Cervidae antlers exploited to manufacture prehistoric tools and hunting implements as a reliable source of ancient DNA.

Antler is one of the primary animal raw materials exploited for technical purposes by the hunter-gatherer groups of the Eurasian Upper Palaeolithic (UP) all over the ecological range of deers, and beyond. It was exhaustively employed to produce one of the most critical tools for the survival of the UP societies: hunting weapons. However, antler implements can be made from diverse deer taxa, with different ecological requirements and ethological behaviours. Identifying the antler's origin at a taxonomic level is thus essential in improving our knowledge of humans' functional, practical and symbolic choices, as well as the human-animal interface during Prehistoric times. Nevertheless, palaeogenetics analyses have focused mainly on bone and teeth, with genetic studies of antler generally focused on modern deer conservation. Here we present the results of the first whole mitochondrial genome ancient DNA (aDNA) analysis by means of in-solution hybridisation capture of antlers from pre-Holocene archaeological contexts. We analysed a set of 50 Palaeolithic and Neolithic (c. 34-8ka) antler and osseous objects from South-Western Europe, Central Europe, South-Western Asia and the Caucasus. We successfully obtained aDNA, allowing us to identify the exploited taxa and demonstrate the archaeological relevance of those finds. Moreover, as most of the antlers were sampled using a minimally-invasive method, further analyses (morphometric, technical, genetic, radiometric and more) remain possible on these objects.

Cultural Dynamics in the Levantine Upper Paleolithic, ca. 40-33 ky BP: Insights Based on Recent Advances in the Study of the Levantine Aurignacian, the Arkov-Divshon, and the Atlitian.

The chrono-cultural sequence of the Levantine Upper Paleolithic went through several major revisions during approximately a century of focused research, each revision contributing to shedding light on the mosaic of cultural entities and the complex social and cultural dynamics composing the Levantine Upper Paleolithic. The current state of research suggests the co-inhabitance of two cultural groups: the Early Ahmarian and the Levantine Aurignacian. Two other cultural entities, the Arkov-Divshon and the Atlitian, are regarded as younger manifestations and were tentatively suggested to relate to the Levantine Aurignacian. This paper presents a research synthesis of two case studies: Manot Cave, located in western Galilee, Israel, and Nahal Rahaf 2 Rockshelter in the Judean Desert. The application of high-resolution excavation methods, alongside detailed documentation of the stratigraphy and site-formation processes and wide-scale radiocarbon-based absolute dating, marked these sites as ideal for chrono-cultural study through the analyses of flint industries. The results indicate a clear distinction between the Levantine Aurignacian and the Arkov-Divshon/Atlitian industries and a chronological overlap between the Arkov-Divshon, Levantine Aurignacian, and possibly with the Early Ahmarian. Subsequently, we suggest another revision of the currently accepted chrono-cultural model: not two, but at least three cultural entities co-inhabited the Levant at ca. 40-30 ky cal BP. This study further suggests an evolvement of the Atlitian flint industries from the Arkov-Divshon and stresses the foreign cultural features of the Levantine Aurignacian. These results were used to construct an updated model of migration and possible interaction patterns.

Early Upper Paleolithic cultural variability in the Southern Levant: New evidence from Nahal Rahaf 2 Rockshelter, Judean Desert, Israel.

The Levantine Early Upper Paleolithic (ca. 45-30 ka) has been a focus of research because of its unique position as a conduit of human, flora, and fauna species between Africa and Eurasia. Studies have mainly focused on the Early Ahmarian and Levantine Aurignacian, two entities, the former endemic and the latter foreign, which are considered to have coinhabited the region during that period. However, other cultural entities, such as the Atlitian in the Mediterranean region and the Arkov-Divshon in the arid regions of the southern Levant received less attention, and accordingly, suffer from broad definitions and chronological insecurity. These cultures hold potential insights regarding nuanced adaptations, reciprocal influences, and diachronic assimilation processes. The recently discovered site of Nahal Rahaf 2 Rockshelter in the Judean Desert provides integral information on one of these entities-the Arkov-Divshon. Two excavation seasons revealed a sequence of archaeological layers, with lithic assemblages in which laterally carinated items were prominent. Alongside rich faunal assemblages, other components of the material culture include perforated marine shells and bone tools, marking the first association of these elements with Arkov-Divshon and implying some degree of contact with the Mediterranean regions of the Levant. Good preservation of organic materials allowed radiocarbon dating of the human occupation at the site to ca. 37.5-34.0 ka cal BP, indicating chronological overlap with the Levantine Aurignacian, and possibly also with the latest phases of the Early Ahmarian. Thus, challenging the validity of the widely accepted 'Two Tradition' Model of the Levantine Upper Paleolithic. Lithic analyses suggest the use of one main reduction sequence and the primary production of bladelets from carinated items. Faunal remains suggest targeted hunting of ibex and gazelle. Botanical remains and sedimentary analyses suggest roughly similar environmental conditions, with a possible woodier environment in the surroundings of the site.

Expansion of eastern Mediterranean Middle Paleolithic into the desert region in early marine isotopic stage 5.

Marine Isotopic Stage 5 is associated with wetter climatic conditions in the Saharo-Arabian deserts. This stage also corresponds to the establishment of Middle Paleolithic hominins and their associated material culture in two geographical provinces in southwest Asia-the Eastern Mediterranean woodland and the Arabian Peninsula desert. The lithic industry of the Eastern Mediterranean is characterized by the centripetal Levallois method, whereas the Nubian Levallois method characterizes the populations of the Arabian desert. The Negev Desert, situated between these regions is a key area to comprehend population movement in correlation to climatic zones. This investigation addresses the nature of the Middle Paleolithic settlement in the Negev Desert during MIS 5 by studying the site of Nahal Aqev. High resolution chronological results based on luminescence dating and cryptotephra show the site was occupied from MIS 5e to MIS 5d. The lithic industries at Nahal Aqev are dominated by centripetal Levallois core method. These data demonstrate that Nahal Aqev is much closer in its cultural attributes to the Eastern Mediterranean Middle Paleolithic than to the Arabian Desert entity. We conclude that Nahal Aqev represents an expansion of Middle Paleolithic groups from the Mediterranean woodland into the desert, triggered by better climatic conditions. These groups possibly interacted with hominin groups bearing the Nubian core tradition from the vast region of Arabia.

The earliest Pleistocene record of a large-bodied hominin from the Levant supports two out-of-Africa dispersal events.

The paucity of early Pleistocene hominin fossils in Eurasia hinders an in-depth discussion on their paleobiology and paleoecology. Here we report on the earliest large-bodied hominin remains from the Levantine corridor: a juvenile vertebra (UB 10749) from the early Pleistocene site of 'Ubeidiya, Israel, discovered during a reanalysis of the faunal remains. UB 10749 is a complete lower lumbar vertebral body, with morphological characteristics consistent with Homo sp. Our analysis indicates that UB-10749 was a 6- to 12-year-old child at death, displaying delayed ossification pattern compared with modern humans. Its predicted adult size is comparable to other early Pleistocene large-bodied hominins from Africa. Paleobiological differences between UB 10749 and other early Eurasian hominins supports at least two distinct out-of-Africa dispersal events. This observation corresponds with variants of lithic traditions (Oldowan; Acheulian) as well as various ecological niches across early Pleistocene sites in Eurasia.

The absolute chronology of Boker Tachtit (Israel) and implications for the Middle to Upper Paleolithic transition in the Levant.

The Initial Upper Paleolithic (IUP) is a crucial lithic assemblage type in the archaeology of southwest Asia because it marks a dramatic shift in hominin populations accompanied by technological changes in material culture. This phase is conventionally divided into two chronocultural phases based on the Boker Tachtit site, central Negev, Israel. While lithic technologies at Boker Tachtit are well defined, showing continuity from one phase to another, the absolute chronology is poorly resolved because the radiocarbon method used had a large uncertainty. Nevertheless, Boker Tachtit is considered to be the origin of the succeeding Early Upper Paleolithic Ahmarian tradition that dates in the Negev to ∼42,000 y ago (42 ka). Here, we provide 14C and optically stimulated luminescence dates obtained from a recent excavation of Boker Tachtit. The new dates show that the early phase at Boker Tachtit, the Emirian, dates to 50 through 49 ka, while the late phase dates to 47.3 ka and ends by 44.3 ka. These results show that the IUP started in the Levant during the final stages of the Late Middle Paleolithic some 50,000 y ago. The later IUP phase in the Negev chronologically overlaps with the Early Upper Paleolithic Ahmarian of the Mediterranean woodland region between 47 and 44 ka. We conclude that Boker Tachtit is the earliest manifestation of the IUP in Eurasia. The study shows that distinguishing the chronology of the IUP from the Late Middle Paleolithic, as well as from the Early Upper Paleolithic, is much more complex than previously thought.

In search of modern humans and the Early Upper Paleolithic at Manot Cave: An overview.

Manot Cave is a unique relict karst cave located in the western Galilee, north-western Israel. The cave was inhabited from the Late Middle Paleolithic through the Early Upper Paleolithic (EUP) periods until its main entrance collapsed, ca. 30 ka. The cave consists of an elongated main hall and two side chambers. The topography of the main hall consists of a steep talus inclining from the original entrance of the cave to the center, a plane area at the lowermost point of the main hall, and a smaller talus inclining from the eastern end of the cave. Nine field seasons (2010-2018) have been conducted so far at the cave. The excavations revealed dense accumulations of EUP deposits near the cave entrance (areas E and I), at the center (area D), at the base of the western talus (area C), and in the plane area (area A). This introductory article describes the cave and its characteristics and provides a background for various contributions in the special issue, devoted to Manot Cave.

The dental remains from the Early Upper Paleolithic of Manot Cave, Israel.

This study presents the dental remains discovered at Manot Cave (MC), Western Galilee, Israel. The cave contains evidence for human occupation during the Early Upper Paleolithic period (46-33 ka) mainly of Early Ahmarian (∼46-42 ka) and Levantine Aurignacian (∼38-34 ka) cultural levels. Six teeth (three deciduous and three permanent) were found at the site, of which four could be thoroughly analyzed. The morphology of the teeth was qualitatively described and analyzed using traditional and geometric morphometric methods. A large comparative sample was used in order to assess the morphological affiliation of the Manot specimens with other Homo groups. The results provided equivocal signals: the upper first premolar (MC-9 P3) is probably modern human; the upper deciduous second molar (MC-10 dm2) and the upper second permanent molar (MC-8 M2) might be modern humans; the lower second deciduous molar (MC-7 dm2) might be Neanderthal. Owing to the small sample size and the almost total lack of distinctive characteristics, our outcome could not supply conclusive evidence to address the question of whether Manot Aurignacian population came from Europe or descended from the local Ahmarian population.

Early Upper Paleolithic human foot bones from Manot Cave, Israel.

The transition from the Middle Paleolithic to the Upper Paleolithic in the Levant represents a major event in human prehistory with regards to the dispersal of modern human populations. Unfortunately, the scarcity of human remains from this period has hampered our ability to study the anatomy of Upper Paleolithic populations. This study describes and examines pedal bones recovered from the Early Upper Paleolithic period at Manot Cave, Israel, from 2014 to 2017. The Manot Cave foot bones include a partial, left foot skeleton comprising a talus, a calcaneus, a cuboid, a first metatarsal, a second metatarsal, a fifth metatarsal, and a hallucal sesamoid. All these remains were found in the same archaeological unit of the cave and belong to a young adult. Shape and size comparisons with Neanderthals, Anatomically Modern Human and modern human foot bones indicate a modern human morphology. In some characteristics, however, the Manot Cave foot bones display a Neanderthal-like pattern. Notably, the Manot Cave foot is remarkable in its overall gracility. A healed traumatic injury in the second metatarsal (Lisfranc's fracture) is most likely due to a remote impact to the dorsum of the foot. This injury, its subsequent debility, and the individual's apparent recovery suggest that the members of the Manot Cave community had a supportive environment, one with mutual responsibilities among the members.

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.

Karst terrain in the western upper Galilee, Israel: Speleogenesis, hydrogeology and human preference of Manot Cave.

A karst survey of the western upper Galilee in Israel shows that karst has been a dominant geomorphic factor throughout the Cenozoic. We discuss the geomorphic character of Manot Cave on the background of other karst features of the region, in order to decipher the preferences of the humans who favored this cave over others. Tens of caves distributed over the study area demonstrate that phreatic and hypogene isolated voids and conduit segments are more abundant than vadose shafts, sinking stream caves and spring caves, although all these types are present. Most caves belong to old stages of landform development, prior to Plio-Pleistocene uplift and stream entrenchment. Manot Cave is a relict chamber cave, which corresponds to a plaeo-water table and the erosion plain above it. Subaerial denudation and slope processes have opened the cave to the surface during the mid-late Pleistocene. Manot Cave is compared with other caves in the region, demonstrating its unique character. It may have been selected due to the small entrance facing to the SW, and the large inner chamber which could be used for non-domestic purposes. This suggests a possible role of a unique behavioral and cultural suite of characters which influenced hominin preferences. The cave was used by hominins and animals until being closed again by colluvium and possibly collapse, ∼30 ka. Clastic, chemical, archaeozoological and anthropogenic accumulations reflect the various stages of cave development and gradual sealing of the entrance.

Climatic and environmental conditions in the Western Galilee, during Late Middle and Upper Paleolithic periods, based on speleothems from Manot Cave, Israel.

Early Ahmarian, Levantine Aurignacian and Post-Levantine Aurignacian archeological assemblages show that the karstic Manot Cave, located 5 km east of the Mediterranean coast in the Western Galilee region of Israel, was intensively occupied during the Early Upper Paleolithic. The coexistence of these rich archaeological layers with speleothems in Manot Cave provides a window of opportunity for determining the relationships between climatic conditions and the nature of human activity and mobility patterns in the Western Galilee region during the Early Upper Paleolithic period. This study, based on four stalagmites that grew almost continuously from ∼75 to 26.5 ka, covers most of the last glacial, and overlaps with the human occupation of the cave. The speleothems oxygen (δ18O) and carbon (δ13C) isotopic records indicate that climate and environmental conditions fluctuated during the last glacial, some of which correspond with Dansgaard-Oeschger (D-O) cycles 12, 10, 7 and Heinrich (H) events VI and V. Consistent with independent evidence from botanic and faunal remains, these climatic shifts brought about significant environmental changes in the region, ranging from dominant thick Mediterranean forest to more open landscape. A good correlation with less negative δ13C values is most pronounced during the Early Ahmarian time period, but there was also a change to less negative δ13C values during the Levantine Aurignacian and Post-Levantine Aurignacian industries in the Levant. These positive δ13C shifts suggest that environmental transformation towards a more open grassy landscape dominated by C4 vegetation might have played an important role in the development of these cultural entities (mainly the Early Ahmarian) in Manot Cave region.

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.

Site formation processes at Manot Cave, Israel: Interplay between strata accumulation in the occupation area and the talus.

Manot Cave contains important human fossils and archaeological assemblages related to the origin and dispersal of anatomically modern humans and the Upper Paleolithic period. This record is divided between an elevated in situ occupation area and a connecting talus. We, thus, investigated the interplay between the accumulation of the sediments and their associated artifacts in the occupation areas and the translocation of part of these sediments and artifacts down the talus. We examined the lithostratigraphy of two excavation locations in the occupation area (areas E and I), and two in the talus (areas C and D). We also assessed the diagenetic processes that have affected all these areas. A linear array of stalagmites and stalactites separates the occupation area from the talus, demarcating a major topographic barrier between the two. We infer that during human occupation, sediment accumulation of soil, wood ash, and bone was rapid and that some sediments with their associated artifacts overflowed the barrier and translocated down the talus. During periods of nonoccupation, the ash in the occupation area partially dissolved owing to the release of acid from the degrading bat and bird guano, and the layer thicknesses decreased. The south side of the talus (area C) has a normally stratified archaeological record, with the older archaeological materials underlying the younger materials. This suggests that the barrier between the occupation area and area C was relatively shallow and allowed a fairly continuous sediment accumulation in the talus. In the central part of the talus (area D), the stratigraphy is complex and shows mixing, presumably owing to the steep underlying bedrock topography and the mixing that occurs when sediments move down a steep slope. Finally, the distribution of secondary phosphates is consistent with the location of a main cave entrance to the south of the Paleolithic occupation area.

The Marine Isotope Stage 3 landscape around Manot Cave (Israel) and the food habits of anatomically modern humans: New insights from the anthracological record and stable carbon isotope analysis of wild almond (Amygdalus sp.).

The excavation of Manot Cave (Israel) reveals intensive occupation during the Early Upper Paleolithic and provides the first continuous set of anthracological data available for the Ahmarian, Levantine Aurignacian and post-Levantine Aurignacian periods. The paper aims to study the vegetal landscape around Manot Cave in the context of climate changes that characterized the last part of the Marine Isotope Stage 3 (MIS 3) and to address the issue of firewood and food procurement among Paleolithic hunter-gatherers. Charcoal samples recovered from the archaeological layers at Manot Cave shed light on the fuel and food procurement strategies while radiocarbon dating and stable carbon isotope analysis (Δ13C) of selected charcoals provide information about the ancient climate. The results show that five woody taxa were exploited at the site; Amygdalus sp. was the most common species, whereas Quercus ithaburensis, Tamarix sp., Pomoideae indet., and Pistacia atlantica were relatively rare. The representations of the recovered wooden species suggest that an open forest of almonds and oaks existed in the area during MIS 3. Radiocarbon dating of Amygdalus sp. charcoals, coupled with stable carbon isotope analysis (Δ13C) of modern and archaeological Amygdalus sp. clearly indicate variations in rainfall that could have decreased the density of tree cover. These analyses provide high-resolution data on the climate changes affecting the surroundings of Manot Cave between ∼46 and 28 ka cal BP and indicate two drier phases corresponding to the Ahmarian and post-Levantine Aurignacian cultures while a more humid period identified during the Levantine Aurignacian.

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.