Stone tools and prey animals were essential to the successful adaptation of Paleolithic humans, leading to their embeddedness in an array of practical, cosmological, and ontological conceptions. The archaeological record of the Lower Paleolithic period in the Levant demonstrates that early humans were very familiar with the landscape and the prey animals they depended upon to sustain their dietary needs. Patterns of continuity and change in stone scrapers, tools commonly associated with the intensive processing of animal remains, might indicate technological and ontological adaptations related to shifting hunting practices or faunal fluctuations. An innovation in scraper technology emerged in the Late Lower Paleolithic Levant, with the introduction of scrapers shaped by Quina retouch. Our analysis of stone tools from the sites of Jaljulia (Late Acheulian, 500–3/200 ka) and Qesem Cave (Acheulo-Yabrudian, 400–200 ka) shows that Quina-like scrapers were often made of flint originating from the Samarian highlands to the east of the sites, the home range of fallow deer, an ungulate frequently hunted by Late Lower Paleolithic humans. We suggest that the nexus between fallow deer hunting grounds and the place of origin of the stones used to produce the scrapers employed to process this prey is a testimony to the role these specific stone tools and prey animals played in the cosmology and ontology of these early humans. In our view, following the disappearance of elephants from the Levant, fallow deer were conceived as a major caloric supplier for humans and, therefore, had to be treated with respect to ensure their constant availability. Thus, it was important for these early humans to process fallow deer using stone that originated from the same hunting grounds where these animals were procured. The Samaria hills, the area of origin of both fallow deer and the particular stone, became central to both practical and spiritual existence of late lower Paleolithic humans in the Levant, and interestingly, this specific location retains its holiness until this very day.

The Late Lower Paleolithic Levant (~ 500 to 200 kya) is characterized by two main cultural entities—an earlier Late Acheulian and a later Acheulo-Yabrudian Cultural Complex (AYCC), generally viewed as consecutive despite some unavoidable overlap or continuity (Bar-Yosef 1994; Barkai et al. 2003; Gopher et al. 2010; Shemer et al. 2022). Levantine Late Acheulian sites are mostly open-air, sometimes with thick stratigraphic sequences representing recurrent human activity, while cave contexts are rare (eg., Marom et al. 2022). On the other hand, AYCC sites are found exclusively in caves and rock shelters, an enigmatic pattern yet to be resolved. Synchronic and diachronic cultural and behavioral dynamics between these entities is an intriguing phenomenon awaiting further research. The Late Acheulian is the last phase of the long-lasting Levantine Acheulian Cultural Complex and is relatively well represented in the Levant. Much like Early and Middle Acheulian industries, Late Acheulian assemblages are dominated by flakes and flake tools produced from single to multi-platform cores, accompanied by a persistent biface component, including chopping tools and the iconic handaxes (eg., Shemer et al. 2022). Several technological innovations appear in the Late Acheulian. These include the systematic production of small flakes from cores-on-flakes (Agam and Barkai 2018b) and predetermined flakes from prepared cores resembling the Levallois method (Rosenberg-Yefet et al. 2021, 2022). Nevertheless, the persistence of the Acheulian handaxe, produced and used for over one million years, led some to view this period as one of cultural “stagnation” or “conservatism”. We take a different view, perceiving the Acheulian as an example of a successful mode of adaptation whose persistent tool-making tradition testifies to a sustainable human existence in equilibrium with the landscape (Barkai 2021; Ben-Dor and Barkai 2021; Finkel and Barkai 2018 and references therein).

The Acheulo-Yabrudian Cultural Complex (AYCC) is a regional entity with innovative cultural and behavioral elements, including extensive occupation of caves and rock shelters, habitual use of fire, long-term bone marrow preservation, and the introduction of new lithic technologies (Barkai and Gopher 2013; Shimelmitz et al. 2014; Blasco et al. 2019; Rosell et al. 2018; Zaidner and Weinstein-Evron 2016). Paleodental remains have led some researchers to associate this period with a new hominin lineage (Barkai and Gopher 2013; Ben-Dor and Barkai 2021; Hershkovitz et al. 2016). Studies of lithic procurement (Agam 2020; Agam and Zupancich 2020; Shimelmitz et al. 2020) and subsistence (Rivals et al. 2021) demonstrated an exceptional familiarity with the landscape. At Qesem Cave, all the archaeological contexts are associated with extensive fallow deer exploitation (Stiner et al. 2011) and are completely devoid of megafauna remains (Barkai et al. 2017; Ben-Dor et al. 2011).

We propose that the emergence, spread, and persistence of Quina and demi-Quina scrapers might be attributed to the dietary reliance of Late Lower Paleolithic humans on fallow deer and the development of a new set of relationships with this animal taxon (following Finkel and Barkai 2021). We are guided by the idea that early human paleoecology and perception of the landscape are crucial driving forces behind patterns of cultural continuity and change. The Late Acheulian and AYCC faunal and technological records are reviewed in light of Finkel and Barkai’s (2021) hypothesis of covariance between specific technological innovations and faunal fluctuations. We highlight a principal difference between the two cultural complexes in terms of zooarchaeological taxonomic variability, reflecting differences in Paleolithic adaptation modes. Then we move to lithic technologies, discussing one of many Late Lower Paleolithic lithic innovations: the early appearance of Quina scrapers in the Late Acheulian and their embeddedness in the AYCC. We also touch upon the role of stone scrapers in successfully butchering and processing animal flesh, bone, and hide, suggesting that innovations in scraper technology and changes in animal acquisition patterns should be considered interrelated. We then proceed to discuss the potential role of particular flint types in the production of Quina scrapers and the probable significance of the place of origin of both rocks and fallow deer. These insights give rise to the notion that Lower Paleolithic cultural and technological patterns of persistency and transformation were guided by human-animal, human-stone, and human-landscape relationships (Romagnoli 2021). We complement the Late Lower Paleolithic data with two relevant, however distant in time and place, case studies: an archaeological case study of Quina scrapers from the European Middle Paleolithic Mousterian and a contemporary ethnographic case study of caribou hunting at Caribou Mountain in Labrador, Canada. These case studies shed light on the potential to discern a universal pattern between hunting societies and the animals, stones, and landscapes they are dependent upon.

Elephants and Fallow Deer in Late Lower Paleolithic Faunal Assemblages

Late Acheulian

The faunal spectrum from the Levantine Acheulian is marked by considerable taxonomical diversity, with the recurrent appearance of medium and large African/Palearctic herbivores, including several proboscidean species (Belmaker 2009; Horwitz and Tchernov 1989; Gaudzinski 2004; Pokines et al. 2019; Rabinovich et al. 2012). Although major faunal turnovers occurred throughout the million years of the Acheulian in the Levant (Belmaker 2009), by 500 kya Late Acheulian humans persisted in the acquisition of large game, including the enormous Palaeoloxodon antiquus (Chazan and Horwitz 2007; Lister et al. 2013; Rabinovich et al. 2012). For example, several noteworthy cases of elephant exploitation by Acheulian humans are of special interest. At Gesher Benot Ya’akov (GBY), a Palaeoloxodon antiquus skull was recovered alongside a large oak log and atop a large basalt core in primary context, most probably as part of an Acheulian living floor (Goren-Inbar et al. 1994). The remains were interpreted as an attempt to position the skull using the log, then penetrating it by crushing its basal part to access the elephant’s brain and other choice edible organs (see also Agam and Barkai 2016). It is of note, however, that fallow deer were also hunted and consumed at GBY (Rabinovich et al. 2008), but the caloric contribution provided by the elephant was unparalleled by any other animal species (Ben-Dor et al. 2011; Ben-Dor and Barkai 2020, 2021). Numerous Palaeoloxodon antiquus remains associated with lithic implements were recovered in the Late Acheulian site of Revadim, where fallow deer was also a minor caloric contributor. A handaxe and a scraper (shaped by scalar-stepped retouch) found near an elephant rib showed traces of use and residue related to animal fat, suggesting their use in butchering (Solodenko et al. 2015). It has been suggested that Early/Middle Pleistocene humans targeted the largest, fat-rich prey available, having sufficient biological, cognitive, and technological capabilities for systematic hunting, butchering, and processing of prey animals (Agam and Barkai 2018a; Domínguez-Rodrigo and Pickering 2017; Gaudzinski-Windheuser et al. 2023; Guil-Guerrero et al. 2014, 2018; see Lupo and Schmitt 2023 for an alternate view). However, elephants were not the only species consumed during Lower Paleolithic times. Evidence for butchering smaller animals, such as numerous prime adult fallow deer carcasses, was found at GBY (Rabinovich et al. 2008) and Holon (Horwitz and Monchot 2002). The butchering process consisted of a wide spectrum of tasks: skinning, carcass dismemberment and disarticulation of the skeleton, defleshing and filleting of meat, and bone breaking for marrow extraction. Other larger ungulates, such as Bos primigenius, Cervus elaphas, and various Equus species, were also hunted and butchered (Goren-Inbar et al. 1994; Horwitz and Monchot 2002; Marom et al. 2022; Rabinovich et al. 2012; Stiner et al. 2009, 2011). The human trophic level during the Lower Paleolithic may have evolved towards a hypercarnivorous diet (Ben-Dor and Barkai 2021). Large herbivores, and especially proboscideans, would thus have provided unprecedented fat and protein reserves, fulfilling the dietary needs of human groups for prolonged periods (Ben-Dor and Barkai 2021; Gaudzinski-Windheuser et al. 2023; Guil-Guerrero et al. 2014, 2018). The energetic return of a single elephant equaled that of hundreds of medium-sized ungulates (Ben-Dor and Barkai 2021, 2–5). They were thus essential to human adaptation for as long as they were available. Human hunting pressure, coupled with environmental changes, may have led to the gradual decline of specific Pleistocene taxa, mainly megaherbivores (Dembitzer et al. 2022; Devès et al. 2014; Surovell et al. 2005), until their final disappearance from the Levant at the end of the Acheulian (Ben-Dor et al. 2011).

Acheulo-Yabrudian Cultural Complex (AYCC)

Acheulo-Yabrudian faunal assemblages are primarily dominated by medium-sized fallow deer (Dama mesopotamica) and small-sized mountain gazelle (Gazella gazella), along with smaller numbers of large equids and bovids (Bate 1937; Devès et al. 2014; Stiner et al. 2009; Yeshurun 2016), while elephants are entirely absent. A subsistence based on systematic hunting of medium-sized ungulates might have been gradually adopted by Acheulo-Yabrudian humans armed with new butchery kits. The oldest scraper-dominated layer in Qesem Cave, dated to around 400 kya (Falguères et al. 2022), yielded animal tooth micro and mesowear suggesting regular procurement of equids and bovids alongside seasonal fallow deer hunting (Rivals et al. 2021), while later horizons indicate growing reliance on fallow deer (Blasco et al. 2013, 2019; Stiner et al. 2009). Shifts in fallow deer/gazelle abundance in Pleistocene Mount Carmel were initially interpreted as an indicator for climatic fluctuations (Bate 1937). Recently, such shifts were explained by human cultural preferences and changes in hunting ranges (Blasco et al. 2013; Speth 2012). From here onward, we will focus on fallow deer, a species well represented in AYCC contexts.

Fallow Deer at Qesem Cave

Dama mesopotamica was widely distributed in the Near East throughout the Pleistocene. In the Levant, fallow deer were extirpated from most of their habitats during the Holocene. Fallow deer went entirely extinct at the beginning of the 20th century before being reintroduced to the Carmel and then to other regions some 30 years ago (Mendelson and Yom-Tov 1999). A medium-sized ungulate, the fallow deer grows up to 2 m in length and weighs ca. 100 kg. Males are larger than females and grow antlers that shed in winter (Mendelson and Yom-Tov 1999). Fallow deer prefer to graze in forests and dense woodlands, although the actual range of the reintroduced population is quite variable (Bar-David et al. 2005). Rut season peaks during the late summer-autumn, and fawning season occurs around March–May, having some impact on their annual range (Bar-David et al. 2005). At Qesem Cave, fallow deer constitute up to 79% of the faunal assemblages, while gazelles are absent (Stiner et al. 2009). Fallow deer were systematically hunted and butchered while selected body parts were transported back to the cave from hunting grounds. An over-representation of limb bones, metapodials, and mandibles indicates selective transportation and correlates with marrow utility indices (Stiner et al. 2009, 2011). Delayed consumption of marrow has been demonstrated and shown experimentally by Blasco et al. (2019), who observed a prevalence (80%) of deep and shallow cut marks on fallow deer metapodials, interpreted as removal of stale tissues that were deliberately kept on the bone to preserve the marrow. At 300 kya, fallow deer were obtained by Qesem Cave inhabitants from several distinct hunting grounds and during more than one season (Rivals et al. 2021). This interesting insight arose from micro and mesowear analyses of fallow deer teeth showing dietary differences between specimens excavated from the central hearth area and those found in other parts of the cave.

Stone Scrapers in Late Lower Paleolithic Toolkits and Practices

Stone scrapers are among the oldest tool categories in the archaeological record. These tools are commonly defined by their abrupt or semi-abrupt retouch, which creates a continuous and durable working edge (Bordes 1961). As their typological name suggests, Paleolithic stone scrapers are predominantly associated with scraping activities and processing of hides, following extensive ethnographic and ethno-archaeological literature (Arthur 2018; Binford 1978; Gould et al. 1971; Sahle et al. 2012). However, the role of scrapers in the Paleolithic past is more complicated (Lazuén and González-Urquijo, 2015). Scrapers are repeatedly found in kill and butchery sites, and use-wear analyses revealed the function of certain scrapers as cutting tools during butchery (Clemente-Conte et al. 2020; Peresani et al. 2023; Solodenko et al. 2015). Although scrapers also were used, to some extent, in working other organic materials such as wood (see, eg., Hardy 2004; Zupancich et al. 2016), their primary role in the Paleolithic was in processing animal matter (Figure 1). At the Late Acheulian Revadim site, scrapers may have been used in specific animal processing tasks in conjunction with other flake tools, bifaces, and cortical and double-ventral flakes (Venditti et al. 2022). The lithic assemblage from the “spear horizon” of the Lower Paleolithic Schoeningen site in Germany, where dozens of horses were intensively butchered, is also dominated by scrapers (some of which are highly elaborated) and other flake tools (Serangeli and Conard 2015). Paleolithic stone scrapers were frequently involved in intensive processing of animal remains during multi-staged butchery and hide-working activities, entailing prolonged contact with animal skin, flesh, and bone (Claud et al. 2019; Clemente-Conte et al. 2020; Lemorini et al. 2016, 2022). They were employed with other stone and organic tools and may have been used during specific stages along the carcass and/or hide processing continuum, such as defleshing or hide softening (Tartar et al. 2022; Venditti et al. 2022). We work under the assumption that scrapers were used in routine butchering and processing of animal remains, thus enhancing, for the humans who used these tools, the practical and perceptual bond between the tools and the processed animals. This bond might have led to a need for innovation in the wake of a major faunal turnover and/or an ecological change affecting an animal frequently hunted by humans (Finkel and Barkai 2021).

Figure 1
figure 1

A Quina scraper from Qesem Cave. Residue and use-wear analyses showed that the scraper was wrapped with organic material and used for scraping a dry hide (after Zupancich 2020)

Late Acheulian Scrapers: Continuity and Innovation

Late Acheulian stone scrapers remained largely similar to their predecessors. Their technological and stylistic simplicity were preserved throughout the very long one-million-year Acheulian, and they remained a recurrent component of Acheulian toolkits (eg., Goren-Inbar et al. 2018). However, in the Late Lower Paleolithic in the Levant, new production and maintenance techniques were introduced by Late Acheulian knappers (Malinsky-Buller 2014), with the initial appearance of scrapers shaped by Quina-like retouch representing an outstanding innovation (Figure 2). With its anchoring in the later Levantine AYCC (Barkai and Gopher 2013), this technique then became an important cultural marker, and shaping by Quina retouch is likewise a major technological attribute of the European Middle Paleolithic Quina Mousterian (Bourguignon 1997). Unlike regular blunted scrapers, the scalar-stepped (“écailleuse scalariforme”) Quina retouch creates a rather sharp working edge, having a series of convexities and concavities in cross-section (Lemorini et al. 2016). Quina-like retouch has been reported from a few Late Acheulian contexts (Ruíz et al. 2011; Solodenko et al. 2015), although the number of items is exceedingly small. The scraper assemblage of Late Acheulian Jaljulia, a large-scale open-air site located in the Levantine central coastal plain, offers some insights into these patterns of continuity and change (Shemer et al. 2022). While regular and simple Acheulian scrapers are expectedly the dominant category, scrapers shaped by scalar-stepped retouch are also present (16% of the scrapers), some exhibiting only ‘initial’ or partial Quina retouch, while others closely resemble their later AYCC counterparts (Figure 2a, d, e). Quina-like scrapers appear in all Jaljulia localities, including the pre-AYCC context dated to the MIS13 (Shemer et al. 2022). Detailed analysis of the scrapers is ongoing and will be presented elsewhere.

Figure 2
figure 2

Examples of scrapers shaped by Quina-like retouch from the Late Acheulian Jaljulia: Déjeté scraper (a); transverse scraper (b); Single scrapers (c, d); Convergent scraper fragment (e)

Acheulo-Yabrudian Quina Scrapers

In the Levant, Quina scrapers are omnipresent in the AYCC but completely absent in later Middle Paleolithic contexts. Surprisingly, they reemerge as an important phenomenon in the European Middle Paleolithic (see below). The connection between these two manifestations of Quina technology is yet to be resolved, despite the resemblance of AYCC Quina and demi-Quina scrapers (Figure 3) to their European Middle Paleolithic successors. Several techno-morpho-functional similarities are evident, as are similarities in blank characteristics and the scalar-stepped retouch that characterize this technology (Lemorini et al. 2016; Zupancich 2020). Levantine Quina scrapers were usually made on thick, cortical flakes, asymmetric in cross-section (Agam and Zupancich 2020; Franklin and Kuhn 2021; Shimelmitz et al. 2014; Zaidner and Weinstein-Evron 2016). Demi-Quina scrapers were more variable in dimensions, often made on relatively thin flakes and partially retouched (Agam and Zupancich 2020). The typo-technological characteristics of Quina and demi-Quina blanks were intended to achieve firm prehension (Shimelmitz et al. 2014), while some of the tools showed hafting or wrapping traces (Zupancich et al. 2016). Levantine Quina production demonstrates a geographically fragmented chaîne opératoire pattern. While scrapers are abundant, related cores and initial Quina debitage are extremely rare in AYCC assemblages, although the presence of resharpening spalls indicates that these scrapers were maintained in or close to the caves (Barkai and Gopher 2013; Zaidner and Weinstein-Evron 2016). Functional analysis of Qesem Cave Quina and demi-Quina scrapers indicate versatility with two main trajectories: the more standardized Quina scrapers were often used for scraping activities, while demi-Quina scrapers bearing relatively acute edges were often used for cutting medium and soft materials, mostly of animal origin (Agam and Zupancich 2020; Lemorini et al. 2016). Different Quina scraper types for cutting and/or scraping also have been observed in European Quina contexts (Peresani et al. 2023).

Figure 3
figure 3

Qesem Cave Quina (a, b) and demi-Quina (c) scrapers

Rock Selectivity in the Production of Jaljulia and Qesem Cave Scrapers

Qesem Cave and Jaljulia are located in the western foothills of the Samarian Highlands within the Turonian limestone of the’Bi’na Formation, containing dozens of flint outcrops (Agam 2020). Turonian flint types dominate the Qesem Cave assemblages, and while the import of those flint types from distant sources cannot be ruled out, it is reasonable to assume that the extensive use of Turonian flint types derived from their local availability. However, flint use patterns are more complicated regarding scraper production. Concentrations of Beryllium-10 isotope (10Be) in Qesem Cave artifacts revealed a selective use of durable flint quarried from underground sources hidden from cosmic rays or from recently exposed primary geological sources, specifically for scraper and handaxe production (Boaretto et al. 2009). Recently, Agam and Zupancich (2020) showed that local Turonian flint is less prevalent in Quina and demi-Quina Qesem scrapers than in other lithic categories. Qesem Cave scrapers often were made of exotic and rare flint types unavailable near the site. Flint procurement from distant sources was reported from other Acheulo-Yabrudian sites as well (Shimelmitz et al. 2020; Zaidner and Weinstein-Evron 2016). The semi-translucent homogenous Campanian flint of the Mishash formation (Figures 2d, 3c) is particularly interesting, as the nearest Mishash sources are 15 km to the south and in the Samarian highlands ~ 20 km east of the cave. The Mishash formation is composed of alternating layers of flint and chalk, with some marl and phosphorite (Arkin 1987) throughout the Samarian western slopes (Figure 4). Dedicated geological and archaeological surveys would shed more light on the matter but are currently prevented by the complicated socio-political situation in the region.

Figure 4
figure 4

The location of Jaljulia and Qesem Cave in relation to potential primary flint sources (after Agam and Zupancich 2020), Wadi Qana (marked in a blue line), and Gerizim & Ebal mountains (marked in a blue circle) (Color figure online)

At Jaljulia, Campanian Mishash flint comprises half of the sample studied by Agam et al. (2022). BrecciatedFootnote 1 types constitute the majority (31.7% of the studied sample) of Campanian flint types in Jaljulia and were preferentially selected to produce bifaces and discoid cores (Agam et al. 2022). Despite the absence of Campanian Mishash outcrops in the vicinity of the sites, various flint cobbles and nodules were carried by the meandering Wadi Qana stream and deposited in a fluvial conglomerate upon which Jaljulia cultural horizons were embedded. Brecciated nodules also were found in an ancient streambed revealed by test pits dug in the current Wadi Qana bed (Agam et al. 2022). Non-brecciated semi-translucent Mishash flint was not located during the surveys nor found in the ancient streambed (A. Agam, personal communication). Local availability of non-brecciated Mishash thus remains unproven. Table 1 shows that non-brecciated Campanian flints are relatively common in a sample of 212 Jaljulia scrapers but compose only 19.7% of the general sample studied by Agam et al. (2022). The frequencies of both non-local rock types are surprising, giving rise to the following insights:

  1. 1.

    The fertile wetlands of Jaljulia were not merely an attractive spot for animal herds, but also an area where stone was plentiful, likely a major secondary source for flint procurement. Paleolithic humans at Jaljulia certainly recognized that the abundance of stone lying under their feet was brought by the eastern course of Wadi Qana, flowing from the Samarian highlands down towards the coastal plain. It is safe to assume that early inhabitants of Jaljulia were intimately familiar with the immediate as well as remote landscapes, were well acquainted with the rock outcrops in the eastern Samaria hills, and were aware that nodules and cobbles brought by the stream to their preferred habitat at Jaljulia originated from these sources.

  2. 2.

    Non-brecciated Campanian flint types were preferably selected for scraper production. At present, their local availability at secondary or primary sources has not been proven, leading to the fascinating hypothesis that this flint was directly procured some ~ 15 to 25 km east of the site. The Samarian highlands are one of the main candidates for a suitable source of Campanian flint, with implications for the mobility patterns of Jaljulia humans and their perception of the environment. As such, this region may have played a central role in Late Lower Paleolithic subsistence, and its regional importance was also preserved later (Frumkin et al. 2016).

TABLE 1 Flint types in Jaljulia and Qesem Cave scrapers compared to general samples (comprising all lithic material) from the sites (after Agam and Zupancich 2020; Agam et al. 2022)

Stone, Animals, and Mountains in the Levant from the Lower Paleolithic and Beyond

The early appearance of Quina scrapers in Jaljulia, prior to their wide distribution and embeddedness in the AYCC, suggests practical and most probably ontological continuity between the two Late Lower Paleolithic entities. Many generations of Late Acheulian humans preserved the bulk of the traditional Acheulian mode of adaptation. They were engaged in the systematic acquisition, butchery, and consumption of a wide variety of large game, including megafauna, following the footsteps of their ancestors. The inhabitants of the Jaljulia wetlands repeatedly visited the area, which was characterized by the conspicuous presence of stone and animals. As they continued to practice the Acheulian lifeway, manufacturing handaxes, regular scrapers, and countless flake tools, early humans at Jaljulia were also exploring and experiencing new trajectories in lithic technologies and flint procurement, adhering to changes in human ecology and behavioral complexity (Romagnoli 2021). We suggest that these Late Acheulian innovations, such as the initial application of Quina retouch, are part of a major cultural, ontological, and biological transformation in the Late Lower Paleolithic Levant.

The emergence of Quina scrapers might indicate a response to changing availability in the primary sources of human sustenance, game animals processed by Paleolithic toolkits. One example of this change is the disappearance of a great primordial human counterpart, the elephant, from zooarchaeological assemblages and its replacement by smaller ungulates, such as the fallow deer who made their home in the Samarian highlands. We will present archaeological evidence demonstrating the vital role of the Samarian hills for human subsistence, ontology, and cosmology. Given the known ritual importance of this mountainous region since the Iron Age, we believe that this landscape’s ontological and cosmological role may extend back to the Paleolithic. The prominent role of mountains in many Indigenous cosmologies also supports the plausibility that the interrelationships between landscape, stone procurement, and animals have ancient roots.

The reliance of Late Acheulian groups on large game was likely shattered by the decrease in availability of elephants, who constituted a significant portion of their diet and whose presence in the edaphically productive coastal plain and the valleys to the northeast allowed them to persist in their lifeways and well-being for over one million years (Barkai 2021; Barkai et al. 2017; Ben-Dor and Barkai 2021). It is uncertain when these enormous animals vanished from the Levant. Most evidence indicates their disappearance roughly at 400 kya (Agam and Barkai 2018a; Dembitzer et al. 2022), while late isolated elephant remains in the region are dated to ~ 192 ka (Pokines et al. 2019). The disappearance may have been gradual, with a reduction in elephant populations over time and perhaps the survival of the last elephants in refugia at the end of this process. From the perspective of early humans, the dwindling availability of elephants and related paleoecologic changes may have triggered their additional interest in other biomes from different, rougher terrains (Devès et al. 2014).

Samaria, with its abundance of stone types preferred by Paleolithic flintknappers, is also known to have been one of the hunting grounds visited by Qesem Cave humans (Rivals et al. 2021). Deer taxa were endemic throughout the Pleistocene, as evident by archaeological localities in the region. Emanuel Cave, located in the western Samarian slopes just above Wadi Qana and dated to 191 kya, is marked by gazelle and fallow deer dominance and extensive use of locally available flint (Goder-Goldberger et al. 2012). Fallow deer dominate the assemblage of the MIS6 (~ 140 kya) Rantis Cave, a natural pitfall (Marder et al. 2011). Paleoenvironmental reconstruction placed Rantis Cave in a mixture of shrublands and oak woodlands, supporting a population of fallow deer and forest dwellers like Caucasian squirrels (Marder et al. 2011, 777–778).

The region’s phytogeographic settings were altered by drastic human impact in the last four millennia and today is mostly a xeric Mediterranean shrubland (Frumkin et al. 2016). Thus, while extremely rare in contemporary contexts, fallow deer are thought to have endured continuously in this region throughout the Pleistocene and the Holocene. Fallow deer remains were found at an Iron Age I structure at Mt. Ebal in the Samaria hills, one of the most important altars known in Biblical times. This altar is even thought to be associated with Joshua’s altar mentioned in the Old Testament (Zertal 1987). Fallow deer constitute 21% of the remains in the altar, implying that a woodland habitat probably still existed in the vicinity of the mountains some 3,000 years ago (Horwitz 1987, 181). At most other Iron Age sites, domesticated animals were consumed and sacrificed. The hunting and sacrificing of wild animals, such as fallow deer at this altar, is unique to this context and locality. A few burnt antlers and bones bearing cut marks suggest that fallow deer were selected for ritualized ceremonies at Mt. Ebal during the Iron Age. This highlights their presence in the wild in the Samaria hills and their importance for the site inhabitants. We contend that the significance of this landscape may have persisted for generations, starting from Pleistocene times.

As elevated landforms that rise above their surroundings, mountains affect local climate, environment, and ecosystems while also being shaped by weathering, erosion, and the powerful dynamic tectonic and orogenetic processes in Earth’s lithosphere. The highest peaks in Samaria, Mt. Gerizim, and Mt. Ebal, are not particularly high (881 and 940 m above sea level, 350 m above their surroundings), separated by a valley where the modern city of Nablus lies (Figure 5). They are, however, a prominent feature of the landscape, especially for someone approaching from the western flatlands: “Mount Ebal itself is the most remarkable feature on the Sheet, and a conspicuous object from the plains” (Conder and Kitchener 1882, 147). Mt. Gerizim and Mt. Ebal, along with their surroundings, are deeply embedded in the Abrahamic religions, as the locations where the tribes of Israel held ceremonies of blessings and curses upon their arrival to Canaan described in the Book of Joshua. Gerizim is home to the Samaritan people, who consider the mountain the holiest place on Earth. We believe that the sacred status of Gerizim and Ebal in the past and present is only the tip of the iceberg, the exposed edge of a perceptual phenomenon rooted in prehistoric times.

Figure 5
figure 5

A view of Mt. Gerizim and Mt. Ebal from the east (credit: Dr. Shai Bar)

As mountains overlooking the valleys and the caves that served as sources of flint and grounds for fallow deer hunting, Gerizim and Ebal ultimately played a significant role in the adaptation and perception of the inhabitants of Jaljulia and Qesem. The flow of Wadi Qana, originating just to the southeast of Mount Gerizim, carried flint nodules and cobbles down to the Jaljulia floodplain and must have been another connecting point noticed by early humans (Figure 6). We argue that Late Lower Paleolithic cultural and technological persistency and transformation were guided by human-animal, human-stone, and human-landscape relations. The noteworthy innovation in scraper technology, with the introduction of Quina retouch in the Late Acheulian and its wide embeddedness in the AYCC, is a paramount example. This change might indicate a shift in animal hunting, butchering, and hide-working practices in relation to the changing availability of elephants versus fallow deer. The Samarian highlands were probably an important destination for early humans, who previously inhabited the coastal plains in their search for fallow deer and particular rock formations. The summits of Mt. Gerizim and Mt. Ebal were probably admired as prominent landscape nodes connecting humans, animals, and stones.

Figure 6
figure 6

Artistic depiction of the discussed landscape. Mt. Gerizim & Mt. Ebal with flint-bearing outcrops (upper right corner), Wadi Qana and Qesem Cave (lower left corner). Drawing by Zhao Ying

Comparative Cosmologies Concerning Mountains, Stone Tools, and Cervids

In support of our claim for a universal phenomenon, we will diverge from the Levant and briefly examine prehistoric Mousterian and Indigenous Arctic populations, who hunted cervids and procured stone from mountainous landscapes. We contend that whenever and wherever humans were dependent on animals and stones, strong practical and ontological bonds between the three were established, setting the stage for what we see as processes of cultural and technological continuity and change. We now wish to extend the scope of this claim by presenting two case studies, one archaeological and one ethno-archaeological/ethnographic, in which we can trace the very same phenomenon.

In Indigenous societies, mountains are often described as ancient entities containing liminal niches and divine supernatural beings, large body masses enabling the existence of the universe, reaching the upper and lower worlds (eg., Brumm 2010, 186–188; Carreño 2017; Jacobson 1993, 181; Reimer 2018; Tinker 2004). One example of the prominent role of mountains in native cosmology, associated with stone tool production and transportation, is the Aboriginal “falling sky” myth, according to which the skies were supported by invisible poles ascending from mountain summits and maintained by supernatural stone axes. When the sky was believed to be in the process of collapse, large numbers of stone axes were manufactured and transported to specific mountains via designated directions (Brumm 2010, 188–191). Stone procurement among many ethnographic societies often occurred in remote, mountainous regions identified with ancestral and supernatural presence (Brumm 2010; Jones and White 1988; Loring 2002; Reimer 2018). It was an arduous and dangerous physical and spiritual task, usually completed by a group of men led by experienced elders (Jones and White 1988; Reimer 2018). Stones were believed to grow in the quarry as in a mother’s womb (Arthur 2018, 81–83; Jones and White 1988). Their procurement was identified with childbirth, a feminine task in front of men in the absence of women, and thus filled with existential danger (Arthur 2018, 83). Being prominent landforms, mountains were barriers and passways in both practical and perceptual meaning.

Thus, we see how different landscapes have connected human groups, animals, and stones in different times and places. We contend that whenever and wherever humans were dependent on animals and stones, strong practical and ontological bonds between the three were established, setting the stage for what we see as processes of cultural and technological continuity and change. We now wish to extend the scope of this claim by presenting two case studies, one archaeological and one ethno-archaeological/ethnographic, in which we can trace the very same phenomenon.

Case Study 1: Animals, Stones, and Quina Scrapers in the European Middle Paleolithic Mousterian

The Quina method is a specific facies or techno-complex of the European Middle Paleolithic, the Quina Mousterian (“Le Moustérien de type Quina”) (Bourguignon 1997). Quina technology is characterized by long-distance transportation of lithic materials across the landscape, production of thick blanks with an asymmetric cross-section systematically obtained via different “volumetric” core reduction strategies, combined use of several types of hammerstones and gesture techniques, frequent use of lithic ramification, and high investment in tool shaping and curation, resulting in long life-history potential (Bourguignon 1997, 2001; Costamagno et al. 2015; Delagnes and Rendu 2011; Hiscock et al. 2009; Meignen et al. 2009; Niven et al. 2012; Peresani et al. 2023; Romagnoli et al. 2016; Turq 1992). The organization and properties of Quina technology were interpreted as a form of Neanderthal adaptation to highly mobile lifeways specialized in the acquisition of migratory prey (Delagnes and Rendu 2011; Rendu et al. 2023; Romagnoli et al. 2016). Quina Mousterian sites are concentrated in southwestern France, in the Périgord, Dordogne, and Charentes regions, and dated to MIS5-3. However, Quina Mousterian contexts have also been discovered in adjacent regions, such as the Pyrenees and northern Italy (Peresani et al. 2023), in the Middle Rhône Valley (a local variant) (Moncel and Daujeard 2012), as well as in other parts of Europe. The Mesozoic and Cenozoic sedimentary environments of Southwestern France are rich in primary and secondary depositions of various flint types, resulting in assemblages dominated by local flint obtained from riverbeds, valleys, and mountain foothills near Quina sites (Hiscock et al. 2009). But there is also clear evidence for the use of non-local stone by Quina knappers, especially for retouched tools and Quina scrapers (Costamagno et al. 2015; Hiscock et al. 2009; Jéquier et al. 2015; Meignen et al. 2009, 19). Long-range transportation of stone tools has been associated with high mobility and extended movement range, “…a highly structured system of resource use that involved regular, well-defined selectivity of items to be transported, and repeated transfers of blanks and tools across the landscape in the same directions” (Hiscock et al. 2009, 235).

The hallmark of this system is the Quina scraper, typically made on a flake that is asymmetric in cross-section with a cortical back or blunted thick margin opposing the working edge, which is shaped by continuous, regular, and invasive scalar-stepped (“écailleuse scalariforme”) retouch (Bourguignon 1997). This retouch was particularly adapted to the volume of the scraper (Peresani et al. 2023), suitable for scraping and cutting, recurrent resharpening, and maintenance (Bourguignon 1997; Lemorini et al. 2016; Romagnoli et al. 2016; Turq 1992; Zupancich 2020). Quina scrapers were functionally versatile, with an emphasis on working hide or soft material, such as flesh (Claud et al. 2019; Hardy 2004; Geneste and Plisson 1996; Lin and Marreiros 2021; Niven et al. 2012; Peresani et al. 2023; Tartar et al. 2022). The link between Quina scrapers and butchery activities was not exclusively derived from use-wear and residue analyses but also, as Lin and Marreiros (2021) noted, from their contextual appearance in scraper-dominated assemblages alongside butchered faunal remains [which was also noted for AYCC industries (Lemorini et al. 2016)]. The emergence of Quina scrapers in a context far from the Levant is intriguing, especially given that production and transportation of Middle Paleolithic Quina scrapers intensified during a period of environmental (climate) change that resulted in reliance on a new source of prey and greater Neanderthal mobility (Delagnes and Rendu 2011).

As in the Paleolithic Levant, Middle Paleolithic hunter-gatherers in Europe may have viewed mountains, massifs, and highlands as very special landscapes associated with lithic transportation, migratory routes, and the arrival of reindeer. The Quina-reindeer nexus was recognized long ago by Mellars (1965) and Chase (1986, 69–70) and more recently by Delagnes and Rendu (2011), who investigated a possible connection between Mousterian lithic systems and the faunal spectrum. The authors claim that Quina core reduction strategies and productivity, extensive use of ramification and recycling, blank versatility, and extended tool life-histories correspond to high technological mobility adapted for annual movement and targeting large migratory ungulates, predominantly reindeer herds. In the heartlands of the Quina Mousterian in southwestern France, Quina technology is recurrently associated with reindeer-dominated faunal assemblages (Costamagno et al. 2006; Discamps and Royer 2017; Frouin et al. 2017; Guérin et al. 2017; Morin et al. 2014; Niven et al. 2012; Richter et al. 2013). Studies addressing the European Middle Paleolithic variability in a biochronological framework highlighted the co-occurrence of reindeer remains and Quina technology (Discamps and Royer 2017; Discamps et al. 2011; Morin et al. 2014). These studies attribute the emergence of the Quina Mousterian to cold episodes, such as the Heinrich 6event, marked by a decline in ungulate biomass and the increased availability of reindeer, an animal well-adapted to the cold. However, absolute ages from several Quina sites suggest a longer duration of this entity, not strictly bounded by cold phases (Guerin et al. 2017; Richter et al. 2013). In southwestern France, reindeer dominate 24 out of 29 Quina stratigraphic units containing lithics and fauna, and 18 units correspond to specialized reliance on this taxon (Rendu et al. 2023). Reindeer zooarchaeological remains from Quina contexts indicate rigorous butchery, skinning, defleshing, and sinew removal. Bone shafts were broken for marrow extraction or shaped as retouchers (Castel et al. 2017; Costamagno et al. 2015).

Pleistocene reindeer ecology varied; some ecotypes were sedentary, and others annually migrated (Britton et al. 2011, Fontana 2017). However, it is important to note that migratory behavior, inherited by genetic transmission, may occur even among individuals in sedentary populations (Cavedon et al. 2022). A study of enamel strontium isotope ratios of MIS3 reindeer from Quina Mousterian site Abri du Maras demonstrated that reindeer populations with distinct seasonal ranges passed through the area before being succeeded by a more sedentary ecotype. The isotopic profiles of Quina Mousterian reindeer recovered from Chez-Pinuad Jonzac are very consistent indicating either acquisition during a single hunting event or similar migration routes (Britton et al. 2011, 182). The latter possibility is unsurprising in light of our knowledge of Alaskan caribou herds, which always returned to the same calving grounds annually before anthropogenic intrusions affected their routes (Miller et al. 2021). The herd engaged by Neanderthals at Jonzac migrated long distances across different geological terrains. The site is in a calcareous Turonian landscape, and its environment contains lower strontium values than those recorded in the reindeer remains. Therefore, Jonzac reindeer must have arrived from an environment with higher values. A brief macro-view of a geological survey reveals that mountainous and geomorphologically elevated areas, such as the Alps, Pyrenees, Massif Armoricain, Massif Centra, and the Central Mountain range, are all evident by exposed metamorphic and igneous formations of the Paleozoic Era. Modern caribou will often locate their calving grounds in either treeless tundra plains, inland islands, or elevated terrain to minimize predation risk (Bergerud et al. 2008). Since reindeer hunting at Jonzac occurred during fall and winter (Niven et al. 2012), we may roughly speculate that the site was situated within the reindeer winter range, with calving grounds located elsewhere, perhaps in one of the elevated regions.

The use of animal bones for retouching lithic implements is documented during the Quina Mousterian, where bone retouchers made of long ungulate (reindeer, but also other large ungulates) bones were employed for scraper maintenance operations (Castel et al. 2017; Costamagno et al. 2015; Rendu et al. 2023; Tartar et al. 2022). Costamagno et al. (2015) suggest that Neanderthals at Les Pradelles were familiar with the intrinsic characteristics of reindeer bones and intentionally selected suitable shaft fragments during the butchery. Moreover, they argue that Quina scrapers were usually retouched with a particular type of bone retoucher (Gr1), while non-Quina scrapers were maintained by other types of retouchers. Considering the evident use of Quina scrapers for butchering and processing reindeer, it is fair to expect that stone scrapers were involved in creating bone retouchers, which were consequently used to resharpen the scrapers. While bone retouchers remained at the butchery site, some scrapers were carried away (Costamagno et al. 2015), continuing to the next destination within the planned movement across the landscape. They could, for instance, be transported back to the campsite with edible reindeer portions or carried to another reindeer butchering event. It is noteworthy that bone retouchers also appeared in the late Lower Paleolithic Qesem Cave, together with fully-fledged Quina scrapers and specialized adaptation for specific ungulates (Blasco et al. 2013, 2016; Rosell et al. 2015, 2018), a combination of technological and behavioral novelties discussed above. We believe that the recurrent circulation of Quina groups, knapped stone, bone tools, and edible shares across the landscape may have formed an ontological relationship between these agencies.

Case Study 2: Caribou Mountain and Lithic Procurement in Labrador, Canada

Our second case study highlights the practical and perceptual connection between knapped stone, caribou, and mountains in the Holocene Eastern Canadian Arctic. The supreme importance of caribou deer for prehistoric and contemporary circumpolar Indigenous groups cannot be underestimated. Although prehistoric and historic Arctic cultures targeted a wide array of prey, certain groups (eg., Kivallirmiut [Caribou Inuit] and Innu [Montagnais-Naskapi] during historic times) relied almost exclusively on caribou predation for their existence (Loring 1988, 1997). Caribou provided fat and protein, bone, and antlers for tool-making, hide and fur for clothes, furniture, containers, structures, and other items, including materials for ritual purposes, such as shaman cloaks (Tanner 2021: Figure 1.12). The shaking tent, an Arctic ritual practiced in total darkness to communicate with animals and supernatural beings (Stépanoff 2019), was performed in a structure covered by caribou hides (Armitage 1992). In return, people who depended on caribou hunting perceived them as non-human persons with whom they shared the world and treated them with deep respect (Hill 2011). Inuit hunters donned precisely decorated caribou skin garments “to please the caribou” and attract the herds (Burnham 1992). Caribou also were butchered and shared with care, bringing groups together and boosting information exchange (Armitage 1992; Loring 1988; Tanner 2021). Caribou were and still are inextricably linked with the cosmology and ontology of Indigenous nations. It was reported that a decline in caribou availability, coupled with an issued ban on caribou hunting, severely harmed the social integrity, sharing habits, knowledge transmission mechanisms, and the collective well-being of current native populations in Eastern Canada (Borish et al. 2022).

The Innu on the Labrador Peninsula traditionally relied on caribou hunting as part of their subsistence and devoted great efforts to maintaining good relations with the caribou (Loring 1988; Tanner 2021, 34). At the beginning of the hunting season, the first hunted caribou individual was brought into the Innu dwellings for the whole group to admire. To ensure a successful hunt, the Innu carefully cleaned and scraped off the fat from caribou limb bones and prepared a substance made of oil and marrow extracted from the bones. This substance was imbued with “potency” and consumed during a “fat-oriented” feast (Tanner 2021, 33–34, 36). The Innu of Eastern Ungava believed that the master of all caribou lives on top of a mountain somewhere in the Torngat mountain range, north of today’s Nain (Figure 7). “According to Indian belief, there is a high mountain in the barren grounds of northeastern Labrador that is called the Caribou House, ah-tee-which-oo-ap, where the chief of all the deer lives. It is here they believe the caribou spend the summer” (Strong 1930, 2). Interestingly, the calving grounds of the George River herd are located next to the proposed “Caribou House” mountain and were respectfully referred to as “Caribou House calving grounds” by the zoological research group who studied the herd (Bergerud et al. 2008, 40). For thousands of years, caribou herds moved annually into and out of the “Caribou House” region through mountainous passes. Innu hunters who lived west of the George River used to observe the herds coming from the east or north and would plan their group’s movements according to caribou migratory patterns (Bergerud et al. 2008, 107–108; Loring 1997). Fundamental to Innu cosmology is the concept that harmful or disrespectful behavior towards the caribou would depress the availability of the whole species, and the caribou master would withdraw the herds back to the mountain (Tanner 2021). According to Strong (1930), the hunters he met claimed that scattered caribou bones angered the animals. The caribou then applied to their master, who in turn conveyed them into mountains unreachable to human hunters (Strong 1930, 2). The Innu used to separate bones and body parts of different animals and prevent them from getting loose by placing them on special elevated platforms (Tanner 2021, 30, 36). The perceptual connection between caribou and the mountain is manifested by a view shared by different Montagnais-Naskapi bands regarding a pure white mountainous range made of caribou fur. Other Montagnais-Naskapi believed the mountains were made of caribou fur and stone (Bergerud et al. 2008, 108–109).

Figure 7
figure 7

Map of Labrador. The circle marks the approximate zone of calving ground locations of the George River herd (modified after Bergerud et al. 2008; Taillon et al. 2012)

The cosmological connection between caribou herds and mountains, including the specific “Caribou House” peak, might have been complemented by another agency—stone. As Tanner (2021) noted, the “Caribou House” is located within the vicinity of Ramah Bay, where the homogenous semi-translucent grayish-white Ramah chert is available along the walls of a glacial cirque within a geological bed of the rare Nullataktok Formation (Loring 2002). This stone was transported 3,000 km far from the Labrador coast and used by various protohistoric and Indigenous cultures for a wide range of lithic trajectories, serving as a synchronic and diachronic cultural marker (Tanner 2021; Anstey and Renouf 2011; Loring 1988, 2002, 2017). It has been suggested that people of the Maritime Archaic complex five thousand years ago planned a seasonal long-distance journey to procure stone and hunt caribou on the coast of Ramah Bay (Fitzhugh 1985). Accessing the quarrying localities was not easy. The quarrying expedition would have to follow a stream and ascend a rigid mountainous terrain, eventually arriving at an iron-stained rocky creek painted in “brilliant blood red” (Loring 2002, 184). According to Loring, the combination of a saturated red landscape with the lithic source for hunting gear created a powerful association with the climax of the hunt. Ramah chert artifacts, including scrapers and caribou remains, were excavated from numerous historic and prehistoric sites in Labrador (Loring 1988, Stopp 1997). It can be safely assumed that tools made of Ramah chert were used for cutting, scraping, and cleaning long caribou bones, perhaps during feasts and other occasions. Given the all-around importance of animal fat for Arctic societies, the semi-translucent grayish-white Ramah chert was also endowed with related qualities since its appearance was associated with animal subcutaneous fat, preferably of caribou (Loring 1997, Tanner 2021, 37–38). Thus, using Ramah chert tools in the butchering and processing fat-bearing caribou must have been perceived as a reminder of the reciprocal relation between animal and stone. Another stone with “striking optical properties” is the snow-white Mistassini Quartzite, quarried from Lake Albanel in Central Quebec (McCaffrey 2011). Everything said about Ramah Chert is true for Mistassini, and it is easy to confuse the two (McCaffrey 2011, 161; Loring 2017). As McCaffrey (2011) writes, the Mistassini Quartzite is mentioned in the 16th-century report of a French cartographer, A. Thevet, who explored eastern Canada and described a native belief regarding a smoking mountain bearing stones gleaming with the color of the sky. McCaffrey suggests that if the identification is correct, this stone might have been associated with a unique mountain releasing/surrounded by smoke ascending to the skies and possibly treated as a means to access the upper world (McCaffrey 2011, 161).

For Indigenous societies dependent on animals and stone tools for their successful adaptation, stone acquisition, and use must have required intimate knowledge of the environment. As Loring writes regarding the use of Ramah chert by the Innu and the Inuit, “the souls of animals demanded respect, and one way to show respect was by crafting and incorporating beautiful elements into the construction of tools that were used upon them” (Loring 2017, 207).


Early humans may have formed existential relationships with the animals processed by their stone tools. In this paper, we examined the plausible notion that this human-animal-stone nexus extended into deep prehistoric times and was a pivotal factor in Paleolithic technological and cultural transformations. The Late Lower Paleolithic Levant is characterized by substantial technological innovations and by shifts in prey selection, which we believe were the result of fluctuations in the local availability of hunted game. In this paper, we examined a particular component in the Paleolithic toolkit, scrapers, which underwent a significant transformation with the initial development of Quina-like scrapers in the Late Acheulian, leading to a fully-fledged Quina cultural marker in the AYCC. The use of Quina scrapers in animal butchering and processing accompanied a growing dependence on systematic hunting of medium and small ungulates and further changes in local animal communities. Scraper production at Jaljulia and Qesem Cave involved continual use of non-local stone, quite possibly procured and transported from the Samarian highlands. The region may have acquired cosmological significance for Late Lower Paleolithic humans, recognized as the source of the wadi stream, a specific rock type, and fallow deer populations. Contemporary hunter-gatherers and traditional societies perceive the environment as a conglomerate of interrelated living beings, human and non-human persons, who share the world. Abundant evidence suggests that this worldview may be traced back as early as the Lower Paleolithic.