Prey choice and changes in site occupation intensity during the Middle and Upper Paleolithic at Ghar-e Boof (southern Zagros Mountains, Iran)

Archaeological and Anthropological Sciences https://doi.org/10.1007/s12520-025-02191-w (2025) 17:95 RESEARCH Prey choice and changes in site occupation intensity during the Middle and Upper Paleolithic at Ghar-e Boof (southern Zagros Mountains, Iran) Mario Mata-González1,2 · Britt M. Starkovich1,3 · Mohsen Zeidi3,4 · Nicholas J. Conard1,3,4 Received: 3 August 2024 / Accepted: 17 February 2025 © The Author(s) 2025 Abstract Ghar-e Boof represents an exceptional Paleolithic site in the southern Zagros Mountains. Due to its long Late Pleistocene sequence that spans from ca. 81 ka until the Epipaleolithic, the site offers a unique opportunity to investigate long-term hominin behavioral patterns on a local scale. In this paper, we examine diachronic trends in prey choice and site occupation intensity during the Middle Paleolithic (MP) through early Upper Paleolithic (UP) at Ghar-e Boof as determined from zooarchaeological data, find densities, accumulation rates, and frequencies of retouched tools. To better understand foraging conditions, variation (or the lack thereof) in species representation and relative abundances are analyzed following the prey choice model of optimal foraging theory. Based on energetic return rates and procurement costs, we distinguish between high-ranked (large and small, slow-moving game) and low-ranked (small-bodied or small, fast-moving game) resources. The occupants of Ghar-e Boof preferentially hunted large game during the MP and early UP and relied on caprines as the main source of meat and marrow. However, there is an increase in the exploitation of fast-moving animals, mostly partridges, relative to small, slow-moving tortoises through the sequence. In addition, site occupation intensity also increased with time. A more intense use of Ghar-e Boof during the early UP may reflect larger groups of people living at the site, more frequent visits, longer periods of occupation, or a combination of some, if not all, of these possibilities. The increased economic importance of lower-ranked prey does not appear to track major environmental or climatic changes, and most likely is tied to higher hunting pressures. The archaeological record of Ghar-e Boof is currently the only example in the Zagros that illustrates the complex interactions between demography, site use, and socioeconomic decisions during the Late Pleistocene, a crucial time period in human evolution. Keywords Middle to early Upper Paleolithic · Zagros Mountains · Zooarchaeology · Prey choice model · Occupation intensity · Paleodemography Introduction Mario Mata-González 1 Institute for Archaeological Sciences, University of Tübingen, Hölderlinstr. 12, 72074 Tübingen, Germany 2 Department of Classics and Archaeology, University of Malta, Msida, MSD 2080, Malta 3 Senckenberg Centre for Human Evolution and Paleoenvironment (SHEP), Hölderlinstr. 12, 72074 Tübingen, Germany 4 Department of Early Prehistory and Quaternary Ecology, University of Tübingen, Schloss Hohentübingen, D-72070 Tübingen, Germany Occupation intensity refers to the length of stay, frequency of visits, and number of people living at a given site (Munro 2004; Stiner et al. 2012). All these variables strongly depend on human population size or density, and different proxy measures of site occupation intensity are commonly used in studies of Paleolithic demography (French 2015). Population size is among the proposed drivers of some of the major evolutionary events that occurred in Eurasia during the Late Pleistocene, such as the Neanderthals’ extinction (Vaesen et al. 2021), the appearance of cultural innovations, and increased technological complexity (Shennan 2001; Powell et al. 2009). Over the last two decades, archaeologists have 13 95 Page 2 of 23 documented an increase in occupation intensity at numerous Eurasian sites across the MP to UP transition and throughout the initial/early UP (e.g., Grayson and Delpech 2003; Münzel and Conard 2004; Kuhn et al. 2009; Marín-Arroyo 2011; Mellars and French 2011; Conard et al. 2012; Starkovich 2017; Yeshurun et al. 2019; Smith et al. 2021; Real and Villaverde 2022). There are also deviations from this pattern (e.g., Morin 2008), and some authors have even questioned the methods and assumptions applied to the analysis of site occupation intensity as a ‘paleodemographic barometer’ (Dogandžić and McPherron 2013; French 2015). Yet the available evidence for changes in occupation intensity between the MP to UP and across the initial/early UP points to either more recurrent visits, longer stays on site and less mobility, larger population densities, or even a combination of some or all of these factors (Grayson and Delpech 2003; Münzel and Conard 2004; Kuhn et al. 2009; Marín-Arroyo 2011; Mellars and French 2011; Conard et al. 2012; Starkovich 2017; Smith et al. 2021; Yeshurun et al. 2021; Real and Villaverde 2022). From a zooarchaeological standpoint, occupation intensity and population pressure are key aspects to consider when investigating Paleolithic diets and subsistence patterns, alongside many other variables, both internal and external, such as climatic and environmental constraints, and technological, cultural, and social adaptations. All else held constant, the human pressure on local food resources would be greater as site occupation intensity increases (Stiner and Munro 2002; Munro 2004). In many regions of Eurasia, zooarchaeological data indicate changes in prey choice and resource intensification during the Late Pleistocene (Stiner et al. 1999, 2000; Stiner and Munro 2002, 2011; Speth and Clark 2006; Kuhn et al. 2009; Stiner 2009; Starkovich 2012, 2014, 2017; Fernández-López de Pablo et al. 2014; Baumann et al. 2020; Romandini et al. 2020; Yeshurun et al. 2021), which has generally been attributed to higher hunting pressures and population growth. Alternatively, some researchers have also found evidence for shifts in prey selection related to climatic fluctuations (Discamps et al. 2011), sites that show either no signs of intensification (Morin 2012; Yravedra-Sainz de los Terreros et al. 2016) or only marginal changes in the way certain prey were used (e.g., birds, Laroulandie et al. 2020), early exploitation of small, fast-moving game during the MP (Blasco et al. 2022), and intensified processing of ungulate carcasses without any apparent modifications in diet breadth during the UP (Manne et al. 2012; Manne 2014). Moreover, the increased use of small game taxa since the initial/early UP onwards could also be associated with the adoption of new technological developments, for example, traps, nets, or snares (Baumann et al. 2020; Bertacchi et al. 2021), which might have allowed hunter-gatherers to lower their 13 Archaeological and Anthropological Sciences (2025) 17:95 procurement and pursuit costs (Morin et al. 2020, 2022). These contributions, therefore, highlight the importance of evaluating overarching behavioral and demographic shifts, while takin (...truncated)