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

Journal of Human Evolution xxx (xxxx) xxx

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Early Upper Paleolithic subsistence in the Levant: Zooarchaeology of the AhmarianeAurignacian sequence at Manot Cave, Israel Reuven Yeshurun a, *, Nehora Schneller-Pels a, Omry Barzilai b, Ofer Marder c a

Zinman Institute of Archaeology, University of Haifa, Mount Carmel, Haifa, 3498838, Israel Archaeological Research Department, Israel Antiquities Authority, POB 586, Jerusalem, 91004, Israel c Department of Bible, Archaeology and the Ancient Near East, Ben-Gurion University of the Negev, PO Box 653, Beer-Sheva, 8410501, Israel b

a r t i c l e i n f o

a b s t r a c t

Article history: Received 5 November 2018 Accepted 10 May 2019 Available online xxx

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 (46e42 ka) and Levantine Aurignacian (38e34 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. © 2019 Elsevier Ltd. All rights reserved.

Keywords: Early Ahmarian Levantine Aurignacian Gazelle Fallow deer Two-tradition model Taphonomy

1. Introduction The Early Upper Paleolithic (EUP) in the Levant, spanning ca. 46e33 ka (Alex et al., 2017) is composed of (at least) two cultural traditions, the Early Ahmarian and the Levantine Aurignacian. The recognition, definition and interpretation of these two entities known as the “two-tradition model” have undergone revisions during more than 40 years of research (e.g., Gilead, 1981, 1991; Marks, 1981, 2003; Williams, 2006; Bar-Yosef and Belfer-Cohen, 2010; Belfer-Cohen and Goring-Morris, 2017). It is currently

* Corresponding author. E-mail address: [email protected] (R. Yeshurun).

accepted that the Early Ahmarian entity commenced before the Levantine Aurignacian and was much more widespread geographically and temporally. Within this model, the Early Ahmarian has been said to represent an autochthonous Levantine entity, possibly developed from the Initial Upper Paleolithic, the MIS3 expansion of modern humans from Africa to the rest of the world (e.g., Kuhn et al., 2009; Kadowaki, 2013; Hublin, 2015; Goring-Morris and Belfer-Cohen, 2018). Conversely, the Levantine Aurignacian sensu stricto, which shares some cultural traits and symbolic “emblems” with the European Aurignacian, was seen as a relatively brief habitation of the Levant by human groups coming from the northern Mediterranean Basin (e.g., Belfer-Cohen and BarYosef, 1981; Bar-Yosef and Belfer-Cohen, 2010; Belfer-Cohen and Goring-Morris, 2014; Tejero et al., 2016, 2018).

https://doi.org/10.1016/j.jhevol.2019.05.007 0047-2484/© 2019 Elsevier Ltd. All rights reserved.

Please cite this article as: Yeshurun, R et al., Early Upper Paleolithic subsistence in the Levant: Zooarchaeology of the AhmarianeAurignacian sequence at Manot Cave, Israel, Journal of Human Evolution, https://doi.org/10.1016/j.jhevol.2019.05.007

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R. Yeshurun et al. / Journal of Human Evolution xxx (xxxx) xxx

The cultural division and chronology of the Levantine EUP record was a subject of recent debate due to mismatching results from northern and southern Levantine sites (e.g., Marks, 2003; Douka et al., 2013; Kadowaki et al., 2015; Bretzke et al., 2017; but see; Bosch et al., 2015a; Alex et al., 2017). Be that as it may, the ecological and technological adaptations of the Early Ahmarian and Levantine Aurignacian have not figured prominently in the EUP literature (see papers in Goring-Morris and Belfer-Cohen, 2003). A few studies have considered ecological models incorporating adaptations (e.g., raw material procurement, lithic economizing behaviors and prey-choice patterns) in interpreting Levantine Upper Paleolithic occurrences (e.g., Kaufman, 1988; Williams, 2006; Kuhn et al., 2009; Stiner, 2009; Stutz et al., 2015; Parow-Souchon, 2016; Leder, 2018). This situation partly results from the fact that wellpreserved, fully collected and contextualized faunal assemblages from Levantine EUP contexts are rare for various historical and taphonomic reasons (surveyed in Rabinovich, 2003). In this article we employ the above-mentioned version of the “two-tradition model” as a working hypothesis and compare the subsistence and mobility of the Early Ahmarian (henceforth Ahmarian) and the Levantine Aurignacian (henceforth Aurignacian) in Manot Cave, northern Israel (Fig. 1), using zooarchaeological remains. We further aim to place both assemblages in evolutionary perspective, i.e., to compare the poorly known EUP zooarchaeological patterns to those in the Middle Paleolithic and Epipaleolithic of the same region. Manot Cave is an excellent case study for deciphering subsistence behavior in relation to the changing environments. First, the site yielded rich stratified archaeofaunas dated to the Ahmarian and Aurignacian entities (Hershkovitz et al., 2015; Barzilai et al., 2016; Alex et al., 2017; Marder et al., 2017), making it one of the few EUP localities in the Levant with good faunal preservation, and the only one where well-collected faunal assemblages from both traditions are currently available for research (see Rabinovich,

2003, 2017). Second, independent paleoenvironmental proxies are available from the cave itself, enabling the reconstruction of the local environment from multiple sources (Caracuta et al., 2019 submitted; Comay et al., 2019 submitted; Yasur et al., 2019). Lastly, the cave yielded hyena den deposits that are mostly spatially and taphonomically separated from the anthropogenic deposits (Fig. 1b). Studying the hyena hunting patterns provided an independent proxy for the availability of game animals around the cave during the EUP (Orbach and Yeshurun, 2019). Altogether, EUP hunting patterns at Manot Cave can be contrasted with the hyena prey choice and the paleoenvironmental indicators to tease out human prey choice and subsequently test the following questions: 1) do the Ahmarian and Aurignacian of the Mediterranean Levant represent adaptations to different environments?; and 2) do the two entities differ in mobility patterns and site use? These hypotheses may or may not be mutually exclusive. To examine these questions, we first evaluate the taphonomy of the remains, to verify that the inter-tradition comparison is valid and to reconstruct bone depositional histories. We then consider the spectrum of prey animals, indications for carcass transport and processing, the age and sex profiles and trends in body-size to shed light on human hunting patterns in the two EUP entities in the cave. 1.1. Theoretical framework The mobility patterns of human foragers, including landscape use and duration of stay in base camps, are a central component in their way of life (Binford, 1980; Kelly, 1992). Cultural remains from Levantine UP sites have been assumed to indicate brief and repetitive occupations, reflecting a high degree of mobility in a residential, circulating pattern (e.g., Marks and Friedel, 1977; BelferCohen and Goring-Morris, 2014). Archaeofaunal remains constitute an important source of information on mobility, intensity of habitation, land use and

Figure 1. (a) Location of Manot Cave and other sites mentioned in the text; (b) Plan of the cave showing the location of Area C, with the sample squares highlighted red. Area D is the hyena den deposits; (c) The studied profile.

Please cite this article as: Yeshurun, R et al., Early Upper Paleolithic subsistence in the Levant: Zooarchaeology of the AhmarianeAurignacian sequence at Manot Cave, Israel, Journal of Human Evolution, https://doi.org/10.1016/j.jhevol.2019.05.007

R. Yeshurun et al. / Journal of Human Evolution xxx (xxxx) xxx

environmental adaptation. Tracking these patterns via zooarchaeology has long been recognized as a powerful explanatory factor by which to interpret changes in the Paleolithic record. The zooarchaeology of the late Pleistocene Mediterranean Basin has made extensive use of human behavioral ecology models, especially prey ranking, to assess the degree of mobility and siteoccupation intensity (Stiner et al., 1999; Stiner and Munro, 2002, 2011; Stiner, 2005; Stutz et al., 2009; Starkovich, 2017). The model that has emerged ranks resources according to the benefits obtained in taking them against the costs incurred. High-ranked resources may simply have been the game animals that provided the most meat and fat. Various factors that increased these species’ cost of capture and processing, such as agility, aggressive behavior, decreasing encounter rates or a socially derived ban, may have lowered the ranking accordingly and caused a shift in hunting patterns. The model posits that in the long run and over the timeaveraged units that are usually used in archaeology, human foragers tended to take the highest-ranked resources first, ignoring the low-ranked ones. Only when high-ranked game were less available or became costlier, did lower-ranked resources become regularly integrated in the diet (MacArthur and Pianka, 1966; Stephens and Krebs, 1986). Here we follow a well-known application of the diet-breadth model to the Levantine Paleolithic context (Stiner, 2005). It posits that larger ungulates (fallow deer, red deer and aurochs) were preferred to smaller ones (gazelle), prime-aged individual ungulates were preferred to juveniles, ungulates in general were preferred to most small game species, and slow, small game species (tortoises) were preferred to fast game species (hares and birds). Deviations from these assumptions during the late Pleistocene have been interpreted as foraging intensification. Depending on context, such intensification was considered to reflect regional human population increase from the earlier to later Middle Paleolithic and during the Upper Paleolithic (Stiner et al., 1999), overhunting in the late Middle Paleolithic (Speth and Clark, 2006), shifts in the sexual division of labor in the Upper Paleolithic (Kuhn and Stiner, 2006) and a profound change in settlement patterns in the late Epipaleolithic (Munro, 2009; Yeshurun et al., 2014a). This concept has been used to examine not only species choice but also the intensity of carcass processing, as humans will only invest in costly, laborintensive processing when the nutrients cannot be obtained from less costly sources (Munro and Bar-Oz, 2005; Manne and Bicho, 2009; Hodgkins et al., 2016). Additional lines of evidence for higher site-occupation intensity are increased quantities of faunal refuse and recurring post-discard occupation damage to the consumption refuse (Stiner and Munro, 2011; Yeshurun et al., 2014b). Thus, archaeological evidence for regular capture of low-ranked game (e.g., small animals entailing high capture cost, or young individuals from gregarious species with less meat and fat) or intensified processing (e.g., exploiting small bones for marrow, rendering grease or developing pounding technology), may indicate intensification and reflect longer stays in the camp or a larger population inhabiting it in a given time. This, in turn, sheds light on mobility and society patterns. Zooarchaeological tests for intensification are not independent from environmental shifts, which may alter the hunting selection of prey taxa because of changing environmental availability. Especially telling is the case of the two most abundant ungulates hunted in the Mediterranean region in the late Pleistocene Levant, mountain gazelle (Gazella gazella) and Mesopotamian fallow deer (Dama mesopotamica). While the former is much smaller in size than the latter (ca. 20 and 80 kg, respectively), hypothetically assigning a lower rank to it, these species thrive on different biomes: shrubland vs. woodland, respectively (Bate, 1937). Shifts in rainfall and forest cover should have changed their availability and encounter rates

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and therefore their respective “rank”, meaning that their ratios may represent environmentally-forced patterns rather than intensification (Bar-Oz et al., 2013). Separating the conflating effects of reduced mobility, demographic rise, and climate change is a major challenge. To decipher the causes of archaeofaunal shifts several independent lines of evidence are therefore required. We approached this problem by attempting to correlate archaeofaunal changes between the Ahmarian and Aurignacian assemblages (representing hunters' game choice) with the local environmental records (estimating game availability). The latter include charred wood remains, micromammals deposited by owl predation, stable isotope records of the cave's speleothems and, specifically, the animals hunted by the hyenas dwelling in the cave (see below). Intensification should become clear when human hunting choice does not track natural availability, as reconstructed by these independent proxies. 1.2. Manot Cave and its paleoenvironmental setting Manot is a large karstic cave located in Israel's Western Galilee, about 5 km east of the Mediterranean shoreline, on the southern slope of a limestone hill, 220 m above sea level (Fig. 1). The cave is situated in a Mediterranean woodland setting with mean annual precipitation of 600e700 mm. Systematic excavations commenced in 2010 and 12 areas have been excavated, labeled AeL (Fig. 1b). Apart from scattered Middle Paleolithic artifacts and the Manot 1 calvaria (Hershkovitz et al., 2015), the main archaeological record of the cave dates to the EUP period. Two excavation areas, C and E, provided well-preserved and -dated EUP sequences (Barzilai et al., 2016; Alex et al., 2017). Area E, near the postulated cave entrance, includes numerous stratified levels exhibiting Levantine Aurignacian and post-Levantine Aurignacian characteristics. Area C, the focus of the current work, is located toward the bottom of the western talus. It consists of eight sedimentological units rich in lithic, faunal, botanical and osseous finds. While the sediments may have originated from anthropogenic primary contexts up the slope, they kept the original sequence (Alex et al., 2017; Berna et al., 2019 submitted) and are securely assigned to the EUP according to their clean assemblages of lithic and osseous artifacts (Barzilai et al., 2016; Abulafia et al., 2019 submitted). The cultural phases include Early Ahmarian, dated 46e42 thousand years BP in sedimentological units 6e7, and Levantine Aurignacian, dated 38e34 thousand years BP in units 3e5 (Fig. 1c; Alex et al., 2017). While the cave lies in a Mediterranean woodland setting today, there are good indications that a more open park and grassy environment prevailed during the EUP, with some fluctuations mainly during the Aurignacian period of deposition (Caracuta et al., 2019 submitted; Comay et al., 2019 submitted; Yasur et al., 2019). The Manot micromammal assemblage, which was mostly deposited by owls, is dominated by Microtus guentheri and other species that inhabit Mediterranean grasslands. Woodland species do exist throughout the sequence in small numbers, as do species that are adapted to a somewhat cooler climate than at the present-day lowelevation western Galilee. The Aurignacian sample seems to represent rather cooler and more open conditions (though possibly with higher water availability) than the Ahmarian one (Comay et al., 2019 submitted). The diversity of the landscape is reflected in the presence of both steppe and woodland dwellers in the macrofaunal assemblages of the cave (Orbach and Yeshurun, 2019; in press, this study). Charred wood remains in the archaeological deposits of both entities consist almost exclusively of wild almond (Prunus cf. amygdalus), which today is more characteristic of the dryer Eastern Galilee. Differences in 13C isotopes of the almond specimens generally indicate more negative values in the Aurignacian, reflecting more episodes of higher rainfall compared to the

Please cite this article as: Yeshurun, R et al., Early Upper Paleolithic subsistence in the Levant: Zooarchaeology of the AhmarianeAurignacian sequence at Manot Cave, Israel, Journal of Human Evolution, https://doi.org/10.1016/j.jhevol.2019.05.007

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Ahmarian (Caracuta et al., 2019 submitted). Similarly, the continuous isotopic record from the cave's speleothems indicates primarily C3 vegetation with some dryer alternations (i.e., C3 vegetation leaning toward C4) throughout the EUP sequence but mainly in the Ahmarian. The Ahmarian corresponds to a major shift toward open grassy landscape, whereas somewhat more mixed woody and grassy landscapes were observed in the Aurignacian. Significantly, the continuous growth of speleothems in the cave throughout the entire EUP occupation indicates that water was always available (Yasur et al., 2019). 2. Materials and methods The analyzed archaeofaunal assemblages were obtained from stratified excavation units (“baskets”), in Squares J65e66 in Area C (Fig. 1b, c). This column displayed a clear stratigraphy with minimal disturbances and provided preserved materials for archaeobotany, osseous, lithic and micromorphological analyses as well as radiocarbon dating (Tejero et al., 2016; Alex et al., 2017; Abulafia et al., submitted; Berna et al. submitted; Caracuta et al., 2019 submitted). In this study we analyzed only baskets that were securely assigned to the Ahmarian (Units 6e7) or Aurignacian (Units 3e5) phases. We first studied a “detailed sample” of these baskets, which involved a multivariate taphonomic and zooarchaeological analysis, including the entire taxonomic spectrum, the identification of all possible bone fragments and the systematic documentation of bone-surface modifications and bone-fracture patterns. This sample was used to infer taxonomic abundances, skeletal-element profiles and taphonomic processes. Additionally, to obtain a sufficient sample for the demographic and osteometric analyses, we scanned all of the excavation units in this column beyond our “detailed sample” (but still with secure cultural affiliation) for specimens that could be identified to the species level and yield information on age, sex and body size of the two main prey taxa (see below). The material obtained was used only for the demographic analysis and was not included in the relative taxonomic abundances or taphonomic results (except for the ageable/sexable specimens that were already part of the “detailed sample”). This flexible strategy facilitated the time-consuming recording of taphonomic properties of a representative sample, along with obtaining one of the largest mensural samples of the Levantine UP. Faunal analysis procedures followed Yeshurun et al. (2007) and hence are compatible with other archaeofaunal studies at Manot Cave (the Area D faunal assemblage; Orbach and Yeshurun, 2019) and other Paleolithic sites in Israel (e.g., Yeshurun et al., 2007, 2014a; Marder et al., 2011). Faunal remains in Area C were systematically collected in the field using wet sieving through 5-mm mesh and 1.5-mm mesh. The analysis included all mammals except rodents, insectivores and bats. We also included all birds larger than Passeriformes, and all reptiles. The anatomical and taxonomic identification was carried out in the comparative collections of the Laboratory of Archaeozoology, University of Haifa. Identifiable elements (henceforth NISP e Number of Identified Specimens) included long-bone articular ends, long-bone shaft fragments with diagnostic zones or indicative characteristics such as thickness and morphology of the cross section and medullary cavity, teeth, cranial fragments, ribs, vertebrae, and all other recognizable bone fragments. The inclusion of these elements was conditioned on the possibility of determining and quantifying the fragments’ precise location in the skeletal element, or portion thereof, and that it could be assigned to species or size class. Size classes are “small mammal” (fox- or hare-sized), “small ungulate” (gazelle-sized), “medium ungulate” (fallow/red deer-sized), and “large ungulate” (aurochs-sized). Body-part profiles were quantified by the Minimum Number of Elements (MNE) and standardized

by Minimum Animal Unit (MAU). All identified specimens (NISP) were considered when calculating the MNE counts. Taking bone side and age into account produced the Minimum Number of Individuals (MNI). All identified specimens were inspected for bone-surface modifications, following Blumenschine et al.‘s (1996) procedures and using a stereoscopic microscope (Olympus SZX7) with highintensity oblique light. We searched for carnivore damage such as pits, scores, punctures, notches, crenellation, channeling and digestion, as well as rodent gnaw marks (Binford, 1981; Brain, 1981). We also looked for human modifications including cutmarks (Binford, 1981; Domínguez-Rodrigo et al., 2009), hammerstone percussion marks (Pickering and Egeland, 2006), bone working (Tejero et al., 2016; Yeshurun et al., 2018), and burning (Stiner et al., 1995). The butchery marks were classified (i.e., skinning, dismemberment, evisceration, and filleting) according to their anatomical placement and appearance, following Binford's (1981) scheme. A variety of post-depositional modifications was also documented e weathering stages (Behrensmeyer, 1978), trampling striations (Domínguez-Rodrigo et al., 2009), biochemical marks (Domínguez-Rodrigo and Barba, 2006), and abrasion (Shipman and Rose, 1988). In addition, we recorded long-bone and first-phalanx breakage patterns, including fracture angle, fracture outline, and fracture edge (Villa and Mahieu, 1991) in order to assess if breakage occurred when the bone was fresh/‘green’ vs. dry; green fractures were assigned when fracture angle was oblique/acute and fracture outline was curved. Dry fractures were defined when only right angles and transverse outlines were present. A combination of these features was categorized as “intermediate”. We also recorded shaft circumference (Bunn, 1983) and measured the greatest length of each identified specimen fragment (excluding excavation breaks) to the nearest millimeter (Bar-Oz, 2004). Ungulate mortality profiles were recorded by tooth eruption and wear patterns and by bone fusion. Gazelle dental and fusion data were recorded and interpreted according to Munro et al.‘s (2009) study of modern, known-age gazelles from Israel. Fallow deer dental remains were aged according to Stiner's (2005) system, while the interpretation of bone fusion data followed Carden and Hayden (2006). Sexing of gazelle remains made use of character traits, i.e., certain skeletal elements that are very dimorphic and can be readily assigned to either male or female (Munro et al., 2011). In addition, we explored sex-related and non-sex-related trends in ungulate body size by measuring selected skeletal elements (Davis, 1981; Munro et al., 2011: Table 7; Yeshurun et al., 2014a). The measurements were performed with digital calipers to the nearest 0.1 mm, and excluded unfused, burned or abraded specimens (von den Driesch, 1976). The small degree of sexual dimorphism in gazelles (Horwitz et al., 1990) does not allow sexual distinction of unfused epiphyses. In some cases, we transformed the measurements to Log Size Index (LSI) values, using the formula, LSI ¼ Log10(XeY), in which X is an archaeological measurement and Y is the corresponding measurement from a ‘standard animal’ (Meadow, 1999; see below for the standard and measurements used in each case). This scaling technique was employed to boost our sample sizes and allow comparisons between the Ahmarian and Aurignacian samples. We statistically compared the relative taxonomic abundances of the Ahmarian and Aurignacian samples by employing the c2 distribution. Adjusted Residuals (AR) were calculated for each cell using SPSS version 23 and presented along the composite c2 to discern which cells most significantly differed from the expected values. AR values are standard normal deviates, indicating the probability that a single-cell comparison is statistically significant.

Please cite this article as: Yeshurun, R et al., Early Upper Paleolithic subsistence in the Levant: Zooarchaeology of the AhmarianeAurignacian sequence at Manot Cave, Israel, Journal of Human Evolution, https://doi.org/10.1016/j.jhevol.2019.05.007

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Significant ARs have values equal to or greater than ±1.96 (Everitt, 1977; Grayson and Delpech, 2008). Mean values of anatomical measurements (raw or LSI values) were compared by ANOVA, Tukey's post-hoc test and Student's t-test, while index values (such as the MNE/NISP Completeness Index) were compared using the KruskaleWallis test. We quantified ungulate carcass completeness by the Shannon Evenness Index (Shannon H divided by the natural log of the number of categories; Faith and Gordon, 2007) applied to MAU tallies of all bones (see a detailed explanation of our assumptions in Yeshurun and Bar-Oz, 2018). Skeletal-element evenness values were bootstrapped to produce 95% confidence intervals and then statistically compared using the Diversity Permutation Test, which compares the diversities using random permutations with PAST software (version 3.07); 9999 random matrices with two columns (samples) are generated, each with the same row and column totals as in the original data matrix (Hammer al., 2001). All other statistical tests were also performed by the same version of PAST. Statistical significance was tested at a ¼ 0.05 level. 3. Results The faunal samples that were subjected to the detailed analysis consisted of 476 and 1066 identified specimens (NISP) in the Ahmarian and Aurignacian assemblages, respectively. Our “demographic sample” included 55 and 492 additional specimens in the Ahmarian and Aurignacian samples, respectively. Faunal remains were abundant in all excavation units and no bone-poor zones were encountered (see Supplemental Online Materials [SOM] Table S1 for breakdown of taxa and major taphonomic variables across the units). However, the two samples markedly differed in the abundance of bone remains. The Aurignacian sediments were much denser with bone finds when standardized by excavation volume, irrespective of whether NISP or all bone specimens were considered (Fig. 2; Table 1). 3.1. Taxonomic composition The Ahmarian and Aurignacian samples display similar taxonomic spectra (Table 2). Both assemblages are dominated by ungulate remains, constituting 88% and 81% of NISP, respectively. The most abundant species in both samples is the mountain gazelle (Gazella gazella), followed by the Mesopotamian fallow deer

12 Ahmarian

Aurignacian

Bone mass / Excavaon volume

10

8

6

4

2

0 Total mass

NISP mass

Figure 2. Comparison of the volumetric density of bone finds between the Ahmarian and Aurignacian sediments, by total bone mass and NISP mass, divided by excavation volume (data from Table 1).

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(D. mesopotamica). Other ungulate remains were found in small numbers: roe deer (Capreolus capreolus), red deer (Cervus elaphus), wild boar (Sus scrofa) and aurochs (Bos primigenius). A number of carnivore species were also found in the assemblages, the most common of which was the red fox (Vulpes vulpes). Isolated remains of leopard (Panthera pardus), wildcat (Felis silvestris), Egyptian mongoose (Herpestes ichneumon), Cape hare (Lepus capensis) and Indian crested porcupine (Hystrix indica) complete the macromammalian spectrum. Birds larger than Passeriformes were found in considerable numbers (Table 2); their precise taxonomic identification will be the subject of on-going study. Reptile remains include bones and shell fragments of spurthighed tortoise (Testudo graeca), as well as squamate (snake and lizard) remains, most of which are vertebrae. One specimen was attributable to a legless lizard, Pseudopus apodus. Other faunal remains, which were not part of this study, were plentiful micromammals (Comay et al., 2019 submitted), as well as large numbers of land snails Levantina caesareana, which could have been consumed, along with some marine gastropod remains (Bar-Yosef Mayer, pers. comm.). Fish remains were extremely rare. While the same animal species dominate both assemblages, the relative abundance of animal groups is significantly different (Fig. 3; c2 ¼ 43.13, p < 0.001). NISP counts of the major animal groups were transformed into Adjusted Residuals (AR) in order to assess which counts significantly diverge from the expected values (SOM Table S2). The Ahmarian sample is significantly richer in medium ungulate remains (mostly representing Mesopotamian fallow deer) while the Aurignacian sample is significantly richer in small ungulates (gazelle) and bird remains. The relative proportion of other animal groups is similar in both samples (Fig. 3b). 3.2. Agents of deposition and modification The examination of bone-surface modifications in both samples established that they both represent human hunting, butchery and consumption. This assertion is based on the ample presence of cutmarks and hammerstone percussion marks (Tables 3 and 4). Cutmarks were found on 24 bones in the Ahmarian sample (5.7% of NISP excluding isolated teeth) and on 30 bones in the Aurignacian sample (3.0%). The proportion of cutmarked specimens is statistically similar between the samples (small ungulate, c2 ¼ 1.68, p ¼ 0.2; medium ungulate, c2 ¼ 1.88, p ¼ 0.17) and both samples exhibit evidence of skinning, dismemberment and filleting, representing the entire butchery process (Table 5). Aside from ungulate bones, cutmarks were noted on a mongoose sacrum in the Ahmarian sample and on a crane-sized avian scapula in the Aurignacian sample (Table 5), suggesting that the presence of at least some of the small mammals and birds originated with human exploitation as well. Lower numbers of cutmarked specimens is to be expected on small animal bones, because less force and cutting action are needed for their butchery. Hammerstone percussion marks appear on 7.5e13.6% of relevant NISP (mammal limb bone shaft fragments, mandibles and phalanges). There was no statistically significant difference in the frequency of the percussion marks between the two samples, for both the small ungulate and medium ungulate groups (c2 ¼ 0.13, p ¼ 0.72 and c2 ¼ 1.36, p ¼ 0.24, respectively). Human butchery activities notwithstanding, the impact of another depositional agent is present, namely carnivore activity. Manot Cave repeatedly served as a spotted hyena den during the late Pleistocene, with denning activities concentrated mainly in the middle of the talus, in Area D, but remnants of their activities, in the form of gnawed ungulate bones, hyena skeletal remains and coprolites, are evident in other places in the cave as well (Orbach and Yeshurun, 2019). Hyena bones and coprolites are extremely rare in

Please cite this article as: Yeshurun, R et al., Early Upper Paleolithic subsistence in the Levant: Zooarchaeology of the AhmarianeAurignacian sequence at Manot Cave, Israel, Journal of Human Evolution, https://doi.org/10.1016/j.jhevol.2019.05.007

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Table 1 Volumetric densities of bone finds in the Ahmarian and Aurignacian sediments.

Ahmarian Aurignacian

NISPa

NISP mass (gr)

Non-NISP mass (gr)

Total bone mass (gr)

volume (liter)

Total bone mass / volume

NISP / volume

476 495b

1715 1170

1647 704

3362 1874

880 180

3.82 10.41

0.54 2.75

a

Abbreviations: NISP ¼ number of identified specimens, gr ¼ grams. We included in this calculation only the finds from excavation units that had a recorded volume, and therefore had to omit some of the Aurignacian excavation units with their recorded specimens. b

our samples, but gnawed bones appear as 3.3% and 1.4% of NISP in the Ahmarian and Aurignacian samples, respectively (Tables 3 and 4). Comparing the abundance of gnawed bones in the small ungulate and medium ungulate groups between samples, no statistically significant difference emerges (c2 ¼ 0.04, p ¼ 0.85 and c2 ¼ 3.38, p ¼ 0.07, respectively). The proportions of gnawed bones are much lower than those expected from assemblages accumulated by “carnivores only” or any combination of hominins and carnivores reported in Marean et al. (2000: Table 3). Additionally, a marked difference exists in gnawing frequency between the Area C assemblages and the Area D hyena den. In almost all cases, the human modifications are more abundant in Area C whereas the carnivore modifications are much more abundant in Area D, creating a different taphonomic profile and emphasizing the prominent agent of accumulation in each area (Fig. 4). Therefore, carnivore activity was not a major contributor of faunal remains in our sample. Despite the anthropogenic character of the assemblages, burned bones are rare in both samples, ranging between 2.1 and 2.3% of NISP. Of these 34 specimens, just four displayed calcination (graycoloring) and all the rest were carbonized. No consistent trend in the anatomical representation of burning was found. All taxonomic groups were impacted by burning in the larger Aurignacian sample, while only the small and medium ungulate remains displayed burned specimens in the Ahmarian, possibly because of the smaller sample size (Tables 3 and 4).

Post-depositional damage in the samples was generally mild and bone surfaces are well-preserved. Some weathering damage (specimens that correspond to Behrensmeyer's [1978] stage 3 and above) was observed on 9.0% of NISP in the Ahmarian and 2.2% of NISP in the Aurignacian. No specimens were assigned to stage 5 and the assemblage was not particularly weathered relative to other cave assemblages in the region (compare Yeshurun et al., 2007). Examined per ungulate size group, the proportion of weathered specimens in the Ahmarian ungulate remains is significantly higher than the Aurignacian ones (small ungulate, c2 ¼ 10.04, p ¼ 0.002; medium ungulate, c2 ¼ 12.85, p < 0.001). Rodent gnaw marks and root (biochemical) marks are infrequent in both samples. Since the assemblages were deposited in the context of a talus, within a closed cave with extensive karstic activity (Berna et al., submitted), it is important to note that striations from trampling and rolled edges related to abrasion are rare in both samples (Tables 3 and 4). Thus, the processes that impacted the samples post-depositionally are generally similar in type and magnitude. Only weathering and the proportion of root marks differ somewhat, being consistently higher in the Ahmarian (Fig. 5). The analysis of bone fracture patterns complements the taphonomic picture. As expected in a Paleolithic assemblage that exhibits hammerstone percussion, a considerable portion of the ungulate limb bones shaft fragments carry “green” breaks (24e32% of relevant NISP) and the majority (75e86%) preserve less than half of their original circumference (Tables 3 and 4). Breaks classified as

Table 2 Taxonomic composition of the Ahmarian and Aurignacian samples in Area C of Manot Cave (NISP). MNI tallies are given for mutually exclusive taxa. Ahmarian a

Ungulates Gazella gazella Capreolus capreolus Small ungulate Dama mesopotamica Cerus elaphus Sus scrofa Medium ungulate Bos primigenius Large ungulate Other game Panthera pardus Vulpes vulpes Felis silvestris Herpestes ichneumon Lepus capensis Hystrix indica Small mammal Testudo graeca Pseudopus apodus Lizard Snake Bird Total a

Aurignacian

NISP

MNI

NISP

MNI

73 3 154 39 2 1 144 2 2

4 1 – 5 1 1 – 1 –

166 2 434 32 3 4 222 0 5

6 1 – 3 1 1 – 0 –

1 2 0 1 0 1 8 8 0 1 14 20 476

1 1 0 1 0 1 – 1 0 –

0 10 1 0 2 0 15 36 1 0 35 98 1066

0 1 1 0 1 0 – 1 1 –

Abbreviations: NISP ¼ number of identified specimens, MNI ¼ minimum number of individuals.

Please cite this article as: Yeshurun, R et al., Early Upper Paleolithic subsistence in the Levant: Zooarchaeology of the AhmarianeAurignacian sequence at Manot Cave, Israel, Journal of Human Evolution, https://doi.org/10.1016/j.jhevol.2019.05.007

Number of Idenfied Specimens

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60% Ahmarian

50%

Aurignacian

40% 30% 20% 10% 0% Small Medium Large Small Tortoise Lizard & ungulate ungulate ungulate mammal snake

a

Bird

8 Ahmarian

Adjusted Residuals

6

Aurignacian

4 2 0 -2 -4 -6 -8

b

Small Medium Large Small Tortoise Lizard & ungulate ungulate ungulate mammal snake

7

specimens in the post-discard stage. There is little difference in these respects between the Ahmarian and Aurignacian samples; rather, they cluster tightly together (Fig. 6). We tested our taphonomic data for different intensity of carcass exploitation between samples, using the proxies suggested by Munro and Bar-Oz (2005) and Hodgkins et al. (2016). The intensity of carcass exploitation may have been somewhat higher in the Ahmarian. The small ungulate Ahmarian samples displayed a higher percentage of bones with cutmarks and percussion marks and more phalanx I items with percussion marks, indicating the exploitation of marrow-poor bones. The same pattern occurred with the medium ungulates, except for higher proportion of percussion marks (Fig. 7a, b). However, bone fragment lengths, which may serve as a proxy for the intensity of taphonomic processes including carcass processing (Bar-Oz, 2004), are identical in the Ahmarian and Aurignacian samples (Fig. 7c; Table 6). Finally, the completeness index (MNE/NISP) of the marrow-bearing bones is statistically similar in both samples (KruskaleWallis, F ¼ 0.17, p ¼ 0.92) indicating a similar degree of fragmentation (Fig. 7d). Given the generally similar intensity of post-depositional attrition in both samples (see above), the similar bone fragment lengths and degree of fragmentation of marrow-bearing elements probably indicate similar intensity of bone nutrient exploitation.

Bird

Figure 3. Comparison of the taxonomic breakdown of the Ahmarian and Aurignacian samples: (a) presentation by %NISP; (b) comparison by AR values. Positive AR values denote overrepresented traits, while negative AR values denote underrepresented traits. Significant values are AR ¼ ±1.96. Data are from SOM Table S2. Taxonomic groups shown include all specimens (NISP), whether identified to species or to sizeclass, i.e., the “small ungulate” group includes specimens identified as gazelle, roe deer, and small ungulate-sized.

“dry” or intermediate” were abundant, raising the possibility that while the primary breakage had occurred for nutritional reasons, leaving green breaks, further cycles of breakage impacted many

3.3. Skeletal-element profiles The observed distribution of skeletal elements of small and medium ungulates in the Ahmarian vs. Aurignacian samples was presented by MAU, i.e., it was compared against a complete skeleton model (Fig. 8; SOM Table S3). Generally, all skeletal parts were represented with no major cases of under-representation except the vertebrae and ribs. Head parts, limb parts, the pelvis and foot parts are well-represented in most cases. Bone-portion survival (MAU) is positively and significantly correlated with Bone Mineral Density (BMD1, 2 values following Lam et al., 1999) in all samples (Table 7), providing a likely explanation for the low survival of the

Table 3 Bone-surface modifications and bone fracture patterns for all taxonomic groups in the Ahmarian sample.

NISPa Burning Green fracture Dry fracture Intermediate Limb Shaft circumference

Weathering (stage 3-4)

cutmarks Percussion marks

Gnawing (carnivore) Gnawing (rodent) Root marks Trampling striations Abrasion a

N % N N N <50 50 100 N of % N % N of % N % N % N % N % N %

Small ungulate

Medium ungulate

230 5 2.2% 8 17 9 53 8 10 15 208 7.2% 11 5.3% 8 71 11.3% 3 1.4% 1 0.5% 15 7.2% 16 7.7% 4 1.9%

187 5 2.7% 5 7 9 55 4 8 22 164 13.4% 12 7.3% 5 67 7.5% 9 5.5% 1 0.6% 18 11.0% 17 10.4% 7 4.3%

Large ungulate

Small mammal

4 0 0%

12 0 0%

0 0 0 1 3 33.3% 0 0% 0 0 0% 1 33.3% 0 0% 0 0% 0 0% 0 0%

0 0 2 0 10 0% 1 10.0% 0 0 0% 0 0% 0 0% 1 10.0% 0 0% 0 0%

Snake/ lizard 15 0 0%

Tortoise

Birds

Total

8 0 0%

20 0 0%

476 10 2.1%

0 9 0% 0 0%

0 8 0% 0 0%

1 0 3 0 18 0% 0 0%

109 12 23 38 420 9.0% 24 5.7%

0 0% 0 0%

0 0% 0 0% 1 12.5% 0 0% 0 0%

1 5.6% 0 0% 3 16.7% 1 5.6% 0 0%

0% 0 0% 0 0%

14 3.3% 2 0.5% 38 9.0% 34 8.1% 11 2.6%

Abbreviations: NISP ¼ number of identified specimens.

Please cite this article as: Yeshurun, R et al., Early Upper Paleolithic subsistence in the Levant: Zooarchaeology of the AhmarianeAurignacian sequence at Manot Cave, Israel, Journal of Human Evolution, https://doi.org/10.1016/j.jhevol.2019.05.007

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Table 4 Bone-surface modifications and bone fracture patterns for all taxonomic groups in the Aurignacian sample.

NISPa Burning Green fracture Dry fracture Intermediate Limb shaft circumference

Weathering (stage 3-4)

Cutmarks Percussion marks

Gnawing (carnivore) Gnawing (rodent) Root marks Trampling striations Abrasion a

Small ungulate

Medium ungulate

Large ungulate

Small mammal

Snake/ lizard

Tortoise

Birds

Total

602 16 2.7% 26 44 38 175 12 17 13 551 2.4% 18 3.3% 20 205 9.8% 9 1.6% 2 0.4% 15 2.7% 33 6.0% 10 1.8%

261 4 1.5% 10 10 11 64 7 9 9 240 3.8% 10 4.2% 11 81 13.6% 5 2.1% 0 0% 13 5.4% 12 5.0% 4 1.7%

5 0 0% 0 1 0 1 0 1 0 5 0% 1 20.0% 0 2 0% 0 0% 0 0% 0 0% 0 0% 0 0%

28 0 0% 0 1 1 2 0 6 0 22 0% 0 0% 0 8 0% 0 0% 0 0% 1 0% 1 4.5% 0 0%

36 1 2.8%

36 1 2.8%

98 2 2.0%

1066 24 2.3%

0 36 0% 0 0%

0 36 0% 0 0%

0 1 53 0 98 0% 1 1.0%

0 0% 0 0% 0 0% 0 0% 0 0%

0 0% 0 0% 0 0% 0 0% 0 0%

0 0% 0 0% 5 5.1% 2 2.0% 0 0%

242 20 86 22 988 2.2% 30 3.0% 31 296 10.5% 14 1.4% 2 0.2% 34 3.4% 48 4.9% 14 1.4%

N % N N N <50 50 100 N of % N % N of % N % N % N % N % N %

Abbreviations: NISP ¼ number of identified specimens.

lowest-density elements such as vertebrae and ribs. However, the correlation coefficients (r2) are quite low (0.11e0.29), suggesting that density-mediated attrition does not account for most of the observed patterns. This seems to be especially true for the fluctuations seen in the relative representation of the medium- and highdensity elements. The observed skeletal-element profiles (Fig. 8) were quantitatively compared by calculating skeletal-element evenness (Faith and Gordon, 2007). Given the mild density-mediated attrition, we considered all bones in calculating skeletal-element evenness, but also provided the evenness values for the high-survival set (Table 7; see Yeshurun and Bar-Oz [2018] for detailed explanation). The representation of bones (MAU) is quite even in most cases, with Shannon Evenness Index values ranging between 0.75 and 0.83 (Fig. 9). The small ungulate evenness is statistically similar in the Ahmarian vs. Aurignacian samples (Diversity Permutation Test, p ¼ 0.16), while in the medium ungulate group, the Ahmarian evenness is significantly lower than the Aurignacian pattern (p < 0.01). However, the Ahmarian medium ungulate is the smallest assemblage in terms of NISP, and the one with the strongest relationship between survivorship and bone density, although both Table 5 Classification of cutmarked specimens (NISP)a in the ungulate groups in the Ahmarian and the Aurignacian samples. Skinning Ahmarian Small ungulate Medium ungulate Large ungulate Total Aurignacian Small ungulate Medium ungulate Large ungulate Total a

Dismemb.

Filleting

Eviscer.

Indet.

Total 11 12 0 23

2 2

1 5

4 4

1

3 1

4

6

8

1

4

6 1

4 6 1 11

6 1

7

7

Abbreviations: NISP ¼ number of identified specimens.

2 2 0

5

18 10 1 30

Figure 4. Comparison of selected bone-surface modifications indicating human activity (cutmarks and percussion marks) and carnivore gnawing, between the Ahmarian and Aurignacian samples (this study) and the Manot Cave hyena den (Orbach and Yeshurun, 2019, Table 4a). Top: Small ungulate, bottom: medium ungulate.

Please cite this article as: Yeshurun, R et al., Early Upper Paleolithic subsistence in the Levant: Zooarchaeology of the AhmarianeAurignacian sequence at Manot Cave, Israel, Journal of Human Evolution, https://doi.org/10.1016/j.jhevol.2019.05.007

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the meaty humerus, radius, femur and tibia and the bulky pelvis. In addition, cranial parts outnumber the mandible, contrasting with a scenario of dismembering the head in the field (Fig. 8). Therefore, the lower evenness of this group relative to the Aurignacian may be a methodological artifact. In the same vein, bone survival (MAU) presents no correlation to bone meat and fat content (Food Utility Index following Metcalfe and Jones, 1988) or to bone-marrow content (Marrow Index, following Bar-Oz and Munro, 2007 and Madrigal, 2004 for gazelle and deer, respectively) in any sample, reinforcing the notion of near-complete or complete, non-selective, carcass transport in both the Ahmarian and Aurignacian (Table 7).

3.4. Prey demography and body-size trends

Figure 5. Comparison of post-depositional bone-surface modifications in the Ahmarian vs. Aurignacian samples. Top: small ungulate, bottom: medium ungulate.

figures are not very far apart from the other samples (Table 7). Additionally, these differences disappear when only the highsurvival elements are considered in the evenness comparison (small ungulate, p ¼ 0.53; medium ungulate, p ¼ 0.07). Looking at the detailed skeletal-element profile of the Ahmarian medium ungulates, this skeletal-element profile is unlikely to represent selective transport for meat, marrow, bulk or raw materials. This is evidenced by the nearly identical representation of high-density elements: the mandible, which contains low nutritional value,

We examined the demography (age and sex distribution) of the two most abundant game animals in the cave, gazelle and fallow deer, and explored changes in their body size. The discussion and comparison will focus on mountain gazelle, as it is the main prey animal in Manot and in most other southern Levantine Upper Paleolithic sites. Gazelle age-at-death distributions are generally similar in both samples (Fig. 10; SOM Tables S4eS5). Examination of the juvenile age classes by bone fusion yielded very similar survival curves, in which almost all animals survived their first few months of life, and then a considerable portion was culled in the 7e18-month stage. Only about two-thirds of the animals survived to adulthood. The tooth eruption and wear sequence, which captures the entire life span of the animal, shows a different picture for the Aurignacian sample, with more intense culling of the 2e7-month age class followed by reduced culling of older juveniles and then culling of all adult age classes. Comparing three methods: fusion status of the latest-fusing elements (corresponding to the juvenileeadult transition of ca. 18 months), dental sequence of the dP4 to M3 and dP4 to P4 (corresponding to the same transition), the picture in the Aurignacian is clear, with non-adult animals making up 42e49% (Fig. 10). The Ahmarian methods do not conform as well to each other, possibly because of the smaller sample. Here, the sequence with the largest sample (n ¼ 9; dP4eM3) indicates only 22% juveniles. The juvenile proportion in the larger Aurignacian sample is virtually identical to the year-round average of juveniles in the gazelle population recorded by Baharav (1974: Fig. 1) in the lower Galilee: ca. 55% adults and 45% juveniles (c2 ¼ 0.00, p ¼ 0.99).

Figure 6. (a) Comparison of limb fracture types (green, dry and intermediate) in the studied assemblages (Tables 3 and 4); (b) comparison of limb shaft circumference types (less than 50% of complete circumference, more than 50%, or 100%) in the studied assemblages (Tables 3 and 4). In both cases the Manot hyena den (Orbach and Yeshurun, 2019) is shown for comparison. AHM, Ahmarian; AUR, Aurignacian; HYN, Hyena den; SU, Small ungulate; MU, Medium ungulate.

Please cite this article as: Yeshurun, R et al., Early Upper Paleolithic subsistence in the Levant: Zooarchaeology of the AhmarianeAurignacian sequence at Manot Cave, Israel, Journal of Human Evolution, https://doi.org/10.1016/j.jhevol.2019.05.007

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Figure 7. Comparison of carcass processing intensity in the Ahmarian (AHM) and Aurignacian (AUR) samples: (a) the proportion of NISP bearing cutmarks and hammerstonepercussion marks, and the proportion of phalanx 1 items that bear percussion marks, in the small ungulate group (SU) and (b) the medium ungulate (MU) group; (c) comparison of bone fragment lengths. Data from Table 6; (d) the completeness index for marrow-bearing bones (mandible, humerus, radius, femur, tibia, calcaneus, metapodials, phalanges 1 and 2; specimens with NISP<10 were excluded). Data from SOM Table S3.

Since most of the variability between assemblages concerns the pre-adult stages, bone fusion data were presented by survival curves to demonstrate the survival of different age classes (Fig. 11; SOM Table S6). Using this technique, the magnitude of the Ahmarian-Aurignacian difference appears to be small in comparison to the earlier Middle Paleolithic case of Hayonim Cave (Stiner, 2005) and to the later Natufian cases (Yeshurun et al., 2014a). The Manot samples are quite similar to the Kebaran of Hayonim (Fig. 11). Robust comparison to other EUP sites in the Levant is hindered because of the lack of published, detailed bone-fusion data. In the meantime, we can suggest an EUP hunting pattern that avoided fawns but did target gazelles toward the end of their first year of life and the beginning of their second year, as well as

Table 6 Comparison of bone fragments lengths (measurements in mm). Ahmarian Small ungulate

Medium ungulate

N Mean Variance t-test N Mean Variance t-test

218 31.459 235.48 t¼-0.18, p¼0.85 178 40.713 468.91 t¼-0.54, p¼0.58

Aurignacian 569 31.682 214.92 244 39.643 339.16

adults. This pattern holds in the Kebaran example, too. In contrast, the preceding MP seems to have had a greater emphasis on adults, while the sedentary hunter-gatherers in the Natufian had begun culling fawns in considerable numbers (Fig. 11). The sex ratio of the Manot gazelles may be revealed by counts of sexually dimorphic elements and also by osteometry of skeletal elements whose size is affected by sex (usually larger size for males). However, anatomical measurements may also be influenced by a variety of sex-independent factors, primarily climateinduced body-size shifts in the ancient population. This is especially true for late Pleistocene gazelle, as this small ungulate displays small but significant sexual dimorphism (Horwitz et al., 1990) that can easily be conflated with increasing or decreasing body size of the entire population (Bar-Oz et al., 2004; Munro et al., 2011). Thus, prior to the sexing analysis we explored gazelle body-size trends in Manot. The full measurement database is given in SOM Table S7. Comparing Ahmarian vs. Aurignacian bone measurements resulted in statistically similar results (LSI values for limb breadth or depth measurements: t ¼ 1.53, p ¼ 0.13; LSI values for sexsensitive measurements: t ¼ 0.36, p ¼ 0.72; see SOM Table S7 for details), attesting to similar body size and probably also to a similar sex make-up. Interestingly, both samples display significantly larger body size compared to modern-day gazelles in Israel according to most measurements (Fig. 12; see also Davis, 1981; Ducos and Kolska Horwitz, 1997). Of the skeletal elements we

Please cite this article as: Yeshurun, R et al., Early Upper Paleolithic subsistence in the Levant: Zooarchaeology of the AhmarianeAurignacian sequence at Manot Cave, Israel, Journal of Human Evolution, https://doi.org/10.1016/j.jhevol.2019.05.007

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Figure 8. Skeletal-element profiles (%MAU) of the small ungulate and the medium ungulate groups in the Ahmarian and Aurignacian samples.

measured, no element was smaller than modern females, and in several cases the elements’ mean measurement was larger than modern males (SOM Table S8). The larger body size of the Manot gazelle effectively prevents us from implementing Munro et al.‘s (2011) equations and cutting points to sex each specimen because of the risk of misidentifying large females as males. Gazelle sexing therefore relied on character traits, the horns and pubis that are unmistakably dimorphic, indicating more females than males in both samples (62e63% female; Table 8). Much of this proportion in the Ahmarian sample stems from horn-cores (n ¼ 7), while just one pubis (a male) was found. In contrast, the Aurignacian sample produced both sexable horn-cores (n ¼ 8) and 10 female and 8 male pubises. This sex ratio in both samples is similar to the year-round average of 81 males for every 100 females, obtained by Baharav (1974) during gazelle drive counts in the Lower Galilee in the 1970s (Aurignacian vs. Baharav's data: c2 ¼ 0.37, p ¼ 0.55). Interestingly, an age-related sex bias may be present in our results. The character traits record both sub-adult and adult animals, because the horn-cores and pubises of sub-adults are also easily

sexed. Hence, these traits provide a sex ratio for the entire captured population. In contrast, limb-bone measurements include only fused bones and therefore only skeletally mature animals (the small degree of sexual dimorphism in gazelle does not allow sexual distinction of unfused epiphyses). The distal breadth of the latefusing metacarpal bone, known to be sensitive to sex in gazelles (Munro et al., 2011) shows some bimodality that is skewed toward the larger group, i.e. males, while the distal humerus breadth, an early-fusing element that is less sensitive to sex, does not show any bimodality (Fig. 13). Thus, the histograms in Figure 13, together with the more inclusive sex ratios obtained by character traits, suggest that more females were taken as sub-adults, while more males were hunted as adults. The second most abundant prey in Manot, Mesopotamian fallow deer, likewise displays a considerable proportion of juveniles in the Ahmarian (33e50%) and the Aurignacian (46e71%; Table 9). High agreement was observed between the bone-fusion method (late-fusing elements including the proximal tibia, proximal and distal femur, distal metapodials and distal radius; Carden and Hayden, 2006) and the dP4eM3 dental series. Both

Table 7 NISP, skeletal-element evenness (Shannon Evenness Index), and results of bone survival to bone density, bone survival to Food Utility Index, and bone survival to Marrow Index Spearman's correlation tests of the faunal assemblages.a

Ahmarian, small ungulate Aurignacian, small ungulate Ahmarian, medium ungulate Aurignacian, medium ungulate a

NISP

Skeletal-element Evenness

Skeletal-element Evenness (high-survival set)

227 595 187 260

0.8046 0.8262 0.7536 0.8118

1.074 1.034 0.9923 1.096

Bone MAU to bone density r r r r

¼ ¼ ¼ ¼

0.40, 0.34, 0.54, 0.49,

p p p p

¼ 0.002 ¼ 0.002 < 0.001 < 0.001

Bone MAU to Food Utility Index r r r r

¼ ¼ ¼ ¼

-0.45, -0.05, -0.29, -0.27,

p p p p

¼ ¼ ¼ ¼

0.08 0.84 0.28 0.14

Bone MAU to Marrow Index r ¼ 0.30, r ¼ 0.23, r ¼ 0.46, r ¼ -0.10,

p p p p

¼ ¼ ¼ ¼

0.42 0.53 0.24 0.80

Abbreviations: NISP ¼ number of identified specimens, MAU ¼ minimum animal unit.

Please cite this article as: Yeshurun, R et al., Early Upper Paleolithic subsistence in the Levant: Zooarchaeology of the AhmarianeAurignacian sequence at Manot Cave, Israel, Journal of Human Evolution, https://doi.org/10.1016/j.jhevol.2019.05.007

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4. Discussion Our study of the Ahmarian and Aurignacian archaeofaunas in Area C of Manot Cave indicated that they represent human hunting, carcass transport, consumption and discard. Preliminary observations on the Aurignacian faunal assemblage of Area E indicated very similar patterns in taxonomic representation, age profile and taphonomic signature (Marder et al., 2019 submitted). Clear Ahmarian deposits have not been attained in Area E. In this discussion, we sum up and interpret the zooarchaeological and taphonomic results, and compare between the two cultural entities. We then place the identified EUP hunting patterns in their evolutionary context and discuss the possible factors that make the Aurignacian archaeological record stand out in the Levantine sequence. 4.1. EUP hunting behavior at Manot

Figure 9. Comparison of the skeletal-element evenness values in the Ahmarian (AHM) vs. Aurignacian (AUR) samples, in the small ungulate (SU) and medium ungulate (MU) groups. An index value approaching 1 means complete skeletal representation. The evenness index values were bootstrapped to produce 95% confidence intervals.

series indicate 33% juveniles in the Ahmarian and rising to 46% juveniles in the Aurignacian. When the dental series was divided into three age cohorts (following Stiner, 2005), the counts of old adults were shown to be negligible, but again indicated the importance of sub-adult animals alongside prime-aged ones, especially so in the larger Aurignacian sample (Table 9). LSI values of breadth and depth measurements of fallow deer limb bones showed a similar average size in both samples (t ¼ 1.56, p ¼ 0.14; see SOM Table S9 for the raw measurements and summary statistics). The samples were too small to osteometrically evaluate sex ratios.

EUP hunters in Manot Cave took mainly ungulates, first and foremost gazelle, followed by fallow deer. While fallow deer were an important dietary staple in the Ahmarian, Aurignacian hunting relied on gazelle to a greater extent and also supplemented ungulate hunting with fowling. During both periods, hunting of gazelle sub-adult females and adult males likely prevailed. A possible scenario accounting for this pattern relies on modern observations of mountain gazelle ethology in the Mediterranean region of Israel (e.g., Baharav, 1974, 1983; Geffen et al., 1999; Yom-Tov, 2016). Gazelles employ resource-defense polygyny; the males maintain territories and attempt to draw females to their resources. Females typically congregate with other females and their offspring to form relatively large herds consisting of three generations: fawns (both sexes), sub-adult females (some of which already breed), and adult females. Male fawns accompany their mothers until about six months of age, when they disperse and sometimes form bachelor herds with other sub-adult males. Adult males in prime condition

Figure 10. Age profiles of gazelle in Area C of Manot Cave. Light gray represents juveniles. (a) Comparison of three age presentation techniques, bone fusion and dental sequences (latest-fusing elements following Munro et al., 2011), in the Ahmarian (top) and Aurignacian (bottom); (b) age distribution according to the most detailed dental sequence, that of dP4eM3. Data are from SOM Table S4 and S5. Age groups are: A, Fetus/neonate (0e1 months); B, Juvenile-young (2e7); C, Juvenile-old (7e18); D, Adult-young (18e36); E, Adultolder (36e96); and E, Adult-very old (96þ).

Please cite this article as: Yeshurun, R et al., Early Upper Paleolithic subsistence in the Levant: Zooarchaeology of the AhmarianeAurignacian sequence at Manot Cave, Israel, Journal of Human Evolution, https://doi.org/10.1016/j.jhevol.2019.05.007

R. Yeshurun et al. / Journal of Human Evolution xxx (xxxx) xxx

Figure 11. Comparison of gazelle culling patterns by survival curves, based on bone fusion. The Ahmarian and Aurignacian data from Manot are shown alongside examples from the Middle Paleolithic of Haynoim (HAYeMP), the Epipaleolithic Kebaran of Hayonim (HAY-KEB) and the late Epipaleolithic Early and Late Natufian of el-Wad Terrace (EWT-EN and EWT-LN, respectively). Age stages follow Munro et al. (2009). Data are from SOM Table S6.

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(ca. 20% of males in the population) maintain a territory and mate with the females that reach it. Females reach sexual maturity faster and may give birth at 12e18 months of age, before they are skeletally mature, while males that hold territories are prime adults. Thus, it seems that the Manot hunters targeted mainly the female herds, capturing sub-adult and adult females. Males could have been captured on the same occasions, as prime-adult males holding territories. If this interpretation is accepted, a possible explanation for preferentially targeting the female herds (and accompanying territorial males) is that the female herds may be more numerous and found in predictable places. For example, presentday observations of female gazelles in the xeric Mediterranean environment of Ramat Hanadiv (southwestern Mount Carmel) show that they maintain home ranges of ca. 0.1e0.2 km2 throughout their lives (Geffen et al., 1999). Therefore, female herds are easier to locate than the more dispersed male bachelor herds. Nevertheless, the still-considerable proportion of males means that, in addition to targeting territorial males that accompany the female herds, male bachelor herds may have been attacked upon encounter. Such a reconstruction may imply that gazelles were hunted in the summer and autumn, when adult males hold territories and are joined by a herd of females from all age classes. Some support for this can be found in the age classes that contain a considerable proportion of “yearlings”; assuming that gazelles gave birth mainly

Figure 12. Comparison of selected Manot gazelle bone measurements (Ahmarian [AHM] and Aurignacian [AUR] samples are shown where the former's sample size permits) to modern known-sex gazelle measurements from Israel (Munro et al., 2011). See SOM Table S8 for statistics.

Please cite this article as: Yeshurun, R et al., Early Upper Paleolithic subsistence in the Levant: Zooarchaeology of the AhmarianeAurignacian sequence at Manot Cave, Israel, Journal of Human Evolution, https://doi.org/10.1016/j.jhevol.2019.05.007

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Table 8 Summary of sexing results by character traits for the Manot gazelles. Ahmarian

a

Horn-core NISP Pubis NISP Total Proportion of sex a

Aurignacian

Female

Male

Female

Male

5 0 5 63%

2 1 3 38%

6 10 16 62%

2 8 10 38%

Abbreviations: NISP ¼ number of identified specimens.

in the late spring, greater availability of animals between the end of their first year of life and the first half of their second year could match a warm-season habitation. This also conforms to Speth's, (2019) suggestion that females would be preferentially hunted in the warm season, because the males are in rut and their physical condition is very poor. However, our aging resolution is not high enough to determine the season with any certainty, and younger juveniles (which could indicate winter kills) may have been avoided for reasons other than

seasonal availability. Also, an amalgamation of hunting episodes spanning several seasons, each with its own demographic make-up, cannot currently be ruled out. More lines of evidence for seasonality will be gathered in future research to shed light on this point. Meanwhile, the similar demographic patterns of the Ahmarian and Aurignacian could be taken to mean that the human habitation of the cave occurred in similar seasons/s during both periods. Our skeletal-element profiles shed some light on where these kills were made. The most important factors that determine carcass

Figure 13. Histograms of gazelle bone measurements, Manot-Aurignacian vs. modern male and modern female (from Munro et al., 2011): Metacarpal Bd, a sexually dimorphic elements (right) and Humerus Bd, which greatly overlaps in both gazelle sexes (left). Kernel density plots are shown.

Please cite this article as: Yeshurun, R et al., Early Upper Paleolithic subsistence in the Levant: Zooarchaeology of the AhmarianeAurignacian sequence at Manot Cave, Israel, Journal of Human Evolution, https://doi.org/10.1016/j.jhevol.2019.05.007

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Table 9 Aging data for fallow deer in Manot. Dental counts are NISP and bone fusion counts are MNE-based.a Ahmarian

dP4-worn M3 dP4-P4 Bone fusion a

Aurignacian

Juvenile

Adult

%Juvenile

Juvenile

Adult

%Juvenile

1 1 5

2 (2 prime-age, 0 old) 1 10

33.3% 50.0% 33.3%

5 5 13

6 (5 prime-age, 1 old) 2 15

45.5% 71.4% 46.4%

Abbreviations: NISP ¼ number of identified specimens, MNE ¼ minimum number of elements.

transport decisions are the animal's size and the distance of the kill site to the camp (Metcalfe and Barlow, 1992; O'Connell et al., 1988, 1990; Schoville and Otarola-Castillo, 2014). Gazelle and fallow deer hunting probably occurred in the cave's immediate proximity, because there is no evidence for field butchery and selective transport of body parts, not even for the larger fallow deer (see Yeshurun et al., 2007 for a different pattern at Misliya Cave). The cave is located in an ecotonal setting that allowed humans to exploit different biomes within a short walking distance. The even skeletal-element profiles point to Manot as having served as an EUP base-camp, in the sense of the final destination of the prey for processing and consumption. This is also evident from the numerous repeated occupations, high quantity and diversity of lithics and osseous items, and the presence (in Area E) of combustion features (Marder et al., 2019 submitted). Consumption of gazelle and deer was similar in both samples and routinely included meat removal and fracturing of many limb bones and phalanges for marrow. The bones were discarded already as defleshed and demarrowed fragments. No patterns of “wasteful” behavior were observed here, in contrast to open-air kill sites in Europe where many individuals were not completely exploited (e.g., Gaudzinski and Roebroeks, 2000; White et al., 2016). Coupled with the even skeletal-element profiles, this is consistent with the notion of a base-camp to which carcasses were imported, and does not support a mass-kill scenario as recently suggested for late Pleistocene gazelle in the Levant (Speth, 2019). Following consumption and discard, some skeletal elements were fashioned into tools, most notably ungulate tibia and metapodial shafts that were slightly modified to make awls in both samples, and antler-working to make projectile points in the Aurignacian (Tejero et al., 2016). The refuse from butchery and consumption activities, along with toolmaking and other processes, was discarded in the cave and underwent mild destruction from hyena activity and chemical diagenesis. These processes mainly caused the attrition of the least dense skeletal elements but probably did not have a strong effect on the denser ones. Assuming that the sediment deposition rate was similar in the Ahmarian and Aurignacian units, as implied by the radiocarbon chronology (Alex et al., 2017), then the much greater quantity of bones in the Aurignacian, coupled with the fact that bones were less exposed to post-discard damage by carnivore ravaging and weathering, could represent more frequent visits to the cave, or habitations of longer duration. 4.2. EUP animal subsistence in the southern Levant Our results verify and expand on older observations of EUP hunting patterns in the southern Levant (summarized in Rabinovich, 2003, 2017; and see Clark and Stutz, 2014; Shimelmitz et al., 2018; Speth, 2019). EUP archaeofaunas in the Mediterranean zone of the southern Levant are always dominated by small and medium ungulates: gazelle and fallow deer in Mount Carmel and Galilee (e.g., Davis, 1982; Marín-Arroyo, 2013; Speth, 2019), supplemented by wild goat in the Jordan Valley (Clark and Stutz, 2014).

Larger ungulates such as red deer, equids, aurochs and rhinos are rare or absent at most sites, especially in relation to the preceding late Middle Paleolithic (Davis et al., 1988; Stiner, 2005; Speth, 2019). Mountain gazelle is always the dominant ungulate species. The Aurignacian layer at Hayonim Cave is heavily gazelle-dominated (Rabinovich, 1998) and, to a lesser extent, so is the Aurignacian layer at Sefunim Cave (Shimelmitz et al., 2018). Hunting of small to medium-sized ungulate prey is also evident in the few EUP case studies from the Mediterranean zone of the central and northern Levant, where subsistence was based on hunting fallow deer, roe deer, wild boar and caprines (Stiner, 2009; Bosch et al., 2015a). An unresolved issue is the impact of hyena denning on the assemblages. Hyenas seem to have preferred medium-sized ungulates (i.e., fallow and red deer) rather than gazelle, and so their activities may have biased some assemblages in favor of deer (Orbach and Yeshurun, 2019). Thus, in the EUP of Qafzeh Cave the ungulate assemblage may well reflect hyena hunting as much, if not more, than human choice (Rabinovich et al., 2004). The impact of hyenas on the EUP of Kebara may also be considerable, especially for the remains that were retrieved from the center of the cave (Speth, 2019). This problem is largely (but not completely) avoided at Manot due to the spatial segregation of human and hyena activities in the cave. The marked hyena presence in the EUP of the Levant should be considered when attempting to compare assemblages that have varying degrees of carnivore influence. Furthermore, the existence of such a formidable competitor targeting medium-sized ungulates may have caused humans to increasingly focus on smaller ungulates and small game (Orbach and Yeshurun, 2019). The relative contribution of non-ungulate species to diets is unknown at many sites because small game was not systematically collected or counted. At the Aurignacian levels of Hayonim Cave small game taxa, including reptiles, birds and hares, do not exceed 25% of NISP (Stiner, 2005). As in the Manot Aurignacian, small mammal remains are scarce, but bird remains are relatively abundant at 8% of total NISP (Stiner, 2005). Many of these remains are large birds that bear substantial evidence for human exploitation of meat and feathers (Rabinovich, 2003). A small assemblage of mostly medium-sized and large birds was published from the EUP layers of Ksar Akil (Kersten, 1991). Increased bird exploitation may have been a hallmark of the Aurignacian in the Levant. An additional under-studied issue is the role of aquatic resources in the Levantine EUP. Fish remains seem to have been rare in this period. Some late Upper Paleolithic sites located near lakes or wetlands reveal the capture of fish and waterfowl in addition to gazelle and deer (Simmons and Nadel, 1998; Martin et al., 2013; Zohar et al., 2018), but these are the exceptions that prove the rule; the exploitation of aquatic resources in sites that are not located directly on the lakeshore was negligible. However, marine mollusks may have played a considerable economic and symbolic role at some sites close to the coast. Shells of edible mollusks bearing taphonomic evidence for consumption are abundant in the
ızlı Caves EUP levels of Ksar Akil (Bosch et al., 2015b) and Üçag (Stiner, 2009), located more to the north of the Levantine coast and

Please cite this article as: Yeshurun, R et al., Early Upper Paleolithic subsistence in the Levant: Zooarchaeology of the AhmarianeAurignacian sequence at Manot Cave, Israel, Journal of Human Evolution, https://doi.org/10.1016/j.jhevol.2019.05.007

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closer to rocky shores. The isotopic study of the consumed EUP mollusks at Ksar Akil showed that they were gathered in all seasons and hence provided a year-round source of food (Bosch et al., 2017). As in the taxonomic abundances, the demographic profiles of gazelle in Manot also seem to accord well with other EUP occurrences. Sub-adult animals constituted between one quarter to one half of hunted individuals in the EUP of Kebara (Speth, 2019), Hayonim (Rabinovich, 1998) and Mughr el-Hamamah (Clark and Stutz, 2014). Our survivorship curves put these EUP (and Kebaran) patterns in perspective and show that more sub-adults were taken in Manot compared to the more adult-dominated Middle Paleolithic, a picture that seems to hold for other sites as well (Rabinovich, 2003). The curves also demonstrate a different pattern in the terminal Pleistocene Natufian, when adult males and fawns were often targeted (Davis, 1983; Cope, 1991; Yeshurun et al., 2014a). The unique demographic patterns seen in the Natufian contrast with the generally non-selective gazelle-hunting patterns of the EUP. While the faunal patterns of the EUP in the Mediterranean southern Levant are slow to be unveiled, we do have the luxury of rich zooarchaeological evidence from the preceding and succeeding periods to put our results in broader temporal perspective. The hunting patterns in Manot, when viewed in the Levantine Paleolithic sequence, were generally very similar to later Upper Paleolithic and pre-Natufian Epipaleolithic patterns in the same region (Rabinovich, 2017; Munro et al., 2018). Gazelles and to a lesser extent fallow deer, were the most important prey taxa, with an emphasis on sub-adult hunting. Some small game species were sporadically taken, with fowling gaining some importance by the Aurignacian (e.g., Davis, 1974; Hovers et al., 1988; Bar-Oz et al., 1999, 2002, 2003; Bar-Oz, 2004; Kuhn et al., 2004; Rabinovich and Nadel, 2005; Marom and Bar-Oz, 2008; Napierala, 2011). The EUP, like the later Upper Paleolithic and earlier Epipaleolithic cases, occupies a middle position between the Middle Paleolithic and the Natufian of the Mediterranean southern Levant sequence. In the Middle Paleolithic, less intensified and narrower diets consisting of hunting prime-age ungulates and capturing tortoises, with little contribution from other small game animals, is noted in several caves and rock shelters (Stiner, 2005; Yeshurun et al., 2007; Speth, 2012). In contrast, in the terminal Pleistocene Natufian ungulates larger than gazelle were almost completely “replaced” by an array of small game species including fast small game and aquatic taxa, and data for the only remaining ungulate staple, gazelle, reveals the hunting of fawns and a bias toward the taking of male adults (Stiner et al., 1999; Bar-Oz, 2004; Davis, 2005; Munro, 2009; Edwards and Martin, 2013; Yeshurun et al., 2014a). If the breadth of the taxonomic spectrum and the proportion of sub-adults and fawns are used as a marker of intensification, then the EUP societies were somewhat more intensified than the preceding Middle Paleolithic (Stiner et al., 1999; Stiner, 2005), but still maintained similar foraging lifeways. Hunting patterns remained relatively unchanged until the onset of the Natufian at ca. 15,000 BP. 4.3. Ahmarian vs. Aurignacian adaptations Turning back to the Manot sequence, our results provide the first Ahmarian vs. Aurignacian faunal comparison, yielding a broadly similar picture of animal exploitation but with several interesting differences. Preference for gazelle can be argued for both entities, but is more significant in the Aurignacian. The null hypothesis, that the abundance of gazelle signals higher availability of this animal in the cave's surroundings and therefore is a function of environmental change, can be tested against the paleoenvironment reconstructed from other proxies as well as the nearcontemporaneous hyenas' hunting choice in Manot. The latter

evidence strongly points to high natural availability of fallow deer, as the hyenas that inhabited the cave during the EUP hunted mainly fallow deer, probably near the site (Orbach and Yeshurun, 2019). The speleothem and charred wood isotopic records hint at wetter oscillations and slightly more woody coverage during the Aurignacian (Caracuta et al., 2019; Yasur et al., 2019), again suggesting an expansion of fallow deer-suited biomes. On the other hand, the micromammals study indicates somewhat reduced woodland coverage in the Aurignacian (Comay et al., 2019). There is no doubt that a mosaic of biomes existed near the cave in EUP times, allowing humans and hyenas to select their favorite hunting grounds based on an array of considerations. Evidently, EUP and especially Aurignacian humans intentionally opted for taking gazelle, irrespective of simple availability or encounter-rate considerations. Gazelle preference in the Mediterranean southern Levant has previously been argued for the Middle Paleolithic (Yeshurun, 2013) and is now demonstrated for the Upper Paleolithic Levant as well, based on our comparison of the Manot human archaeofauna and hyena den (Orbach and Yeshurun, 2019). Mesopotamian fallow deer, which were undoubtedly abundant in the landscape in the immediate vicinity of the sites as evident in hyena dens and natural traps, were secondary to gazelle even though they provide considerably more meat and fat. Thus, the Middle and Upper Paleolithic preference for gazelle demands some explanation. The major difference between these two ungulates, besides their size (ca. 20 kg for gazelle, 80 kg for deer) is their environmental adaptation. Mountain gazelles are grazers that thrive in open landscape (shrubland rather than garrigue and maquis), and are sensitive to low temperatures and can tolerate aridity quite well (Yom-Tov, 2016). On the other hand, Mesopotamian fallow deer are browsers that spend much of their time in woodland settings (BarDavid et al., 2005). Since the scenario of environmental change that caused the contraction of woodlands in the Aurignacian is less likely, other potential explanations for the gazelle abundance are the intensification of hunting due to higher site-occupation intensity (which led to taking the lesser ranked species), or preference for hunting in open environments because of a technological adaptation for killing at a distance. Evolved projectile technology could have played a role in favoring open environments in the EUP sequence, because prey can be located from afar and killed at a considerable distance (see discussion in Yeshurun, 2013). Indeed, Yasur et al., (2019) demonstrated close correlation between the proliferation of open environments and the timing of EUP habitations in Manot. The scenario of Aurignacian hunting specialization in open biomes may seem attractive, given the evidence for the abundance (perhaps prevalence) of such biomes in the immediate vicinity of the cave (Comay et al., 2019) and the conspicuous appearance of osseous projectile points (Tejero et al., 2016). Such an adaptation would almost certainly result in gazelle-dominated hunting. However, the Ahmarian hunters in Manot lived in similarly open environments and, while lacking osseous projectile points, they did possess an array of lithic projectiles (Abulafia et al., submitted) that could have been used as dart or atlatl points (e.g., Shea, 2006). Hence, we cannot argue for less evolved projectile technology or other adaptations to exploiting grassy landscapes in the Ahmarian compared to the succeeding period, and this scenario does not explain the shift away from exploiting woody environments in the Manot sequence. In fact, it is the intensification scenario that can explain the patterns we see. Our behavioral ecology assumptions showed that subsistence intensification can be tracked by increased emphasis on lower-ranked resources: shifting from larger-bodied ungulates to smaller ones, culling more juvenile ungulates and investing more

Please cite this article as: Yeshurun, R et al., Early Upper Paleolithic subsistence in the Levant: Zooarchaeology of the AhmarianeAurignacian sequence at Manot Cave, Israel, Journal of Human Evolution, https://doi.org/10.1016/j.jhevol.2019.05.007

R. Yeshurun et al. / Journal of Human Evolution xxx (xxxx) xxx

in the capture of fast small game. With the exception of a higher proportion of juveniles, these are exactly the patterns that set the Aurignacian sample apart from the underlying Ahmarian assemblage. Such intensification should result either from environmental deterioration (for which we have no unequivocal evidence) or from increased numbers of people that need to be fed. Thus, Aurignacian occupations in the cave may have been more frequent, longer, or involved a larger group of people. Additional lines of evidence that suggest greater occupation intensity in the Aurignacian are the much higher quantity of bones (scaled by excavation volume) and less weathering compared to the Ahmarian. Both suggest faster build-up of habitation refuse as a result of more intensive occupations (compare Stiner and Munro, 2011). Significantly, the Aurignacian occupations at Manot accumulated to a thickness of >2 m and produced a large variety of flint artifacts, shells, bone tools and incised bones and grinding stones, as well as the remains of several human individuals. This again points to base-camps of some duration and a range of activities that were performed in the cave (Marder et al., 2019 submitted). Thus, we suggest that the Levantine Aurignacian cultural entity is generally one of higher site-occupation intensity compared to the “local” Ahmarian entity. Increased site-occupation intensity in the Aurignacian may explain some of the variability seen in the Levantine EUP record. Specifically, the presence of Aurignacian camps mostly in the Mediterranean region where longer duration of occupation (perhaps by more people) is feasible when compared to the lower carrying capacity of the arid zones. More indications of flint economizing behaviors, such as greater reuse and curation, were observed in the Aurignacian (Parow-Souchon, 2016). Also, more investment is seen in the production of personal ornaments in the form of perforated deer canines (Belfer-Cohen and Bar-Yosef, 1981; Tejero et al., 2019), which may have stemmed from intensifying social relations in a larger group, or more interactions between groups (compare Stiner, 2014 for the northern Levant Ahmarian patterns). The early Ahmarian and Levantine Aurignacian of the southern Levant probably do not reflect adaptations to different environmental conditions, but they do exhibit some differences that can be attributed to differing mobility patterns and group organization. Whether these arose as a result of migration of forager groups into the Levant who captured the most productive environments and maintained relatively small territories with higher social cohesion, is an open question. 5. Conclusions This study focused on EUP adaptive behavior in the Levant. It has contributed by, first, contextualizing the EUP hunting patterns in the regional sequence and second, by conducting the first zooarchaeological study of the “two-tradition model”. The EUP patterns as a whole seem to concur with other Upper Paleolithic and pre-Natufian Epipaleolithic cases in the southern Levant; taxonomic abundances and game demographic profiles situate them in a middle position between the less intensified diets of the Middle Paleolithic and the broad-spectrum foraging seen in the terminal Pleistocene Natufian of the same region. Additionally, we found no evidence to suggest that the Ahmarian and Aurignacian of the Mediterranean zone of the Levant represent (or were caused by) adaptations to different environments. On the contrary, both samples exhibit similar patterns of animal exploitation, and the differences that do exist do not conform to the independent paleoenvironemntal trends in the same cave. Rather than environmentally derived differences, we found support for the hypothesis that the two entities differed in mobility patterns and site use. Somewhat greater site-occupation intensity seems to have

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occurred in the Aurignacian, as evidenced by greater reliance on smaller game, increased bone deposition and perhaps shorter abandonment periods. This could represent a certain decrease in mobility or somewhat larger groups inhabiting the cave. More studies oriented toward the adaptations of these two EUP traditions in the Levant are needed to determine if this conclusion can be generalized to characterize the Levantine Aurignacian cultural entity. Acknowledgements This study is based on N.S.-P.‘s M.A. thesis research in the Department of Archaeology, University of Haifa, supervised by Guy Bar-Oz and Ofer Marder. We thank Guy Bar-Oz for his help and support in the course of this research and for his comments on a previous draft. We thank the Manot Cave team for their assistance, especially Israel Hershkovitz, Mae Goder-Goldberger, Ron Lavi, Talia Abulafia and Meir Orbach. Miriam Feinberg Vamosh provided editing assistance and Patrice Kaminsky provided graphic assistance. We thank Meir Orbach and four anonymous reviewers for helpful comments on an earlier draft of the paper. Fieldwork at Manot Cave was supported by the Dan David Foundation, Binational Science Foundation (grant no. 2015303), Israel Science Foundation (grant no. 2632/18), and The Leakey Foundation. Supplementary Online Material Supplementary online material to this article can be found online at https://doi.org/10.1016/j.jhevol.2019.05.007. References Abulafia, T., Goder-Goldberger, M., Berna, F., Barzilai, O., Marder, O., 2019. A technotypological analysis of the Ahmarian and Levantine Aurignacian assemblages from Manot Cave (Area C) and the interrelation with site formation processes. Journal of Human Evolution submitted to special issue on Manot Cave. Alex, B., Barzilai, O., Hershkovitz, I., Marder, O., Berna, F., Caracuta, V., Abulafia, T., Davis, L., Goder-Goldberger, M., Lavi, R., Mintz, E., Regev, L., Bar-Yosef Mayer, D., Tejero, J.-M., Yeshurun, R., Ayalon, A., Bar-Matthews, M., Yasur, G., Frumkin, A., Latimer, B.G., Hans, M.G., Boaretto, E., 2017. Radiocarbon chronology of Manot Cave, Israel and Upper Paleolithic dispersals. Science Advances 3 e1701450. Baharav, D., 1974. Notes on the population structure and biomass of the mountain gazelle, Gazella gazella gazella. Israel Journal of Zoology 23, 39e44. Baharav, D., 1983. Observation on the ecology of the mountain gazelle in the Upper Galilee, Israel. Mammalia 47, 59e69. Bar-David, S., Saltz, D., Dayan, T., 2005. Predicting the spatial dynamics of a reintroduced population: the Persian fallow deer. Ecological Applications 15, 1833e1846. Bar-Oz, G., Dayan, T., Kaufman, D., 1999. The Epipalaeolithic faunal sequence in Israel: a view from Neve-David. Journal of Archaeological Science 26, 67e82. Bar-Oz, G., Dayan, T., 2002. “After 20 Years”: a taphonomic re-evaluation of Nahal Hadera V, an Epipalaeolithic site on the Israeli coastal plain. Journal of Archaeological Science 29, 145e156. Bar-Oz, G., Dayan, T., 2003. Testing the use of multivariate inter-site taphonomic comparisons: the faunal analysis of Heftziba in its Epipalaeolithic cultural context. Journal of Archaeological Science 30, 885e900. Bar-Oz, G., 2004. Epipaleolithic Subsistence Strategies in the Levant: A Zooarchaeological Perspective. ASPR Monograph Series. Brill, Boston. Bar-Oz, G., Dayan, T., Kaufman, D., Weinstein-Evron, M., 2004. The Natufian economy at el-Wad Terrace with special reference to gazelle exploitation patterns. Journal of Archaeological Science 31, 217e231. Bar-Oz, G., Munro, N.D., 2007. Gazelle bone marrow yields and Epipalaeolithic carcass exploitation strategies in the southern Levant. Journal of Archaeological Science 34, 946e956. Bar-Oz, G., Yeshurun, R., Weinstein-Evron, M., 2013. Specialized hunting of gazelle in the Natufian: cultural cause or climatic effect? In: Bar-Yosef, O., Valla, F.R. (Eds.), Natufian Foragers in the Levant: Terminal Pleistocene Social Changes in Western Asia. International Monographs in Prehistory, Ann Arbor, pp. 685e698. Bar-Yosef, O., Belfer-Cohen, A., 2010. The Levantine Upper Palaeolithic and Epipaleolithic. In: Garcia, E.A.A. (Ed.), South-Eastern Mediterranean Peoples Between 130,000 and 10,000 Years Ago. Oxbow, Oxford, pp. 44e167. Barzilai, O., Hershkovitz, I., Marder, O., 2016. The early upper Paleolithic period at Manot Cave, western Galilee, Israel. Human Evolution 31, 85e100.

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