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Evidence for the cooking of fish 780,000 years ago at Gesher Benot Ya’aqov, Israel



Although cooking is regarded as a key element in the evolutionary success of the genus Homo, impacting various biological and social aspects, when intentional cooking first began remains unknown. The early Middle Pleistocene site of Gesher Benot Ya’aqov, Israel (marine isotope stages 18–20; ~0.78 million years ago), has preserved evidence of hearth-related hominin activities and large numbers of freshwater fish remains (>40,000). A taphonomic study and isotopic analyses revealed significant differences between the characteristics of the fish bone assemblages recovered in eight sequential archaeological horizons of Area B (Layer II-6 levels 1–7) and natural fish bone assemblages (identified in Area A). Gesher Benot Ya’aqov archaeological horizons II-6 L1–7 exhibited low fish species richness, with a clear preference for two species of large Cyprinidae (Luciobarbus longiceps and Carasobarbus canis) and the almost total absence of fish bones in contrast to the richness of pharyngeal teeth (>95%). Most of the pharyngeal teeth recovered in archaeological horizons II-6 L1–7 were spatially associated with ‘phantom’ hearths (clusters of burnt flint microartifacts). Size–strain analysis using X-ray powder diffraction provided evidence that these teeth had been exposed to low temperature (<500 °C), suggesting, together with the archaeological and taphonomic data, that the fish from the archaeological horizons of Area B had been cooked and consumed on site. This is the earliest evidence of cooking by hominins.

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Fig. 1: Relative abundance of fish remains in GBY areas A and B by archaeological horizons.
Fig. 2: Characteristics of assemblages of fish remains from Lake Kinneret (natural death) and GBY Area B (archaeological horizons II-6 L1–7).
Fig. 3: XRD and apatite CS analysis of Cyprinidae (L. longiceps and M. piceus) molariform pharyngeal teeth enameloid.
Fig. 4: Ranges, mean values and standard deviations of Cyprinidae teeth enameloid CS for modern heated and unheated teeth and for the GBY teeth retrieved from areas A and B.
Fig. 5: Kernel density maps from archaeological horizons at GBY.
Fig. 6: Oxygen and carbon isotope analyses of extant and ancient cyprinid molariform pharyngeal teeth enameloid.

Data availability

The archaeological and palaeontological materials used in this study are held at the National Natural History Collections of the Hebrew University of Jerusalem, Edmond J. Safra Campus, Givat Ram, Israel; the fish reference collection and the XRD samples are held at the SMNH, Tel Aviv University; and the teeth sampled for the isotope analysis are held at the Institute of Geosciences, Johannes Gutenberg University of Mainz, Germany. All data are available in the main text or the Supplementary Information.


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This study was supported by the Israel Science Foundation (ISF) Center of Excellence for the study of ‘Climate change in the Upper Jordan Valley between ca. 800 Ma and 700 Ma ago—its impact on the environment and hominins and its potential as a prediction for future scenarios’ (grant no. 300/06 and grant no. 858/09); ISF grant ‘The nature, scope and interpretation of the Acheulian variability at GBY’ (grant no. 27/12); Irene Levi Sala CARE Archaeological Foundation (grant nos. 178/09, 5/14 and 206/20); and the Dan David Foundation grant for ‘The search and study of modern humans’. Access to the NHM in London was also supported by the SYNTHESYS+ project, with funding from the European Union’s Horizon 2020 research and innovation programme under grant agreement 823827. Isotope analysis was supported by funding from the European Research Council under the European Union’s Horizon 2020 research and innovation programme (grant no. 681450). The Israel Antiquities Authority provided the permit for the archaeological excavations (G-101/97). This study was performed at the National Natural History Collections of the Hebrew University of Jerusalem, Edmond J. Safra Campus, Givat Ram; at the Institute of Archaeology of the Hebrew University, Mount Scopus, Jerusalem, Israel; at the SMNH, Tel Aviv University, Israel; at the Department of Archaeology, University of Western Australia, Perth, Australia; and at the NHM in Brussels, Belgium. We thank T. Vennemann (University of Lausanne) and M. Maus (University of Mainz) for performing the phosphate oxygen isotope analysis of the silver phosphate samples. Micro-CT analyses were performed at the Dan David Center for Human Evolution and Biohistory Research and the Shmunis Family Anthropology Institute, Tel Aviv University, Israel. We thank H. May and A. Pokhojaev for their help and support. Part of this study was performed by I.Z. as a visiting researcher at the Department of Archaeology and Anthropology at the University of Western Australia. We thank M. Goren from the SMNH, who provided us with tremendous help and support in the taxonomic study and the collection of freshwater fish; and W. Van Neer (NHM, Brussels; Royal Belgian Institute of Natural Sciences), M. Richter, P. Campbell and O. Crimmen (NHM, UK) for providing access to their laboratories and osteological collections. We thank the late O. Bar-Yosef for providing us with access to the ‘Ubeidiya fish remains, A. Belfer-Cohen for scientific advice, E. Geffen for statistical assistance, K. Stewart for her constructive review and scientific suggestions to improve this manuscript, and also N. Paz and S. Gavrieli for the English editing.

Author information

Authors and Affiliations



I.Z., N.G.-I. and N.A.-A. conceived the main conceptual ideas. I.Z. and M.P. performed the taxonomic identification and analyses of GBY fish remains. I.Z. and J.N. designed and performed the XRD laboratory experiments and analysed and interpreted the data. I.Z. and I.H. conducted the micro-CT scans and their interpretations. G.S.-V. and T.T. performed the stable isotope analysis, the isotope data evaluation and interpretation. N.A.-A. performed the spatial analyses. I.Z. and I.H. constructed the structure of manuscript, processed all of the presented data and wrote the manuscript, from the initial to the final version. All authors discussed the results and contributed to the final version of the manuscript.

Corresponding author

Correspondence to Irit Zohar.

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The authors declare no competing interests.

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Nature Ecology & Evolution thanks Patricia Eichler, Helen Coxall, Christopher Lowery and the other, anonymous, reviewer(s) for their contribution to the peer review of this work.

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Extended data

Extended Data Fig. 1 Geographic location of the Gesher Benot Ya’aqov (GBY) site on the paleo shore of Lake Hula (Jordan Rift Valley, Israel).

a, Geographic location of GBY, ‘Ubeidiya and Erq-el-Ahmar (red dots). b, Lake Hula and its surrounding swamps are marked according to their size before the drainage in 1954 (modified from83). Selected archaeological sites (Pleistocene to Holocene) located in the vicinity of palaeo-Lake Hula are denoted by black dots. Rivers and springs draining to Lake Hula appear as blue lines. c, Location of GBY on the modern eastern bank of the Jordan River; main excavated areas appear in coloured squares (Area A = brown, Area B = purple, Area C = orange). d, Archaeological excavation of one of the tilted consecutive archaeological horizons recovered in Area B Layer II-6, with rich material culture. JRD = Jordan River Dureijat84.

Extended Data Fig. 2 Micro-CT scans of Luciobarbus longiceps (Cyprinidae, Jordan barbel) head, showing the location of the pharyngeal jaws and teeth.

a, Schematic representation of Luciobarbus longiceps (Jordan barbel) elongated body, with two pairs of barbells below the mouth opening. The circle marks the scanned area (TL = total length; HL = head length; Scanned specimen #13.125SMNH). b – c, Micro-CT scans reconstructing the anatomical location of L. longiceps pharyngeal jaw and teeth (fifth ceratobranchials), exhibiting heterodont dentition organized in three rows (teeth formula: 2,3,4-4,3,227). The first molariform tooth (on the 1st row) is adapted for feeding on hard prey, such as mollusks. The teeth are replaced continuously throughout the fish’s life. The enameloid (gold colour) forms only in the penultimate step of differentiation, prior to the attachment of the tooth to the underlying bone, and following complete resorption of the previous functional tooth.

Extended Data Fig. 3 The association between the number (NISP) of fish remains recovered at GBY, by archaeological horizons in three excavated areas (A, B, C) and the level of taxonomic richness and diversity.

a, A summary table showing the total number of fish remains (NISP), the number of remains identified to genus or species level, species richness (S’), and diversity in three excavated areas (A, B, C), and by archaeological horizon. Area A exhibits highest species richness (S’ = 16), while the lowest values for species richness and diversity are observed at the eight sequential AH of Area B II-6 L1–7 (marked in the light blue background; S’ range between 2 to 6 species). b, Rarefaction curves (calculated with PAST) comparing fish remains NISP and species richness identified at GBY, from areas A (red line), B (blue line), and C (green line). On the graph, in each area, the centre line marks the mean change in species richness according to sample size (NISP), whereas the upper and lower lines display the 95% confidence limit of the sample size and species richness. The rarefaction curves show that in areas A, B, and C, fish remains sample size (NISP) did not influence our ability to reconstruct fish population structure (species richness, diversity, and evenness).

Extended Data Fig. 4 Correspondence analyses graphs of excavated areas and fish taxa (a), and of excavated areas and representation of skeletal elements (b).

a, Correspondence analysis graph showing the association between archaeological horizons and fish taxa (Area A; NISP = 9,206, Area B; NISP = 30,318). The two dimensions, the location of the fish remains (51.4%), and the cause of death (natural - Area A, or cultural- Area B; 45%), almost equally account for the data set variation. b, Correspondence analysis graph showing the association between AH and skeletal elements. The deposition agent (natural or cultural) accounts for most of the variation in the data set (80.8%). This derives from the unique preservation pattern observed in the eight superimposed AHs of Area B (II-6 L1–7), that is, comprises almost exclusively cyprinid pharyngeal teeth. Bottom: Representative fish remains recovered at GBY areas A and B (from left to right): Cyprinidae pharyngeal teeth, L. longiceps 1st molariform tooth, C. canis pharyngeal jaw, Clariidae frontal bone, Cyprinidae pectoral spine, and Cichlidae dorsal fin spine.

Extended Data Fig. 5 X-ray diffraction (XRD) analysis of bioapatite lattice parameters, microstrain, and crystallite size (CS) in fish tooth enameloid.

a, Association between lattice parameters ratio (c/a) and estimated stoichiometric coefficient (x) in cyprinid teeth used in this study (n = 49) and in Palaeozoic and Mesozoic (400‒200 Ma) fossil fish teeth (n = 10)85,86,87,88. b, Lattice parameters of cyprinid pharyngeal teeth analysed in this study and in Palaeozoic and Mesozoic fossil fish teeth85,86,87,88. c, Association between enameloid microstrain and CS in: fresh unheated molariform teeth (n = 4), teeth from whole fresh fish experimentally heated to 200‒600 °C (n = 11), fossil cyprinid teeth of GBY (n = 31), and Erq-el-Ahmar (n = 2). Note that moderate heating (200‒500 °C) increases CS to values of 18 to 23 nm (compared to CS ≈ 17 nm in unheated teeth), whereas natural diagenesis decreases CS and increases microstrain. d, Relation between enameloid microstrain and CS of molariform teeth in: fresh unheated fish, fish cooked in the oven (low heat), and fish cooked in the fire (up to 900 °C). The curved stippled red line marks the trend between microstrain and CS. Note that above a temperature of 600 °C the CS, value strongly increases, reaching a maximum value of up to 68 nm in calcinated teeth, while microstrain decreases below a value of 0.20%.

Extended Data Fig. 6 δ18OPO413C values of ancient and modern Cyprinidae tooth enameloid from Gesher Benot Ya’aqov (GBY), ‘Ubeidiya (UB), Lake Kinneret, and the Jordan River.

a, The phosphate and the carbonate fraction isotope composition (δ18OPO413C) in fossil L. longiceps teeth from ‘Ubeidiya (1.5 Ma; n = 9) and GBY (0.78 Ma; n = 29), compared with that observed for modern L. longiceps (Lake Kinneret; n = 5), reveal that diagenesis did not alter the fossil teeth and that the isotopic values represent changes in water salinity level and temperatures; b, Quadratic discriminant analysis of the phosphate and the carbonate fraction isotope composition (δ18OPO413C) in the sampled teeth, in various aquatic habitats: Lake Kinneret (black); Jordan River (light blue) ‘Ubeidiya (orange); GBY A (green) and GBY B (Blue). The ellipse for each habitat is the 95% confidence interval. The dot markers indicate teeth and the “+” marker indicates the multivariate mean of each group. The discrimination diagram shows distinct separation between aquatic habitats due to water evaporation rate and salinity level (Supplemantery Tables 78; Wilk’s lambda = 0.033, F = 20.33, p < 0.0001, n = 24).

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Supplementary Text A–F (A, the GBY site; B, Cyprinidae pharyngeal teeth; C, natural versus cultural assemblages of GBY fish remains; D, tooth mineral component, XRD analysis; E, spatial analyses of fish remains and burnt flint microartifacts; and F, stable oxygen and carbon isotope analyses of fish teeth), references, Tables 1–8 and Figs. 1 and 2.

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Zohar, I., Alperson-Afil, N., Goren-Inbar, N. et al. Evidence for the cooking of fish 780,000 years ago at Gesher Benot Ya’aqov, Israel. Nat Ecol Evol (2022).

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