Sulfur isotopes evidence spatial ecology of Late Pleistocene ungulates in southwestern France

Sarah Barakat; Elodie-Laure Jimenez; Raija Katarina Heikkilä; Aurélien Royer; Jamie Hodgkins; Laura Niven; Marie-Cécile Soulier; Susan Lagle; Christelle Dancette; Teresa E. Steele; Jean-Christophe Castel; Shannon McPherron; Jean-Jacques Hublin; Karen Ruebens; Emmanuel Discamps; Kate Britton

doi:10.1016/j.quascirev.2025.109744

Sulfur isotopes evidence spatial ecology of Late Pleistocene ungulates in southwestern France

Sarah Barakat^* (Corresponding Author)
, Elodie-Laure Jimenez
, Raija Katarina Heikkilä
, Aurélien Royer
, Jamie Hodgkins
, Laura Niven
, Marie-Cécile Soulier
, Susan Lagle
, Christelle Dancette
, Teresa E. Steele
, Jean-Christophe Castel
, Shannon McPherron
, Jean-Jacques Hublin
, Karen Ruebens
, Emmanuel Discamps^* (Corresponding Author)
, Kate Britton^* (Corresponding Author)

^*Corresponding author for this work

Archaeology

Research output: Contribution to journal › Article › peer-review

Abstract

Understanding prey species spatial behaviour is key to unravelling contemporary hunter-gatherer subsistence and movement patterns. Here, we use sulfur (δ34S), carbon (δ 13C) and nitrogen (δ 15N) isotope compositions of bone collagen extracted from Rangifer, Equus, Bovinae, Capreolus, and Cervus (n = 244) to explore isotope spatial and dietary niches of several key hunter-gatherer prey species from three Late Pleistocene sites in southwestern France between MIS 5 and MIS 3. Alongside morphological identification, Zooarchaeology by Mass Spectrometry (ZooMS) was used to confirm deer taxa (n = 125) allowing for a better interpretation of the isotopic data. δ34S analysis identifies differences in ranging sizes between ungulates with known large ranging sizes from those with small to medium size ranges. Rangifer, who, today, have large home ranges, have the largest range of δ34S values and lowest δ34S values with low to no overlap with other ungulate species through time. This indicates that Rangifer had a larger range size with a distinct isotopic area that is different to other ungulates, potentially farther inland and farther north. Bovines and Equus have largely overlapping δ34S values that are similar to the modern local δ34S value of the sites, and Cervus and Capreolus have δ34S values that overlap but are slightly higher than the modern δ34S values, potentially indicating a ranging area to the west of the sites. These results reveal that Neanderthals hunted three different groups of game who occupied different areas of the landscape.

Original language	English
Article number	109744
Number of pages	21
Journal	Quaternary Science Reviews
Volume	374
Early online date	17 Dec 2025
DOIs	https://doi.org/10.1016/j.quascirev.2025.109744
Publication status	E-pub ahead of print - 17 Dec 2025

Bibliographical note

Open Access via the Elsevier agreement

Sampling permits were given to E.D. and E.L.J. by the Conseil Départemental de Dordogne (Roc de Marsal), and the Musée National de la Préhistoire des Eyzies (Roc de Marsal, Combe Grenal, Pech de l’Azé IV). We would like to thank Pauline Raymond at the Collège de France for her help with ZooMS sample preparation, and Emmanuelle Demey-Thomas and Joelle Vinh at the ESPCI Paris for their assistance with the MALDI-TOF mass spectrometry, Orsolya Czére (Aberdeen) for assistance with collagen extractions, and the staff from the Musée National de la Préhistoire for their help during initial sampling of faunal collections. We would like to thank Mael Le Corre for his assistance with statistical analysis and model building. We would also like to thank Clément Bataille for his random forest R code to help rebuild a cropped version of the modern sulfur isoscape.

Data Availability Statement

The isotopic and ZooMS data are available in Supplementary Tables 1 and 2 The ZooMS spectra are available in mzxml format on Zenodo: 10.5281/zenodo.17738202. Access to codes to build the figures can be made available upon request.

Funding

Funding for isotopic and proteomic analysis was funded by PleistoHERD (ref: RPG-2017-410), Leverhulme Prize (ref: PLP-2019-284), DeerPal (ANR-18-CE03-0007), and the University of Aberdeen. Zooarchaeological morphological analysis at Pech de l’Azé IV was funded in by the Max Planck Society and the Archaeological Research Centre and Museum for Human Behavioural Evolution (Neuwied, Germany). We thank UKRI (NERC QUADRAT DTP studentship ref: NE/S007377/1 to SB; ERC-selected/EPSRC-funded project ref: EP/Y023641/1 to KB) for salary support during production of this manuscript.

Funders	Funder number
The Leverhulme Trust	PLP-2019-284, RPG-2017-410
UK Research and Innovation	NE/S007377/1, EP/Y023641/1
Agence nationale de la recherche	ANR-18-CE03-0007

Keywords

Multi-isotope analysis
ZooMS
Palaeoecology
Biogeography
Neanderthal
Home range
Western Europe

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Barakat_etal_QSR_Sulfur_Isotopes_Evidence_VoR
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Cite this

Barakat, S., Jimenez, E.-L., Heikkilä, R. K., Royer, A., Hodgkins, J., Niven, L., Soulier, M.-C., Lagle, S., Dancette, C., Steele, T. E., Castel, J.-C., McPherron, S., Hublin, J.-J., Ruebens, K., Discamps, E., & Britton, K. (2026). Sulfur isotopes evidence spatial ecology of Late Pleistocene ungulates in southwestern France. Quaternary Science Reviews, 374, Article 109744. Advance online publication. https://doi.org/10.1016/j.quascirev.2025.109744

Barakat, S, Jimenez, E-L, Heikkilä, RK, Royer, A, Hodgkins, J, Niven, L, Soulier, M-C, Lagle, S, Dancette, C, Steele, TE, Castel, J-C, McPherron, S, Hublin, J-J, Ruebens, K, Discamps, E & Britton, K 2026, 'Sulfur isotopes evidence spatial ecology of Late Pleistocene ungulates in southwestern France', Quaternary Science Reviews, vol. 374, 109744. https://doi.org/10.1016/j.quascirev.2025.109744

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title = "Sulfur isotopes evidence spatial ecology of Late Pleistocene ungulates in southwestern France",

abstract = "Understanding prey species spatial behaviour is key to unravelling contemporary hunter-gatherer subsistence and movement patterns. Here, we use sulfur (δ34S), carbon (δ 13C) and nitrogen (δ 15N) isotope compositions of bone collagen extracted from Rangifer, Equus, Bovinae, Capreolus, and Cervus (n = 244) to explore isotope spatial and dietary niches of several key hunter-gatherer prey species from three Late Pleistocene sites in southwestern France between MIS 5 and MIS 3. Alongside morphological identification, Zooarchaeology by Mass Spectrometry (ZooMS) was used to confirm deer taxa (n = 125) allowing for a better interpretation of the isotopic data. δ34S analysis identifies differences in ranging sizes between ungulates with known large ranging sizes from those with small to medium size ranges. Rangifer, who, today, have large home ranges, have the largest range of δ34S values and lowest δ34S values with low to no overlap with other ungulate species through time. This indicates that Rangifer had a larger range size with a distinct isotopic area that is different to other ungulates, potentially farther inland and farther north. Bovines and Equus have largely overlapping δ34S values that are similar to the modern local δ34S value of the sites, and Cervus and Capreolus have δ34S values that overlap but are slightly higher than the modern δ34S values, potentially indicating a ranging area to the west of the sites. These results reveal that Neanderthals hunted three different groups of game who occupied different areas of the landscape.",

keywords = "Multi-isotope analysis, ZooMS, Palaeoecology, Biogeography, Neanderthal, Home range, Western Europe",

author = "Sarah Barakat and Elodie-Laure Jimenez and Heikkil{\"a}, \{Raija Katarina\} and Aur{\'e}lien Royer and Jamie Hodgkins and Laura Niven and Marie-C{\'e}cile Soulier and Susan Lagle and Christelle Dancette and Steele, \{Teresa E.\} and Jean-Christophe Castel and Shannon McPherron and Jean-Jacques Hublin and Karen Ruebens and Emmanuel Discamps and Kate Britton",

note = "Open Access via the Elsevier agreement Sampling permits were given to E.D. and E.L.J. by the Conseil D{\'e}partemental de Dordogne (Roc de Marsal), and the Mus{\'e}e National de la Pr{\'e}histoire des Eyzies (Roc de Marsal, Combe Grenal, Pech de l{\textquoteright}Az{\'e} IV). We would like to thank Pauline Raymond at the Coll{\`e}ge de France for her help with ZooMS sample preparation, and Emmanuelle Demey-Thomas and Joelle Vinh at the ESPCI Paris for their assistance with the MALDI-TOF mass spectrometry, Orsolya Cz{\'e}re (Aberdeen) for assistance with collagen extractions, and the staff from the Mus{\'e}e National de la Pr{\'e}histoire for their help during initial sampling of faunal collections. We would like to thank Mael Le Corre for his assistance with statistical analysis and model building. We would also like to thank Cl{\'e}ment Bataille for his random forest R code to help rebuild a cropped version of the modern sulfur isoscape.",

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AU - Barakat, Sarah

AU - Jimenez, Elodie-Laure

AU - Heikkilä, Raija Katarina

AU - Royer, Aurélien

AU - Hodgkins, Jamie

AU - Niven, Laura

AU - Soulier, Marie-Cécile

AU - Lagle, Susan

AU - Dancette, Christelle

AU - Steele, Teresa E.

AU - Castel, Jean-Christophe

AU - McPherron, Shannon

AU - Hublin, Jean-Jacques

AU - Ruebens, Karen

AU - Discamps, Emmanuel

AU - Britton, Kate

N1 - Open Access via the Elsevier agreement Sampling permits were given to E.D. and E.L.J. by the Conseil Départemental de Dordogne (Roc de Marsal), and the Musée National de la Préhistoire des Eyzies (Roc de Marsal, Combe Grenal, Pech de l’Azé IV). We would like to thank Pauline Raymond at the Collège de France for her help with ZooMS sample preparation, and Emmanuelle Demey-Thomas and Joelle Vinh at the ESPCI Paris for their assistance with the MALDI-TOF mass spectrometry, Orsolya Czére (Aberdeen) for assistance with collagen extractions, and the staff from the Musée National de la Préhistoire for their help during initial sampling of faunal collections. We would like to thank Mael Le Corre for his assistance with statistical analysis and model building. We would also like to thank Clément Bataille for his random forest R code to help rebuild a cropped version of the modern sulfur isoscape.

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N2 - Understanding prey species spatial behaviour is key to unravelling contemporary hunter-gatherer subsistence and movement patterns. Here, we use sulfur (δ34S), carbon (δ 13C) and nitrogen (δ 15N) isotope compositions of bone collagen extracted from Rangifer, Equus, Bovinae, Capreolus, and Cervus (n = 244) to explore isotope spatial and dietary niches of several key hunter-gatherer prey species from three Late Pleistocene sites in southwestern France between MIS 5 and MIS 3. Alongside morphological identification, Zooarchaeology by Mass Spectrometry (ZooMS) was used to confirm deer taxa (n = 125) allowing for a better interpretation of the isotopic data. δ34S analysis identifies differences in ranging sizes between ungulates with known large ranging sizes from those with small to medium size ranges. Rangifer, who, today, have large home ranges, have the largest range of δ34S values and lowest δ34S values with low to no overlap with other ungulate species through time. This indicates that Rangifer had a larger range size with a distinct isotopic area that is different to other ungulates, potentially farther inland and farther north. Bovines and Equus have largely overlapping δ34S values that are similar to the modern local δ34S value of the sites, and Cervus and Capreolus have δ34S values that overlap but are slightly higher than the modern δ34S values, potentially indicating a ranging area to the west of the sites. These results reveal that Neanderthals hunted three different groups of game who occupied different areas of the landscape.

AB - Understanding prey species spatial behaviour is key to unravelling contemporary hunter-gatherer subsistence and movement patterns. Here, we use sulfur (δ34S), carbon (δ 13C) and nitrogen (δ 15N) isotope compositions of bone collagen extracted from Rangifer, Equus, Bovinae, Capreolus, and Cervus (n = 244) to explore isotope spatial and dietary niches of several key hunter-gatherer prey species from three Late Pleistocene sites in southwestern France between MIS 5 and MIS 3. Alongside morphological identification, Zooarchaeology by Mass Spectrometry (ZooMS) was used to confirm deer taxa (n = 125) allowing for a better interpretation of the isotopic data. δ34S analysis identifies differences in ranging sizes between ungulates with known large ranging sizes from those with small to medium size ranges. Rangifer, who, today, have large home ranges, have the largest range of δ34S values and lowest δ34S values with low to no overlap with other ungulate species through time. This indicates that Rangifer had a larger range size with a distinct isotopic area that is different to other ungulates, potentially farther inland and farther north. Bovines and Equus have largely overlapping δ34S values that are similar to the modern local δ34S value of the sites, and Cervus and Capreolus have δ34S values that overlap but are slightly higher than the modern δ34S values, potentially indicating a ranging area to the west of the sites. These results reveal that Neanderthals hunted three different groups of game who occupied different areas of the landscape.

KW - Multi-isotope analysis

KW - ZooMS

KW - Palaeoecology

KW - Biogeography

KW - Neanderthal

KW - Home range

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M1 - 109744

ER -

Sulfur isotopes evidence spatial ecology of Late Pleistocene ungulates in southwestern France

Abstract

Bibliographical note

Data Availability Statement

Funding

Keywords

Access to Document

Fingerprint

Cite this