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Evidence for hunter-gatherer impacts on raven diet and ecology in the Gravettian of Southern Moravia

Abstract

The earlier Gravettian of Southern Moravia—the Pavlovian—is notable for the many raven bones (Corvus corax) documented in its faunal assemblages. On the basis of the rich zooarchaeological and settlement data from the Pavlovian, previous work suggested that common ravens were attracted by human domestic activities and subsequently captured by Pavlovian people, presumably for feathers and perhaps food. Here, we report independent δ15N, δ13C and δ34S stable isotope data obtained from 12 adult ravens from the Pavlovian key sites of Předmostí I, Pavlov I and Dolní Věstonice I to test this idea. We show that Pavlovian ravens regularly fed on larger herbivores and especially mammoths, aligning in feeding preferences with contemporaneous Gravettian foragers. We argue that opportunistic-generalist ravens were encouraged by human settlement and carcass provisioning. Our data may thus provide surprisingly early evidence for incipient synanthropism among Palaeolithic ravens. We suggest that anthropogenic manipulation of carrion supply dynamics furnished unique contexts for the emergence of human-oriented animal behaviours, in turn promoting novel human foraging opportunities—dynamics which are therefore important for understanding early hunter-gatherer ecosystem impacts.

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Fig. 1: Pavlovian common ravens conform to a broad dietary niche.
Fig. 2: Pavlovian common ravens regularly consumed large herbivores and mammoths and were commensal to large carnivores and Gravettian humans.
Fig. 3: Mammoth and large non-reindeer herbivores constitute the most important dietary sources of Pavlovian common ravens.
Fig. 4: Sulfur stable isotopes in Pavlovian common ravens display elevated variability when compared to large herbivore baseline.
Fig. 5: AMS radiocarbon dating of analysed Pavlovian ravens shows that they are associated with advanced stages of human settlement.
Fig. 6: Pavlovian ravens partake in the emergence of an Upper Palaeolithic synanthropic niche.

Data availability

All data generated and analysed in this paper are provided as raw values in the main text, as Extended Data figures or can be found in the Supplementary Information.

Code availability

The R code used in this paper is provided in the Supplementary Code.

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Acknowledgements

Long-term support in the development of research infrastructures and osteological collections at the Moravian Museum was provided by the Ministry of Culture of the Czech Republic (ref. MK000094862). S.T.H. and F.R. acknowledge funding received from the European Research Council under the European Union’s Horizon 2020 research and innovation programme (grant agreement no. 817564). C.B. was supported by the University of Helsinki (Academy of Finland project no. 341622). Data collection was made possible through the Materials, Culture and Heritage seed funding scheme of the School of Culture and Society, Aarhus University. We thank K. Wertz from the Polish Academy of Sciences for sharing zooarchaeological expertise during bone sampling in Budišov. We are grateful to P. Tung, D. Drucker and V. García-Huidobro from the Biogeology Working Group (Senckenberg-HEP) in Tübingen for their support in laboratory work.

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Study design and conceptualization were jointly conducted by S.T.H. and C.B. Data collection, formal analysis and interpretation were undertaken by C.B. and S.T.H. The original manuscript was prepared by S.T.H. and C.B. Review and revisions of the manuscript were undertaken by S.T.H., C.B., M.R., F.R., M.A.M. and H.B. Curation of zooarchaeological collections was facilitated by M.R. Funding and research infrastructure were arranged by S.T.H., M.A.M., F.R. and H.B.

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Correspondence to Shumon T. Hussain.

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Extended data

Extended Data Fig. 1 Quantification of occupied raven niche space suggests adaptive-generalist feeding behaviours.

A: Boxplot of Bayesian Standard Ellipse Area (SEAb) of species-specific feeding niches. The black dot indicates the median niche size, while the vertical extension of the boxplot indicates consumptive precision (2.5, 5%, 25%, 75%, 95%, and 97.5% quartiles based on the Bayesian mixing model). Taxa with smaller boxplots have more precisely defined feeding niches than taxa with extensive boxplot coverage. Niche size provides important information on the relative prey specialization of considered species: smaller niche sizes represent food-specialized species, while higher values indicate generalist dietary strategies. B: Boxplot of Bayesian Standard Ellipse Area (SEAb) of species-specific feeding niches, distinguishing between high δ34S ravens and ravens within the baseline defined by large Pavlovian herbivores (Hb; the sample size of low δ34S ravens was too small for this analysis). The niche space of Hb ravens is distinctly larger than that of high δ34S ravens, underscoring intraspecific feeding differences in mid-Upper Palaeolithic raven populations. The black dot indicates the median niche size, while the vertical extension of the boxplot indicates consumptive precision (2.5, 5%, 25%, 75%, 95%, and 97.5% quartiles based on the Bayesian mixing model).

Extended Data Fig. 2 Empirical structure of the Pavlovian isospace suggests three main resources of primary consumers corresponding to three secondary feeding preferences.

A: Main prey resources inferred from Pavlovian herbivore δ13C and δ15N isotopic data using Ward Hierarchical Cluster analysis (Euclidean distance). The three groups are named according to the most common taxon in the cluster (‘reindeer’, ‘large herbivore’ and ‘mammoth’ resource). B: Main consumer groups inferred from the available Pavlovian isotopic data of predators and raven using Ward Hierarchical Cluster analysis (Euclidean distance), revealing three principal feeding preferences (Cluster 1–3). Hb ravens = ravens within the herbivore sulfur baseline.

Extended Data Fig. 3 The three clusters of secondary consumers are mainly defined by the relative importance of mammoth vs. large herbivore intake.

Overview of Bayesian mixed-model (MixSIAR) estimates of dietary compositions for all Pavlovian predators (including H. sapiens) and ravens organized by main consumer groups inferred from the isotope data (cf. Extended Data Fig. 2b). Boxplots show the median value (thick line) and the quartiles (5%, 25%, 75%, and 95%) based on the Bayesian mixing model. A: Dietary profiles of secondary consumers in Cluster 1. This cluster is characterized by ‘mammoth’ as the dominant dietary contributor and includes most Pavlovian humans as well as both ravens from Pavlov I. B: Dietary profiles of secondary consumers in Cluster 2. The cluster is characterized by a high proportion of ‘mammoth’ intake paired with the increased importance of the ‘large herbivore’ resource and contains the majority of Hb ravens (δ34S within the baseline of large herbivores) and one Gravettian human. C: Dietary profiles of secondary consumers in Cluster 3. Specimens in this cluster show the lowest proportional ‘mammoth’ intake and the cluster hosts most high δ34S Pavlovian ravens as well as most large carnivores and Arctic foxes.

Extended Data Fig. 4 The combination of archaeological and isotopic data points to an integrated foraging niche based on novel, carrion-mediated foraging opportunities.

Hypothetical role of Pavlovian people in the assembly and maintenance of regional, patchy micro-ecologies anchored in the accumulation of high-caloric carrion. These secondary food patches provide scavenging and food-steeling opportunities for a range of predators, in turn opening up novel low-cost hunting and trapping opportunities for human foragers. The anthropogenic facilitation of terrestrial mesoscavengers and opportunistic ravens can alter the payoff structure of human subsistence behaviour from an optimal foraging perspective: pursue and failure costs for attracted small to medium-sized mammalian scavengers and ravens are substantially reduced while ensuring elevated encounter rates. The otherwise costly hunting of large proboscideans may in this way become an attractive subsistence strategy.

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Supplementary Tables 1–15, Figs. 1–21 and Discussion.

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R code used for the analysis and data visualization.

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Baumann, C., Hussain, S.T., Roblíčková, M. et al. Evidence for hunter-gatherer impacts on raven diet and ecology in the Gravettian of Southern Moravia. Nat Ecol Evol 7, 1302–1314 (2023). https://doi.org/10.1038/s41559-023-02107-8

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