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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Barton, C. M., Riel-Salvatore, J., Anderies, J. M. & Popescu, G. Modeling human ecodynamics and biocultural interactions in the Late Pleistocene of Western Eurasia. Hum. Ecol. 39, 705–725 (2011).
Bird, R. B. Disturbance, complexity, scale: new approaches to the study of human–environment interactions. Annu. Rev. Anthropol. 44, 241–257 (2015).
Moritz, M. et al. Emergent sustainability in open property regimes. Proc. Natl Acad. Sci. USA 115, 12859–12867 (2018).
Bliege Bird, R. et al. Fire mosaics and habitat choice in nomadic foragers. Proc. Natl Acad. Sci. USA 117, 12904–12914 (2020).
Foley, S. F. et al. The Palaeoanthropocene—the beginnings of anthropogenic environmental change. Anthropocene 3, 83–88 (2013).
Erlandson, J. M. & Braje, T. J. Archeology and the Anthropocene. Anthropocene 4, 1–7 (2013).
Boivin, N. L. et al. Ecological consequences of human niche construction: examining long-term anthropogenic shaping of global species distributions. Proc. Natl Acad. Sci. USA 113, 6388–6396 (2016).
Fox, T., Pope, M. & Ellis, E. C. Engineering the Anthropocene: scalable social networks and resilience building in human evolutionary timescales. Anthr. Rev. 4, 199–215 (2017).
Bocherens, H. The rise of the anthroposphere since 50,000 years: an ecological replacement of megaherbivores by humans in terrestrial ecosystems? Front. Ecol. Evol. 6, 3 (2018).
Hussain, S. T. & Riede, F. Paleoenvironmental humanities: challenges and prospects of writing deep environmental histories. WIREs Clim. Change 11, e667 (2020).
McCorriston, J. & Field, J. S. Anthropocene: A New Introduction to World Prehistory (Thames and Hudson, 2020).
Boivin, N. & Crowther, A. Mobilizing the past to shape a better Anthropocene. Nat. Ecol. Evol. 5, 273–284 (2021).
Edgeworth, M. Transgressing time: archaeological evidence in/of the Anthropocene. Annu. Rev. Anthropol. 50, 93–108 (2021).
Riel-Salvatore, J. A niche construction perspective on the Middle–Upper Paleolithic transition in Italy. J. Archaeol. Method Theory 17, 323–355 (2010).
Raymond, H. The ecologically noble savage debate. Annu. Rev. Anthropol. 36, 177–190 (2007).
DeLancey An ecological concept of wilderness. Ethics Environ. 17, 25 (2012).
Bliege Bird, R., Bird, D. W., Codding, B. F., Parker, C. H. & Jones, J. H. The ‘fire stick farming’ hypothesis: Australian Aboriginal foraging strategies, biodiversity, and anthropogenic fire mosaics. Proc. Natl Acad. Sci. USA 105, 14796–14801 (2008).
Smith, B. D. General patterns of niche construction and the management of ‘wild’ plant and animal resources by small-scale pre-industrial societies. Philos. Trans. R. Soc. B 366, 836–848 (2011).
Bliege Bird, R. & Nimmo, D. Restore the lost ecological functions of people. Nat. Ecol. Evol. 2, 1050–1052 (2018).
Moran, E. F. People and Nature: An Introduction to Human Ecological Relations (John Wiley Blackwell, 2017).
Wehi, P. M. & Lord, J. M. Importance of including cultural practices in ecological restoration: biocultural restoration. Conserv. Biol. 31, 1109–1118 (2017).
Comberti, C., Thornton, T. F., Wyllie de Echeverria, V. & Patterson, T. Ecosystem services or services to ecosystems? Valuing cultivation and reciprocal relationships between humans and ecosystems. Glob. Environ. Change 34, 247–262 (2015).
Rowley-Conwy, P. & Layton, R. Foraging and farming as niche construction: stable and unstable adaptations. Philos. Trans. R. Soc. B 366, 849–862 (2011).
Riede, F. Adaptation and niche construction in human prehistory: a case study from the southern Scandinavian Late Glacial. Philos. Trans. R. Soc. B 366, 793–808 (2011).
Stiner, M. C. & Kuhn, S. L. Are we missing the ‘sweet spot’ between optimality theory and niche construction theory in archaeology? J. Anthropol. Archaeol. 44, 177–184 (2016).
Thompson, J. C., Wright, D. K. & Ivory, S. J. The emergence and intensification of early hunter‐gatherer niche construction. Evol. Anthropol. Issues N. Rev. 30, 17–27 (2021).
O’Connor, T. Animals as Neighbors: The Past and Present of Commensal Species (Michigan State Univ. Press, 2013).
Brown, A. G., Basell, L. S. & Farbstein, R. Eels, beavers, and horses: human niche construction in the European Late Upper Palaeolithic. Proc. Prehist. Soc. 83, 1–22 (2017).
Baumann, C., Bocherens, H., Drucker, D. G. & Conard, N. J. Fox dietary ecology as a tracer of human impact on Pleistocene ecosystems. PLoS ONE 15, e0235692 (2020).
Bocherens, H. & Drucker, D. G. in Human–Elephant Interactions: From Past to Present (eds Konidaris, G. E. et al.) 249–262 (Tübingen Univ. Press, 2021).
Roebroeks, W. et al. Landscape modification by Last Interglacial Neanderthals. Sci. Adv. 7, eabj5567 (2021).
Grayson, D. K. The archaeological record of human impacts on animal populations. J. World Prehist. 15, 1–68 (2001).
Jöris, O. & Weninger, B. Coping with the cold: on the climatic context of the Moravian Mid Upper Palaeolithic. The Gravettian along the Danube. Dolnověstonické Stud. 11, 57–70 (2004).
Fewlass, H. et al. Direct radiocarbon dates of mid Upper Palaeolithic human remains from Dolní Věstonice II and Pavlov I, Czech Republic. J. Archaeol. Sci. Rep. 27, 102000 (2019).
Svoboda, J., Novák, M., Sázelová, S. & Demek, J. Pavlov I: a large Gravettian site in space and time. Quat. Int. 406, 95–105 (2016).
Wilczyński, J. et al. New radiocarbon dates for the Late Gravettian in Eastern Central Europe. Radiocarbon 62, 243–259 (2020).
Svoboda, J. Dolní Věstonice--Pavlov: explaining Paleolithic settlements in central Europe (Texas A&M Univ. Press, 2020).
Oliva, M. Dolní Věstonice I (1922–1942): Hans Freising—Karel Absolon—Assien Bohmers (Moravské Zemské Muzeum, 2014).
Pavlov I Southeast: A Window into the Gravettian Lifestyles (Academy of Sciences of the Czech Republic, Institute of Archaeology, 2005).
Svoboda, J. et al. Pavlov VI: an Upper Palaeolithic living unit. Antiquity 83, 282–295 (2009).
Absolon, K. & Klíma, B. Předmostí, ein Mammutjägerplatz in Mähren (Archeologický ústav ČSAV v Brnë, 1977).
Svoboda, J. A. The Gravettian on the Middle Danube. Paléo https://doi.org/10.4000/paleo.607 (2007).
Svoboda, J. Dolní Věstonice II: Chronostratigraphy, Paleoethnology, Paleoanthropology (Academy of Sciences of the Czech Republic, 2016).
Svoboda, J. et al. Pleistocene landslides and mammoth bone deposits: the case of Dolní Věstonice II, Czech Republic. Geoarchaeology 34, 745–758 (2019).
Revedin, A. et al. Thirty thousand-year-old evidence of plant food processing. Proc. Natl Acad. Sci. USA 107, 18815–18819 (2010).
Revedin, A. et al. New technologies for plant food processing in the Gravettian. Quat. Int. 359–360, 77–88 (2015).
Adovasio, J. M., Soffer, O. & Klíma, B. Upper Palaeolithic fibre technology: interlaced woven finds from Pavlov I, Czech Republic, c. 26,000 years ago. Antiquity 70, 526–534 (1996).
Farbstein, R. Technologies of art: a critical reassessment of Pavlovian art and society, using chaîne opératoire method and theory. Curr. Anthropol. 52, 401–432 (2011).
Verpoorte, A. Places of Art, Traces of Fire. A Contextual Approach to Anthropomorphic Figurines in the Pavlovian (Archaeological Studies Leiden Univ., 2000).
Svoboda, J. et al. Dolní Věstonice IIa: Gravettian microstratigraphy, environment, and the origin of baked clay production in Moravia. Quat. Int. 359–360, 195–210 (2015).
Goutas, N. From stone flaking to grinding: three original Pavlovian antler tools from Moravia (Pavlov I, Czech Republic). Quat. Int. 359–360, 240–260 (2015).
Nývltová Fišáková, M. Seasonality of Gravettian sites in the Middle Danube Region and adjoining areas of Central Europe. Quat. Int. 294, 120–134 (2013).
Beresford-Jones, D. G. et al. Burning wood or burning bone? A reconsideration of flotation evidence from Upper Palaeolithic (Gravettian) sites in the Moravian Corridor. J. Archaeol. Sci. 37, 2799–2811 (2010).
Hussain, S. T. Gazing at owls? Human–strigiform interfaces and their role in the construction of Gravettian lifeworlds in East-Central Europe. Environ. Archaeol. 24, 359–376 (2019).
Bochenski, Z. M. et al. Fowling during the Gravettian: the avifauna of Pavlov I, the Czech Republic. J. Archaeol. Sci. 36, 2655–2665 (2009).
Wojtal, P., Wilczyński, J., Bocheński, Z. M. & Svoboda, J. A. The scene of spectacular feasts: animal remains from Pavlov I south-east, the Czech Republic. Quat. Int. 252, 122–141 (2012).
Wojtal, P., Wilczyński, J., Wertz, K. & Svoboda, J. A. The scene of a spectacular feast (part II): animal remains from Dolní Věstonice II, the Czech Republic. Quat. Int. 466, 194–211 (2018).
Wertz, K., Wilczyński, J. & Tomek, T. Birds in the Pavlovian culture: Dolni Vestonice II, Pavlov I and Pavlov II. Quat. Int. 359–360, 72–76 (2015).
Wertz, K., Wilczyński, J., Tomek, T., Roblickova, M. & Oliva, M. Bird remains from Dolni Vestonice I and Predmosti I (Pavlovian, the Czech Republic). Quat. Int. 421, 190–200 (2016).
Kost, C. & Hussain, S. T. Archaeo-ornithology: towards an archaeology of human–bird interfaces. Environ. Archaeol. 24, 337–358 (2019).
Hussain, S. T. in The Situationality of Human–Animal Relations. Perspectives from Anthropology and Philosophy (eds Breyer, T. & Widlok, T.) 83–112 (transcript Verlag, 2018).
Klegarth, A. R. in The International Encyclopedia of Primatology (eds Bezanson, M. et al.) 1–5 (John Wiley, 2017). https://doi.org/10.1002/9781119179313.wbprim0448
Froiland, S. Finding nature in the unnatural: toward a philosophy of synanthropy. Eukaryon 13, 41–43 (2017).
Johnston, R. F. in Avian Ecology and Conservation in an Urbanizing World (eds Marzluff, J. M., Bowman, R. & Donnelly, R.) 49–67 (Springer, 2001). https://doi.org/10.1007/978-1-4615-1531-9_3
Povolný, D. in Flies and Disease (ed. Greenberg, B.) 16–55 (Princeton Univ. Press, 2019).
Gade, D. W. Shifting synanthropy of the crow in Eastern North America. Geogr. Rev. 100, 152–175 (2010).
Marzluff, J. M. & Angell, T. In the Company of Crows and Ravens (Yale Univ. Press, 2005).
Bocherens, H. & Drucker, D. G. in The Encyclopedia of Quaternary Science (ed. Elias, S.) 304–314 (Elsevier, 2013).
Krajcarz, M. T., Krajcarz, M. & Bocherens, H. Collagen-to-collagen prey–predator isotopic enrichment (Δ13C, Δ15N) in terrestrial mammals—a case study of a subfossil red fox den. Palaeogeogr. Palaeoclimatol. Palaeoecol. 490, 563–570 (2018).
Craig, O. E. et al. Stable isotope analysis of Late Upper Palaeolithic human and faunal remains from Grotta del Romito (Cosenza), Italy. J. Archaeol. Sci. 37, 2504–2512 (2010).
Bocherens, H. et al. Reconstruction of the Gravettian food-web at Předmostí I using multi-isotopic tracking (13C, 15N, 34S) of bone collagen. Quat. Int. 359–360, 211–228 (2015).
Drucker, D. G. et al. Tracking possible decline of woolly mammoth during the Gravettian in Dordogne (France) and the Ach Valley (Germany) using multi-isotope tracking (13C, 14C, 15N, 34S, 18O). Quat. Int. 359–360, 304–317 (2015).
Drucker, D. G., Bridault, A., Ducrocq, T., Baumann, C. & Valentin, F. Environment and human subsistence in Northern France at the Late Glacial to early Holocene transition. Archaeol. Anthropol. Sci. 12, 194 (2020).
Britton, K. et al. Multi-isotope zooarchaeological investigations at Abri du Maras: the paleoecological and paleoenvironmental context of Neanderthal subsistence strategies in the Rhône Valley during MIS 3. J. Hum. Evol. 174, 103292 (2023).
Bataille, C. P. et al. Triple sulfur-oxygen-strontium isotopes probabilistic geographic assignment of archaeological remains using a novel sulfur isoscape of western Europe. PLoS ONE 16, e0250383 (2021).
Stevens, R. E. et al. Iso-Wetlands: unlocking wetland ecologies and agriculture in prehistory through sulfur isotopes. Archaeol. Int. 25, 168–176 (2022).
DeNiro, M. J. Postmortem preservation and alteration of in vivo bone collagen isotope ratios in relation to palaeodietary reconstruction. Nature 317, 806–809 (1985).
van Klinken, G. J. Bone collagen quality indicators for palaeodietary and radiocarbon measurements. J. Archaeol. Sci. 26, 687–695 (1999).
Nehlich, O. & Richards, M. P. Establishing collagen quality criteria for sulphur isotope analysis of archaeological bone collagen. Archaeol. Anthropol. Sci. 1, 59–75 (2009).
Parnell, A. C. et al. Bayesian stable isotope mixing models. Environmetrics 24, 387–399 (2013).
Phillips, D. L. et al. Best practices for use of stable isotope mixing models in food-web studies. Can. J. Zool. 92, 823–835 (2014).
Stock, B. C. et al. Analyzing mixing systems using a new generation of Bayesian tracer mixing models. PeerJ 6, e5096 (2018).
Wilczyński, J., Wojtal, P., Robličková, M. & Oliva, M. Dolní Věstonice I (Pavlovian, the Czech Republic)—results of zooarchaeological studies of the animal remains discovered on the campsite (excavation 1924–52). Quat. Int. 379, 58–70 (2015).
Brugère, A. Not one but two mammoth hunting strategies in the Gravettian of the Pavlov Hills area (southern Moravia). Quat. Int. 337, 80–89 (2014).
Guiry, E. J., Orchard, T. J., Needs-Howarth, S. & Szpak, P. Freshwater wetland-driven variation in sulfur isotope compositions: implications for human paleodiet and ecological research. Front. Ecol. Evol. 10, 953042 (2022).
Zawadzka, D. & Zawadzki, G. Synanthropisation and synurbisation of raven Corvus corax in Poland: a review. Int. Stud. Sparrows 38, 11–16 (2014).
West, E. H., Henry, W. R., Goldenberg, W. & Peery, M. Z. Influence of food subsidies on the foraging ecology of a synanthropic species in protected areas. Ecosphere 7, e01532 (2016).
Savage, C. S. & Croll, J. Bird Brains: The Intelligence of Crows, Ravens, Magpies, and Jays (Greystone Books, 2018).
Heinrich, B. Mind of the Raven: Investigations and Adventures with Wolf-birds (Harper Perennial, 2006).
Van Dooren, T. The Wake of Crows: Living and Dying in Shared Worlds (Columbia Univ. Press, 2019).
Marzluff, J. M. & Angell, T. Cultural coevolution: how the human bond with crows and ravens extends theory and raises new questions. J. Ecol. Anthropol. 9, 69–75 (2005).
Svoboda, J. et al. Paleolithic hunting in a southern Moravian landscape: the case of Milovice IV, Czech Republic. Geoarchaeology 26, 838–866 (2011).
Holyoak, D. A comparative study of the food of some British Corvidae. Bird Study 15, 147–153 (1968).
Gołdyn, B., Książkiewicz-Parulska, Z. & Zduniak, P. Freshwater molluscs in diet of hooded crow (Corvus cornix). Wilson J. Ornithol. 128, 459–462 (2016).
Tome, D., Krofel, M. & Mihelic, T. The diet of the raven Corvus corax in south-west Slovenia. Ann. Ser. Hist. Nat. 19, 161–166 (2009).
Boarman, W. I. & Berry, K. H. in Our Living Resources: A Report to the Nation on the Distribution, Abundance, and Health of U.S. Plants, Animals, and Ecosystems (eds LaRoe, E. T. et al.) 73–75 (National Biological Service, 1995).
Wojtal, P., Svoboda, J., Roblíčková, M. & Wilczyński, J. Carnivores in the everyday life of Gravettian hunters-gatherers in Central Europe. J. Anthropol. Archaeol. 59, 101171 (2020).
Wilczyński, J. et al. Friend or foe? Large canid remains from Pavlovian sites and their archaeozoological context. J. Anthropol. Archaeol. 59, 101197 (2020).
Svoboda, J. A. The Upper Paleolithic burial area at Předmostí: ritual and taphonomy. J. Hum. Evol. 54, 15–33 (2008).
Pettitt, P. The Palaeolithic Origins of Human Burial (Routledge, 2011).
Martin, D. On the cultural ecology of sky burial on the Himalayan Plateau. East West 46, 353–370 (1996).
Moreman, C. M. On the relationship between birds and spirits of the dead. Soc. Anim. 22, 481–502 (2014).
Maring, R. & Riede, F. Possible wild boar management during the Ertebølle Period. A carbon and nitrogen isotope analysis of Mesolithic wild boar from Fannerup F, Denmark. Environ. Archaeol. 24, 15–27 (2019).
Musil, R. Gravettian environmental changes in a N–S transect of central Europe. Open Geosci. 3, 147–154 (2011).
Svoboda, J. A. Dolní Věstonice—Pavlov: Ort: Südmähren, Zeit: 30000 Jahre v.Chr. = Unter-Wisternitz und Pollau (Regionalmuseum, 2010).
Borgia, V. The mammoth cycle. Hunting with ivory spear-points in the Gravettian site of Pavlov I (Czech Republic). Quat. Int. 510, 52–64 (2019).
Oliva, M. (ed.) Sidlíště Mamutího Lidu u Milovic pod Palávou: Otázka Struktur s Mamutími Kostmi. Milovice: Site of the Mammoth People Below the Pavlov Hills (Moravské Zemské Muzeum, 2009).
Maher, L. A. Persistent place-making in prehistory: the creation, maintenance, and transformation of an epipalaeolithic landscape. J. Archaeol. Method Theory 26, 998–1083 (2019).
Trinkaus, E., Sázelová, S. & Svoboda, J. Pieces of people in the Pavlovian. Hum. Remains Violence 5, 70–87 (2019).
Pryor, A. J. E., Pullen, A., Beresford-Jones, D. G., Svoboda, J. A. & Gamble, C. S. Reflections on Gravettian firewood procurement near the Pavlov Hills, Czech Republic. J. Anthropol. Archaeol. 43, 1–12 (2016).
Douglas, M. S. V., Smol, J. P., Savelle, J. M. & Blais, J. M. Prehistoric Inuit whalers affected Arctic freshwater ecosystems. Proc. Natl Acad. Sci. USA 101, 1613–1617 (2004).
Gomo, G., Mattisson, J., Hagen, B. R., Moa, P. F. & Willebrand, T. Scavenging on a pulsed resource: quality matters for corvids but density for mammals. BMC Ecol. 17, 22 (2017).
Loretto, M.-C., Reimann, S., Schuster, R., Graulich, D. M. & Bugnyar, T. Shared space, individually used: spatial behaviour of non-breeding ravens (Corvus corax) close to a permanent anthropogenic food source. J. Ornithol. 157, 439–450 (2016).
Rösner, S., Selva, N., Müller, T., Pugacewicz, E. & Laudet, F. in Corvids of Poland (eds Jerzak, L. et al.) 385–405 (Bogucki Wydawnictwo Naukowe, 2005).
Selva, N. et al. in Carrion Ecology and Management (eds Olea, P. P. et al.) 71–99 (Springer International, 2019).
Temple, S. A. Winter food habits of ravens on the Arctic slope of Alaska. Arctic 27, 41–46 (1974).
Strømseng, E. Environmental Determinants of Spatio-temporal Variation in a Scavenger Guild on Sub-Arctic Tundra. MSc thesis, Univ. Tromsö (2007).
Killengreen, S. T., Strømseng, E., Yoccoz, N. G. & Ims, R. A. How ecological neighbourhoods influence the structure of the scavenger guild in low arctic tundra: neighbourhood effects on tundra scavenger guild. Divers. Distrib. 18, 563–574 (2012).
Blázquez, M., Sánchez-Zapata, J. A., Botella, F., Carrete, M. & Eguía, S. Spatio-temporal segregation of facultative avian scavengers at ungulate carcasses. Acta Oecologica 35, 645–650 (2009).
Cortés-Avizanda, A., Selva, N., Carrete, M. & Donázar, J. A. Effects of carrion resources on herbivore spatial distribution are mediated by facultative scavengers. Basic Appl. Ecol. 10, 265–272 (2009).
Stahler, D., Heinrich, B. & Smith, D. Common ravens, Corvus corax, preferentially associate with grey wolves, Canis lupus, as a foraging strategy in winter. Anim. Behav. 64, 283–290 (2002).
Selva, N., Jędrzejewska, B., Jędrzejewski, W. & Wajrak, A. Factors affecting carcass use by a guild of scavengers in European temperate woodland. Can. J. Zool. 83, 1590–1601 (2005).
Kaczensky, P., Hayes, R. D. & Promberger, C. Effect of raven Corvus corax scavenging on the kill rates of wolf Canis lupus packs. Wildl. Biol. 11, 101–108 (2005).
Bugnyar, T. & Kotrschal, K. Scrounging tactics in free-ranging ravens, Corvus corax. Ethology 108, 993–1009 (2002).
Lamoureux, R. Winter population dynamics between the Eastern Wolf (Canis lycaon) and the Common Raven (Corvus corax) in Algonquin Park, Ontario. J. Undergrad. Stud. Trent. 4, 61–66 (2016).
Ripple, W. J. & Beschta, R. L. Wolves and the ecology of fear: can predation risk structure ecosystems? BioScience 54, 755–766 (2004).
Hammerschlag, N. et al. Evaluating the landscape of fear between apex predatory sharks and mobile sea turtles across a large dynamic seascape. Ecology 96, 2117–2126 (2015).
Fielding, M. W. et al. Dominant carnivore loss benefits native avian and invasive mammalian scavengers. Proc. R. Soc. B https://doi.org/10.1098/rspb.2022.0521 (2022).
Marzluff, J. M. & Neatherlin, E. Corvid response to human settlements and campgrounds: causes, consequences, and challenges for conservation. Biol. Conserv. 130, 301–314 (2006).
Webb, W. C., Marzluff, J. M. & Hepinstall-Cymerman, J. Linking resource use with demography in a synanthropic population of common ravens. Biol. Conserv. 144, 2264–2273 (2011).
Webb, W. C., Boarman, W. I. & Rotenberry, J. T. Common raven juvenile survival in a human-augmented landscape. Condor 106, 517–528 (2004).
O’Neil, S. T. et al. Broad‐scale occurrence of a subsidized avian predator: reducing impacts of ravens on sage‐grouse and other sensitive prey. J. Appl. Ecol. 55, 2641–2652 (2018).
Baltensperger, A. P. et al. Seasonal observations and machine-learning-based spatial model predictions for the common raven (Corvus corax) in the urban, sub-arctic environment of Fairbanks, Alaska. Polar Biol. 36, 1587–1599 (2013).
Restani, M., Marzluff, J. M. & Yates, R. E. Effects of anthropogenic food sources on movements, survivorship, and sociality of common ravens in the Arctic. Condor 103, 399–404 (2001).
Backensto, S. Common Ravens in Alaska’s North Slope Oil Fields: An Integrated Study Using Local Knowledge and Science. MSc thesis, Univ. Fairbanks Alsk. (2010).
Oosten, J. & Laugrand, F. The bringer of light: the raven in Inuit tradition. Polar Rec. 42, 187–204 (2006).
Zahara, A. R. D. & Hird, M. J. Raven, dog, human: inhuman colonialism and unsettling cosmologies. Environ. Humanities 7, 169–190 (2016).
Kalof, L., Whitley, C., Vrla, S. & Rizzolo, J. B. in Shared Lives of Humans and Animals (eds Räsänen, T. & Syrjämaa, T.) Ch. 11 (Routledge, 2017).
Bijlsma, R. G. & Seldam, Hten Impact of focal food bonanzas on breeding ravens Corvus corax. Ardea 101, 55–59 (2013).
Arnold, Z. J., Wenger, S. J. & Hall, R. J. Not just trash birds: quantifying avian diversity at landfills using community science data. PLoS ONE 16, e0255391 (2021).
Ellis, E. C. Ecology in an anthropogenic biosphere. Ecol. Monogr. 85, 287–331 (2015).
Lupo, K. D. & Schmitt, D. N. When bigger is not better: the economics of hunting megafauna and its implications for Plio-Pleistocene hunter-gatherers. J. Anthropol. Archaeol. 44, 185–197 (2016).
Stahl, P. W. in Encyclopedia of Global Archaeology (ed. Smith, C.) 4433–4439 (Springer, 2020).
Neusius, S. W. in Case Studies in Environmental Archaeology (eds Reitz, E. J. et al.) 297–314 (Springer, 2008).
Guiry, E., Orchard, T. J., Needs-Howarth, S. & Szpak, P. Isotopic evidence for garden hunting and resource depression in the Late Woodland of Northeastern North America. Am. Antiq. 86, 90–110 (2021).
Acosta, A., Carbonera, M. & Loponte, D. Archaeological hunting patterns of Amazonian horticulturists: the Guarani example. Int. J. Osteoarchaeol. 29, 999–1012 (2019).
Linares, O. F. ‘Garden hunting’ in the American tropics. Hum. Ecol. 4, 331–349 (1976).
Sugiyama, N., Martínez-Polanco, M. F., France, C. A. M. & Cooke, R. G. Domesticated landscapes of the neotropics: isotope signatures of human–animal relationships in pre-Columbian Panama. J. Anthropol. Archaeol. 59, 101195 (2020).
Dunn, R. R., Nunn, C. L. & Horvath, J. E. The Global Synanthrome Project: a call for an exhaustive study of human associates. Trends Parasitol. 33, 4–7 (2017).
Guimarães, P. R., Pires, M. M., Jordano, P., Bascompte, J. & Thompson, J. N. Indirect effects drive coevolution in mutualistic networks. Nature 550, 511–514 (2017).
Thornton, T. F., Deur, D. & Adams, B. in Language and Toponomy in Alaska and Beyond: Papers in Honor of James Kari (eds Holton, G. & Thornton, T. F.) 39–55 (Alaska Native Language Center Press, 2019).
Sax, B. Crow (Reaktion Books, 2003).
Chowning, A. Raven myths in Northwestern North America and Northeastern Asia. Arct. Anthropol. 1, 1–5 (1962).
Laugrand, F. & Oosten, J. Hunters, Predators and Prey: Inuit Perceptions of Animals (Berghahn Books, 2016).
Thomas, R. The comparative osteology of european corvids (Aves: Corvidae), with a key to the identification of their skeletal elements. Int. J. Osteoarchaeol. 11, 448–449 (2001).
Cohen, A. & Serjeantson, D. A Manual for the Identification of Bird Bones from Archaeological Sites (Archetype, 1996).
Bocherens, H., Drucker, D., Billiou, D. & Moussa, I. Une nouvelle approche pour évaluer l’état de conservation de l’os et du collagène pour les mesures isotopiques (datation au radiocarbone, isotopes stables du carbone et de l’azote). L'Anthropologie 109, 557–567 (2005).
Bocherens, H. et al. Paleobiological implications of the isotopic signatures (13C, 15N) of fossil mammal collagen in Scladina Cave (Sclayn, Belgium). Quat. Res. 48, 370–380 (1997).
Wickham, H. ggplot2: Elegant Graphics for Data Analysis (Springer, 2009).
Hamilton, N. E. & Ferry, M. ggtern: ternary diagrams using ggplot2. J. Stat. Softw. 87, 1–17 (2018).
Stock, B. & Semmens, B. MixSIAR GUI User Manual v3.12016. (Zenodo, 2016).
Richards, M. P., Pettitt, P. B., Stiner, M. C. & Trinkaus, E. Stable isotope evidence for increasing dietary breadth in the European mid-Upper Paleolithic. Proc. Natl Acad. Sci. USA 98, 6528–6532 (2001).
Pettitt, P. & Trinkaus, E. Direct radiocarbon dating of the Brno 2 Gravettian human remains. Anthropologie 38, 149–150 (2000).
Trinkaus, E., Svoboda, J. A., Wojtal, P., Fišákova, M. N. & Wilczyński, J. Human remains from the Moravian Gravettian: morphology and taphonomy of additional elements from Dolní Vĕstonice II and Pavlov I: morphology and taphonomy of additional Pavlovian human remains. Int. J. Osteoarchaeol. 20, 645–669 (2010).
Gelman, A. Bayesian Data Analysis (Chapman and Hall/CRC, 2013).
Cheung, C. & Szpak, P. Interpreting past human diets using stable isotope mixing models—best practices for data acquisition. J. Archaeol. Method Theory 29, 138–161 (2022).
Jackson, A. L., Inger, R., Parnell, A. C. & Bearhop, S. Comparing isotopic niche widths among and within communities: SIBER—Stable Isotope Bayesian Ellipses in R: Bayesian isotopic niche metrics. J. Anim. Ecol. 80, 595–602 (2011).
Layman, C. A., Arrington, D. A., Montaña, C. G. & Post, D. M. Can stable isotope ratios provide for community-wide measures of trophic structure? Ecology 88, 42–48 (2007).
Robinson, J. R. Investigating isotopic niche space: using rKIN for stable isotope studies in archaeology. J. Archaeol. Method Theory 29, 831–861 (2022).
Reade, H. et al. Deglacial landscapes and the Late Upper Palaeolithic of Switzerland. Quat. Sci. Rev. 239, 106372 (2020).
Reade, H. et al. Magdalenian and Epimagdalenian chronology and palaeoenvironments at Kůlna Cave, Moravia, Czech Republic. Archaeol. Anthropol. Sci. 13, 4 (2021).
Wißing, C. et al. Stable isotopes reveal patterns of diet and mobility in the last Neandertals and first modern humans in Europe. Sci. Rep. 9, 4433 (2019).
Hajdas, I. Radiocarbon dating and its applications in Quaternary studies. E&G Quat. Sci. J. 57, 2–24 (2008).
Hajdas, I., Bonani, G., Furrer, H., Mäder, A. & Schoch, W. Radiocarbon chronology of the mammoth site at Niederweningen, Switzerland: results from dating bones, teeth, wood, and peat. Quat. Int. 164–165, 98–105 (2007).
Bronk Ramsey, C. OxCal Software, version 4.4 (2023); https://c14.arch.ox.ac.uk/oxcal.html
Reimer, P. J. et al. The IntCal20 Northern Hemisphere radiocarbon age calibration curve (0–55 cal kBP). Radiocarbon 62, 725–757 (2020).
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.
The authors declare no competing interests.
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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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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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