The effect of terrain on Neanderthal ecology in the Levant

Quaternary International xxx (2015) 1e12

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The effect of terrain on Neanderthal ecology in the Levant Donald O. Henry a, *, Miriam Belmaker a, Sean M. Bergin b a b

Department of Anthropology, Harwell Hall, The University of Tulsa, Tulsa, OK 74104, USA School of Evolution and Social Change, Arizona State University, PO Box 872402, Tempe, AZ 85287, USA

a r t i c l e i n f o

a b s t r a c t

Article history: Available online xxx

Our study assesses the influence of differences in terrain and locomotor energetics on the land-use strategies and settlement patterns of Levantine Neanderthals and Modern Human e Early Upper Paleolithic groups through a digital application of site catchment analysis. Our findings indicate that Neanderthals habitually commanded smaller site exploitation territories (SETs), principally situated in the rugged Mediterranean Woodlands of the Levant, whereas early Upper Paleolithic groups generally enjoyed larger SETs and displayed a more generalized, wider settlement range encompassing both rugged woodland and more regular, level steppe landscapes. The broader implications of these findings may explain the biogeographic limits on the Neanderthal dispersal into Southwest Asia. © 2015 Elsevier Ltd and INQUA. All rights reserved.

Keywords: Neanderthal Ecology Terrain Site catchment analysis

1. Introduction Variation in terrain is generally accepted as one of the fundamental factors in understanding ecological adaptation, even to the extent that the common names of many taxa are often reflective of their habitats as defined by terrain; e.g., mountain gazelle, upland and lowland gorillas and so forth. While terrain is recognized as an important variable in general ecologic research, paleoanthropologists have given little attention to the adaptive responses of hominins to variations in terrain or landscape features, and this is especially relative to Neanderthal ecology. The interrelationship of Neanderthals and their environment has been examined largely from the perspective of biotic evidence of floral and faunal remains and attendant climatic reconstructions. Given their inherent energetic and locomotor differences from modern humans, an understanding of the effect of terrain on Neanderthal foraging patterns is especially relevant as to how variations in landscape influenced Neanderthal land-use and settlement-procurement patterns and, in turn, their biogeography (Burke, 2006; Miller and
n, 2007; Uthmeier et al., 2008; Barton, 2008; Finlayson and Carrio Churchill, 2014). Advances in satellite imagery and digital science have prompted a wide-range of researchers to use terrain measures as a means of better defining the biogeography of plant and animal populations and to correlate their distributions to other landscape features. In

* Corresponding author. E-mail address: [email protected] (D.O. Henry).

the research reported upon here, we adopted a similar strategy using satellite imagery and geo-spatial digital data to assess the degree to which variation in terrain influenced the behavioral ecology of Neanderthals within the Levantine Near East. At a more specific level, our approach involved site catchment analysis (Vita-Finzi and Higgs, 1970; Bailey and Davidson, 1983) as a means of integrating site specific, digital data and assessing the degree to which terrain may have played a role in shaping the foraging strategies of Neanderthals and in producing any regional adaptive differences between Neanderthals and Anatomically Modern Human (AMH) groups. In comparing the effects of terrain on Neanderthals and AMH groups, we focused on site-specific data associated with the latest proposed pulse of modern human expansion into the Levant from Africa and/or Arabia during MIS 4 e early MIS 3, reflected in early Upper Paleolithic (UP) sites. 2. Site catchment analysis The underlying concept of site catchment analysis (SCA) is that prehistoric groups exploited resources habitually within a nearby site exploitation territory (SET) and occasionally from more distant sources within a site catchment (SC) conditioned by the competing factors of the economic value of resources and the energy expended in procuring them, essentially following the law of diminishing returns. The term site territory was initially used by Vita-Finzi and Higgs (1970), but Bailey and Davidson (1983) suggested the term site exploitation territory (SET) to avoid confusion with social, defended site territories. Bailey and Davidson (1983) also proposed that SCA and STA be distinguished, a suggestion not followed here

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in that we view a SC as a larger spatial scale that encompasses one, or perhaps, several site territories. In any event, this is a topic beyond the scope of our paper. In many ways, at a more detailed level, this notion parallels the fundamental ecological concepts of optimal foraging theory as expressed in equations including such specific variables as search time, energetic expenditures and food yields. In operationalizing SCA, researchers have adopted different approaches in determining the time-distance limits of a SET and the manner in which the configuration of a SET is reconstructed around a site. While it is given that there must be some finite spatial limit to the area about a site that was exploited, the question is, how is this to be determined? In their seminal study, Vita-Finzi and Higgs (1970) used ethnographic observations to establish time (2 h) and distance (10 km) limits in configuring their SETs, measures that most researchers have subsequently followed. Although the 2 h outbound time for foraging from a base-camp has been traditionally employed in site catchment studies, this likely represents the lower range of foraging time in that more comprehensive ethnographic data indicate a range of 2.5e9 h and a clear difference between average female (2.8 h) and male (5.9 h) foraging times (Binford, 2001, p. 235e38). In reconstructing a SET for a foraging group there are numerous variables such as ground conditions (sand, snow, rough landscapes, thick vegetation), carried load, and even social considerations that might affect forage time-distance, but topography and locomotor energetics are the most important factors, at least the ones that are reasonably knowable. In practice, SCA was initially based on actualistic assessments of SETs by walking out from a site along a number of arbitrarily selected, 2-hour transects and then visually interpolating these distant points on a map (Higgs, 1975; Bailey and Davidson, 1983). Given the difficulty in conducting actualistic assessments of SETs in many archaeological projects, researchers have adopted another approach that simply involves overlaying a circle of 10 km radius on a map around a site. While this “magic circle” approach is quick and convenient, it fails to reflect the importance of terrain in regulating energetic expenditures and ultimately in spatially defining a SET. In recognizing the importance of terrain variations to SET reconstructions, Bailey and Davidson (1983) devised a technique based upon Naismith's Rule that calculated hiking time/distance in consideration of variation in elevation. Their technique allowed for a relatively precise reconstruction of a SET using the time-distanceelevation variables provided in the Naismith equation applied on a topographic map. In the research program described here, we generally follow the approach introduced by Bailey and Davidson (1983), but introduce a digitally-based methodology that is sensitive to variations in terrain as well as differences in hominin locomotor energetics. A SET can be examined using two complimentary approaches. The first is the area (km2) of a SET and the second is the Accessibility Index (AI) that compares this area to the area of a hypothetical SET around a site that displayed level, unvarying terrain. An AI, factoring in the differences in hominin energetics, is seen as a proxy of the ruggedness of a SET and its actual effect on the foraging limits of Neanderthals and modern humans. The AI is calculated for each site by comparing the actual SET based on topography to the SET area generated with no topography (a completely flat plain). This flat digital elevation model (DEM) functions as a ‘hypothetical maximum’ that can be walked within 2 h without any influence from topography. The AI is calculated by dividing the area of a SET conditioned by topography by the hypothetical maximum SET to demonstrate the influence of site specific topography. For example, a catchment with an AI of 0.70 would mean that 70% of a potential 100% of the SET area surrounding a site with unvarying terrain could be exploited within a 2 h walking time, based on specific hominin energetics. Coastal sites, especially those framed by very

rugged inland terrain, often displayed exceptionally small SET areas and AI's because of their limited access to exploitable terrestrial landscape and in these cases the implied ruggedness of the AI is likely exaggerated. However, in such cases the SET area and AI nevertheless provide an actualistic representation of the foraging parameters for the prehistoric occupants of these shoreline sites. 3. Neanderthal locomotor energetics When studying how Neanderthals related to their environment it is also important to keep in mind that Neanderthal locomotor energetics differed from those of modern humans. Looking at gross morphology, Neanderthals had much shorter lower limbs, both absolutely and also relative to the trunk height, than modern humans as well as higher brachial and crural indices (Weaver and Steudel Numbers, 2005) and were more heavily muscled. There has been a debate in the literature if the unique morphology of the Neanderthals reflects an adaptation to climate (Steegmann et al., 2002; Aiello and Wheeler, 2003; Churchill, 2006) or a difference in locomotor energetics and mobility efficiency (Steudel-Numbers and Tilkens, 2004; Hora and Sladek, 2014), such as one that focused on a close range hunting strategy over persistence hunting that involved endurance running (Bocherens et al., 1999, 2001, 2005; Richards et al., 2000, 2005; Raichlen et al., 2011). In addition, Neanderthal morphologies (as indicated by both the effective inter-membral index and the length of the Achilles tendon) may have actually conferred a locomotive advantage in a rugged landscape when compared to modern humans who would have had a selective advantage in open habitats where running long distances would have been beneficial (Higgins and Ruff, 2011). Current data also suggests that Neanderthals had a higher daily energetic cost of foraging compared to Early Upper Paleolithic modern humans based on lower limb length, body mass and both (Frohehle and Churchill, 2009; Snodgrass and Leonard, 2009). The lower energy expenditure of AMH is thought to have conferred a selective demographic and reproductive advantage and to have played an important role in the competitive exclusion between the two species. Our understanding of Neanderthal locomotor constraints and high energetic costs has prompted proposals that Neanderthal groups exploited smaller foraging ranges than those of modern humans (Verpoorte, 2006; Macdonald et al., 2009), however, these notions have been largely hypothetical. Similarly, it has been argued that Neanderthals were limited to 0.5e1 h to 3 h average walking time for the acquisition of stable food resources, compared to 2e10 h for AMH hunter gatherers (Uthmeier et al., 2008, p. 487). Hayden (2012, p. 13) has suggested that Neanderthals habitually exploited areas within 5 km of sites based upon procurement dis
blot-Augustins (1997) for tances of lithic resources as reported by Fe Middle Paleolithic occupations. Although some materials were acquired from greater distances, 60e98% of all artifacts were acquired
blot-Augustins, 1997, p.62). from sources within 5 km of the sites (Fe Uthmeier et al. (2008) estimated walking times of 2e3 h, with concomitant distances of 6e8 km, for dedicated forays to chert sources from the site of Kabazi V in the Crimea, but they also note that most daily foraging activities were likely to have involved smaller SETs restricted to 2 h, one-way walking times. These data are noteworthy, in that a 5 km radius for SETs is considerably smaller than the mean distances reported in ethnographic contexts by Binford (2001, p. 238) for six groups of 7.3 km (female) and 8.7 km (male), by Marlowe (2005, p. 63) for 8 groups of 9.5 km (female) and 14 km (male) and by Churchill (2014, p. 308) of 6e17 km (female) and 12e25 km (male). However, most energetic studies (but see Frohehle and Churchill, 2009) have focused only on Western European

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specimens, which maximized the difference between the morphologies of the two populations due to maximum/pronounced climatic differences. Comparison of energetic studies between fossil populations inhabiting the Levant, which exhibits fewer/less pronounced climatic changes between the glacial and inter-glacial periods, and is situated in the mid to low latitudes have not been done. While there is no significant difference in lower limb robusticity between populations of Near Eastern Middle Paleolithic hominins (Trinkaus et al., 1998), it also has been noted that there is a shift in lower limb morphology between Neanderthal populations and Upper Paleolithic modern human populations. Specifically, mean limb length increased ca. 7% and crural indices increased about 10% between the MP and UP (Trinkaus, 1986). This suggests that Neanderthal populations in the MP may have differed in their locomotion stride and efficiency from UP modern humans. In our study, we ask if Levantine Neanderthals were able to minimize their energetic costs by habitually inhabiting the rugged regions of their biogeographic range. While this has been generally suggested based on Neanderthal morphology (Higgins and Ruff, 2011), we ask the question by assessing terrain preferences and SETs of Neanderthals compared to randomly distributed sites and to sites occupied by UP Modern Humans in the Levant. We hypothesize that if Levantine Neanderthals were able to minimize their energetic costs by inhabiting the more rugged settings of the region, their SET areas and Accessibility Index (AI) values, viewed as a proxy of terrain ruggedness as explained earlier, would be lower than the SET areas and AI values of random points and also lower than the SET areas and AI values of the sites of UP modern humans. 4. Reconstructing site exploitation territories Site exploitation territories were configured around four groups of locational points following the concepts advanced in the seminal study of Vita-Finzi and Higgs (1970), but with significant technical modifications. In reconstructing the SETs for the four groups of locational points, a 2-hour walking time was adopted to calculate the spatial limits of foraging in consideration of variations in terrain as computed by the r.walk module of the GRASS GIS software. The SETs were therefore reconstructed weighted largely by topography as in the actualistic method followed by Higgs (1975) and as adapted to time-distance calculations and topographic map applications by Bailey and Davidson (1983). However, our reconstructions of the SETs also involved calculations that included the differences in walking speeds for Neanderthals and modern humans. Another important consideration is the degree to which the current and paleo-topographies differ. Clearly, the Levantine coastal topographies of MIS 4 e Early 3 would have varied from the modern coastline, while interior landscapes are likely to have generally resembled what we see today with exception to the shoreline fluctuations of the ancient Lisan Lake. In an effort to reconstruct realistic, time-dependent paleo-topographies we consulted studies of eastern Mediterranean sea-level (Schattner et al., ^an et al., 2012) and Lisan Lake level changes (Torfstein 2010, Dog et al., 2013) and factored the DEM data into our map constructions. 4.1. Data-sets, study area and site locations In order to better understand the relationship of terrain variability and Levantine Neanderthal settlement-procurement strategies and the degree to which these relationships may have varied from those of early modern humans, the specific locations of sites and their associated SETs were compared across four data-sets. The data-sets included locational points for Neanderthal sites (MP) which were divided into camp sites and kill sites, Initial Upper

Fig. 1. Map of the Levant as defined for the study area (modified after Bar-Yosef, 2000:112).

Paleolithic and Early Ahmarian sites (UP) and those randomly selected from the study area conditioned by Neanderthal energetic parameters. The geographic definition of the Levant varies not only between, but also within disciplines. In this study, we specifically adopted the regional definition used by Bar-Yosef (2000) in his overview of Levantine Middle and Upper Paleolithic site distributions, a regional delineation commonly followed by other researchers (Fig. 1). 4.2. The sites and locational points The specific locations of sites were obtained from primary site reports and from secondary sources that provided detailed maps, locational coordinates (typically as Geographic Coordinate System noting longitude/latitude) or specific descriptions of site settings. These locational points were then positioned on Google earth satellite images using coordinate data, if available, along with prominent landscape features and elevations as additional guides. The site coordinate information (UTM) generated from Google Earth was then imported into the GRASS GIS (GRASS Development Team, 2015) in combination with 90 m resolution DEM (Table 1). Mid-distance, single points were used to denote site clusters in those situations in which sites were separated by < 500 m. Although the coordinate data derived from Google earth is not likely to be as precise as on-site field coordinates established through GPS, for the purposes of our study the locational data are thought to be sufficiently refined.

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Table 1 The sites by period (MP e Middle Paleolithic, UP e Early Upper Paleolithic) showing the associated site exploitation territory (SET) area (km2) and accessibility index (AI). Site names followed by an * relate to site clusters formed by two or more sites in the immediate area. Also, note that the SET areas and AIs will differ for MP and UP occupations because of the differences in walking speeds for Neanderthals and modern humans. MP sites

SET area

AI

UP sites

SET area

AI

Amud Dederiyah Erq el Ahmar Fara II Geula Hummal Kebara Keoue Cave Ksar Akil Quneitra Shubabiq Shukbah Tabun Tor Faraj Tor Sabiha Umm el Tlel Yabrud I

177.714 203.780 178.532 247.479 251.448 250.225 246.127 244.118 118.981 261.087 175.851 194.910 221.000 206.834 191.662 246.872 198.563

0.666 0.764 0.669 0.927 0.943 0.938 0.922 0.941 0.459 0.978 0.659 0.730 0.828 0.775 0.718 0.925 0.744

Abu Halka Abu Noshra I &II Boker Tachtit* El Wad Emireh Erq el Ahmar Kadesh Barnea* Kanal Kebara Ksar Akil Lagaman Sites* Nizzana XIII Qafzeh Thalab al-Buhayra Tor Sadaf Ucagizli Umm el Tiel Wadi Sudr* Yabrud II

226.614 247.925 202.208 238.561 200.588 194.376 247.001 77.622 269.722 125.688 255.523 275.270 252.356 232.559 254.534 103.802 270.370 201.844 223.649

0.841 0.845 0.689 0.813 0.684 0.663 0.842 0.424 0.919 0.554 0.871 0.938 0.860 0.793 0.868 0.495 0.922 0.688 0.762

4.3. Neanderthal, Late Levantine Mousterian Sites (MP) Of the 17 sites in this group, only five (Tabun B, Kebara, Amud, Shuqbah and Dederiyeh) yielded Neanderthal fossil remains (Fig. 2). The stratigraphic position of the Tabun C1 mandible (Tabun I) has been widely questioned, however, a study of the unpublished notes of Garrod in conjunction with her excavation report indicate that the “Tabun woman” most probably came from layer B (Bar-Yosef and Callander, 1999). In absence of fossil

remains, the other sites were placed in the Neanderthal group on the basis of proxies associated with lithic artifact industrial placement and/or chronometry. The occupations containing Neanderthal remains are all associated with the distinctive technotypological signatures of the Late Levantine Mousterian and the dated ones cluster in MIS 4 e early MIS 3 between ~70 and 48 ka. From a technotypologic perspective, the lithic assemblages belong to the Tabun B-type Industry and share high Levallois, blade and faceting indices in conjunction with high frequencies of broadbased, triangular Levallois points often exhibiting classic chapeau de gendarme butts and concorde silhouettes (Meignen, 1995; Bar-Yosef, 2000; Meignen and Bar-Yosef, 2002). The site of Quneitra is the single exception to this Tabun-B Type affiliation, but the age of the site of ~54 ka is consistent with a Neanderthal association (Goren-Inbar, 1990). Although the Tabun technotypologic succession is traditionally followed as a region-wide framework for organizing the Levantine Mousterian (Shea, 2013, pp. 105e107 and references therein), technotypologic variance with the succession has been observed regionally (Hovers, 2009, pp. 228e229; Hauck, 2011, p. 534). And even among the relatively homogeneous Tabun-B assemblages, which concerns us here, technological attributes formerly thought to be time sensitive (e.g., lamellar index, bi-directional removals, cores-on-flakes) are now shown to be related to functional and lithic economizing strategies (Henry, 2003, p. 83). Of the MP group, four sites were identified as containing evidence for short-term, kill and butchery sites: Umm el Tlel, Hummal, Fara II, and Quneitra (Gilead and Grigson, 1984; € eda et al., 2001; Hauck, 2011; Jagher and Goren-Inbar, 1990; Bo Le Tensorer, 2011; Oron and Goren-Inbar, 2013), while the other sites were identified as base-camp sites. Here it is important to note that while Fara II and Quneitra have benefitted from extensive intra-site spatial analyses and been identified as killbutchery sites, both Hummal (Hauck, 2011) and Umm el Tlel

Fig. 2. Map of the study area showing the Middle Paleolithic (MP) sites (left) and early Upper Paleolithic (MP) sites selected for site catchment analysis. The site names followed by an * represent site clusters containing two or more sites in close proximity.

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€eda et al., 2001) present more complicated problems relative (Bo to the nature of their numerous, stratified occupations. Hauck (2011, p. 534) has noted that the Late Levantine Mousterian occupations at Hummal likely varied in activities dependent on fluctuating water levels of the basin with ephemeral, more task specific occupations taking place during low water conditions and repetitive and/or prolonged occupations during wetter episodes. At Umm el Tlel, with its 70 or so Late MP occupational layers, at least three different activity sets were identified including a prolonged occupation with diverse activities and temporary encampments associated with hunting/butchery or other special€ eda et al., 2001). ized tasks (Bo 4.4. Initial Upper Paleolithic and Early Ahmarian sites (UP) Within this group of 20 sites (Fig. 2), hominin fossil associations of AMH were recovered from five sites, Ksar Akil XVII (Bergman and Stringer, 1989; Douka et al., 2013), Kebara (Smith, 1995), el Wad (Smith, 1995), and Qafzeh UP (Vandermeerch, 1981); Unfortunately, all of the fossils are very fragmentary or lost (i.e., Ksar Akil's Egbert). The rest of the sites were placed within the Initial Upper Paleolithic and Early Ahmarian on the basis of distinctive technotypological signatures and chronometry. The technotypologic signatures of the Initial Upper Paleolithic consist of a melding of Late Levantine Mousterian (Levallois point technology) and Upper Paleolithic features in conjunction with some unique elements as expressed in Emireh, Ksar Akil and Umm el Tlel point types (Marks, 2003; Shea, 2013, pp. 140e141). Early Ahmarian assemblages are principally defined by an emphasis on the production of blade/bladelets from parallel sided cores (some opposed platform), often involving crested blade removals for initial core shaping and core tablet production for maintenance, in conjunction with a shift in loading from hard to soft-hammer (Marks, 2003; Shea, 2013). Typologically, the Early Ahmarian is distinguished by the common occurrence of pointed blade/bladelets, especially slightly retouched El Wad points (Coinman, 2003; Marks, 2003). The chronometry of Initial and Early Upper Paleolithic occupations is complicated by ages falling near the early limits of 14 C, coupled with differences in dating techniques (ABA vs ABOx-SC, Uranium-Thorium) and materials (charcoal, shell, calcite). However, in general, the Initial Upper Paleolithic is conventionally dated to ~47/45 ka followed by the Early Ahmarian from ~38/36 ka to 25 ka (Belfer-Cohen and GoringMorris, 2003).

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5. Site exploitation territory (SET) and Accessibility Index (AI) In reconstructing SETs, we used the r.walk module included in the GRASS GIS package (GRASS Development Team, 2015) to calculate a map of walking time costs within 2 h limits using each site as the starting point. We used two variables to compare the SET: Area (km2) and Accessibility Index (AI) (Fig. 4). Least cost maps produced by r.walk are anisotropic, which means that the algorithm takes direction into account when calculating walking time to distance (Fig. 3). For example, it takes more effort to walk up a slope than it does to walk down that same slope, thus the calculated cost of walking along a path in one direction may be lower or higher than it would be to walk along that same path in the opposite direction. In order to calculate a SET we used the variables A, B, C, and D in the GRASS GIS package to estimate the time cost of walking from one location to another based upon specific slope intervals. The

Fig. 3. An example of the site exploitation territory (SET) calculated by the GRASS module, GIS for a 2 h, out-bound walking time around the early UP site of Boker Tachtit conditioned by AMH friction costs. Note the difference between the GRASS generated SET and an arbitrary 5 km radius around the site.

4.5. Multi-component sites with long stratigraphic sequences There are several sites in the Levant, specifically from the rugged region that exhibit a long sequence, which eclipsed both the MP and the UP (e.g., Kebara, Ksar Akil). In these cases, we assigned the site both to the Neanderthal database and the UP modern human database.

4.6. Randomly selected points Within the study area, 60 points were randomly selected to be used in comparison to the groups of MP and UP archaeological sites in an effort to assess the degree to which the terrain of site locations may have differed from the terrain of the region as a whole. Random point locations were generated using the GRASS GIS r.random module which creates a specified number of random locations within a bounded area.

Fig. 4. An example of how an Accessibility Index (AI) is calculated showing a site exploitation territory (SET) calculated by the GRASS module from DEM data for the site of Erq el Ahmar compared to a hypothetical SET calculated for level terrain.

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coefficients used in the equation include (A) 1/walking speed where walking speed is time in seconds it takes to walk for a distance of 1 m over a flat surface, (B) additional walking time in seconds, per meter of elevation gain on uphill slopes, (C) additional walking time in seconds, per meter of elevation loss on moderate downhill slopes and (D) additional walking time in seconds, per meter of elevation loss on steep downhill slopes. The base line values were obtained from studies of modern hikers (Langmuir, 1984) and were (A): 0.72 (B) cost of uphill: 5.281, (C) cost of downhill: 1.278 and (D) cost of steep downhill: -2.781. In order to calculate coefficients that were specific to AMH and Neanderthals, we substituted walking speed with calculated walking speeds from the literature. For AMH we took into consideration both Upper Pleistocene AMH males and females from the Levant for an average value of 1.33 based on Hora and Sladek (2014). For MP Neanderthal population we used an average value of 1.27 based on Hora and Sladek (2014), which used both Levantine Neanderthals (Amud, Kebara and Shanidar) and European fossils. Coefficient A was calculated as 1/walking speed. To calculate the other coefficients we used the following equations: B ¼ A þ cost of uphill; C ¼ A þ cost of downhill and D ¼ A þ cost of steep down hill. The variables used in the calculations for SETs were for AMH: A ¼ 0.75; B ¼ 6.03; C ¼ 2.029; D ¼ 1.966 and for Neanderthals A ¼ 0.787; B ¼ 6.06; C ¼ 2.065; D ¼ 1.930. While we acknowledge that the limb bones of Levantine Neanderthals (specifically Amud 1), differed morphologically from European Neanderthals as well as from modern humans (Arensberg and Belfer Cohen, 1998), recent studies have shown that the important factor in locomotion energetics is total limb length rather than crural index (Pontzer, 2012). There is no difference between European and Levantine Neanderthals in femur, tibia, talar and calcaneal length (femur t test ¼ 0.507 df ¼ 10 p > 0.05; tibia t test ¼ 0.495 df ¼ 10 p > 0.05; talus t test ¼ 0.227 df ¼ 6 p > 0.05; calcaneus t test ¼ 0.333 df ¼ 5 p > 0.5) suggesting we can group European and Levantine Neanderthals into a single group. Moreover, Neanderthal specimens, which include the European specimens and the Levantine specimens, are more than 2 SDs below the range for modern human lower limb length (Pontzer, 2012).

SET area and AI distributions for MP base-camp sites, MP killephemeral sites, UP sites and Random points (Tables 2 and 3, Figs. 5 and 6).

Table 3 Mann Whitney pairwise comparisons of SET areas and AI's. Comparisons SET areas compared MP Base-camp to MP KillEmphemeral sites MP Base-camp to UP sites MP Base-camp to Random points MP Kill-Ephemeral to UP sites MP Kill-Ephemeral to Random points AIs compared MP Base-camp to MP KillEmphemeral sites MP Base-camp to UP sites MP Base-camp to Random points MP Kill-Ephemeral to UP sites MP Kill-Ephemeral to Random points

Results

Differ

z ¼ 2.6 two tailed p value ¼ 0.0092 z ¼ 1.76 two tailed p value ¼ 0.077 z ¼ 3.056 two tailed p value ¼ 0.0022 z ¼ 1.239 two tailed p value ¼ 0.215 z ¼ 0.832 two tailed p value ¼ 0.4054

Yes

z ¼ 2.26 two tailed p value ¼ 0.0235 z ¼ 0.29 two tailed p value ¼ 0.76 (z ¼ 2.94 two tailed p value ¼ 0.0033 z ¼ 2.866 two tailed p value ¼ 0.0042 z ¼ 0.832 two tailed p value ¼ 0.4054

No Yes No No

Yes No Yes Yes No

6.1. Middle Paleolithic SETs and AIs Overall, MP base-camp sites showed smaller SET areas and lower AI values than those of MP kill-ephemeral sites and random points. This implies that MP base-camp locations were situated in more rugged terrain with more compressed SET areas than seen for

6. Results of statistical analyses and discussion Given that none of the variables were randomly distributed we opted to use the pairwise Mann Whitney test for comparison of the Table 2 Descriptive statistics for SET area (km2) and AI for MP sites, UP sites and Random points. SET area (km2)

AI

Mean Median SD

212.658 206.834 38.248

0.799 0.775 0.143

Base-camp Sites N ¼ 13

Mean Median SD

200.732 198.563 35.845

0.755 0.744 0.135

Kill, Ephemeral Sites N¼4

Mean Median SD

251.416 248.852 6.611

0.942 0.933 0.025

Mean Median SD

215.816 229.586 55.082

0.760 0.803 0.144

Mean Median SD

232.200 246.940 38.460

0.870 0.920 0.144

MP sites All MP Sites N ¼ 17

UP sites All UP sites N ¼ 20 Random Random points N ¼ 60

Fig. 5. Box and whiskers plots of site exploitation territory (SET) areas (km2) for Middle Paleolithic and Upper Paleolithic sites and random points within the study area.

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the mean SET area is about 9% smaller than in the UP sites. The absence of clear statistical differences between the AI's is thought to reflect the largely rugged landscape of the study area in general and the overlap in the geographic distributions of MP and UP sites. In fact, four of the sites hold both MP and subsequent UP occupations. Beyond the SET areas, where MP and UP site distributions do differ greatly is in the paucity of MP sites in most of the open, more regular landscapes of the arid zone across the region. Although an MP presence has been recorded in ancient lake and marsh settings (e.g. Azraq, El Kowm) in the steppes of eastern Jordan and Syria, there is a striking absence of Late Levantine Mousterian (Tabun B-Type) occupations in extensively surveyed portions of Sinai, the Negev and west central Jordan. Only Fara II (Gilead and Grigson, 1984), situated in a riverine setting near the coast in the northwestern Negev stands out as an exception to this observation. 7. The influence of terrain on land-use strategies and settlement range

Fig. 6. Box and whiskers plots of Accessibility Indices (AIs) for Middle Paleolithic and Upper Paleolithic sites and random points within the study area.

the region as a whole, while MP kill-ephemeral sites were positioned in more open landscapes with attendant larger SETs. More specifically, MP base-camps were associated with caves or rock shelters located at topographic breaks that separate relatively even landscapes (e.g., coastal plain, broad interior valley) from more rugged uplands. Site specific (Henry, 2003, pp. 43e49) and areawide (Ullman et al., 2013) research in the region has shown that Neanderthal groups placed their base-camps in largeemedium size protected settings that were close to water and easily accessed from valley floors or lower slopes. The sites of base-camps also were positioned near stream confluences and ecotonal terrain settings that allowed for access to diverse environments (Henry, 2003, pp. 43e49,; Ullman et al., 2013) and in some situations appear to have supported multi-season occupations (Lieberman and Shea, 1994; Henry, 2003). In contrast to the MP base-camp data, the less permanent, more specialized occupations of MP kill-ephemeral sites (Fara II, Quneitra, Hummal and Umm el Tlel) are all open-air sites located in much more regular terrain with mean SET areas some 20% larger than their base-camp counterparts. Moreover, their placement on the landscape differed greatly from that of base-camps in that they were positioned in settings adjacent to lakes, springs and rivers where such water sources may have been especially attractive to ungulates and thus enhanced Neanderthal hunting strategies. Although the very high AIs (mean 0.94) of the MP kill-ephemeral sites indicate a potential for the nearly maximum use of the SETs, the regular terrain surrounding the sites would have commanded limited environmental diversity, especially beyond the areas immediately adjacent to the water sources. 6.2. Comparison of Middle and Upper Paleolithic SETs and AIs The SET areas of MP base-camps were significantly different from those of UP sites, but relative to AI's, only MP kill sites differed from the UP sites. Although not significantly different, the lower median of AI's of MP base-camps suggests a tendency for placement in more rugged terrain than that associated with UP sites and

Neanderthals appear to have been largely restricted to the more rugged terrain of the Levant that during MIS 4 and early 3 would have been principally associated environmentally with the Mediterranean woodland zone. Although there is evidence for forays into the more regular landscapes of the adjacent steppe zone, these appear to be primarily associated with more task specific activities (kill sites, quarries) and ephemeral occupations near permanent water sources, lacking evidence of habitual occupations as basecamps. 7.1. Environmental background Although the Mediterranean woodland is heavily vegetated by evergreen shrubs and trees, the warm-temperate biome is composed of a mosaic of different plant communities ranging from true forest to savannah. Within the Levant, woodland refers to regions with dense forest coverage (>80%) as well as maquis, garrige and batha communities associated with decreasing levels of ground coverage, density and height of vegetation (Allen, 2001). The eco-zone is estimated to presently cover some 80,000 km2 (Shea, 2008), but this likely represents a significantly smaller figure than that covered by Mediterranean woodlands during MIS 4 and perhaps even very early MIS 3 as a result of cooler temperatures and greater available moisture (Frumkin et al., 2011) coupled with the degraded landscape of the modern setting (Cordova, 2007). Within the biome, gradients of moisture and temperature produce a range of ground coverage of various heights and density as well as community affiliation. Given this, topography plays a large role in shaping the composition of plant coverage in that slope, exposure aspect and elevation all act to select for specific plant communities and densities (Carmel and Kadmon, 1999; Cordova, 2007, pp. 63e70). During MIS 4 - 3 there is increasing evidence for a humid and cold environment, which was manifested by an increase in woodland and dense coverage of the forest compared to today (Frumkin et al., 2011, Bar-Yosef, 2000; Cordova, 2007; but cf. Shea, 2008). Recently, evidence from Amud cave (Belmaker and Hovers, 2011) has suggested that the presence of squirrel in the site, absent from the region today, points to increased precipitation and woodland coverage. The distribution of Mediterranean woodland extended east of the rift to the slopes of the Aijloun (Belmaker et al., 2014) in northern Jordan and as far south as the edge of the Ma'an Plateau. There, phytolith (Rosen, 2003) and pollen (Emery-Barbier, 1995) studies of Late Levantine Mousterian deposits indicate moister

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conditions and the presence of nearby Mediterranean woodlands. On the other hand, several areas have been shown to have large grassland swaths. Nonetheless, within the mesic region of the Levant, it is important to stress, that current savanna grassland around the Mediterranean basin is regarded as largely anthropogenic (rather than climatic) in origin having been cleared of trees and shrubs for and through grazing of herd animals (Carmel and Kadmon, 1999; Allen, 2001; Cordova, 2007). 7.2. Locations of Neanderthal base-camps Within the context of the rugged landscape of the rich and diverse Mediterranean woodland environment, Neanderthal placement of base-camps may have offered several adaptive benefits that would have acted to off-set the disadvantages associated with their relatively small SET areas. By situating base-camps in settings of high biodiversity and compact, vertically ordered environmental zonation, Neanderthals would have been able to partially compensate for their high Basal Metabolic Rate (BMR) and locomotor energetic costs through reducing search time and increasing diet breadth. An irregular landscape not only tends to produce greater biodiversity and concomitant increased diet breadth, it also organizes environmental zones into elevational belts that are more tightly packed than those associated with more regular terrain. Moreover, subsistence resources in such vertically zoned environments tend to peak at different times, often staggered over several seasons, thus reducing the need for Neanderthal groups to have repositioned their base-camps as often. 7.3. Coping with compressed SETs In order to compensate for their compressed SETs, some 20% smaller than those of modern human groups, Levantine Neanderthals would have been required to: (a) engage in intensive exploitation of resources (Lieberman and Shea, 1994; Lieberman, 1998), (b) limit their residential group size to some 8e16 individuals (Henry et al., 2004; Vallverdú et al., 2010; Carles Lalueza-Fox et al., 2011; Hayden, 2012) attendant to maintaining relatively low population densities (Burke, 2006; Kuhn and Stiner, 2006; Roebroeks et al., 2011; Bocquet-Appel and Degioanni, 2013) and (c) exercise limited residential mobility through an emphasis on logistical as opposed to more mobile, circular settlement-procurement strategies (Lieberman and Shea, 1994; Henry, 1995, 2003; Shea, 2003). The rugged landscape within the Mediterranean woodland environment would have accommodated each of these requirements more fully than would have the regular, open terrain of the Levantine arid zone. While in agreement with the observation that Neanderthals had smaller SETs than modern humans, other researchers have suggested that Neanderthals must have responded to their more limited access to resources derived principally from hunting, by following a highly mobile settlement-procurement strategy that enabled them to reestablish SETs and gain access to new resources more often (Verpoorte, 2006; Macdonald et al., 2009; Roebroeks et al., 2011; Churchill, 2014). Although Churchill (2014, p. 292) argues for a situational mixture of logistical-radiating and more mobile, circular land-use strategies, he emphasizes the latter based upon the importance of transportation technology, bulk harvesting and storage facilities among logistical foragers in ethnographic contexts and an absence or paucity of these features in the context of Neanderthals. Along these lines, it is noteworthy that the Natufian (with occupations in many of the same sites used by Neanderthals) appears to have followed a logistical-radiating strategy within the Levantine Mediterranean Woodland biome during the terminal Pleistocene without benefit of a transportation

technology, but through the intensive exploitation of storable cereals and nuts. The Natufian, however, was obligated to develop substantial stores of resources in order to support much larger residential groups on the order of three-times greater than those suggested for Neanderthals. Whereas, there is no evidence for inground storage facilities in Levantine MP sites, there are a number of kill sites (Fara II, Quneitra, Umm el Tlel, Hummal) reported that may have been connected to a logistical-radiating strategy. Others point to the paucity of base-camps and simple internal site structure as evidence for the high levels of mobility and ephemeral nature of most Neanderthal encampments. Whereas these site signatures appear to hold for many European sites, they differ markedly from the Levantine record. Neanderthal basecamps in the Levant are not only common, some also reveal living floors with evidence of complex site structure and taskspecific activity areas (Henry et al., 2004; Alperson-Afil and Hovers, 2005; Speth, 2006; Henry, 2010; Hayden, 2012; Speth et al., 2012). This complex site structure implies that Levantine Neanderthals maintained long-term occupations of their base-camps, most likely following a logistical procurement strategy, at least for part of their settlement cycle (Lieberman and Shea, 1994; Shea, 2003; Henry, 2003, 2011; Wallace and Shea, 2006). In a regional review of MP settlement patterns Meignen et al. (2006, p. 161e162) concluded that Late MP groups followed a mixed strategy involving both high and low residential mobility and Hovers (2009) came to a similar inference from her research at the site of Qafzeh. In many ways this resembles a central-based wandering pattern (Kelly, 1995, p. 116e117) in which groups would have maintained longterm repetitive (seasonal?) occupations in base-camps and other more ephemeral encampments that were often associated with task-specific activities as evidenced at Kebara, Umm el Tlel, Qafzeh and Tor Faraj/Tor Sabiha. If Levantine Neanderthals did not employ higher mobility levels to compensate for their smaller SETs, how did they cope? In part the rich, diverse environment of the Mediterranean woodland biome would have allowed for a relatively balanced meat/plant diet, but more importantly Neanderthal residential groups appear to have been relatively small, consisting of some 8e16 members (Henry et al., 2004; Vallverdú et al., 2010; Lalueza-Fox et al., 2011; Hayden, 2012), about half the size of those reported ethnographically for warm-climate, non-fishing foragers (Kelly, 1995; Binford, 2001; Marlowe, 2005, but cf. Shea, 2003, p. 349). Of special interest here is the uncanny correspondence between these estimates, derived from alternative and independent calculations of genetic and archaeological field evidence, and those arrived at on the basis of the relative caloric needs of Neanderthals and modern humans in which Neanderthal population densities are thought to have been about half that of modern humans (Bocquet-Appel and Degioanni, 2013, p. 219). The irony here is that both high-mobility and lowmobility/small group size models may well be accurate in that given the great temporal duration, geographic distribution and environmental variability of the Neanderthal settlement range it would be surprising if they had not followed a variety of land-use strategies (Hardy, 2010; Fiorenza et al., 2015). In general, the Neanderthal dispersal from high latitude, cold-temperate settings to lower latitude, warm-temperate environments may have been accompanied by a shift to an intensive, broader, more meat/plant balanced diet supported by a logistically based settlementprocurement strategy and more permanent base-camps. 7.4. Landscape and Neanderthal morphology The landscape contexts of Levantine Neanderthal base-camps would also have better accommodated hunting practices tied to thrusting or short-cast spears and a strategy of close-up, ambush

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hunting best undertaken in rugged, wooded landscapes rather than in open, level terrain. Such hunting techniques would have been consistent with Neanderthal limb structure that is thought to have conferred power for spear thrusting (Steudel-Numbers and Tilkens, 2004; Churchill and Rhodes, 2006), walking over running efficiency (Hora and Sladek, 2014) and surges of strength and speed for short distance pursuit of prey (Finlayson, 2005; Stewart, 2005); activities that would have been more common in closed than open settings. From another perspective, Neanderthal leg morphology, while being relatively disadvantageous in regular landscapes, may actually have conferred locomotive advantages in more rugged landscapes (Higgins and Ruff, 2011). Additionally, the short, compact stature of Neanderthals, traditionally linked to adaptation to cold climatic conditions, also may have been beneficial to maneuvering within and exploiting more closed environments such as found in the Mediterranean woodlands (Bro-Jǿrgensen, 2008, p. 732). Moreover, analyses of spears and lithic armatures, Levallois points, suggest that Neanderthal hunting weapons consisted mainly of robust spears that would have been used for thrusting or casting over relatively short distances, again consistent with notions of intercept over pursuit hunting practices (Shea, 1998; Schmitt et al., 2003). 7.5. Land-use strategies, diet and division of labor The configuration of Neanderthal SETs may also have influenced the ways in which they organized their subsistence activities along the lines of age and gender. In contrast to following a traditionally recognized foraging pattern involving a division of labor in which males hunted and females/children focused on harvesting plant and other less mobile resources, Kuhn and Stiner (2006) have argued that Neanderthals followed a different, mixed-group strategy. This proposal, based mainly on the notion that Neanderthals relied heavily on the hunting of large prey species with little emphasis on the collection of plant resources, is best supported by higher latitude, European dietary evidence from the analyses of lithic artifacts, faunal assemblages, and isotopic signatures. However, there is a growing body of evidence that plant foods formed a substantial portion of Neanderthal diets (Hovers, 2009; Hayden, 2012; Fiorenza et al., 2015). From Europe, examination of dental calculus has identified microfossil evidence of plant foods such as fruits, grass seeds, and underground storage organs (Hardy et al., 2012; Henry et al., 2011, 2014; Fiorenza et al., 2015), analysis of fecal biomarkers indicate significant plant intake (Sistiaga et al., 2014), phytolith studies suggest possible grass seed consumption (Cabanes et al., 2010) and based upon unique bone artifact forms, Sandgathe and Hayden (2003) have proposed that Neanderthals may have been using the calorie-rich inner bark of some trees for food. Within the warm-temperate Levant, the recovery of macrobotanic remains from Kebara (Lev et al., 2005) and phytoliths from Amud and Tor Faraj (Albert et al., 2000; Madella et al., 2002; Rosen, 2003) denotes an extensive, habitual exploitation of a wide range of plants (such as grass seeds, acorns, fruits and legumes) as foods by Neanderthals across a wide geographic range stretching from coastal and inland northern Israel to southern Jordan. Whereas the subsistence evidence alone is not especially supportive of the notion of mixed-group foraging, the compact SETs of Levantine Neanderthals in conjunction with their logistical procurement and ambush hunting strategies may well have favored cooperative, full-group efforts in some of their subsistence activities. The rich environments of the compressed SETs would have likely displayed relatively closely spaced resources, which in turn, would have reduced the effectiveness of a division in labor and its attendant focus on different widely spaced resources. While division of labor characteristically results in an overall, group reduction in search time coupled with enhanced diet breadth, these

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relationships may not have been as significant in the context of the compressed Levantine Neanderthal SETs as they would in larger exploitation territories. Ambush or intercept hunting may also have benefitted from cooperative, mixed-group efforts in which females and young principally participated as drivers of large prey species into predetermined traps where they were killed at close quarters by males (Kuhn and Stiner, 2006). This practice may also have increased efficiency by having a larger work force to reduce handling costs and increase returns related to butchering, preliminary processing and portaging of meat back to a base-camp. In other contexts, however, gender/age compositions of special task groups were likely to have followed the more typical pattern of division of labor. The settlement-procurement systems discussed for Levantine Neanderthals by several researchers (Lieberman and Shea, 1994; Bar-Yosef, 2000; Henry et al., 2004) are thought to have involved an habitual, intensive exploitation of a SET and an occasional, dedicated exploitation of a distant resource falling beyond the limits of the SET, but within a site catchment. At Tor Faraj, for example, chert was principally imported from a distance of 22 km downslope from the edge of the Ma'an Plateau and more rarely 16e18 km upslope from the Rift Valley along with date fruit (Henry et al., 2004). These more distant task-specific, subsistence efforts were also likely to have been partial, rather than full-group efforts and as such gender/age specific as well. 8. Conclusions and broader implications In our study, we assessed the influence of differences in terrain and accompanying environments on the adaptive strategies and settlement patterns of Levantine Neanderthals and AMH groups through site catchment analysis. Our findings indicate that Neanderthals habitually exploited smaller areas around their sites, principally situated in the rugged Mediterranean Woodlands of the Levant, whereas early Upper Paleolithic groups generally enjoyed larger SETs and displayed a wider settlement range encompassing both rugged woodland and more regular, level steppe landscapes. This is consistent with the observations of others (Finlayson, 2004, pp. 206e8; Stewart, 2005, p. 38; Dusseldorp, 2009, p. 30) that Neanderthals preferred intermediate, elevationally diverse habitats across Eurasia in contrast to the exploitation of the more regular landscape of the open steppe by modern human groups. The preference for the occupation of a rugged terrain as a facet of Neanderthal ecology may also explain why Neanderthals in their dispersal during MIS 4 e early MIS 3, were confined to the irregular landscape of the Mediterranean woodlands of the southern Levant and deterred from expanding further east and south into the Arabian Peninsula or west across the Negev and Sinai into North Africa. East of the Levant, the central Arabian Peninsula is dominated by the Nedj, the plateau of central Arabia, which is composed of vast sand seas, small isolated mountains and plains within the virtually continuous desert stretching from the Nafud in the north to the Rub al-Khali in the south. The most rugged terrain in Arabia follows the mountain ranges formed along the eastern edge of the Syrian-East African Rift and across the southern edge of the peninsula. Although archaeological research in Arabia has been concentrated in the southern and eastern parts of the peninsula, recent research (Groucutt and Petraglia, 2012; Scerri et al., 2014) in the northern Nafud, the closest area to the Levant, has complemented pioneering efforts (Garrard et al., 1981) to show a robust Middle Paleolithic presence associated with the moister conditions of MIS 4-3. The lithic assemblages suggest considerable autochthonous development, with none exhibiting the specific technological characteristics associated with the Late Levantine Mousterian nor, surprisingly, the Nubian Complex signatures reported from southern Arabia (Rose et al., 2011).

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To the west across the Negev and Sinai, the rugged landscape of the high central Negev is framed by more regular terrain northward toward the coast and westward into northern and central Sinai. The narrow strip of uplands along the western margin of the Gulf of Aqaba forms a discontinuous broken landscape before joining the mountains and rugged topography of southern Sinai. Our understanding of the paleoenvironments of the Negev and Sinai during MIS 4- early 3 largely comes from speleothem, geomorphic and palynological studies. Speleothem data from the northern Negev point to a southern extension of the Mediterranean zone, dominated by steppic vegetation (Vaks et al., 2006, p. 394e395), but further south in the central and southern Negev Desert dry conditions prevailed as evidenced by the last speleothem deposition during MIS 6a-5e (Frumkin et al., 2011) and none occurring after 86 ka (Vaks et al., 2010). Beginning with MIS 5, the Negev appears to have been dominated by an eolian environment with the deposition of sand and silt deposits inter-calculated with paleosols (Crouvi et al., 2009; Roskin et al., 2013, p. 149) and given parallel deposits in northern (Goldberg, 1977) and northeastern Sinai (Gladfelter, 2000) a similar environment is suspected. Within MIS4, during Late MP times, Goldberg (1986, p. 239) observed an erosional episode thought to reflect arid conditions similar to today. Palynological studies of archaeological deposits in the central Negev Highlands (Horowitz, 1979, pp. 245e247) and of travertines in the Wadi Arava (Weinstein, 1987) generally paralleled the speleotherm records. The Negev Highlands revealed a surprisingly high proportion of arboreal pollen (25%) for the 80 ka, Early MP (Tabun type-D) deposit of Nahal Aqev and declining AP percentages (16e17%) for subsequent Early UP deposits (Horowitz, 1979, p. 245e247), but this is consistent with the proposed southern extension of the Mediterranean zone based on speleothem data, especially given the higher elevations. The Wadi Arava study also shows a higher proportion of AP than that of the area at present (9%) for MIS 5b (38%) and late MIS 4 (20%) samples, but Weinstein (1987, p. 87) cautions that the travertines selectively represent episodes of high humidity and may not be a signature of the entire interval. What is so striking relative to site distributions is the absence of sites with Tabun B-Type signatures for this vast region of the Negev and Sinai despite extensive research efforts. In contrast, virtually all of these major surveys in the area (Bar-Yosef and Phillips, 1977; Marks, 1983; Phillips and Gladfelter, 1989) have yielded evidence of an early UP presence dating to early MIS 3 (Belfer-Cohen and Goring-Morris, 2003) in addition to an AMH cranium from a burial at Taramsa Hill on the west bank of the Nile dated to ~ 55 ka (Van Peer et al., 2010). From these data we argue that during MIS 4 and early MIS 3 Neanderthals were restricted to the rugged Levant because the richness, diversity and vertical arrangement of environments of this setting enabled Neanderthal groups to compensate for their compressed SETs induced by their inherent locomotor deficiencies and high energetic costs. Moreover, occupation of the Levantine Mediterranean woodlands allowed Neanderthal groups to follow similar adaptive strategies and land-use patterns that had served them in the similar biome of southern Europe and extending around the Mediterranean basin. Our proposal also draws support from the observation that the regions surrounding the Levantine Mediterranean woodland zone lack any indication of a Neanderthal presence, despite abundant evidence of hominin occupation by contemporary and/or immediately successive groups. The selective adaptation of Neanderthals to rugged, wooded landscapes beyond the region may go a long way in understanding their dispersal in Eurasia and their terminal presence restricted largely to mountainous refugia. Beyond these biogeographic constraints, the definition of significantly smaller SETs than those enjoyed by early

modern human groups furnishes additional support for the notion of a Broserupian demographic trap in which Neanderthals maintained marginal group sizes and regional population densities, suffered bottlenecks and hovered dangerously close to the threshold of viability (Bocquet-Appel and Degioanni, 2013, p. 219). In contrast to the constraints that a rugged Mediterranean woodland setting placed on Neanderthal settlement-procurement strategies, AMH groups appear to have enjoyed far fewer restrictions relative to landscape and environment as they occupied both open, level steppic settings and rugged, Mediterranean woodlands of the southern Levant. Moreover, this apparent adaptive flexibility may have been one facet of the behavior ecology of AMH populations that enabled them to expand so rapidly across the varied landscapes of Eurasia, at the expense of Neanderthals (Beyin, 2011; Boivin et al., 2013).

Acknowledgements
n, Knut We would like to thank the guest editors Gema Chaco Bretzke, Florent Rivals and Nicholas Conard for their assistance in addition to the organizers of the XVII USIPP Congress. Moreover, we are grateful to the Center for Global Education, the Henry Kendall College of Arts and Sciences and the Office of Research of the University of Tulsa for their support.

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