageTips","volume":"2","author":"Rappsilber","year":"2007","journal-title":"Nat. Protoc."},{"key":"2026102020180475900_RSPB20253300C75","doi-asserted-by":"crossref","first-page":"e0251061","DOI":"10.1371\/journal.pone.0251061","article-title":"Distinguishing African bovids using zooarchaeology by mass spectrometry (ZooMS): new peptide markers and insights into Iron Age economies in Zambia","volume":"16","author":"Janzen","year":"2021","journal-title":"PLoS One"},{"key":"2026102020180475900_RSPB20253300C76","volume-title":"R: a language and environment for statistical computing","author":"R Core Team","year":"2023"},{"key":"2026102020180475900_RSPB20253300C77","volume-title":"rDeamidation (v. 1.0.1)","author":"Tsutaya","year":"2025"},{"key":"2026102020180475900_RSPB20253300C78","doi-asserted-by":"crossref","first-page":"7369","DOI":"10.1002\/anie.201713020","article-title":"Palaeoproteomic profiling of conservation layers on a 14th century Italian wall painting","volume":"57","author":"Mackie","year":"2018","journal-title":"Angew. Chem. Int. Edn Engl."},{"key":"2026102020180475900_RSPB20253300C79","doi-asserted-by":"crossref","first-page":"D543","DOI":"10.1093\/nar\/gkab1038","article-title":"The PRIDE database resources in 2022: a hub for mass spectrometry-based proteomics evidences","volume":"50","author":"Perez-Riverol","year":"2022","journal-title":"Nucleic. Acids. Res."},{"key":"2026102020180475900_RSPB20253300C80","doi-asserted-by":"crossref","first-page":"2056","DOI":"10.1021\/ac1027275","article-title":"Deamidation at asparagine and glutamine as a major modification upon deterioration\/aging of proteinaceous binders in mural paintings","volume":"83","author":"Leo","year":"2011","journal-title":"Anal. Chem."},{"key":"2026102020180475900_RSPB20253300C81","doi-asserted-by":"crossref","first-page":"251","DOI":"10.1002\/rcm.7445","article-title":"Glutamine deamidation: an indicator of antiquity, or preservational quality?","volume":"30","author":"Schroeter","year":"2016","journal-title":"Rapid Commun. Mass Spectrom."},{"key":"2026102020180475900_RSPB20253300C82","doi-asserted-by":"crossref","first-page":"2319","DOI":"10.1002\/rcm.6351","article-title":"Site-specific deamidation of glutamine: a new marker of bone collagen deterioration: site-specific deamidation of glutamine","volume":"26","author":"van Doorn","year":"2012","journal-title":"Rapid Commun. Mass Spectrom."},{"key":"2026102020180475900_RSPB20253300C83","doi-asserted-by":"crossref","first-page":"105454","DOI":"10.1016\/j.jas.2021.105454","article-title":"Examining collagen preservation through glutamine deamidation at Denisova Cave","volume":"133","author":"Brown","year":"2021","journal-title":"J. Archaeol. Sci."},{"key":"2026102020180475900_RSPB20253300C84","doi-asserted-by":"crossref","first-page":"29","DOI":"10.1016\/j.jas.2016.02.002","article-title":"The effects of demineralization and sampling point variability on the measurement of glutamine deamidation in type I collagen extracted from bone","volume":"69","author":"Simpson","year":"2016","journal-title":"J. Archaeol. Sci."},{"key":"2026102020180475900_RSPB20253300C85","doi-asserted-by":"crossref","first-page":"15678","DOI":"10.1038\/s41598-019-51680-1","article-title":"Mechanisms of soft tissue and protein preservation in Tyrannosaurus rex","volume":"9","author":"Boatman","year":"2019","journal-title":"Scient. 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