The hunters or the hunters: Human and hyena prey choice divergence in the Late Pleistocene Levant.

Hunting preferences reveal a great deal about the life of Paleolithic humans, and may reflect changes in human demography, technology, and adaptations to changing environments. However, the effects of hunting preferences and environmental availability are often conflated, stressing the need for comparisons to other predators that exploited the same environment. Manot Cave (Israel), preserved rich Early Upper Paleolithic (46-33 ka) human occupations, along with repeated spotted hyena (Crocuta crocuta) occupations, allowing us to compare anthropogenic and biogenic bone assemblages within the same space and time frame. We focused on the faunal remains retrieved in the middle of the cave (Area D), and conducted detailed taphonomic and zooarchaeological analyses. The Area D archaeofauna was dominated by Mesopotamian fallow deer (Dama mesopotamica) and mountain gazelle (Gazella gazella), which showed high abundance of carnivore damage. This and the carnivore-to-ungulate ratio, the presence of juvenile hyena and numerous coprolites match the criteria of a hyena den, confirming that the bone assemblage was created mainly by hyenas. Manot Area D thus reveals hyena prey choice in the Upper Paleolithic Galilee, which we then compared with human prey choice. Our results showed that hyena prey assemblages in Manot and elsewhere in the Levant were Dama-dominated while human assemblages were dominated by Gazella, demonstrating that hyenas and humans hunted different animal size groups, possibly in different habitats. We interpret this phenomenon as resulting from two possible scenarios: the emergence of projectile technology which may favor hunting in open environments, and niche partitioning derived by human-hyena competition. Hyenas were abundant and hunted unselectively while the anthropogenic record presents population turnovers and some dietary diversification. Whereas both scenarios are difficult to test directly with the available evidence, we argue that either one explains quite well the late Pleistocene archaeofaunal patterns in the Levantine record.

Early Upper Paleolithic subsistence in the Levant: Zooarchaeology of the Ahmarian-Aurignacian sequence at Manot Cave, Israel.

The Early Upper Paleolithic period in the Levant is essential in the studies of the establishment of modern human communities outside Africa, and corresponding archaeological evidence may be used to shed light on human ecology, economy and demography. Specifically, cultural differences between two Early Upper Paleolithic entities, the Early Ahmarian and the Levantine Aurignacian, raise the question of differing adaptations. In this article we use archaeofaunal remains from the Early Upper Paleolithic sequence at Manot Cave (Western Galilee, Israel), to track human hunting patterns, carcass transport and processing within the Early Ahmarian (46-42 ka) and Levantine Aurignacian (38-34 ka) phases. We test two hypotheses: 1) the Ahmarian and Aurignacian represent adaptations to different environments; and 2) the two entities differ in mobility patterns and site use. Our multivariate taphonomic analysis showed subtle differences in depositional processes between the two phases and demonstrated a primarily anthropogenic complex. In both phases, human subsistence was based on two ungulate species, mountain gazelle (Gazella gazella) and Mesopotamian fallow deer (Dama mesopotamica), with some contribution from birds, tortoises and small mammals. Among the gazelles, it appears that female herds were targeted, and that hunting took place close to the cave. The results of the research show great similarity in environmental exploitation between the Ahmarian and Aurignacian phases concerning prey spectrum and choice, carcass transport and processing. These patterns occupy a middle position between the Middle Paleolithic and the late Epipaleolithic of the region. Despite this, there are also several significant differences between the phases such as increased exploitation of small game (especially birds) and faster accumulation and higher densities of material in the Aurignacian. This may indicate greater occupation intensity during the Aurignacian compared to the Ahmarian, and thus could explain the outstanding character of this entity in the Levant.

Preliminary observations on the Levantine Aurignacian sequence of Manot Cave: Cultural affiliations and regional perspectives.

A well-preserved sequence of Early Upper Paleolithic (EUP) occupations has been revealed in the past decade in Manot Cave, the studies of which shed light on the cultural dynamics and subsistence patterns and paleoenvironment. Most intriguing is the series of overlying Levantine Aurignacian occupation layers, exposed near the entrance to the cave. Area E is considered the inner part of the main activity area in Manot Cave. Remains of intact combustion features, as well as numerous flint artifacts and faunal remains, were found, indicating a high level of preservation. Within a 2.5 m sequence, nine distinct occupation layers were defined. The presence of characteristic flint and osseous industries alongside a rich mollusk assemblage led to the initial association of the sequence as a whole to the Levantine Aurignacian. However, as research advanced and variability in the material culture became apparent, it became clear that a division of the sequence into two phases, early and late, is required. A preliminary study of the assemblage variability implies distinct changes in human behavior between the two phases. Most prominently, these are indicated by a change in bladelet production method and morphology alongside an increase in the significance of the bladelet component within the flint assemblage, the disappearance of composite osseous industries, and a steep decrease in mollusk shell representation in the late occupation phase. Radiocarbon dating indicates a short time span between the two phases. The earlier phase defined as, Levantine Aurignacian, was ascribed an age range of 38-34 ka cal BP with a more constrained age range of 37-35 ka cal BP suggested based on Bayesian models. In the late phase, which is temporarily referred to as "post-Levantine Aurignacian," an age range of 36-33 ka cal BP is suggested.

Personal ornaments from Hayonim and Manot caves (Israel) hint at symbolic ties between the Levantine and the European Aurignacian.

Situated at the crossroads of Africa and Eurasia, the Levant is a crucial region for understanding the origins and spread of Upper Paleolithic (UP) traditions associated with the spread of modern humans. Of the two local Early Upper Paleolithic technocomplexes, the Ahmarian and the Levantine Aurignacian, the latter appears to be unique in the endemic UP sequence, exhibiting greater similarity to the West European 'classic' Aurignacian than to the local preceding and proceeding UP entities. Previous publications have mostly focused on the similarities between the two lithic industries and less on studies conducted on Levantine Aurignacian bone tools and ornaments. Here, we present an archaeozoological, technological and use-wear study of ornaments on animal teeth from the Levantine Aurignacian layers at Manot and Hayonim caves (the Galilee, Israel). The selection of taxa, the choice of teeth, the mode of modification, and the use-wear analysis exhibit clear similarities with the European Aurignacian. This, with the technology of the osseous raw material exploitation, the presence of antler simple-base points, and some lithic typotechnological features, suggest a link between the symbolic spheres of the Levantine and the European Aurignacian cultural entities. Such similarity also supports some contribution of European Aurignacians groups to the local cultural entities, intermingling with the local material culture features.

Levantine cranium from Manot Cave (Israel) foreshadows the first European modern humans.

A key event in human evolution is the expansion of modern humans of African origin across Eurasia between 60 and 40 thousand years (kyr) before present (bp), replacing all other forms of hominins. Owing to the scarcity of human fossils from this period, these ancestors of all present-day non-African modern populations remain largely enigmatic. Here we describe a partial calvaria, recently discovered at Manot Cave (Western Galilee, Israel) and dated to 54.7 ± 5.5 kyr bp (arithmetic mean ± 2 standard deviations) by uranium-thorium dating, that sheds light on this crucial event. The overall shape and discrete morphological features of the Manot 1 calvaria demonstrate that this partial skull is unequivocally modern. It is similar in shape to recent African skulls as well as to European skulls from the Upper Palaeolithic period, but different from most other early anatomically modern humans in the Levant. This suggests that the Manot people could be closely related to the first modern humans who later successfully colonized Europe. Thus, the anatomical features used to support the 'assimilation model' in Europe might not have been inherited from European Neanderthals, but rather from earlier Levantine populations. Moreover, at present, Manot 1 is the only modern human specimen to provide evidence that during the Middle to Upper Palaeolithic interface, both modern humans and Neanderthals contemporaneously inhabited the southern Levant, close in time to the likely interbreeding event with Neanderthals.

Site occupation dynamics of early modern humans at Misliya Cave (Mount Carmel, Israel): Evidence from the spatial taphonomy of faunal remains.

Space use in Middle Paleolithic (MP) camps has been suggested as a source of information on the intensity and repetition of occupations and, by extension, of demographics. In the Levant, clear evidence for differential intrasite use and maintenance was important in viewing the late MP Neanderthal sites as base camps inhabited for a significant duration, relative to the Early MP (EMP). We test this model with the rich faunal assemblage from the EMP (>140 ka) site of Misliya Cave, Mount Carmel, Israel. Excavations in Misliya yielded a large and diverse lithic assemblage, combustion features, and a modern human maxilla, together with a large archaeofaunal assemblage that we use as a spatial marker. We analyzed the distribution of bone items with variable taphonomic properties (anthropogenic, biogenic, and abiotic bone-surface modifications) in a hearth-related context, both by comparing grid squares and point patterns. Both analyses are largely congruent. They indicate repeated and consistent use of the site's space that includes hearth-related consumption activities and peripheral activities further away, albeit with little evidence for site maintenance. Thus, the Misliya results display a mixed signal of clear around-the-fire pattern and differential space use in the EMP that is reminiscent of the much later MP sites in the Levant, but without the more elaborate camp maintenance. More case studies and detailed proxies are needed before we can fine-tune our understanding of camp structure in the Levantine MP and its bearing to site occupation dynamics.

Earliest floral grave lining from 13,700-11,700-y-old Natufian burials at Raqefet Cave, Mt. Carmel, Israel.

Flowering plants possess mechanisms that stimulate positive emotional and social responses in humans. It is difficult to establish when people started to use flowers in public and ceremonial events because of the scarcity of relevant evidence in the archaeological record. We report on uniquely preserved 13,700-11,700-y-old grave linings made of flowers, suggesting that such use began much earlier than previously thought. The only potentially older instance is the questionable use of flowers in the Shanidar IV Neanderthal grave. The earliest cemeteries (ca. 15,000-11,500 y ago) in the Levant are known from Natufian sites in northern Israel, where dozens of burials reflect a wide range of inhumation practices. The newly discovered flower linings were found in four Natufian graves at the burial site of Raqefet Cave, Mt. Carmel, Israel. Large identified plant impressions in the graves include stems of sage and other Lamiaceae (Labiatae; mint family) or Scrophulariaceae (figwort family) species; accompanied by a plethora of phytoliths, they provide the earliest direct evidence now known for such preparation and decoration of graves. Some of the plant species attest to spring burials with a strong emphasis on colorful and aromatic flowers. Cave floor chiseling to accommodate the desired grave location and depth is also evident at the site. Thus, grave preparation was a sophisticated planned process, embedded with social and spiritual meanings reflecting a complex preagricultural society undergoing profound changes at the end of the Pleistocene.

Intensification and sedentism in the terminal Pleistocene Natufian sequence of el-Wad Terrace (Israel).

Measuring subsistence intensification in the archaeofaunal record has provided strong evidence for socioeconomic shifts related to sedentarization in the terminal Pleistocene Mediterranean Basin, but the precise timing and scale of the intensification trend and its place in the evolution of settled societies remain contentious. New archaeofaunal data from the key Natufian sequence of el-Wad Terrace (Mount Carmel, Israel, ca. 15.0-11.7 ka [thousands of years ago]) is used here to clarify and contextualize paleoeconomy and mobility trends in the latest Pleistocene Levant, representing the culmination of Epipaleolithic subsistence strategies. Taphonomic variables serve as supplementary indicators of habitation function and occupation intensity along the sequence. At el-Wad, a very broad range of animals, mostly small to medium in size, were captured and consumed. Consumption leftovers were discarded in intensively occupied domestic spaces and suffered moderate attrition. The Early (ca. 15.0-13.7/13.0 ka) and Late (ca. 13.7/13.0-11.7 ka) Natufian phases display some differences in prey exploitation and taphonomic markers of occupation intensity, corresponding with other archaeological signals. We further set the intra-Natufian taxonomic and demographic trends in perspective by considering the earlier Epipaleolithic sequence of the same region, the Israeli coastal plain. Consequently, we show that the Early Natufian record constituted an important dietary shift related to greater occupation intensity and sedentarization, rather than a gradual development, and that the Late Natufian record appears to be maintaining, if not amplifying, many of these novel signals. These conclusions are important for understanding the mode and tempo of the transition to settled life in human evolution.

A series of Mousterian occupations in a new type of site: the Nesher Ramla karst depression, Israel.

We report the discovery of a new type of hominin site in the Levant, inhabited during MIS 6-5. The site, found within a karst depression at Nesher Ramla, Israel, provides novel evidence for Middle Paleolithic lifeways in an environmental and depositional setting that is previously undocumented in the southern Levant. The carbonate bedrock in the area is characterized by surface depressions formed by gravitational sagging of the rock into underlying karst voids. In one such depression, an 8 m thick sequence comprising rich and well-preserved lithic and faunal assemblages, combustion features, hundreds of manuports and ochre was discovered. Here we focus on the geological and environmental setting and present optically stimulated luminescence (OSL) ages for the 8 m sequence, aiming to place the site within a firm chronological framework and determine its significance for a more complete reconstruction of cultural developments in the Levantine Middle Paleolithic. To that end, preliminary results of the lithic and faunal studies are also presented.

Identification and quantification of projectile impact marks on bone: new experimental insights using osseous points.

Shifts in projectile technology potentially document human evolutionary milestones, such as adaptations for different environments and settlement dynamics. A relatively direct proxy for projectile technology is projectile impact marks (PIM) on archaeological bones. Increasing awareness and publication of experimental data sets have recently led to more identifications of PIM in various contexts, but diagnosing PIM from other types of bone-surface modifications, quantifying them, and inferring point size and material from the bone lesions need more substantiation. Here, we focus on PIM created by osseous projectiles, asking whether these could be effectively identified and separated from lithic-tipped weapons. We further discuss the basic question raised by recent PIM research in zooarchaeology: why PIM evidence is so rare in archaeofaunal assemblages (compared to other human-induced marks), even when they are explicitly sought. We present the experimental results of shooting two ungulate carcasses with bone and antler points, replicating those used in the early Upper Paleolithic of western Eurasia. Half of our hits resulted in PIM, confirming that this modification may have been originally abundant. However, we found that the probability of a skeletal element to be modified with PIM negatively correlates with its preservation potential, and that much of the produced bone damage would not be identifiable in a typical Paleolithic faunal assemblage. This quantification problem still leaves room for an insightful qualitative study of PIM. We complement previous research in presenting several diagnostic marks that retain preservation potential and may be used to suggest osseous, rather than lithic, projectile technology. Supplementary Information: The online version contains supplementary material available at 10.1007/s12520-024-01944-3.

Dating the Lower to Middle Paleolithic transition in the Levant: a view from Misliya Cave, Mount Carmel, Israel.

The transition from the Lower to the Middle Paleolithic in the Levant is a crucial event in human evolution, since it may involve the arrival of a new human population. In the current study, we present thermoluminescence (TL) dates obtained from 32 burnt flints retrieved from the late Lower Paleolithic (Acheulo-Yabrudian) and Early Middle Paleolithic (Mousterian) layers of Misliya Cave, Mount Carmel, Israel. Early Middle Paleolithic industries rich in Levallois and laminar products were assigned mean ages ranging from ~250 to ~160 ka (thousands of years ago), suggesting a production of this industry during MIS 7 and the early part of MIS 6. The mean ages obtained for the samples associated with the Acheulo-Yabrudian (strengthened by an isochron analysis) indicate a production of this cultural complex ~250 ka ago, at the end of MIS 8. According to the Misliya TL dates, the transition from the Lower to the Middle Paleolithic in the site took place at the limit MIS 8/7 or during the early part of MIS 7. The dates, together with the pronounced differences in lithic technology strongly suggest the arrival of a new population during this period.

Lower Paleolithic hominin ecology at the fringe of the desert: Faunal remains from Bizat Ruhama and Nahal Hesi, Northern Negev, Israel.

The Southern Levant is a pivotal area for the study of hominin paleoecology during the Lower Paleolithic, because of its location on the out-of-Africa dispersal route and its significant ecological diversity. Important information has been gained by archaeofaunal studies, which usually reveal that exploitation of diverse Mediterranean environments with woodlands, marshes and lake margins, represents a dominant subsistence strategy for Lower Paleolithic hominins. Here, we present new taxonomic and taphonomic data from two sites in the southern coastal plain of the Southern Levant, at the fringe of the Negev Desert: Bizat Ruhama (Early Pleistocene) and Nahal Hesi (Middle Pleistocene). The sites preserve anthropogenic faunas, with the former signaling a marrow-exploitation strategy, perhaps related to scavenging from carnivore kills, and the latter showing evidence for primary access to fleshed ungulate carcasses. The species composition of these Northern Negev sites is unique for the Levantine Lower Paleolithic in that these sites lack typical woodland and riparian species, probably indicating an open, relatively uniform environment with patchy water sources and trees, much like this semiarid region today. Bizat Ruhama and Nahal Hesi are among the only Levantine Lower Paleolithic faunas associated with such a setting, thereby widening the known spectrum of environments exploited by hominins in the region. It is suggested that the two sites, coupled with the nearby Late Pleistocene evidence, reflect a largely stable semiarid environment on the northwestern fringe of the Negev Desert throughout much of the Pleistocene.

Exploring exchange and direct procurement strategies for Natufian food processing tools of el-Wad Terrace, Israel.

We present the results of a detailed geochemical provenance study of 54 Natufian (ca. 15,000-11,700 cal. BP) basalt pestles from the site of el-Wad Terrace (EWT), Israel. It is the first time precise locations from where basalt raw materials were derived are provided. The results indicate that the Natufian hunter-gatherers used multiple sources of basaltic rocks, distributed over a large area surrounding the Sea of Galilee. This area is located at a considerable distance from EWT, ca. 60-120 km away, in a region where contemporaneous Natufian basecamps are few. We consider two possible models that suggest vehicles for the transportation of these artifacts to EWT, namely the exchange obtaining model (EOM) and the direct procurement model (DPM). We argue that these mechanisms are not mutually exclusive and may have operated together. We also suggest that at a time of increasing Natufian territoriality, a large area around the Sea of Galilee remained unclaimed. The paper concludes with a brief discussion of the implications for the two models. In particular, we note that the DPM implies that technological know-how for pestle production was maintained within the EWT community.

Squamate bone taphonomy: A new experimental framework and its application to the Natufian zooarchaeological record.

Squamate (lizard and snake) remains are abundant in the terminal Pleistocene Natufian archaeological sites of the Levant, raising the question of whether they constitute part of the broad-spectrum diet characteristic of this period. However, the role of squamates in Natufian diets remains unclear, as they are taphonomically under-studied. We conducted a series of experiments and actualistic observations that tested the impact of pre- and post-depositional processes on squamate vertebrae. We emphasized the multiple destruction processes that leave overlapping or altered marks on the bones, such as digestion marks that were modified by trampling. The resulting bone modification typology provides a tool for studying archaeological squamate remains. The experimental data were compared to the archaeological bone samples of the Natufian sequence of el-Wad Terrace (Mount Carmel, Israel, 15,000-12,000 cal BP). The Natufian squamate samples deviate from all actualistic ones in their lesser evidence of digestion and much greater indications for trampling, erosion and breakage. The taphonomic study, coupled with intra-site analysis, has unraveled the complex depositional history of el-Wad Terrace, enabling us to differentiate between cultural and non-cultural contexts and to identify possible human consumption of the European glass lizard and the large whip snake in the Natufian.

Versatile use of microliths as a technological advantage in the miniaturization of Late Pleistocene toolkits: The case study of Neve David, Israel.

The miniaturization of stone tools, as reflected through the systematic production of bladelets and bladelet tools (microliths), characterized many industries of the Late Pleistocene, with the Levantine Epipalaeolithic serving as a well-studied example. It is commonly held that microliths were used as modular inserts in composite projectiles, while their incorporation in other tools for different tasks is generally overlooked, the latter aspect being the main focus of this paper. We present here a more inclusive approach through a case study of the Geometric Kebaran (Middle Epipalaeolithic, ca. 18,500-15,000 cal BP) site of Neve David, Mount Carmel, Israel. Recent excavations at the site exposed a variety of features, and one well-preserved shallow pit provided a large lithic assemblage with ca. 90 microliths. We studied this assemblage using both the low- and high- magnification use-wear protocols, accompanied by a range of experiments. Our results show that a) the fragmentation rate is very high in this assemblage (ca. 90%), b) most of the microliths have identifiable use-wear, c) the microliths were commonly used as inserts in composite projectiles, d) many microliths were used for functions not related to weaponry and hunting, such as wood-working, weed harvesting and meat processing. These findings strongly support the suggestion that the small insets, regardless of their specific type (trapeze, rectangle, backed/retouched bladelet), were used in a wide variety of composite tools. We argue that such a versatile approach and flexibility in the use of microliths reflect a technological advantage where a minimal set of microlithic types, produced in large numbers, could provide the required elements for weapons, as well as for a variety of cutting, processing and harvesting tools needed for mundane tasks at a large Middle Epipalaeolithic camp.

Persistent Neanderthal occupation of the open-air site of 'Ein Qashish, Israel.

Over the last two decades, much of the recent efforts dedicated to the Levantine Middle Paleolithic has concentrated on the role of open-air sites in the settlement system in the region. Here focus on the site of 'Ein Qashish as a cases study. Located in present-day northern Israel, the area of this site is estimated to have been >1300 m2, of which ca. 670 were excavated. The site is located at the confluence of the Qishon stream with a small tributary running off the eastern flanks of the Mt. Carmel. At the area of this confluence, water channels and alluvial deposits created a dynamic depositional environment. Four Archaeological Units were identified in a 4.5-m thick stratigraphic sequence were dated by Optically Stimulated Luminescence (OSL) to between-71 and 54 ka, and probably shorter time span-~70-~60 ka. Here we present the diverse material culture remains from the site (lithics, including refitted sequences; modified limestone pieces; molluscs; faunal remains) against their changing paleogeographic backdrop. Skeletal evidence suggests that these remains were associated with Neanderthals. The large-scale repeated accumulation of late Middle Paleolithic remains in the same place on the landscape provides a unique opportunity to address questions of occupation duration and intensity in open-air sites. We find that each occupation was of ephemeral nature, yet presents a range of activities, suggesting that the locale has been used as a generalized residential site rather than specialized task-specific ones. This role of 'Ein Qashish did not change through time, suggesting that during the late Middle Paleolithic settlement system in this part of the southern Levant were stable.

The Emergence of Animal Management in the Southern Levant.

Our compilation of zooarchaeological data from a series of important archaeological sites spanning the Epipaleolithic through Pre-Pottery Neolithic B periods in the Mediterranean Hills of the southern Levant contributes to major debates about the beginnings of ungulate management in Southwest Asia. The data support an onset of ungulate management practices by the Early PPNB (10,500-10,000 cal. BP), more than 500 years earlier than previously thought for this region. There is a clear developmental connection between reduced hunting intensity and the uptake of ungulate management, confirming that this process began in response to local, density-dependent demographic factors. The early process of goat domestication in the southern Levant appears to have been overwhelmingly local. This may have been true for cattle and pigs as well. Nevertheless, the loose synchrony of animal management trends across Southwest Asia was undoubtedly enabled by large-scale social networks that transmitted knowledge. The results add to growing evidence that animal management processes followed multiple regional evolutionary pathways within the Fertile Crescent.

The earliest modern humans outside Africa.

To date, the earliest modern human fossils found outside of Africa are dated to around 90,000 to 120,000 years ago at the Levantine sites of Skhul and Qafzeh. A maxilla and associated dentition recently discovered at Misliya Cave, Israel, was dated to 177,000 to 194,000 years ago, suggesting that members of the Homo sapiens clade left Africa earlier than previously thought. This finding changes our view on modern human dispersal and is consistent with recent genetic studies, which have posited the possibility of an earlier dispersal of Homo sapiens around 220,000 years ago. The Misliya maxilla is associated with full-fledged Levallois technology in the Levant, suggesting that the emergence of this technology is linked to the appearance of Homo sapiens in the region, as has been documented in Africa.

Radiocarbon chronology of Manot Cave, Israel and Upper Paleolithic dispersals.

The timing of archeological industries in the Levant is central for understanding the spread of modern humans with Upper Paleolithic traditions. We report a high-resolution radiocarbon chronology for Early Upper Paleolithic industries (Early Ahmarian and Levantine Aurignacian) from the newly excavated site of Manot Cave, Israel. The dates confirm that the Early Ahmarian industry was present by 46,000 calibrated years before the present (cal BP), and the Levantine Aurignacian occurred at least between 38,000 and 34,000 cal BP. This timing is consistent with proposed migrations or technological diffusions between the Near East and Europe. Specifically, the Ahmarian could have led to the development of the Protoaurignacian in Europe, and the Aurignacian in Europe could have spread back to the Near East as the Levantine Aurignacian.