Intensification and sedentism in the terminal Pleistocene Natufian

Transcription

Journal of Human Evolution 70 (2014) 16e35
Contents lists available at ScienceDirect
Journal of Human Evolution
journal homepage: www.elsevier.com/locate/jhevol
Intensification and sedentism in the terminal Pleistocene Natufian
sequence of el-Wad Terrace (Israel)
Reuven Yeshurun a, b, *, Guy Bar-Oz a, Mina Weinstein-Evron a
a
Zinman Institute of Archaeology, University of Haifa, Mount Carmel, Haifa 3498838, Israel
Program in Human Ecology and Archaeobiology, National Museum of Natural History, Smithsonian Institution, PO Box 37012, MRC 112, Washington,
DC 20013-7012, United States
b
a r t i c l e i n f o
a b s t r a c t
Article history:
Received 10 August 2013
Accepted 21 February 2014
Available online 21 March 2014
Measuring subsistence intensification in the archaeofaunal record has provided strong evidence for
socioeconomic shifts related to sedentarization in the terminal Pleistocene Mediterranean Basin, but the
precise timing and scale of the intensification trend and its place in the evolution of settled societies
remain contentious. New archaeofaunal data from the key Natufian sequence of el-Wad Terrace (Mount
Carmel, Israel, ca. 15.0e11.7 ka [thousands of years ago]) is used here to clarify and contextualize paleoeconomy and mobility trends in the latest Pleistocene Levant, representing the culmination of Epipaleolithic subsistence strategies. Taphonomic variables serve as supplementary indicators of habitation
function and occupation intensity along the sequence. At el-Wad, a very broad range of animals, mostly
small to medium in size, were captured and consumed. Consumption leftovers were discarded in
intensively occupied domestic spaces and suffered moderate attrition. The Early (ca. 15.0e13.7/13.0 ka)
and Late (ca. 13.7/13.0e11.7 ka) Natufian phases display some differences in prey exploitation and
taphonomic markers of occupation intensity, corresponding with other archaeological signals. We
further set the intra-Natufian taxonomic and demographic trends in perspective by considering the
earlier Epipaleolithic sequence of the same region, the Israeli coastal plain. Consequently, we show that
the Early Natufian record constituted an important dietary shift related to greater occupation intensity
and sedentarization, rather than a gradual development, and that the Late Natufian record appears to be
maintaining, if not amplifying, many of these novel signals. These conclusions are important for understanding the mode and tempo of the transition to settled life in human evolution.
Ó 2014 Elsevier Ltd. All rights reserved.
Keywords:
Epipaleolithic
Levant
Zooarchaeology
Broad-spectrum revolution
Mobility
Contextual taphonomy
Introduction
The process of settling down by hunter-gatherer groups in the
terminal Pleistocene (ca. 20,000e11,700 years cal. BP [calibrated
before present]) was an important milestone in human evolution,
entailing a series of changes in mobility, economy and society.
Sedentary or semi-sedentary groups were emerging in the Mediterranean Basin in the millennia following the Last Glacial
Maximum (LGM), displaying novel adaptations and the roots of
social complexity. Specifically in the Levant region, the cultural
period bridging the LGM and the end of the Pleistocene, the Epipaleolithic, has been a major subject of investigation concerning
the formation of complex foraging societies, which eventually laid
* Corresponding author.
E-mail addresses: [email protected] (R. Yeshurun), guybar@research.
haifa.ac.il (G. Bar-Oz), [email protected] (M. Weinstein-Evron).
http://dx.doi.org/10.1016/j.jhevol.2014.02.011
0047-2484/Ó 2014 Elsevier Ltd. All rights reserved.
the foundations for the subsequent Neolithic Period (e.g., Kaufman,
1992; Bar-Yosef and Meadow, 1995; Henry, 1995; Stiner and Kuhn,
2006; Watkins, 2010; Belfer-Cohen and Goring-Morris, 2011;
Maher et al., 2012a). Some of the earliest and most conspicuous
manifestations of the pre-agricultural shift to sedentary living
appear ca. 15,000e11,700 years cal. BP, in the Late Epipaleolithic
Natufian culture of the Levant. These include stone structures and
terraces, large cemeteries, diverse groundstone assemblages and
hewn bedrock features, plus numerous personal adornments and
art items, as well as remains of commensal animals. These traits are
much better represented, quantitatively and qualitatively, relative
to the earlier Epipaleolithic record, indicating increased permanence of occupation and probably increasingly complex societies
and greater human impact on their surroundings (e.g., Garrod,
1957; Henry, 1991; Tchernov, 1993a,b; Valla, 1995; Bar-Yosef,
1998; Belfer-Cohen and Bar-Yosef, 2000; Munro, 2004, 2009;
Byrd, 2005; Goring-Morris and Belfer-Cohen, 2008).
R. Yeshurun et al. / Journal of Human Evolution 70 (2014) 16e35
Changes in subsistence go hand in hand with sedentarization in
the Mediterranean Basin. While most pre-LGM groups habitually
hunted ungulates and paid less attention to small game species,
thereby continuing earlier Paleolithic traditions of big-game
hunting as the main source of animal food, it is generally
accepted that Epipaleolithic, and particularly Natufian groups,
intensified their resource base, habitually exploiting juvenile, small
or non-terrestrial animals as important dietary components (Davis,
1991, 2005; Tchernov, 1993a,b,c; Stiner et al., 1999, 2000; Bar-El and
Tchernov, 2000; Stiner, 2001; Hockett and Haws, 2002; Bar-Oz,
2004; Munro, 2004; Atici, 2009; Munro and Atici, 2009; Stutz
et al., 2009; Bar-Yosef Mayer and Zohar, 2010; Stiner and Munro,
2011; Starkovich, 2012; Zeder, 2012). The addition of a suite of
animal taxa, many of which are small-bodied or non-terrestrial, to
the regular human diet in the millennia just preceding the onset of
food production is commonly referred to as the Broad Spectrum
Revolution (BSR) (Flannery, 1969).
Subsistence intensification and the BSR are important in the
wider context of human evolution because of their bearing on siteoccupation intensity in the Epipaleolithic. A strong link exists between increasing sedentism and intensifying subsistence. It is often
conceptualized using a behavioral ecology approach, where game
taxa may be ranked according to their dietary gains versus search
and handling costs. Foragers are likely to habitually procure lowerranked game only when higher-ranked resources become less
available (Winterhalder and Smith, 2000; Stiner and Munro, 2002;
Munro, 2004, 2009). Economic intensification means that lowerranked prey is regularly included in the diet, presumably because
encounter rates with higher-rank prey decrease or because demand
for prey increases. During the Epipaleolithic, the ability to reside in
a site for longer periods would have meant relying on such intensification, i.e., extracting more nutrients from the environment
(Munro, 2009), all the more so if the number of people inhabiting a
settlement was higher than before, or if they maintained smaller
territories (Rosenberg, 1998). In the Levantine context, low-ranked
prey include small and fast-escaping mammals and birds, which
provide small quantities of edible material for a high capture cost,
as well as juvenile ungulates, which provide less meat and fat than
adults (Stiner, 2001; Munro, 2004). Hence, economic intensification
can be related to increases in human population in a given territory
(Stiner et al., 1999, 2000), but the precise timing and scale of the
post-LGM intensification process and its place in the evolution of
sedentary societies remain contentious (see below).
This study investigates sedentism and economic intensification
in the key Natufian base-camp of el-Wad Terrace (Mount Carmel,
Israel). This is the classic Natufian sequence where the long-held
view of this culture, as a complex and sedentary society at the
threshold of farming, was first conceived (Garrod, 1932, 1957). We
aim to clarify, refine and contextualize trends in Natufian economy
and sedentism by employing detailed zooarchaeological data from
our new excavations at the site. Taxonomic abundances, gazelle
culling patterns and taphonomic indicators are used to evaluate
habitation type, the magnitude of intensification and site occupation intensity. Our results are set in context by comparing the
Natufian animal economy with the earlier, well-studied Epipaleolithic sequence of the same region (Fig. 1A). Ultimately, we
aim to shed light on Epipaleolithic socioeconomic developments by
pinpointing precisely when and how sedentism-related intensification occurred within the long process of settling down in the
terminal Pleistocene of Southwest Asia.
Intensification and sedentarization processes in the Epipaleolithic
The Natufian record of the late Epipaleolithic Levant figures
prominently in all discussions of pre-agricultural intensification
17
and sedentarization. Scholarly opinions differ regarding the Natufian phenomenon. Since the early days of research, the large Early
Natufian (EN) hamlets displaying architecture, cemeteries, art and
abundant groundstone items caused the Natufian to be viewed as a
major break from preceding Paleolithic cultures (Garrod, 1957;
Valla, 1995; Bar-Yosef, 1998). Based on archaeological criteria such
as changes in architecture, mortuary practices and art, an important intra-Natufian difference was noted by some, in that the EN
phase (ca. 15.0e13.7/13.0 ka [thousands of years ago]) was
considered the classic sedentary phase and the Late Natufian (LN)
phase (ca. 13.7/13.0e11.7 ka) was interpreted as having a retreat to
greater mobility (Garrod, 1957; Belfer-Cohen and Bar-Yosef, 2000;
Bar-Yosef and Belfer-Cohen, 2002). Growing archaeological evidence in the last three decades has placed the Natufian culture in
context, demonstrating its Epipaleolithic roots (Kaufman, 1989,
1992; Maher et al., 2012a,b). Recent data from Israel and Jordan
indicate prolonged site habitation and the presence of defined huts
that were rich in symbolic meaning, echoing the hallmark Natufian
features, as early as 23e20 ka, and therefore the EN was viewed as
gradually evolving from the preceding Epipaleolithic cultures
(Nadel et al., 2004; Maher et al., 2012b).
Interpretations of Natufian zooarchaeological data are crucial
for understanding the nature of Epipaleolithic sedentarization.
However, the timing, mode and tempo of the terminal Pleistocene
shift to intensified economy and BSR are often contested or loosely
defined. Regarding the well-studied Levantine record, several
scholars maintained that the Natufian is exceptional relative to the
preceding cultures in the high proportion of small mammals and
sometimes birds and fish in conjunction with diminishing proportions of medium and large ungulates (e.g., Davis et al., 1988;
Davis, 1991; Pichon, 1991; Bar-Oz, 2004). In contrast, it has been
suggested that diversification of Levantine animal economies
occurred millennia before the terminal Pleistocene (Edwards,
1989), although this analysis was countered on the basis of inappropriate statistical methods, poor sampling quality and lack of
taphonomic consideration (Neeley and Clark, 1993; Bar-Oz, 2004).
Several fine-grained archaeofaunal analyses pertaining to Epipaleolithic intensification have recently been published. Combined
with multiple lines of archaeological evidence for increased sedentism, the Natufian subsistence trends were interpreted as evidence for intensification and diversification of animal exploitation
due to greater permanence of site occupation (Bar-Oz, 2004;
Munro, 2004, 2009; Davis, 2005; Stutz et al., 2009). The Natufians not only exploited small game in unprecedented proportions,
they began in particular to exploit less cost-effective but resilient
animals such as lagomorphs and birds, hinting at novel capture
techniques, elevated pursuit costs and rising occupation intensity
(Stiner et al., 1999, 2000; Stiner and Munro, 2002; Munro, 2004,
2009). The exploitation of the main ungulate prey, the mountain
gazelle (Gazella gazella), was geared towards culling of lower-yield
younger individuals, which was taken as another sign of foraging
intensification (Munro, 2004, 2009; see also; Davis, 1983, 2005;
Bar-Oz, 2004). In some of these studies, it was possible to investigate intra-Natufian trends, which showed that more fast small
game was exploited in the EN phase, while in the LN phase slow
small game was more dominant (Munro, 2004, 2009; Stutz et al.,
2009). Zooarchaeological evidence suggesting Natufian economic
intensification was recently viewed as the product of gradual
trends throughout the Epipaleolithic, culminating in the Natufian
(Munro, 2009; Stutz et al., 2009). Proponents of the ‘gradual
transition’ downplayed the BSR hallmark of the Natufian, claiming
similarity to earlier Epipaleolithic subsistence (Maher et al., 2012a),
or altogether ignoring economic factors when comparing preNatufian and Natufian adaptations (Richter et al., 2011; Maher
et al., 2012b).
18
Figure 1. Provenance of the faunal samples: (A) Location map showing el-Wad and other sites mentioned in the text; (B) Plan of el-Wad in the late Early Natufian (LEN) phase, the
main ‘architectural’ phase of the site (after Weinstein-Evron et al., 2013); (C) plan of the NE Terrace excavation e the Late Natufian layer; (D) plan of the NE Terrace excavation e late
Early Natufian layer. Note the ‘domestic’ (Structure II) and ‘non-domestic’ (Locus 25) areas.
The key sequence of el-Wad Terrace (EWT) is ideally positioned
in time and space to shed light on the mode and tempo of terminal
Pleistocene socioeconomic shifts pertaining to economic and sedentism. These new data, put in regional context, allow us to
contribute to two of the pressing issues in the study of Levantine
sedentarization:
1. Did Early Natufian subsistence and mobility patterns constitute
a significant break from preceding Epipaleolithic cultures?
2. Was the Late Natufian fundamentally different from the Early
Natufian in mobility and subsistence?
Framework of analysis
Here we provide a detailed taphonomic and zooarchaeological
analysis of EWT, identifying the economically important animals
and their deposition and preservation patterns. We then put our
results in wider diachronic perspective by considering the pre-
Natufian sites. In such large and complex sites it is quite possible
that intra-site depositional mechanisms, type of habitation and
topographic factors blur the sought paleoeconomic signals through
time (Munro and Grosman, 2010; Yeshurun et al., 2013a, submitted
for publication). Thus, we narrow the geographic, ecological and
‘functional’ (e.g., site type) scope by focusing exclusively on a series
of Epipaleolithic ‘base-camps’ on the Israeli coastal plain (Fig. 1A).
This sequence contains the well-published archaeofaunas of Nahal
Hadera V (NHV, Early Epipaleolithic, Kebaran Culture), Hefzibah 7e
18 (HEF) and Neve David (NVD, Middle Epipaleolithic, Geometric
Kebaran Culture) and the new results from EWT, presented below
(Late Epipaleolithic, Early and Late Natufian Culture). The coastal
plain series forms a natural geographic cluster during the Epipaleolithic (Bar-Oz et al., 1999; Bar-Oz and Dayan, 2002, 2003; BarOz, 2004; Bar-Oz et al., 2004).
Stiner and colleagues (Stiner et al., 1999, 2000; Munro, 2004)
argued that intensification needs to be measured by categorizing
animals according to their locomotion and resilience, rather than
traditional taxonomic groups. Therefore, the first avenue of investigation examines trends in the taxonomic abundance (based on
Number of Identified Specimens, NISP) of small versus big game,
larger (medium-size) ungulates versus small ungulates, and fast
small game versus slow small game, following Stutz et al. (2009:
their Table 4). Stutz et al. (2009) predicted, based on the behavioral
ecology approach explained above, that economic intensification
would be reflected by: (1) elevated hunting of small game to
augment dwindling ungulates; (2) decrease in larger ungulates
(primarily fallow deer and aurochs) relative to the more resilient
small ungulates (gazelle); and (3) acquisition of fast-escape small
animals (e.g., hare and partridge) rather than easily caught but less
resilient slowly escaping small animals (e.g., tortoise). These trends
may reflect human predation pressure at different scales. As ungulates have larger home ranges than most small game species,
they provide a measure of hunting pressure on the regional scale,
while small game response to predation may be more local-scale,
around the sites themselves (Stiner and Munro, 2002; Munro,
2009, 2012).
The second line of inquiry investigates culling patterns (age and
sex profiles) of mountain gazelle, the most abundant ungulate prey
in the Epipaleolithic of the Mediterranean southern Levant. It has
been suggested, based on the same set of considerations as above,
that intensifying foragers inflicting predation pressure on gazelle
herds will progressively broaden their diet by hunting more lowreturn individuals, such as juveniles and eventually fawns, which
provide less meat and fat than adults (Munro, 2004, 2009).
The third line of inquiry, which constrains our paleoeconomic
inferences, employs taphonomic measures of site occupation intensity to evaluate the habitation type of the different phases in
EWT (Yeshurun et al., submitted for publication). These measures
include volumetric bone densities and frequencies of post-discard
damage such as weathering, animal gnawing, trampling and nonnutritional burning and fragmentation. Volumetric densities serve
as rough correlates for the intensity of site use, assuming that, all
else being equal, higher quantities of refuse will be discarded when
more people inhabit the site for a given time. The magnitude of
post-discard destruction processes among the EN and LN samples
will serve as an estimate of occupation intensity; a greater number
of repeated occupations or more prolonged occupations would
have a different taphonomic effect on the faunal remains than short
and sporadic occupations. The latter would theoretically display
less non-nutritional burning (Stiner and Munro, 2011), reduced
rates of trampling and subsequent dry breakage (Haynes, 1983),
and more weathering and animal damage as a result of decreased
buildup of cultural refuse, leaving the bones exposed for a longer
duration (Behrensmeyer, 1978; Kent, 1993; Magdwick and Mulville,
19
2012). These multiple measures, though rough and indirect proxies,
are tied to the type of habitation, which we strive to keep as constant as possible when comparing paleoeconomic indices through
time.
The site and its setting
El-Wad Cave is part of the UNESCO World Heritage Site complex
of Nahal Me’arot/Wadi el-Mughara that also includes the caves of
Tabun, Jamal and Skhul (Garrod and Bate, 1937). The site, a large
cave with an adjacent terrace containing a long and rich Early, Late
and Final Natufian sequence, is situated on the western face of
Mount Carmel, Israel, where the mountain cliff meets the open
expanses of the Mediterranean coastal plain, 45 m above modern
sea level, within the Mediterranean climatic zone of the Levant
(Fig. 1A). The site was first investigated by Lambert in 1928
(Weinstein-Evron, 2009), but became well-known as a result of
Garrod’s 1929e1933 excavation campaign (Garrod and Bate, 1937).
Garrod’s finds from el-Wad were the foundation of her subsequent
definition and interpretation of the Natufian culture as a transitional phase between foraging and fully agricultural life-styles
(Garrod, 1932, 1957). The terrace was later revisited (Valla et al.,
1986), as was the cave (Weinstein-Evron, 1998). The renewed and
on-going excavation was initiated in 1994 and has been focused on
the north-eastern part of the terrace (Fig. 1B). An area of ca. 70 m2
was exposed and the attained thickness of Natufian sediments
ranges between ca. 0.5 and 1.5 m. Structures, burials and a high
density of finds, specifically chipped lithic and groundstone tools,
bone tools, bone and shell ornaments, ochre and a rich faunal
assemblage were retrieved during the renewed excavation
(Weinstein-Evron et al., 2007, 2012, 2013; Yeshurun et al., 2013b).
A composite stratigraphy of the site, based on a compilation of
data from all excavations (Weinstein-Evron, 2009; WeinsteinEvron et al., 2013), suggests an ephemeral occupation at the base
of the Early Natufian (designated Early Early Natufian or EEN),
followed by a prolific burial phase comprising almost 100 individuals (Middle Early Natufian or MEN) and culminating with the
Late Early Natufian (LEN), the ‘classic’ Early Natufian layer of the
site, with its varied architectural features (Figs. 1D and 2). This
phase appears as a massive, >0.5 m thick accumulation of repeated
occupations. Overlying this architectural phase are thick Early
Natufian living levels with a few stone features but normally lacking structures. The Natufian sequence ends with a thinner Late
Natufian layer devoid of architecture, but displaying several concentrations of graves (Weinstein-Evron et al., 2007; Bachrach et al.,
2013). The Early Natufian occupations (Garrod’s layer B2, our Unit
2) are much thicker and richer than the Late and Final Natufian
layers throughout the site (Garrod’s layer B1; here, the upper part of
Unit 2 and base of Unit 1).
The renewed excavation has exposed an architectural complex
in the LEN phase, composed of a 9-m long curvilinear wall (Wall I)
encompassing a sequence of at least nine architectural sub-phases,
each defined by a thin stony floor. In the area enclosed by Wall I,
several partially preserved stone structures and stone-rich ‘living
floors’ have been excavated, of which the best preserved is Structure II (Fig. 1D). Several discrete types of Early Natufian architectural contexts were defined: inside Structure II, including stony
floors and between-floor fills; outside Structure II, in levels corresponding to some or all of the dwelling floors; the area of Locus 67,
a massive stone pile lying northwest of the Structure II area; and
Locus 25, an amalgamation of stones outside the living compound
demarcated by Wall I (Fig. 1D). A contextual taphonomic analysis
indicated meaningful differences in the tempo and mode of deposition of the faunal remains among these contexts, ranging from
recurrent deposition of consumption refuse in domestic space in
20
Figure 2. (A) Section through the NE terrace excavation, corresponding to the NeO line in Fig. 1D. Note the stone architecture (Walls I and II) in the lowest attained levels. The inset
depicts the proportions of Helwan-retouched lunates, abruptly-retouched lunates and microburins in each phase (after Weinstein-Evron et al., 2012); (B and C) examples of the EN
architecture in the renewed excavation.
the first three contexts to an occasional toss area in the last one
(Yeshurun et al., submitted for publication). Importantly, the study
concluded that the Wall I living area at the heart of the excavation
contains discarded fauna in primary deposition in the context of
well-preserved architecture, and that post-discard damage was
inflicted on the bones as a result of repeated human occupation and
a fast rate of accumulation in the same spot. This depositional
circumstance caused indirect burning, trampling and nonnutritional fragmentation, in tandem with low weathering and
animal gnawing. In contrast, the volumetric density of faunal remains, as well as burning, trampling and fragmentation, is markedly reduced outside of the Wall I compound (Locus 25) and
weathering and gnawing appear in elevated proportions, all indicating a non-domestic locale (Yeshurun et al., 2013b, submitted for
publication). The distribution of groundstone items, bone tools and
micromammal remains generally show similar patterns of intrasite use, namely the discard of used, broken pestles and bone
points and consumed rodents in the Wall I complex rather than
outside it (Yeshurun, 2011; Rosenberg et al., 2012; Weissbrod et al.,
2012).
Thirteen radiocarbon measurements on charcoals and ungulate
bones yielded a calibrated age range of 14,660e14,030 years BP
(1s) for the architecture-bearing phases and a range of ca.
15,000e13,000 years BP (1s) for the entire Unit 2 accumulation
(Eckmeier et al., 2012; Weinstein-Evron et al., 2012). The dating of
the Late Natufian contexts is underway. The density of finds in the
Wall I compound is extremely high, but human remains are
virtually absent. The stone structures, numerous living floors,
density and diversity of finds and the absence of burials indicate
that this part of the site was used primarily for habitation and daily
activities in the later parts of the Early Natufian. The character of
the site in LN times is less clear; our ensuing comparison to both
kinds of EN depositions (dwelling contexts versus Locus 25) will
help in tracing the nature of the LN settlement at this part of the site
and illuminate changes in site function and intensity of occupation.
Materials and methods
Sampling
The faunal assemblage retrieved by the renewed excavation
is vast, and therefore had to be sampled with special attention
to the chronological and contextual proveniences. Diachronic
archaeofaunal variability was investigated by comparing two subassemblages: an EN sample from the lowest attained phases (the
Wall I compound; 9141 identified specimens) against an LN sample
from the same spatial location (Unit 1e2 and Phase W-0 in Unit 2,
squares OeP/6e7; see Fig. 1C; 2420 identified specimens). In the EN
lithic sample, >70% of all lunates are Helwan-retouched, microburins are very rare, and all radiocarbon dates are older than
13,300 years cal BP, in line with other EN occurrences (WeinsteinEvron et al., 2012). In the LN sample, >64% of the lunates are
abruptly retouched, and microburins are present in ratios of 1:2 to
1:4 compared with the lunates. In these squares, located in the
middle of the excavation, mid-way between the cliff wall and the
fall of the talus (Figs. 1 and 2), the deposits attain maximum
thickness and are minimally disturbed. Importantly, in these
squares no burials were cut into the layer, making in situ preservation more likely than in the east area of the excavation, where
eleven LN graves were dug, creating a significant potential for
mixture of sediments. Still, the EN sample is much larger than the
LN sample by virtue of the much larger magnitude of the former
layer. The material in the phases vertically sandwiched between
our two samples was left for future investigations, in order to
ensure that the EN versus LN samples were as clean and distinct as
possible. The good architectural preservation of the lower layers
allows the comparison of the LN material with two kinds of EN
deposits: inside Structure II, interpreted as a domestic area with
consumption refuse in primary deposition, and Locus 25, interpreted as an occasional toss-zone outside the living compound
(Fig. 1D; Yeshurun et al., submitted for publication). The geogenic
processes contributing to the formation of the strata appear to be
similar in both the EN and LN samples in this work. Both are hardly
differentiated sedimentologically (Weinstein-Evron et al., 2007),
suggesting a similar rate of sedimentation in the central part of the
terrace excavation (i.e., the Structure II area in the EN sample, and
the studied LN sample). Care was taken not to sample the LN
cemetery or other contexts conspicuously different in their
’behavioral’ mode of deposition compared with the EN sample in
this study.
Faunal analysis procedures
All faunal remains in this study were either plotted and packed
in the field or retrieved by wet sieving employing 5 mm sieves. The
5 mm sieves were used to separate the zooarchaeological assemblage described here from most of the micromammal remains,
which were captured by the underlying 1 mm sieve and are undergoing detailed study (Weissbrod et al., 2005, 2012, 2013).
Extremely few recognizable remains of animals larger than
micromammals passed through the 5 mm sieve, in line with
empirical studies comparing sieving methods (Lyman, 2008). The
analysis included reptiles, birds larger than Passeriformes and all
mammals except rodents, insectivores and bats. The NISP (defined
as fragments whose precise location in the skeletal element, or
portion thereof, can be determined and quantified, and can be
assigned to species or size class) was as inclusive as possible (see
Yeshurun, 2011 for details of skeletal-element identification and
counting). Intra- and inter-site taxonomic comparisons in this work
are based on the NISP counts (Lyman, 2008). Due to the high
fragmentation of the bone assemblage, the majority of identified
specimens were assigned to size class rather than species (Table 1).
While the assemblage includes large numbers of unidentified
fragments (a sample of which was counted and divided by bone
type, size and burning for the sake of bone-fracture analyses
(Yeshurun, 2011)), all of the following procedures concern the NISP.
Body-part profiles were assessed by calculating Minimum Number
of Element (MNE; Lyman, 1994) values for each bone portion.
Standardizing the MNE by also taking bone side and age into account produced the Minimum Number of Individuals (MNI) figures.
All identified specimens except isolated teeth and squamate
vertebrae were systematically examined for bone-surface
Table 1
Animal groups used in this study.
Group
Small mammal
Small ungulate
Medium ungulate
Large ungulate
Tortoise
Lizard/snake
Bird
Species in the Epipaleolithic assemblages
Lepus capensis, Vulpes vulpes, Martes foina,
Felis sp., Meles meles, Herpestes incheumon,
Vormela peregusna, Canis lupus
Gazella gazella, Capreolus capreolus
Dama mesopotamica, Sus scrofa
Bos primigenius
Testudo graeca
Ophisaurus apodus
Alectoris chukar, Grus grus, Ardeidae, Falconiformes,
Strigiformes
The dominant species in each group appears in bold.
21
modifications, using a stereoscopic microscope (Olympus SZX7)
with a high intensity oblique light source, at 8e56 magnification,
following the procedure described in Blumenschine et al. (1996).
We searched for cutmarks (Binford, 1981) and hammerstone percussion marks, including conchoidal notches (Bunn, 1981; Capaldo
and Blumenschine, 1994; Pickering and Egeland, 2006) and percussion pits and striations (Blumenschine and Selvaggio, 1988;
Blumenschine et al., 1996; Pickering and Egeland, 2006). Evidence
for bone-working, ubiquitous in many Natufian contexts, was
recorded (Campana, 1989). We also looked for carnivore punctures,
scoring and digestion marks (Binford, 1981), as well as rodent gnaw
marks (Brain, 1981) and biochemical (root) marks (DomínguezRodrigo and Barba, 2006). Trampling striations (Behrensmeyer
et al., 1986; Domínguez-Rodrigo et al., 2009; de Juana et al., 2010;
Gaudzinski-Windheuser et al., 2010) and abrasion of bone edges
(Shipman and Rose, 1988) were sought, and weathering was noted
(Behrensmeyer, 1978). Burning presence and intensity were
recorded by bone color, generally following Stiner et al. (1995).
Bones with external and internal faces were assigned two burning
codes according to their external and internal surfaces, in order to
discern when in the life history of the specimen burning took place
(i.e., fleshed bone, defleshed bone, or cracked bone, following Cain,
2005). The mode of bone fragmentation was assessed by recording
shaft fracture-plane typology, shaft circumference and fragment
lengths (Villa and Mahieu, 1991; Bar-Oz, 2004),
Age-at-death profiles of gazelle, the most prominent game animal in the assemblages, were reconstructed by tooth eruption and
wear patterns and by bone fusion, following Munro et al. (2009).
We recorded the occlusal wear stages of isolated lower M3, P4 and
dP4 teeth and of M3 through P4/dP4 teeth in mandibles (where the
M2 and M1 could be distinguished). The tooth eruption and wear
data, and epiphyseal fusion data were assigned actual ages according to Munro et al.’s (2009) study of modern mountain gazelle
specimens from Israel, and also according to Davis’ (1983) scheme,
to ensure the compatibility with older studies. The proportion of
gazelle fawns was measured by the unfused proximal epiphysis of
the phalanx I, which fuses at approximately five to eight months of
age (Munro, 2009).
Gazelle sexing was performed using applicable character traits
and by multiple measurements, as defined by Munro et al. (2011)
for modern, known-sex mountain gazelles from Israel. The character traits used here included horn cores, the caudal wing tip of
the atlas and the cross-section of the pubis shaft. Measurements
included the relevant portions in the pelvis, atlas, axis, phalanx II,
calcaneum, metapodials, tibia and radius (see Munro et al., 2011:
their Table 7 for details of the measurements taken on each bone).
Additionally, we measured the distal humerus (BT*HDH, following
Davis, 1981) to facilitate comparisons with older studies, where it
was widely used for reconstructing both sex ratios and body-size
(e.g., Davis, 1981; Bar-Oz et al., 2004). Even though this measure
is not as sexually dimorphic in gazelles as previously thought
(Munro et al., 2011), it may still yield important information on sexindependent body-size patterns.
Statistical significance was tested at a ¼ 0.05 level (Shennan,
1988), using PAST freeware (Hammer et al., 2001) and SPSS
version 19 software. We employed the c2 distribution for
comparing nominal variables, such as the relative abundance of
species or taphonomic variables among selected samples. When
comparing many variables, Adjusted Residuals (AR) were calculated
for each cell and presented along with the composite c2, in order to
discern which cells most significantly differ from the expected
values. Adjusted Residual values are standard normal deviates,
indicating the probability that a single-cell comparison is statistically significant. Significant ARs have values equal to or greater
than 2 (Everitt, 1977; see Grayson and Delpech, 2008 for a
22
zooarchaeological application). We compared anatomical measurements and bone fragment lengths using Student’s t-test and
analysis of variance (ANOVA). For comparing taxonomic diversity
and similarity we used the ShannoneWiener Index for Heterogeneity (H), which considers both taxonomic richness and evenness
(Krebs, 1989). We used the Evenness Index e (Shannon H divided by
the natural log of the number of taxa or categories) for assessing the
level of evenness of the values of a given variable across anatomical
elements, or the evenness of the relative frequencies of age classes
(Faith and Gordon, 2007). The latter analysis was done to quantitatively discern even culling of age-classes from uneven culling, a
strategy focusing on the exploitation of one or two age classes
(Davis, 1983).
Results and preliminary discussion
Zooarchaeology and taphonomy of el-Wad Terrace
The Natufian of EWT is extremely rich in faunal remains (and
other classes of finds), yielding 2100e2500 identified specimens
per cubic meter of sediment.
Taxonomic spectrum The sample is based on 9141 identified specimens in the EN layer, and 2420 identified specimens in the LN layer,
representing a minimum number of 159 and 43 individual animals,
respectively (Table 2). The MNI tallies are strongly correlated with
the NISP counts (Spearman’s rho: r ¼ 0.92, p < 0.001 and r ¼ 0.90,
p < 0.001 for the EN and LN samples, respectively). This tight fit
means that the proportional representation of animals of very
different build (e.g., ungulates versus tortoises) is not biased due
to our identification and counting procedure; for example, tortoise
shell counts did not inflate the frequency of this animal. The EN
and LN samples are similar in their taxonomic spectrums. Both
assemblages are diverse and very uneven (Evenness Index:
e ¼ 0.27 and 0.29, respectively). Ungulates are the most numerous
group in NISP terms, followed closely by squamates (lizards and
snakes). Tortoises and small mammals are frequent, and some
birds are also found (Fig. 3).
The squamates are very well represented in the assemblages, by
their skull pieces and especially vertebrae (Table 2). The only
squamate that was identified to species in this study, based on the
dentary bone, is the legless lizard (Ophisaurus apodus), a large lizard
that may reach 1.2 m in length and weighs 300e600 g (Amitai and
Bouskila, 2003). Ungulates make up 42e44% of the assemblages
with mountain gazelle (Gazella gazella) being by far the most
common ungulate species in both periods. Larger ungulates are rare
(Table 2). The spur-thighed tortoise (Testudo graeca) is abundant in
both periods. It is represented by numerous carapace, plastron and
limb remains. Small mammals are well represented by the Cape
hare (Lepus capensis) and by four to seven species of small carnivores, predominantly red fox (Vulpes vulpes) (Table 2). Birds are
infrequently represented in both periods (Table 2, Fig. 3).
Complementing the faunal spectrum of Natufian EWT, but not
included in this study, are numerous micromammals (Weissobrod
et al., 2005, 2013), and some fish and marine shellfish. Among
the rodents, mole-rats (Spalax ehrenbergi) are very abundant, and
their taphonomic, contextual and demographic traits suggest regular consumption by humans (Weissbrod et al., 2012). Fish are
present along the sequence. Desse (in Valla et al., 1986) identified
four families of Mediterranean fish (Sparidae, Serranidae, Mullidae
and Muglidae) from the 1980s soundings at EWT. The fish from the
recent excavation are yet to be studied; thus far Sparidae have been
tentatively identified (I. Zohar, Personal communication). The
presence of these taxa suggests exploitation of the Mediterranean
coastal plain and estuarine zones (Bar-Yosef Mayer and Zohar,
2010). Large and rich assemblages of marine shells were
Table 2
Taxonomic composition of the EWT assemblages.
Early Natufian
Late Natufian
NISP
MNI
NISP
MNI
Ungulates
Gazella gazella
Capreolus capreolus
Small ungulate
Dama mesopotamica
Sus scrofa
Medium ungulate
Bos primigenius
Large ungulate
Fetus/neonate ungulate
1068
2
2873
24
42
88
5
6
3
43
1
300
1
697
3
6
7
7
1
1
11
1
Small mammals
Lepus capensis
Smal mammal-indet. (1)
Vulpes vulpes
Canis lupus
Martes foina
Felis sp.
Meles meles
Herpestes incheumon
Vormela peregusna
Small carnivore-indet.
231
212
186
20
19
25
11
2
1
112
17
30
32
43
3
6
3
2
2
1
1
43
1
2
3
2
3
6
7
14
1
10
1
1
1
1
1
1
1134
78
481
2424
12
9141
38
13
Birds
Alectoris chuckar
Grus grus
Ardeidae
Falconiformes-small
Falconiformes-medium
Falconiformes-large
Strigiformes
Raptor (indet.)
Bird-medium
Bird-large
Reptiles
Testudo graeca
Ophisaurus apodus (2)
Lizard (3)
Snake (3)
Squamata (indet.) (4)
Total
2
2
1
7
2
2
3
1
1
1
1
1
2
3
27
5
1
1
1
1
1
2
9
303
10
94
834
4
2420
11
2
1
Notes: (1) May include remains of small carnivores; (2) All skull and jaw fragments;
(3) All vertebrae; (4) All jaws.
retrieved, including edible mollusks, such as the Mediterranean
limpet Patella caerulea (Weinstein-Evron et al., 2007; Bar-Yosef
Mayer and Zohar, 2010).
Vertebrate taphonomy Counts of bone-surface modifications in the
EN and LN samples are grouped by the four mammal size classes
(small mammal, small ungulate, medium ungulate, and large
ungulate), the tortoises, birds and squamates (Table 3). Since the
squamates were not subjected to systematic microscopic
inspection, only burning data are given for them. Bone-surface
modification data indicate that the assemblages overwhelmingly
constitute butchered, consumed and discarded animal remains,
which consequently underwent some pre- and post-burial
destruction processes. Cutmarks related to all butchery stages, and
especially dismemberment and filleting, attest to the exploitation
of all ungulates, hare, fox and tortoises as food. A modest amount
of carnivore ravaging affected the assemblages, probably by a
small-medium carnivore. Subsequently, some of the skeletal
elements suffered moderate exposure related damage attested by
weathering, physical abrasion, rodent gnawing, and trampling.
Burning (indicated by bone color) appears on 27% of NISP in the
EN sample and on 24% of NISP in the LN sample. A sample of unidentified small ungulate specimens from the EN layer yielded an
identical proportion of burned bones, 27% (1553 of 5670 specimens
Figure 3. Taxonomic composition based on lumping species and size-classes into
higher taxonomic groups: (a) EN sample; (b) LN sample.
>8 mm in maximum dimension). In spite of the fact that bone
burning is abundant in EWT, burning intensity is not high. To
quantify bone burning, we used the Combustion Index (CI) defined
by Costamagno et al. (2005). The index values range from 0 (no
burned bones) to 1 (when all specimens are calcined), and thus
measures both burning frequency and intensity. The CI of the entire
assemblages is 0.10e0.11, reflecting moderately burned bones in
both periods. Looking at burning intensity by taxonomic group, the
squamates display the highest burning intensity (highest CI) during
both periods, with as much as 7e8% calcined bone. Birds display the
least burning, while small mammals, small ungulates and tortoises
display similar CI values (Table 4).
In the gazelle size class, the distribution of burning by skeletal
element is quite even and does not correlate with food utility.
Numerous cortical and compact bones (limb shafts, toes, teeth,
carpals/tarsals) display burning, and therefore preferential burning
of fat-rich portions cannot be demonstrated (Table 5). Additionally,
in both the EN and LN samples, most shaft fragments were burned
to a similar intensity on their exterior and interior surfaces,
meaning that the bone was burned when already defleshed and
fractured. Nevertheless, a significant difference exists between the
two samples in that during the LN a considerable number of shafts
exhibited burning with greater intensity on the exterior (25% in
the LN versus only 10% in the EN; c2 ¼ 12.49, p ¼ 0.002; Table 5).
Hence, it seems that the majority of ungulate gazelle bone burning
in EWT, especially in the EN sample, was unintentional, in accordance with lighting of hearths on pre-existing refuse, thereby
unintentionally inflicting secondary burning on bones buried underneath (Stiner et al., 1995; see also; Hardy-Smith and Edwards,
2004). The averaging pattern of such an activity in the faunal assemblages would eventually lead to near-uniform burning intensity
23
and frequency on different skeletal parts, probably masking some
of the effects of roasting body-parts or heating bones for marrow
extraction.
The analysis of breakage patterns was conducted on mammal
limb bones and phalanx I specimens, yielding largely similar results
for both periods (Table 3). Green, or fresh, fractures, attributed to
deliberate fracturing by humans or carnivores to access the bone
marrow, appear on 42e44% of the small mammal and ungulate
bones. This sizeable proportion is in line with a processed assemblage that subsequently suffered additional cycle/s of breakage
while the bones were already dry, resulting in intermediate and dry
breaks amounting to more than half of the specimens (Villa and
Mahieu, 1991).
Gazelle mortality profiles Mortality profiles were constructed for
gazelles by three tooth eruption and wear series and by two bone
fusion methods (following Davis, 1983; Munro et al., 2009). The raw
tooth-wear and bone fusion data are presented in Supplementary
Online Material [SOM] Tables A.1 and A.2, and five different
methods for presenting age data are compared in Fig. 4.
In the EN sample, the three tooth eruption and wear series
indicate a low number of juvenile (<18 months) gazelles,
amounting to 10e18% of ageable specimens. A more detailed
mortality profile, covering the entire life-span of the animal, indicates the overwhelming preponderance of adult animals, 18e96
months of age, along with a few fawns and juveniles (2e18 months)
and one very old animal (>96 months) in the EN sample (Fig. 4).
However, the picture changes when bone fusion is examined,
indicating a much larger proportion of juveniles (34e53%). The
most recently published and refined bone fusion method, based
upon a large sample of known-age gazelles from Israel, diverges in
the most marked way from the values obtained by tooth eruption
and wear (both following Munro et al., 2009). The skeletal elements
advocated by Munro et al. (2009) as being comparable to the dP4P4/M3 replacement and the attainment of adult body size are the
proximal humerus, proximal tibia and distal radius (MNE ¼ 17 in
the EN sample), while the older and widely used method of Davis
(1983) employs the summation of fused and unfused distal radii,
metapodials, tibiae, femora and the tuber calcis (MNE ¼ 104 in the
EN sample).
The results of the LN sample show a much sounder agreement
among tooth eruption and wear and bone fusion methods, even
though the samples are smaller than the EN sample (Fig. 4). All age
presentation methods indicate that a significant portion (31e50%)
of the culled gazelles were juveniles. Notably, Davis’ (1983) and
Munro et al.’s (2009) bone-fusion methods generate nearly identical figures here (31% and 33% juveniles, respectively) and the
proportion of juveniles is lower when presented by bone fusion
than by tooth eruption and wear, in contrast to the situation in the
EN sample. The detailed dentition-based sequence points to culling
of just two age groups, in equal proportions: fawns (two to seven
months of age) and young adults (18e36 months).
The marked discrepancy between the dental eruption and wear
and the bone fusion methods in the EN sample is difficult to
reconcile. Juvenile epiphyses could be less well preserved because
of their greater porosity, and epiphyses in general may be underrepresented in relation to teeth in an assemblage that suffered
density-mediated attrition and carnivore ravaging. This bias is expected to reduce the number of juveniles in the bone fusion
method compared with the tooth wear method, but the age profiles
exhibit the reverse pattern, namely a much larger representation of
juveniles by epiphyseal fusion.
Looking at the EN bone fusion results more closely (SOM
Table A.2), it appears that early-fusing elements are usually found
fused and that the high juvenile ratio stems from the late-fusing
elements. For example, the fusion state of the proximal phalanx I
24
Table 3
Bone-surface modifications and bone fracture patterns for all taxonomic groups: (a) Early Natufian sample; (b) Late Natufian sample.
A
(a)
NISP
Burning
Green fracture
Dry fracture
Intermediate
Limb shaft circumference
Weathering (stage 3e5)
Cutmarks
Percussion marks
Working
Gnawing (carnivore)
Gnawing (rodent)
Root-marks
Trampling striations
Abrasion
(b)
NISP
Burning
Green fracture
Dry fracture
Intermediate
Limb shaft circumference
Weathering (stage 3e5)
Cutmarks
Percussion marks
Working
Gnawing (carnivore)
Gnawing (rodent)
Root-marks
Abrasion
Small
mammal
Small
ungulate
Medium
ungulate
Large
ungulate
Tortoise
Lizard and snake
Birds
Total
n
%
n
n
n
<50
>50
100
n
of
%
n
%
n
of
%
n
%
n
%
n
%
n
%
n
%
n
%
819
212
25.9%
20
18
10
22
0
56
6
749
0.8%
21
2.8%
0
78
0.0%
0
0.0%
17
2.3%
8
1.1%
215
28.7%
23
3.1%
27
3.6%
3943
1027
26.0%
328
233
172
900
8
42
70
3775
1.9%
239
6.3%
108
1616
6.7%
91
2.3%
212
5.6%
63
1.7%
1238
32.8%
299
7.9%
169
4.5%
154
32
20.8%
10
14
10
39
0
4
8
136
5.9%
14
10.3%
3
71
4.2%
2
1.3%
8
5.9%
3
2.2%
55
40.4%
15
11.0%
2
1.5%
11
3
27.3%
1
0
0
1
0
0
2
10
20.0%
1
10.0%
0
1
0.0%
0
0.0%
1
10.0%
0
0.0%
5
50.0%
2
20.0%
0
0.0%
1134
279
24.6%
2995
929
31.0%
83
11
13.3%
9139
2493
27.3%
4
1132
0.4%
6
0.5%
0
0
30
1
83
1.2%
3
3.6%
0
0.0%
14
1.2%
4
0.4%
185
16.3%
11
1.0%
13
1.1%
1
1.2%
0
0.0%
0
0.0%
17
20.5%
2
2.4%
5
6.0%
962
8
132
91
5885
1.5%
284
4.8%
111
1766
6.3%
94
1.0%
252
4.3%
78
1.3%
1715
29.1%
352
6.0%
216
3.7%
n
%
n
n
n
<50
>50
100
n
of
%
n
%
n
of
%
n
%
n
%
n
%
n
%
n
%
n
%
143
31
21.7%
2
3
1
2
0
7
0
132
0.0%
5
3.8%
0
9
0.0%
0
0.0%
2
1.5%
3
2.3%
41
31.1%
3
2.3%
7
5.3%
998
267
26.8%
83
71
42
190
1
11
48
965
5.0%
39
4.0%
13
346
3.8%
16
1.6%
61
6.3%
14
1.5%
326
33.8%
76
7.9%
39
4.0%
16
2
12.5%
0
1
1
2
1
0
1
15
6.7%
2
13.3%
0
7
0.0%
0
0.0%
0
0.0%
0
0.0%
8
53.3%
1
6.7%
1
6.7%
8
1
12.5%
0
0
0
0
0
0
0
4
0.0%
1
25.0%
0
1
0.0%
0
0.0%
1
25.0%
0
0.0%
4
100.0%
1
25.0%
(which fuses at five to eight months) indicates 13% fawns, closely
replicating the lower proportion of juveniles obtained by the
dental methods. Thus, it may be that near-adult specimens were
misidentified as young-adults in the tooth wear methods. Moreover, the modern tooth eruption and wear, and bone fusion sequences were derived from captive animals (for which known
ages at death exist), and these could exhibit somewhat different
wear patterns due to different diets, and/or varying rates of growth
and development due to their captive status. This suggestion is
0.0%
303
57
18.8%
942
217
23.0%
9
0
0.0%
0
303
0.0%
0
0.0%
0
0
4
0
9
0.0%
0
0.0%
1
0.3%
4
1.3%
2
0.7%
84
27.7%
4
1.3%
6
2.0%
0
0.0%
0
0.0%
0
0.0%
3
33.3%
0
0.0%
1
11.1%
2419
575
23.8%
85
75
44
194
2
22
49
1428
3.4%
47
3.3%
13
363
3.6%
17
0.7%
68
4.8%
19
1.3%
466
32.6%
85
6.0%
54
3.8%
obviously hard to test without more actualistic data. Presently, we
draw on the fact that the high juvenile ratio is caused by late-fusing
elements to suggest that adults, as well as animals that nearly
reached adulthood and were osteologically immature, were primarily culled during the EN. The detailed tooth wear series point to
‘even culling’ of various age classes (e ¼ 0.712), with special
emphasis on adults, during the EN. Conversely, during the LN fewer
age classes were targeted (e ¼ 0.387) and more juveniles were
captured.
Table 4
NISP counts of bones burned to various degrees and the Combustion Index for each
taxonomic group.
Unburned Brown Black Gray White Total Combustion
NISP
index
Early Natufian
Small mammal
Small ungulate
Med. ungulate
Large ungulate
Tortoise
Lizard
and snake
Bird
Total
Late Natufian
Small mammal
Small ungulate
Med. ungulate
Large ungulate
Tortoise
Lizard
and snake
Birds
Total
607
2913
122
8
855
2058
127
607
22
2
160
408
71
328
7
1
99
275
11
62
2
0
11
187
3
40
1
0
8
59
819
3950
154
11
1133
2987
0.10
0.10
0.07
0.09
0.09
0.15
71
6634
6
1332
4
785
1
274
0
111
82
9136
0.05
0.11
112
731
14
7
246
725
16
154
2
0
32
72
14
96
0
0
22
72
0
7
0
0
3
56
1
10
0
1
17
143
998
16
8
303
942
0.08
0.10
0.03
0.13
0.07
0.12
9
1844
0
276
0
204
0
66
0
29
9
2419
0.00
0.10
Using the dP4eM3 series results, the LN sample is much more
biased towards juveniles younger than 18 months than the EN
sample, a difference that is statistically significant (c2 ¼ 5.43,
p ¼ 0.02). These figures can be cautiously interpreted using the
drive counts conducted monthly by Baharav (1974) between
November 1971eNovember 1972 on living gazelle herds in Ramot
Yissakhar (southeast Galilee, Israel; Baharav’s year-round averages
of the proportions of age classes and sexes are used here, because
they are reasonably comparable with the zooarchaeological data).
The modern counts indicated that the proportion of juveniles (18
months of age) in living herds is approximately 45% on a year-round
average. The low juvenile proportion in the EN sample diverges
significantly from the natural age structure (c2 ¼ 5.40, p ¼ 0.02),
highlighting the special emphasis on adult culling in that period.
Conversely, the LN proportion of juveniles is similar to a natural
herd structure (c2 ¼ 0.07, p ¼ 0.78; see also Bar-Oz et al., 2004).
Gazelle sexing The sexing results of the character traits show a
marked male bias in both periods (Fig. 5; SOM Table A.3) stemming
mainly from horn cores. This element is unmistakably dimorphic
but its utility for reflecting the actual sex ratios of the hunted
population has been questioned, because male horn cores are
Table 5
Characteristics of burned specimens in the small ungulate group.
Early Natufian
Evenness index for the
anatomical distribution
of burning
Correlation FUI*element
burning
Limb burning
Two sides equally burned
Exterior more burned
Interior more burned
Burned bone end
Burned bone shaft
Comparison end*shaft
Comparison of fragment
lengths
N:
Mean:
Var.:
T
Late Natufian
0.98
0.99
r ¼ 0.29, p ¼ 0.274
r ¼ 0.03, p ¼ 0.502
NISP
308
37
25
166
36.6%
208
30.0%
2
c ¼ 5.35, p ¼ 0.024
Burned Unburned
NISP
NISP
991
2523
19.662
23.972
131.06
313.44
8.51 (p < 0.001)
NISP
39
15
6
39
31.7%
23
20.7%
2
c ¼ 3.61, p ¼ 0.057
Burned Unburned NISP
NISP
257
627
17.222
19.742
47.853
109.82
4.19 (p < 0.001)
25
much more robust than female horn cores, possibly reflecting a
preservational difference, and because of the possibility that male
horn cores had been used as raw material and therefore might
have been preferentially cached in the habitation area (Munro,
2001; Bar-Oz, 2004). While no evidence exists for horn core
caches in el-Wad, ca. 10% of these elements bear traces of
fabrication (Yeshurun, 2011) and some finished artifacts made of
horn were discovered elsewhere at the site (Garrod and Bate,
1937; Weinstein-Evron, 1998). On the other hand, contextual
taphonomic analysis of the EN layer showed that both bone tool
production waste and unmodified butchery refuse are present in
the living surfaces investigated here, with little evidence of
removal of selected skeletal elements, especially not small and
non-obstructing elements such as female horn cores, from the
habitation area (Yeshurun et al., submitted for publication).
Nevertheless, the male-dominated picture is augmented by the
only other available character trait in the samples, the pubic shaft,
displaying four male elements versus one female element in the
EN sample (SOM Table A.3). Only one sexable (male) pubis shaft
was available in the LN sample.
An additional avenue is sexing by osteometrics, which necessitates an investigation of general body-size variability (see SOM
Table A.4 for the raw gazelle measurement data). Two skeletal elements that provide a sufficient number (n > 5) of measurements
in both the EN and LN samples are the distal humerus (BT and HDH,
following Davis, 1981) and the phalanx II (GL, Bd and Dd, following
Munro et al., 2011). The measurements on these elements are independent of age and taphonomic processes, since only fused,
unburned and unabraded elements were measured (von den
Driesch, 1976). Additionally, no significant intra-individual size
difference is evident for the phalanx II (Munro et al., 2011). All five
measurements either show significantly higher means in the EN
sample, or statistically similar means, while no measurement
provided a higher mean value in the LN sample (SOM Table A.5).
Plotting the phalanx II measurements (greatest length versus distal
depth) in both samples revealed that the EN gazelles are divided
into two size groups, with the LN gazelles corresponding with the
smaller one (Fig. 6). Thus it seems that EN gazelles possessed a
larger mean body-size compared with LN gazelles, and that a potential cause may be shifting sex ratios in favor of females in the LN.
The samples that were measured and sexed using the discriminate functions, following Munro et al. (2011) (altogether 42 specimens in the EN sample and 14 in the LN sample) yielded results
that were mostly larger than the female/male cutting point for the
modern specimens in the EN sample, and mostly smaller than the
cutting point in the LN sample (SOM Table A.4; cutting points
follow Munro et al. (2011)). Had the EWT gazelles been identical in
size to modern Israeli gazelles, the assemblages would be
composed of 81% males in the EN sample and 43% males in the LN
sample, a statistically significant difference (c2 ¼ 7.47, p < 0.01).
However, it is possible that Natufian gazelles differed in size
compared with present-day gazelles, which could cause the cutting
point to be too low and to misidentify large females as males.
A possible solution to separate body-size trends and sex ratios
would be to determine if EN and LN gazelle body-size differed
significantly from the modern Israeli gazelles that were used to
determine the sexing functions. This comparison (Table 6) indicates
that EN gazelles usually possess a similar body-size to modern
males and are significantly larger than modern females, while LN
gazelles exhibit the reverse pattern, being significantly smaller than
modern males but as large as modern females. Importantly, neither
the EN nor LN gazelles display larger body-size compared with
modern males, or smaller body-size than modern females (Table 6).
This indicates that the range of gazelle body-size in the EN and LN
was un-skewed compared with present-day gazelles, lending some
26
Figure 4. Gazelle aging. Left: juvenile-adult ratios based on five bone-fusion or tooth-wear series. Bone fusion 1 refers to the elements fusing at 18 months, equivalent to the dP4-P4/
M3 replacement (following Munro et al., 2009). Bone fusion 2 follows the traditional method by Davis (1983), which includes more elements. Sample size is indicated to the right of
each bar; Right: detailed mortality profile using the dP4eM3 series, based on tooth NISP. The numbers in parentheses indicate age in months.
support to the inter-period body-size difference in EWT as indeed
stemming from shifting sex ratios.
According to phalanx II measurements, male gazelles dominate
the EN assemblage by ratio of approximately 4:1 while males and
females are similarly represented in the LN, perhaps with females
slightly dominating. The inter-period difference in sex ratios is
highly significant (EN versus LN: c2 ¼ 7.47, p < 0.001). Baharav’s
(1974) drive counts indicated that in modern Israeli herds, females outnumbered males by a ratio of 100:81 in a year-round
average (see also Baharav, 1983). Thus, the EN sex ratios differ
significantly from the sex ratios of living herds (c2 ¼ 17.89,
p < 0.001), indicating specific targeting of males. On the other hand,
the LN sample is similar to the living herd in this respect (c2 ¼ 0.02,
p ¼ 0.90), meaning that LN hunters culled gazelles with no preference to sex.
Taphonomic markers for occupation type: Early versus Late
Natufian Several taphonomic markers were defined above for
gauging site-occupation intensity through the Natufian sequence of
EWT. We compare the LN with two different EN accumulations
(described above), displaying clearer contextual (architectural)
patterns: a domestic faunal assemblage in primary deposition
following consumption activities on the spot versus Locus 25, a
non-domestic area beyond the living compound, containing
occasionally-tossed consumption refuse (Yeshurun et al., submitted
for publication).
The comparison results (Table 7) show that the volumetric
density of faunal remains by total bone mass is highest in the domestic EN and about half of that in Locus 25 and in the LN. When
the volumetric densities of NISP counts are taken into account, the
phalanx II Dd (mm)
14
13
12
11
10
9
8
18
Figure 5. Gazelle sexing by character traits.
20
22
24
phalanx II GL (mm)
26
28
Figure 6. Gazelle sexing by body-size: comparison of phalanx II measurements in the
EN (black diamonds) and LN (gray squares) samples.
Table 6
Summary of the comparisons of Early Natufian (EN) and Late Natufian (LN) gazelle
measurements with modern male and modern female gazelle measurements.
BT
GL
Bd
Dd
Dd tibia
humerus phalanx II phalanx II phalanx II
EN compared
with modern
EN compared
with modern
LN compared
with modern
LN compared
with modern
No diff.
No diff.
No diff.
No diff.
No diff.
No diff.
Larger
Larger
Larger
Larger
Smaller
Smaller
No diff.
Smaller
No diff.
No diff.
Larger
No diff.
male
female
male
female
Raw measurements and t-test results are detailed in SOM Tables A.5 and A.6.
Modern gazelle measurements were taken from Munro et al. (2011: their Table 3).
difference among the domestic EN and the LN is less dramatic,
while Locus 25 still lags behind. The NISP-based counts of bone
modifications (e.g., burning, gnawing, etc.) are transformed into
Adjusted Residuals (AR) in order to assess which counts significantly diverge from the expected values. Some significant differences are evident in the statistical comparison (Fig. 7; see SOM
Table A.7 for details, AR values and composite c2 results): sub-areal
weathering is significantly higher in the Locus 25 and LN samples
compared with the domestic EN, attesting to slower rate of sediment build-up. On the other hand, carnivore gnawing is significantly underrepresented in the LN. The slow-accumulating
material in Locus 25 displays both higher weathering and more
intense carnivore gnawing compared with the LN sample. Boneburning measures are similarly represented in the LN compared
with the domestic EN. In contrast, burned bones are significantly
underrepresented in Locus 25. Finally, trampling is significantly
overrepresented in the domestic EN sample and significantly underrepresented in the LN sample. Post-discard fractures (defined
here as the ‘dry’ and ‘intermediate’ fracture categories combined)
are significantly underrepresented in Locus 25 but similarly
distributed in the domestic EN and in the LN samples (Fig. 7,
Table 7).
To conclude, the taphonomic measures of occupation intensity
position the LN as intermediate between the EN domestic and nondomestic samples. Some important characteristics such as bone
burning and breakage align the domestic EN and the LN, highlighting the similarity in post-discard destruction processes and
consequently, the degree of occupation repetitiveness and type of
habitation. This similarity, in turn, helps to keep site type as constant as possible and renders the inter-phase paleoeconomic
comparison more robust.
Table 7
Comparison of taphonomic markers for occupation intensity among the Early
Natufian (EN) and Late Natufian (LN) samples.
Bone mass (gr)/m3
NISP count/m3
Weathering
(long bones)
Carnivore gnawing
Rodent gnawing
Burning (total)
Indirect burning
(shafts)
Dry þ intermediate
fractures
EN domestic area
(inside Structure II)
EN occasional
toss area (Locus 25)
LN
15,267
2716.54
13 (2.5%)
9883.89
1738.61
4 (14.3%)
106 (6.2%)
35 (2.1%)
136 (8.0%)
944 (23.5%)
136 (84.0%)
11 (10.7%)
4 (3.9%)
8 (7.8%)
35 (12.0%)
2 (67.0%)
68 (4.8%)
19 (1.3%)
85 (6.0%)
575 (23.8%)
45 (75.0%)
149 (58.0%)
16 (36.0%)
119 (58.0%)
7708.57
2180.95
25 (8.8%)
Excavation volumes were not available for all units, so the faunal remains from these
units were excluded from calculations.
27
El-Wad in diachronic context
A broader perspective of intra-Natufian economic trends is
achieved by comparing the entire Epipaleolithic sequence of the
Israeli coastal plain, including the Kebaran site of Nahal Hadera V
(Bar-Oz and Dayan, 2002), the Geometric Kebaran sites of Hefzibah
(Bar-Oz and Dayan, 2003) and Neve David (Bar-Oz et al., 1999), and
the Early and Late Natufian faunas of EWT presented above. All of
these sites are positioned in a north-south strip along the Mediterranean coastal plain of Israel, ca. 40 km long, representing dense
anthropogenic accumulations of vertebrate faunas and other cultural refuse. Their zooarchaeology and taphonomy were published
in detail (Bar-Oz, 2004) using compatible methods to the ones
employed here.
Taxonomic abundances in the Epipaleolithic coastal plain The EN
and LN phases of EWT are generally similar in the presence and
abundance of taxa, but some variations do occur. Rather than
focusing on particular differences in relative abundances of certain
taxa, which are sometimes difficult to interpret and may stem from
differing sample sizes or other idiosyncrasies, the taxonomic
abundances are evaluated here using Stutz et al.’s (2009) indices,
described above. Our results indicate no gradual temporal trends
(as interpreted by Stutz et al., 2009); rather, marked differences
characterize the pre-Natufian and Natufian assemblages,
regardless of chronological position of the sample within this
division (Fig. 8, Table 8). The notable differences amongst the preNatufian and Natufian strata, namely the increase in small game
and the dwindling numbers of larger ungulates, overshadow the
subtle differences that exist among the Early and Late Natufian
phases of EWT. The fast small game index declines during the
Natufian of EWT, while the slow small game index remains
constant in the EN-LN comparison.
Gazelle culling patterns in the Epipaleolithic coastal plain Detailed
investigations of gazelle age and sex profiles at EWT provide a
picture of shifting culling patterns through the Natufian sequence.
Putting the EWT results in broader Epipaleolithic perspective, the
pre-Natufian gazelle culling trends are summarized in Table 9. The
proportion of juveniles in the Kebaran and Geometric Kebaran
faunas is 24e39%, higher than the Early Natufian EWT (18%).
However, only the Neve David assemblage significantly differs in
this respect from the natural herd age structure (c2 ¼ 9.73,
p ¼ 0.001). Nahal Hadera V and Hefzibah 7e18 do not differ from
a natural herd, as also Late Natufian EWT (c2 ¼ 1.07, p ¼ 0.30 and
c2 ¼ 0.35, p ¼ 0.55, respectively). The evenness of age classes is
similar across all assemblages (e ¼ 0.671e0.712) except for the
markedly low value of the LN sample from EWT (Fig. 9).
Additionally, the proportion of gazelle fawns (about six months
old or less), measured by the proportion of unfused to fused
proximal phalanx I, is variable, reaching 18% at the most in the LN
assemblage. Notably, fawns increase significantly from the earlier
Epipaleolithic to the Natufian (Fig. 9; pooled pre-Natufian versus
pooled Natufian: c2 ¼ 11.66, p < 0.001). Since the average yearround proportion of fawns in living herds is about one-seventh of
the population (Baharav, 1974), the low pre-Natufian proportion
of fawns significantly differs from living herds (c2 ¼ 10.92,
p < 0.001) while the Early and Late Natufian assemblages sample
fawns according to their living abundance (c2 ¼ 0.25, p ¼ 0.62).
Sex ratios, reconstructed by body-size in comparison to knownsex gazelle collections, again showed an inter-phase difference in
EWT: male culling in the EN versus unselective culling of sexes in
the LN. Comparison with the pre-Natufian Epipaleolithic of the
same ecological region can help separate conflating factors that
affect body-size (Bar-Oz, 2004). Gazelle body-size trends were
evaluated using the humerus BT measurement, a relatively wellpreserved portion that was measured in previous studies. Results
28
Total big game index
Early Natufian - domesƟc
Adjusted Residuals
6
4
2
0
-2
-4
-6
Early Natufian - non-domesƟc
Adjusted Residuals
6
NHV
4
2
-2
-4
-6
Late Natufian
6
Adjusted Residuals
NVD
Large big game index
0
4
2
0
-2
Total small game index
1
0.9
0.8
0.7
0.6
0.5
0.4
0.3
0.2
0.1
0
HEF
EWT-EN
EWT-LN
Medium big game index
1
0.9
0.8
0.7
0.6
0.5
0.4
0.3
0.2
0.1
0
NHV
NVD
HEF
EWT-EN
EWT-LN
-4
-6
Figure 7. Comparison of taphonomic markers for occupation intensity among the EN
(2 different contexts) and LN samples, using Adjusted Residuals (AR). Positive AR
values denote overrepresented traits, while negative AR values denote underrepresented traits. Significant values (AR 2) appear as black columns.
indicate that gazelle body-size underwent significant changes
during the Epipaleolithic (ANOVA F ¼ 5.31, p < 0.001). Tukey’s
pairwise comparisons indicate that this difference derives mostly
from the EN assemblage, which displays significantly larger measurements than the preceding Nahal Hadera V and Neve David
gazelles (Fig. 10A; data from SOM Table A.8).
The trend of larger gazelle body-size in the EN was attributed to
increased proportion of males, but other potential explanations
should be considered, such as cooler climate (e.g., Bergmann’s rule,
see Davis, 1981) and human impact on gazelle populations, as
suggested by Cope (1991) and later countered by Bar-Oz et al.
(2004). If cooling was the reason for the increase in gazelle bodysize, other mammals should also follow this trend (Ducos and
Horwitz, 1997). The only mammal available for comparison was
the hare (humerus BT and scapula glenoid fossa BG measurements).
Although the samples are small (SOM Table A.8.), the hare does not
exhibit a parallel trend of body-size increase (EN versus preNatufian hares: ANOVA F ¼ 2.87, p ¼ 0.07 and F ¼ 0.07, p ¼ 0.959
for the humerus and scapula measurements, respectively)
(Fig. 10B,C). Hence, the most plausible interpretation of the significantly larger-sized gazelles in the EN is change in sex ratios.
Apparently males and females were culled in similar proportions
along the coastal plain sequence, except for Early Natufian EWT.
Slow small game index
Fast small game index
1
0.9
0.8
0.7
0.6
0.5
0.4
0.3
0.2
0.1
0
NHV
NVD
HEF
EWT-EN
EWT-LN
Figure 8. Index values for the Epipaleolithic sequence of the Israeli coastal plain
(abbreviations as in Table 8). Black dashed lines separate the Natufian and pre-Natufian
samples.
In sum, during most of the Epipaleolithic hunters exploited a
wide range of gazelle age classes in the Israeli coastal plain, with
varying degrees of reliance on juveniles, and in most cases did not
seek to capture a particular sex. The Early Natufian stands out in
this series for its emphasis on male culling and its particularly low
proportion of juveniles, albeit with a seemingly contradictory trend
of taking fawns in relation to their natural abundance (Fig. 9).
Discussion
Similar site function through time
The ancient function of excavated localities may affect paleoeconomic inferences. Feasting activities or ephemeral use of parts
of large and complex sites may introduce serious biases when
evaluating the long-term subsistence trends of Epipaleolithic
29
Table 8
NISP relative abundance indices used to measure intensification of animal resources during the Epipaleolithic and index values for the Epipaleolithic series of the coastal plain,
following Stutz et al. (2009: their Table 4).
Formula
Total big game index
Total small game index
Large big game index
Medium big game index
Slow small game index
Fast small game index
NISPbg/(NISPbg þ NISPsg)
NISPsg/(NISPbg þ NISPsg)
NISPlbg/(NISPlbg þ NISPsbg)
NISPmbg/(NISPmbg þ NISPsbg)
NISPssg/(NISPssg þ NISPsbg)
NISPfsg/(NISPfsg þ NISPsbg)
KEB
GKEB
GKEB
EN
LN
NHV
NVD
HEF
EWT-EN
EWT-LN
0.93
0.07
0.01
0.31
0.03
0.08
0.94
0.06
0.01
0.34
0.01
0.07
0.94
0.06
0.03
0.22
0.01
0.06
0.67
0.33
0.00
0.04
0.22
0.18
0.69
0.31
0.01
0.02
0.23
0.13
‘Big game’ refers to ungulates: small, medium and large. ‘Small game’ refers to all other animals included in this study except squamates, divided to fast types (hare, small
carnivores and respective size class, game birds such as partridge) and slow types (tortoise). Note that in Stutz et al. (2009) original indices small carnivores were excluded.
Abbreviations: KEB, Kebaran Culture; GKEB, Geometric Kebaran Culture; EN, Early Natufian; LN, Late Natufian; NHV, Nahal Hadera V; NVD, Neve-David; HEF, Hefzibah 7e18;
EWT, el-Wad Terrace.
Table 9
Summary of gazelle age and sex data for the two assemblages in the present study
(in bold type), augmented by data from the Epipaleolithic sequence of the coastal
plain (Nahal Hadera V, Hefzibah 7e18 and Neve David, all taken from Bar-Oz, 2004).
Proportion of juveniles (dental)
Evenness of age classes
Proportion of fawns
(unfused phalanx I)
Proportion of males
(by body size)
KEB
GKEB
GKEB
EN
LN
NHV
39%
0.671
7.3%
NVD
25%
0.683
8.5%
HEF
24%
0.710
0.5%
EWT
18%
0.712
13.0%
EWT
50%
0.387
18.0%
81%
43%
Under 50%
Abbreviations are as in Table 8.
households (Yeshurun et al., 2013a, submitted for publication; see
also Asouti and Fuller, 2013). We applied taphonomic proxies of
site-occupation intensity to investigate this issue and control for
changing site type. Our intra-Natufian analysis within the key site
of el-Wad Terrace compared domestic and non-domestic EN accumulations with the more enigmatic LN phase. Since the postdiscard damage patterns in the LN sample are intermediate between the domestic and non-domestic EN samples, we suggest that
the difference between the two Natufian chronological phases was
of degree, not of kind. The intensive post-discard damage from
recurrent human activities and the considerable quantities of
faunal remains that were found in the LN probably mean that EWT
continued to serve as an important habitation site during the latter
half of the Natufian. This conclusion is somewhat unexpected,
Figure 9. Inter-phase comparison of gazelle culling patterns in the coastal plain Epipaleolithic: proportions of juveniles (<18 months old); proportion of fawns (<6
months old); and the evenness of age classes (e). The proportion of males, based on
body-size considerations, is given for the sites in this study and approximated values
(based on osteometrics) are given for the pre-Natufian sites (see discussion in Bar-Oz,
2004). The dashed line separates the pre-Natufian and Natufian samples.
given the dissimilarities evident in the Natufian sequence of the
site: the dramatic decrease in the thickness of habitation sediments, the disappearance of stone architecture, the dwindling of
bone ornaments and the higher accumulation rates of naturallydeposited micromammals (Yeshurun, 2011; Weinstein-Evron
et al., 2012; Weissbrod et al., 2012). This discrepancy may be
explained when the entire Mt. Carmel settlement system is
considered (see below).
The general depositional similarity through time in EWT is
reinforced when compared with several conspicuously different
types of Natufian accumulations. The first example comes from the
same site, the EN fauna from Chamber III of el-Wad Cave (EWC;
Weinstein-Evron, 1998; see Fig. 1B for location). The EWC assemblage exhibits a medium big game index almost reaching preNatufian proportions and a very high fast small game index (Stutz
et al., 2009), both very high compared with the EN terrace fauna.
On the basis of the location of this excavation area in the dark part
of the cave, the rarity of built features, lack of burials and composition of the lithic assemblage, it was suggested that it represented
a specialized ‘dumping area’ and not a domestic or a burial locality
(Weinstein-Evron, 1998). Supporting evidence for the nondomestic use of this locality may be found in the scanty evidence
for in situ fire activity; the proportion of burned bone specimens is
very low (4% of NISP; Rabinovich, 1998). These characters are
reminiscent of the ‘occasional-toss area’ of Locus 25 in the EN
terrace. Moving to the LN, Raqefet Cave (Mount Carmel, ca. 10 km
east of el-Wad) that was used primarily for burial and other
communal activities (Nadel et al., 2012, 2013) yielded a faunal
assemblage weakly affected by post-discard damage (notably, low
bone-burning intensity and frequency), as well as a relatively high
value for the medium big game index, clustering with Locus 25 and
the EWC samples in some attributes (Yeshurun et al., 2013a).
Similarly, the LN fauna in the burial cave of Hilazon Tachtit (Galilee,
Israel) yielded a low proportion of burned bones, almost identical
to the EWC and Raqefet cases (Grosman and Munro, 2007; Munro
and Grosman, 2010). These three non-domestic examples contrast
with the intensively-trampled and unintentionally-burned EN and
LN faunas in the current study. The LN fauna sampled here clusters
better with LN habitation sites such as Hayonim Terrace (Munro,
2012) and ‘Eynan (Bridault et al., 2008), which also show extensive post-discard damage and, unlike Late Natufian EWT, preserve
architectural features.
By the same token, the pre-Natufian assemblages in the Israeli
coastal plain Epipaleolithic series, namely, Nahal Hadera V, Hefzibah and Neve David, seem to represent spatially and vertically
extensive amalgamations of human occupations attesting to
various domestic (and other) activities and preserving ‘living
floors’, a hut-like feature (in NHV), spatially defined concentrations
of finds and diverse chipped lithics and groundstone assemblages
30
Figure 10. Comparison of measurements among the coastal plain Epipaleolithic sites: (A) gazelle humerus BT; (B) hare humerus Bd; and (C) hare scapula-glenoid fossa (GLP). The
Geometric Kebaran assemblages were pooled due to sample size limitations. See SOM Table A.8 for statistics.
(Ronen et al., 1975; Saxon et al., 1978; Kaufman and Ronen, 1987;
Kaufman, 1989, 1992; Barkai and Gopher, 2001; Bar-Oz, 2004;
Bocquentin et al., 2011). The general similarity in the function of
habitation through time in our case-study enables a more robust
appreciation of Natufian economic trends, relatively independent
of changes in site function.
Economic intensification through time
Economic trends in the Natufian sequence of EWT were put in a
broader perspective, using the geographically distinctive Epipaleolithic series of the coastal plain (Bar-Oz, 2004) and a behavioral ecology approach (Stiner and Munro, 2002; Munro, 2004). The
results attest to regional-scale intensification of game resource
exploitation in the Natufian, relative to the pre-Natufian Epipaleolithic of the same region. Starting with the EN and lasting
through the LN, ungulates dwindle in numbers and small animals
comprise a much larger portion of Natufian prey. The ungulates that
were still routinely captured were overwhelmingly small ungulates
(i.e., gazelles) as opposed to a significant input of larger ungulates
(i.e., fallow deer) in the preceding Epipaleolithic samples. No difference was observed in the EN versus LN phases of EWT in these
respects. On the more local (site) scale, fast small game is somewhat more abundant in the EN phase, possibly signaling stronger
human prey pressure in the camp’s catchment area (Stiner et al.,
1999, 2000; Munro, 2004).
The observed prey trends were previously identified in the
coastal plain sequence (Bar-Oz, 2004) as well as in the KinneretGolan region (Davis et al., 1988; Davis, 2005), the Wadi Meged series in the lower Galilee (Munro, 2004; Stiner, 2005) and the
Damascus Basin (Napierala, 2011). However, only the Wadi Meged
series had previously included EN samples in the analysis, yielding
significant differences in small game procurement between the EN
and LN (Munro, 2004). Other inter-regional syntheses (Munro,
2009; Stutz et al., 2009) argued for a gradual decline of medium
and large big game and an increase of small game species over time,
between the Early Epipaleolithic and the EN and LN of the Mediterranean zone. While the present results generally support these
assertions, it is important to note that no ‘gradual’ trend is
apparent. Rather, the shift in ungulate proportions relative to small
game, and the near-disappearance of medium and large ungulates,
occur abruptly in the EN and remain largely unchanged in the LN.
Additionally, small game indices (fast versus slow) display the same
marked ‘step’ in the pre-Natufian e EN transition, exhibiting
negligible index values in the Kebaran and Geometric Kebaran
samples as opposed to high values in either the EN or LN. The
striking difference between the pre-Natufian and Natufian assemblages overshadows any minor variations within these two time-
periods, including the modest decline in fast small game abundance in the LN.
The importance of small animals in human diet during the late
Epipaleolithic reaffirms the true broad spectrum nature of Natufian
animal economy. Such common exploitation of tortoises, hares,
foxes and possibly squamates, which comprise around 50% of the
identified specimens, along with mole-rats, Mediterranean fish and
edible mollusks (Yeshurun et al., 2009; Bar-Yosef Mayer and Zohar,
2010; Weissbrod et al., 2012) was rarely observed in the preNatufian Epipaleolithic and Upper Paleolithic record of northern
Israel (e.g., Rabinovich, 2003; Bar-Oz, 2004; Stiner, 2005; Marom
and Bar-Oz, 2008), let alone the more arid areas of the southern
Levant (e.g., Martin et al., 2010). The lakeside settlement of Ohalo II
is currently the only example of pre-Natufian fishing (of freshwater
fish) and one of the rare cases of fowling (Simmons and Nadel,
1998; Zohar, 2003). However, pre-Natufian sites that are not
located in a lacustrine setting have not yielded any significant evidence of fishing, in contrast with the presence of Mediterranean
fish in several Natufian inland sites (Davis et al., 1994; Bar-Yosef
Mayer and Zohar, 2010). Importantly, four families of Mediterranean fishes were identified at EWT (Valla et al., 1986), even though
the site was 8e12 km away from the sea shore. Birds are sporadically present in the pre-Natufian Epipaleolithic, but can usually be
considered to reflect specialized exploitation for raw materials, not
regular and long-term dietary contributions (Martin et al., 2013).
The extremely broad spectrum animal diet of the Natufian lends
strong support to the view of Natufian economy as being intensified. Natufian foragers chose to hunt significant amounts of lowerranked game, probably in response to population packing in a
defined territory (Munro, 2004). Taking a long-term perspective
beyond the Epipaleolithic, ungulates had been the main target of
hunting in the Levant since the Middle Pleistocene and Natufianlike exploitation of many small game taxa was very uncommon
up until this stage (Stiner, 2005; Yeshurun, 2013).
Plant exploitation must have played a very important role in
Epipaleolithic subsistence, and intensified plant exploitation is
commonly mentioned as an Epipaleolithic hallmark. While sporadic charcoal, pollen and phytolith data exist in the Natufian (LevYadun and Weisntein-Evron, 1994; Weinstein-Evron, 1998; Portillo
et al., 2010; Rosen, 2010; Power et al., 2014) macrobotanical evidence from the southern Levant Epipaleolithic is so scarce (Asouti
and Fuller, 2012: their Table 2) that quantitative analyses cannot
realistically be performed. Therefore, until new archaeobotanical
data sets are available, the BSR must be measured by animal
exploitation alone in this period.
Results from gazelle culling patterns augment the economic
‘break’ seen between the EN and its forerunners. The probable
targeting of males during the EN is unique in the Epipaleolithic
series of the coastal plain, paralleling other Natufian assemblages
(including LN ones; Garrard, 1980; Cope, 1991; Bar-Oz et al., 2004).
It was suggested that targeting males was a (possibly seasonal)
adaptation to ecologically sustainable hunting, attempting to
minimize the effects of human predation on the non-migratory
mountain gazelle herds in a territory (Cope, 1991).
The pre-Natufian assemblages exhibit age distributions that are
similar to the year-round average of modern herds and cannot
point to exploitation of specific age groups. An exception is the
avoidance of fawns, presumably due to small meat and fat returns
(Munro and Bar-Oz, 2005). The picture changes in the Early Natufian, where adult male gazelles were targeted, and fawns were
taken in proportions similar to their natural abundance, in line with
the BSR hypothesis. The evenness of age groups in both the earlier
Epipaleolithic and the EN is moderately high. It is possible that the
age group evenness index reflects two different mechanisms in the
pre-Natufian and EN assemblages, namely, culling of both young
and older gazelles in the former and averaging of exceedingly
numerous occupations in the latter, when gazelles were deposited
during several seasons (Davis, 1983). Such a scenario may explain
the seemingly contradictory evidence of adult-focused culling in
conjunction with hunting some fawns in the EN. The picture
changes again in the LN, demonstrating an unselective culling of
sexes, juveniles and fawns. The low evenness of age groups here
may stem from the small sample in comparison to the other assemblages and therefore enlarged samples will be needed to verify
the LN culling evenness trend.
Age profiles generally present a complex and inconsistent picture throughout the Natufian assemblages. The proportion of gazelle fawns in the EN period ranges between 4 and 40%, with older
sub-adult proportions very variable (Munro, 2009; Edwards and
Martin, 2013). High juvenile proportions of 30e50% characterize
several, but not all, LN assemblages (Davis, 1983, 2005; Munro,
2012; Yeshurun et al., 2013a). Hence, ambiguous patterns are
evident in the issue of Natufian gazelle culling, which, although one
of the most researched topics in Natufian archaeology, it is one of
the least understood. Better interpretative frameworks are especially needed if we are to understand the Natufian preferences for
gazelles of certain age classes.
Decreasing mobility and increasing catchment exploitation
The importance of the Natufian economic transformation lies in
its implications for inferring terminal Pleistocene mobility patterns.
Small animal exploitation has been seen as an important marker of
prolonged site occupation and population growth in the Natufian,
because of the need to feed a growing or more permanent population in a given territory (Tchernov, 1993b; Stiner and Munro,
2002; Munro, 2004, 2009; Weissbrod et al., 2012). Thus, the
broad-spectrum economy and the increased culling of immature
gazelles were interpreted as a certain facet of the EN sedentarization phenomenon, which has also been inferred from multiple
archaeological proxies. The notion of EN sedentism is strongly
supported by our findings, as the fauna presented here comes from
a major hamlet, where stone architecture is plentiful and the
density and diversity of finds are very high (Weinstein-Evron et al.,
2013). Both the intensified animal economy and the archaeological
nature of the site contrast sharply with the pre-Natufian Epipaleolithic evidence from the same region (Bar-Oz, 2004). Thus,
keeping the site function, site catchment and analytical methods as
constant as possible convincingly demonstrated the economic
intensification marking the emergence of sedentary life in the EN,
ca. 15e14 ka in the Israeli coastal plain.
In conjunction with decreasing mobility, one of the hallmarks of
the Natufian economy, the regular capture of small mammals, may
31
reflect intensified site catchment exploitation in this period. While
gazelle hunting was probably performed by shooting with
microlith-tipped projectiles (Yeshurun and Yaroshevich, 2014),
albeit with increasingly efficient designs (Yaroshevich et al., 2010),
the regular hunting of small and agile terrestrial animals, primarily
hares and foxes, which are solitary and nocturnal, was likely performed by traps and snares (e.g., Winterhalder, 1980; Holliday and
Churchill, 2006). Modern studies on poaching of Israeli wildlife
showed that most trapping methods, such as simple cable snaring,
would indiscriminately capture a broad spectrum of small and
medium mammal species (Yom-Tov, 2003). Thus, snaring would
have constituted an efficient and predictable hunting method for
the systematic execution of the broad-spectrum animal economy in
a defined Natufian territory. Setting, maintaining and keeping track
of a system of traps and snares requires a long presence of humans
in a given territory and excellent acquaintance with the landscape,
the animals and their local patterns of foraging, and thus may be
related in the ethnographic record to sedentary or semi-sedentary
societies (Holliday, 1998; Holliday and Churchill, 2006).
The inferred patterns of LN sedentism and site-catchment
exploitation do not differ much from EN patterns in this study.
The LN period in the Mediterranean southern Levant was sometimes viewed as one of cultural decline and a return to a more
mobile way of life, compared with the Early Natufian (Garrod, 1957;
Henry, 1991; Valla, 1995; Belfer-Cohen and Bar-Yosef, 2000; BarYosef and Belfer-Cohen, 2002; Grosman, 2003; Munro, 2004).
This was mainly due to the more impoverished and thinner nature
of LN layers in the intensively-researched sites of el-Wad, Hayonim
Cave and ‘Eynan, where architecture, elaborate burials and art objects were considered to be less frequent. However, it has become
increasingly clear that stone-built dwellings and other architectural
features are common at some Late/Final Natufian sites, as apparent
from the new fieldwork at ‘Eynan (Valla et al., 2007), Huzuk Musa
(Rosenberg et al., 2010) and Nahal Ein Gev II (L. Grosman, Personal
communication), as well as other sites in the more arid regions
(Henry, 1976; Goring-Morris, 1991; Conard et al., 2013; Richter
et al., 2013). Elaborate LN funerary customs are now known in
Hayonim Terrace (Tchernov and Valla, 1997), Hilazon Tachtit Cave
(Grosman et al., 2008; Munro and Grosman, 2010) and Raqefet Cave
(Nadel et al., 2012, 2013; Yeshurun et al., 2013a). While at EWT the
nature of the LN layer is indeed impoverished compared with the
EN, it was taphonomically demonstrated here that the site was still
used for diverse domestic activities (as well as for burial, which is
spatially and perhaps temporally segregated from the habitation
sediments; Weinstein-Evron, 2009) and was intensively and
recurrently inhabited. While the taphonomic markers of siteoccupation intensity indicate that the LN habitation were somewhat less intensive compared with the EN, the economic indicators
pertaining to site-occupation intensity do not differ much from the
EN phase, especially when the pre-Natufian sites are used as
reference. Furthermore, just 5 km north of el-Wad there existed a
major LN base-camp at the site of Nahal Oren, displaying an
architectural compound (a terrace wall and associated living surfaces), as well as a spatially distinct cemetery (Stekelis and Yizraely,
1963; Noy, 1991) and evidence of occupations throughout all of the
LN sub-phases (Grosman et al., 2005), much like EN el-Wad. Thus,
the major base-camp of the Carmel Coast may have simply shifted
from EN el-Wad to LN Nahal Oren, hardly suggesting any loss of
mobility or cultural complexity in the Mediterranean Levant
(Weinstein-Evron, 2009). The presence of both base-camps and
specialized sites may suggest stable (or even growing) LN populations in the Mount Carmel-Galilee region ca. 13.7e11.7 ka, corresponding with the evidence for geographic expansion of
settlements and the emergence of novel technological and social
adaptations in the latter half of the Natufian (e.g., Bar-Yosef, 2002;
32
Valla et al., 2007; Grosman et al., 2008; Stutz et al., 2009;
Weinstein-Evron, 2009; Rosen, 2010; Valla, 2012; Nadel et al.,
2013).
Conclusions
We evaluated terminal Pleistocene subsistence change in the
Levantine Epipaleolithic, when mobile foragers were settling down
in the course of the earliest foraging-to-farming transition.
Archaeologically, the salient features of the late Epipaleolithic
Natufian Culture at key sites such as el-Wad (the regular appearance of stone architecture, cemeteries, groundstone and art) stand
out, suggesting remarkable socioeconomic changes. Early Natufian
el-Wad Terrace yielded novel paleoeconomic signals compared
with the pre-Natufian Epipaleolithic of the same region, accompanied by the preponderance of stone-constructed dwellings, high
density of finds, and heavy taphonomic damage from repeated and
intensive occupations. The Late Natufian sample differs in gazelle
culling patterns but nonetheless represents enduring economic
intensification, coupled with relatively reduced, but still considerable post-discard taphonomic damage from repeated occupations.
The LN occupations may have been somewhat less intensive than
the EN occupations in the particular case of el-Wad, but still left
behind habitation deposits of an important hamlet. The use of
contextual taphonomy, focusing on post-discard damage in Epipaleolithic camps, carries a somewhat unexplored potential and
should be implemented in forthcoming studies to distinguish degrees in the intensity of accumulation and repetition of habitations.
It is evident from the Epipaleolithic series of the Israeli coastal
plain that the Early Natufian constituted an economic break,
reflecting decreasing mobility and increasing exploitation of the
site catchment. While the roots of the Early Natufian can surely be
traced back to the preceding cultures, the economic and social
transformation it reflects should not be underestimated. Concurrently, the EN-LN differences should not be overemphasized. The
Natufian economic transformation may be mentioned as a genuine
stage in the evolution of human diet in the latest Pleistocene
Levant, characterized by obtaining a broader spectrum of animal
resources and reflecting the rise of sedentism in this period. The
Natufian strategy, commencing in the EN and maintained in the LN,
was successfully employed for several millennia until it was gradually replaced by food production during the Pre-Pottery Neolithic
Period.
Acknowledgments
This paper is based on R.Y.’s doctoral research at the University
of Haifa, generously funded by the Graduate Studies Authority, the
Hecht Scholarship, the Wolf Foundation Scholarship and the Carmel Research Center Grant. The manuscript was written during his
Fulbright post-doctoral fellowship in the Smithsonian Institution.
We thank D. Kaufman for his help throughout this research and for
commenting on a previous draft and A. Regev for graphic assistance. The paper greatly benefitted from the helpful comments
and suggestions by the editors and two anonymous reviewers. The
renewed excavation at el-Wad Terrace is sponsored by the
Wenner-Gren Foundation, the Care Foundation and the Faculty of
Humanities, University of Haifa. Thanks are also due to the Dan
David Foundation and to Sarah and Avie Arenson for their support.
Appendix A. Supplementary data
Supplementary data related to this article can be found at http://
dx.doi.org/10.1016/j.jhevol.2014.02.011.
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Intensification and sedentism in the terminal Pleistocene Natufian

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