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Journal of Archaeological Science 36 (2009) 1538–1546
Contents lists available at ScienceDirect
Journal of Archaeological Science
journal homepage: http://www.elsevier.com/locate/jas
Gazelle exploitation in the early Neolithic site of Motza, Israel: the last of the
gazelle hunters in the southern Levant
Lidar Sapir-Hen a, *, Guy Bar-Oz b, Hamoudy Khalaily c, Tamar Dayan a
a
Department of Zoology, Tel Aviv University, Tel Aviv 69978, Israel
Zinman Institute of Archaeology, University of Haifa, Haifa 31905, Israel
c
Israel Antiquities Authority, POB 586, Jerusalem 91004, Israel
b
a r t i c l e i n f o
a b s t r a c t
Article history:
Received 7 October 2007
Received in revised form
15 December 2008
Accepted 9 March 2009
We studied the faunal remains from the Early and Middle PPNB site of Motza, Judean Mts., Israel, in order
to gain insight into the economic basis prior to livestock husbandry, with a focus on gazelle hunting.
Taphonomic analysis showed that bone preservation at the site was excellent. The subsistence economy
in Motza was based on a broad spectrum of hunted species, with mountain gazelle (Gazella gazella) as the
dominant prey, similar to many Epipalaeolithic and Pre-Pottery Neolithic sites in the southern Levant.
We studied gazelle exploitation patterns, in order to learn about the interaction of humans with this
species prior to ungulate domestication. Analysis of the demography of the gazelle herd, which included
aging and sexing, revealed no age preferences and no selective culling. Moreover, the PPNB gazelle
population of Motza does not exhibit allometric changes in morphology, that are allegedly correlated to
increased hunting pressure on gazelle populations prior to livestock domestication.
Ó 2009 Elsevier Ltd. All rights reserved.
Keywords:
Neolithic
EPPNB
Gazelle
Levant
Domestication
1. Introduction
The beginning of animal domestication in the early Neolithic of
the southern Levant was an influential stage in the development of
human societies. This process led to the beginning of civilization as
we know it, and changed in its intensity and consequent influences,
irreversibly, the perception of the environment by humans (BarYosef and Belfer-Cohen, 1989; Bar-Yosef and Meadow, 1995; Bellwood, 2005; Smith, 1995). The process of domestication in the
Levant has been the focus of interest for a wealth of archaeological
research, and has raised many questions concerning the reasons,
the roots and the timing of this process (examples of major publications: Bökönyi, 1974; Clutton-Brock, 1989, 1999; Davis, 1987;
Zeuner, 1963). Of special interest are the subsistence strategies of
the latest stages of hunter-gatherers, which may hold the key for
understanding the causes or the flow of events that led to ungulate
domestication.
During the Epipalaeolithic and early Neolithic cultures, prior to
the domestication of the main livestock animals, the dominant prey
species throughout the Mediterranean region of the southern
Levant was the mountain gazelle (Gazella gazella) (Bar-Oz, 2004;
Davis, 1982; Munro, 2004; Tchernov, 1993a). The dominance of
* Corresponding author. Tel.: þ972 3 6409024; fax: þ972 3 6409403.
E-mail address: [email protected] (L. Sapir-Hen).
0305-4403/$ – see front matter Ó 2009 Elsevier Ltd. All rights reserved.
doi:10.1016/j.jas.2009.03.015
gazelle in archeofaunal assemblages ended during the Early/Middle
phase of the Pre-Pottery Neolithic B (PPNB, 10,500–8250 yr BP,
calibrated), when gazelles were virtually replaced by domestic
goats and sheep that may have been introduced from northern
parts of the Levant (Bar-Yosef, 2000; Peters et al., 1999). The
dominance of gazelles in the latest hunter sites, and the fact that
the gazelle was never domesticated, generated extensive research
concerning the interaction of humans with this species, prior to
livestock husbandry (e.g., Bar-Oz, 2004; Bar-Oz et al., 1999, 2004;
Bar-Oz and Munro, 2007; Campana and Crabtree, 1990; Cope, 1991;
Davis, 1982, 1983, 1991; Horwitz et al., 1990; Legge, 1977; Munro,
2001, 2004; Munro and Bar-Oz, 2005; Tchernov, 1993a). Understanding the gazelle-based subsistence economy at this transitional
stage could be key to understanding the roots for the process of
domestication.
Several researchers (Cope, 1991; Tchernov, 1993a,b) have suggested that during the Natufian (at the final stage of the Epipaleolithic) highly selective culling of male gazelles was practiced.
This eventually led to size diminution and increased variation of
gazelle populations, a pattern that reflects some form of cultural
control of gazelles (referred to by Cope, 1991 as ‘‘proto-domestication’’). However, later studies, including a reanalysis of Cope’s
published sample statistics (Dayan and Simberloff, 1995) and an
analysis of new Natufian faunal remains (Bar-Oz et al., 2004) found
no support for the suggested scenario of cultural control of gazelle
populations.
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Despite that, the theory of ‘‘proto-domestication’’ is still
considered valid by some researchers. For example, Verhoeven
(2004) considered Cope’s theory and concludes that ‘‘the selective
culling seems to have caused a degeneration of the species, mainly
resulting in extreme size variability’’ and that ‘‘the gazelle population came under serious stress’’. Mithen (2003, pp. 47–48)
claimed that ‘‘by preferentially selecting the males, the Natufians
were probably attempting to conserve the gazelle populations’’.
Other researchers (Bar-Yosef and Meadow, 1995; Davis et al.,
1988; Davis, 2005) suggested that increasingly intense exploitation
of the environment led to the depression of gazelle populations by
hunters. This increased exploitation pressure is reflected as the
increased frequency of juvenile gazelles in archaeofaunal assemblages, alongside a ‘‘spectrum shift’’ towards exploiting smaller
animals, especially fishes and birds (Davis, 2005; Stiner et al., 1999,
2000). These events led to active human intervention and so to the
beginning of domestication of species that were amenable to this
kind of treatment, in order to compensate for the decline of prey
populations.
In recent years some compelling evidence for the increased
exploitation of low-ranked small prey was published (Davis, 2005),
as another reason for suggesting increased predation pressure on
environmental resources. However, the question of gazelle
exploitation patterns remains intriguing. Davis (2005 and references therein) and Munro (2004) reviewed studies of gazelle
exploitation, focusing on age-at-death data, from the mid-Paleolithic to the Natufian in the southern Levant. They found a gradual
increase in the culling of gazelle young, doubling from 17.3% in the
mid-Paleolithic to 34.5% in the Natufian. As younger animals are
smaller and contain limited body fat, they are considered lowranked food resources. It could be argued that as younger animals
are considered low-ranked resources, their capture should be
avoided when encounters with higher-ranked adult prey suffice
(Speth and Clark, 2006). Davis (2005) and Munro (2004) suggested
that the gradual increase in juvenile frequency is a result of the
overexploitation of gazelle population, as the increased frequency
of juvenile gazelles in the assemblage results from reduced
encounter-rates with more highly ranked adult prey. Davis (2005)
and Munro (2004) concluded that gazelle populations were hunted
in increased intensity in the Natufian, compared to the previous
cultures, leading to their depression.
Bar-Oz (2004) and Bar-Oz et al. (2004) investigated the idea of
overexploitation during the Epipalaeolithic period in the Levant,
and examined the changes in juvenile frequency between the early
Epipaleolithic (Kebaran and Geometric Kebaran) and the late Epipaleolithic (Natufian). They found that aging the gazelle population with fusion data (Davis, 1983) provide similar results to
Davis (2005) and Munro (2004). However, Bar-Oz (2004) suggested that there is a bias in these results, as dental wear survivorship curves are similar across the studied periods, and
concluded that there is no support for increased gazelle hunting
pressures during the Natufian period. Thus, research results to date
provide contradicting support for gazelle exploitation patterns
congruent with overexploitation, that is – human gradual
depression of natural resources as a driver of the shift in subsistence patterns.
Although the replacement of gazelle by domestic sheep and goat
occurred in the early Neolithic, most probably during the MPPNB,
most of the research concerning gazelle exploitation patterns to
date has focused on the Natufian culture. Faunal exploitation
patterns within the PPNA and the EPPNB remain to be investigated,
in order to gain insight into the transition to agriculture.
We studied gazelle exploitation patterns at Motza, an Early and
Middle PPNB site from the Judean Mts., 5 km from Jerusalem. Few
studies of gazelle exploitation have focused on PPN sites. Data on
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gazelle sexing and aging are not provided for PPNA Jericho in the
Jordan Valley, but Clutton-Brock (1979) stated that no osteological
evidence to suggest that gazelle were either herded or tamed was
found. For PPNA Hatoula in the Judean Hills (Davis, 1985, 2005)
a high percentage of juveniles (30%) was found, and gazelles were
not sexed. A preference for hunting of males was suggested for the
PPNA of Gilgal (Noy et al., 1980) and Netiv Hagdud (Tchernov, 1994),
both in the Jordan Valley but the faunal assemblage in these sites is
far too small to be conclusive (NISP ¼ 8 and 121, respectively). For
EPPNB Horvat Galil in the Upper Galilee (Gopher, 1997) and for
Mujahiya in the Golan Heights (Gopher, 1990), only a species list
was published and the gazelle population was not studied. A sex
preference for females (70%), and a preponderance of adults (17%
juveniles) was found in MPPNB Yiftah-el in the Lower Galilee
(Horwitz, 2003a). The gazelles from MPPNB Abu Ghosh in the
Judean Hills were not sexed, and the demographic profile showed
representation of all ages (with 20% juveniles) (Horwitz, 2003b).
Gazelles from PPNC ‘Ain Ghazal in north-eastern Jordan (von den
Driesch and Wodtke, 1997) were neither sexed nor aged. Thus, at
this point there appears no coherent pattern of deliberate herd
management of gazelles during the PPN of the southern Levant.
The large faunal assemblages from two consecutive phases in
Motza provide an excellent opportunity for in-depth study of the
Early and Middle PPNB subsistence economy and for investigating
the interaction of the latest southern Levantine hunters with their
chief prey, the gazelle. We studied patterns of gazelle hunting,
asking whether gazelle remains reflect a preference for a specific
sex or age group, and whether there were any changes in gazelle
body size or body-proportions during the early Neolithic, that could
support a hypothesis of cultural control as suggested by Cope
(1991) and others (see Section 1).
2. The site
Motza is located about 5 km west of Jerusalem and 5 km east of
the Neolithic site of Abu Ghosh. The Neolithic site of Motza was
excavated as a salvage excavation by one of the authors (H.K.) on
behalf of the Israel Antiquities Authority.
Neolithic remains were first noted in Motza in the 1920s when
artefacts including arrowheads, sickle blades and bifaces, were
collected during a cursory survey (Shalem, 1928, 1937). Forty years
later, another survey conducted by Bar-Yosef and others, explored
the southern margins of the site and recovered more Neolithic finds
(Bar-Yosef personal communication). Still both surveys did not
detect the exact location of the Neolithic occupation and it was not
until the early 1990s when Pre-Pottery Neolithic tools were found
in excavated fills of an Iron Age settlement in Area B that the site
was actually located (De Groot and Greenhut, 1996).
The first in situ Neolithic remains were discovered in 2000 when
a test trench was excavated at the southern margin of the site. This
deep sounding revealed a Middle/Late Pre-Pottery Neolithic (PPNB)
layer with no architectural features except for few constructed
small hearths (Eisenberg and Sklar, 2005). Tools, on the other hand,
were abundant, including Byblos and Amuq points, sickle blades,
bone tools, and a remarkble hafted tool (Khalaily et al., 2005).
Two PPNB strata were uncovered in two seasons of excavation
(2002, 2003): Layer V – Middle PPNB, 10,100–9600 yr BP calibrated,
and Layer VI – Early PPNB, dated to 10,500–10,100 yr BP calibrated
(Yizhaq et al., 2005).
The excavation grid was based on 1 m2, with a depth of 10 cm.
Sediment from floors was wet sieved through 1 mm mesh, and
material recovered from other areas was dry sieved through 2 mm
mesh. Zooarchaeological finds were packed in separate bags that
were labeled with the archaeological context.
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3. Methods
All animal remains recovered from the site were examined, and
identified elements were coded according to their stratigraphic
location. The complete zooarchaeological and taphonomic analysis
procedures followed Bar-Oz (2004) and Bar-Oz and Munro (2004)
research protocols. Skeletal elements were identified to the closest
possible taxonomic unit, including cranial fragments, vertebrae,
long bone articular ends and long-bone shafts. The identified bone
elements were documented by describing the specific element, its
side and the portion of the bone (e.g., proximal-distal epiphysis). In
most cases identified bone elements were coded according to their
fraction of completeness (i.e., percentage of complete bone).
Percentages of elements were used to calculate the minimum
number of skeletal elements (MNE) and the minimum number of
individuals (MNI) following Klein and Cruz-Uribe (1984), Lyman
(1994) and Grayson (1984). The number of identified specimens
(NISP) was used as a basic measure of taxonomic abundance
(assuming independence) (Grayson, 1984). Identified elements
were examined for macroscopic surface modifications using a lowresolution magnifying lamp (2.5). Modifications such as bone
weathering (Behrensmeyer, 1978), butchery marks (Binford, 1981),
and evidence of rodent gnawing, carnivore punctures and digestion, (Fisher, 1995; Lyman, 1994) were recorded. The mode of bone
fragmentation was analyzed for a sample of shaft fragments with
attached epiphyses. The morphology of the fracture angle was
assessed in order to determine the stage at which the bones were
broken (i.e., fresh vs. dry; see Villa and Mahieu, 1991). Shaft
circumference was measured to determine the role of carnivores in
assemblage formation (Bunn, 1983; Marean et al., 2004).
The age structure of the hunted gazelle population, was
analyzed on the basis of tooth eruption and wear patterns of the
lower deciduous fourth premolar (dP4) and lower third molar (M3).
The teeth were compared to modern specimens of gazelle skulls
with recorded age at death housed in the mammalian collection at
the Tel Aviv University Zoological Museum. Age structure was also
analyzed by examining the stage of epiphyseal fusion of certain
skeletal elements, according to Davis (1983). Selected post-cranial
elements were measured and compared to recent sexed specimens
from the Department of Evolution, Systematics and Ecology, at the
Hebrew University, Jerusalem. This was done in order to accomplish two goals: (1) to determine the relative abundance of males
and females in the culled gazelle population. Most measurements
of gazelles show much overlap between the sexes, with the
exception of the atlas and axis (Horwitz et al., 1990), but they are
very rare in the assemblage. We followed Davis’s (1985, 2008)
suggestion of measuring the distal humerus as a means for separation (humerus breadth of trochlea [BT] and minimum diameter or
height of trochlea [HTC] measurements). (2) To detect changes in
body size through time (scapula GLP and BG; radius Dp and Bp;
tibia Bd and Dd; pelvic acetabulum LA; 3rd molar width; following
von den Driesch, 1976).
4. The Motza faunal assemblage
A total of 7021 complete and fragmentary bones were identified
in the Early PPNB assemblage, and 913 in the Middle PPNB
assemblage (Table 1). Both assemblages are dominated by mountain gazelle (82% and 68% of total ungulates, respectively) (gazelle
NISPs and MNEs are detailed in Appendix 1). Other ungulates
represented in both assemblages include wild boar (Sus scrofa),
goat (Capra sp.) and aurochs (Bos primigenius). The goat remains
could not be identified to species with much certainty, and are
referred to as Capra sp. alone. Even so, in the Early PPNB they are so
uncommon that they may well still be wild, and the increase in
Table 1
Species abundance (NISP, MNI) of the taxa represented in Early and Middle PPNB
assemblages in Motza.
Species
Gazella gazella
Body size Gazelle
Sus scrofa
Body size Sus
Capra sp.
Ovis/Capra
Bos primigenius
Body size Bos
Dama mesapotamica
Capreolus capreolus
Vulpes vulpes
Body size Vulpesa
Felis silvestris
Martes foina
Canis lupus
Meles meles
Panthera leo
Testudo graeca
Lepus capensis
Scuirus sp.
Spalax sp.
Erinaceus europaeus
Ophisaurus apodus
Aves
Mollusca
Total
a
EPPNB
MPPNB
NISP
MNI
NISP
MNI
2881
1155
358
298
144
57
100
50
3
2
561
412
154
17
5
4
1
313
181
28
3
16
7
91
180
7021
69
/
6
/
4
/
3
/
1
1
17
/
6
5
2
1
1
24
9
10
2
3
5
/
/
289
88
60
50
51
45
18
9
/
1
54
52
18
5
3
2
/
136
7
2
/
3
/
9
11
913
7
/
3
/
4
/
1
/
/
1
4
/
2
1
1
1
/
7
2
1
/
1
/
/
/
Body size vulpes could include vulpes, lepus or felis.
their frequency by the Middle PPNB suggest the appearance of
domestic goats in the southern Levant (Davis, 1987; Horwitz, 1993).
Examining changes in the relative frequency (%NISP) of the main
ungulates (Fig. 1) shows that the relative proportion of gazelles in
the assemblage decreases in the MPPNB (although they are still the
main prey category), while the frequency of goat increases significantly, demonstrating their growing importance in the economy of
the site (c2 ¼ 17.6, p < 0.001).
5. State of bone preservation
Bone preservation is excellent. Yizhaq et al. (2005) showed that
extracted collagen samples from Early PPNB faunal remains are
comparable to modern collagen from various aspects. The exceptional preservation is also indicated in our analysis of bone surface
modifications.
In order to test the skeletal part representation of gazelles, we
grouped skeletal elements into nine carcass parts, following Stiner
(1991, 2002): horn, head, neck, axial, upper hind, upper front, lower
hind, lower front and feet. The skeletal part representation (%MAU)
in both assemblages is quite similar, with a preference for the toes,
and low representation of the axial, neck, head, and horn (Fig. 2)
and is different from the expected frequency in a complete skeleton. However, this difference is not related to density-mediated
attrition or to the economic value of body parts. We found nonsignificant relationships between bone structural density [based on
the BMD1þ2 values of Rangifer tarandus; data from Lam et al. (1999)]
and gazelle bone survivorship (%MNE): Spearman’s r ¼ 0.34,
p ¼ 0.17 for the EPPNB and r ¼ 0.29, p ¼ 0.26 for the MPPNB. In
addition, we found no meaningful relationship between gazelle
bone survivorship (%MAU) and food utility index [FUI; calculated as
the weight of usable tissue of R. tarandus; Metcalfe and Jones
(1988)]: Spearman’s r ¼ 0.40, p ¼ 0.057 for the EPPNB and r ¼ 0.32,
p ¼ 0.88 for the MPPNB. Similarly, we found no meaningful
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L. Sapir-Hen et al. / Journal of Archaeological Science 36 (2009) 1538–1546
1541
Fig. 1. Ungulates representation in Early and Middle PPNB Motza.
relationship between gazelle bone survivorship (MAU) and the
marrow index (dry marrow of gazelles; calculated by Bar-Oz and
Munro (2007)): spearman’s r ¼ 0.21, p ¼ 0.61 for the EPPNB and
spearman’s r ¼ 0.5, p ¼ 0.2 for the MPPNB. Thus it appears that
selective transport of body parts according to their economic value
was also not a major factor in the formation of Motza PPNB
assemblages.
All examined bones (NISP ¼ 3560) from both assemblages were
recorded in weathering stages 0–1, of the six stages of Behrensmeyer (1978). No signs of animal activity (Fisher, 1995), such as
carnivore gnawing or chewing were recorded. It seems that both
PPNB assemblages were not altered appreciably by any pre- or
post-burial attritional processes. The morphology of the fracture
angle was assessed following Villa and Mahieu (1991); the high
representation of bones with an oblique fracture angle in the
EPPNB assemblage (>92%; NISP ¼ 280) suggests that most bone
destruction occurred when the bones were fresh, most probably
while processing the bones for consumption and as part of marrow
extraction processes. Conversely, the EPPNB bone assemblage
contains very low proportions of bones with right fracture angle
(NISP ¼ 17), further demonstrating the negligible role of in situ
bone attrition and destruction.
Butchery marks were found on very few specimens, mostly on
the remains of gazelles (0.9% and 0.35% of the gazelles in EPPNB and
MPPNB, respectively) and aurochs. All gazelle butchery marks that
could be assigned to a specific prey-handling stage were made
during the dismembering or skinning of the carcass, according to
the typology described by Binford (1981) (Table 2). Burning signs
were recorded on 20% and 16% of gazelle bones in the EPPNB and
MPPNB, respectively.
6. Demographic composition of Motza gazelles
Demographic composition of Motza gazelles was studied for the
EPPNB assemblage alone, due to the small sample size of the
MPPNB assemblage. Age classes of gazelle from EPPNB Motza were
estimated for 28 specimens based upon the eruption and wear of
the deciduous fourth premolar (dP4) and lower third molar (M3),
compared to modern aged gazelle mandibles. The percentage of
individuals (%NISP) under the age of 12 months in the Motza
assemblage is 21.4%, and under the age of 18 months is 28% [age
classes, grouped to stages following Payne (1973), are detailed in
Table 3]. The survivorship curve obtained (Fig. 3) does not resemble
a catastrophic curve, nor does it seem to represent culling of
a specific age group.
We also determined age distributions by examining the stage of
epiphyseal fusion (Davis, 1980) of certain skeletal elements,
according to Davis (1983): distal radius, distal metapodial, distal
femur, distal tibia, and calcaneus. The percentage of individuals
(%NISP) under the age of 15 months in the Motza EPPNB assemblage, according to this method, is 19.5%, which closely resembles
the dental profile under the age of 12 months. The frequency of
young individuals under the age of 18 months in the archaeological
herd of Motza, is not significantly different than that of a modern
Fig. 2. Skeletal part representation of gazelle from Early and Middle PPNB Motza pooled into nine carcass parts.
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Table 2
Butchery marks in Early and Middle PPNB Motza and the activities which they may
be associated with (following Binford, 1981).
EPPNB
Species
Bone
Stage
N
Gazella gazella
Capra sp.
Astragal
Axis
Dist humerus
Pelvis
1st Phalanx
3rd Phalanx
Scapula
Prox femur
Calcaneus
1st Phalanx
2nd Phalanx
Scapula
Thoracic vertebra
Dist humerus
1st Phalanx
Dist humerus
Dismembering
Dismembering
Dismembering
Dismembering
Skinning
Skinning
Dismembering
Dismembering
Dismembering
Skinning
Skinning
Dismembering
Dismembering
Dismembering
Skinning
Dismembering
4
1
7
4
3
1
6
1
1
1
1
1
1
1
3
1
Gazella gazella
Sus scrofa
Dist humerus
Astragal
Dismembering
Dismembering
1
1
Bos primigenius
Sus scrofa
MPPNB
gazelle herd of 35% (Baharav, 1974) (c2 ¼ 0.41, p ¼ 0.52). Thus
hunting was not selective for a specific age group.
Sexual dimorphism in gazelles is most pronounced in the atlas
and axis (Horwitz et al., 1990). Unfortunately, these bones are rarely
represented in faunal remains, and are not complete enough to be
measured in the Motza assemblage. The distal epiphysis of the
humerus, which shows some morphological separation of the sexes
(Davis, 1977, 1985), is very common and often complete in the
EPPNB assemblage in Motza. A comparison between measurements
taken from 50 individuals of the archaeological herd, and
measurements taken from a recent sample with equal numbers of
adult males and females from the Mediterranean region of
northern Israel (18 of each sex), does not reveal a significant
difference for the BT (breadth of trochlea) measurements (Student’s
t-test, p ¼ 0.53, t ¼ 0.62), but does so for the HTC (minimum
diameter or height of trochlea) (p ¼ 0.04, t ¼ 2.08) (Fig. 4). Two very
small measurements are extreme outliers that possibly represent
juveniles. Upon their removal from the analysis, the significant
difference for the HTC measurements is lost (p ¼ 0.09, t ¼ 1.71). This
comparison reveals that Motza gazelles are not different in their
size from a modern gazelle herd with an equal proportion of males
and females from the Mediterranean region of northern Israel (see
below).
A discriminant function analysis provides a function that enables
the discrimination between the sexes (Sokal and Rohlf, 1995). A
stepwise discriminant analysis conducted on the recent specimens
shows that the BT and HTC measurements are both good indicators
Fig. 3. Survivorship curve of gazelle from the Early PPNB assemblage according to
dental wear rate of the deciduous lower fourth premolar (dP4) and lower third molar
(M3).
for the separation between the sexes (BT: p < 0.001, F ¼ 28.141; HTC:
p < 0.001, F ¼ 19.195), and that they are strongly correlated (Pearson
correlation ¼ 0.778, p < 0.0001). The BT measurement alone is
enough to enable discrimination between the sexes, and the HTC
measurement does not change the results (F ¼ 0.752). The function
provided by the analysis is y ¼ 311.035 þ 24.75 BT for females
and y ¼ 349.853 þ 26.256 BT for males. The cut-off point
calculated from these functions is BT ¼ 25.74 mm. Although there is
some overlap between the sexes, and there is no certain ‘‘clean cut’’
point to separate between the sexes, we consider this method fairly
indicative for studying sex ratio. Measurements greater than this are
males and smaller than that value are females. Twenty-eight specimens from the archaeological herd are smaller (females), while 22
are larger (males). The ratio of sexes in the archaeological herd from
EPPNB Motza is not different from the ratio of a modern herd
(Baharav, 1974), of 81 males to 100 females (c2 ¼ 0.01, p ¼ 0.92).
Thus we found no sex or age preferences in hunting in the EPPNB
assemblage of Motza. It must be noted, of course, that this
comparison of a modern herd to an archaeological herd is based
upon the assumption that herd demography in prehistoric times is
not different from that of today.
Another way of sexing gazelles is by examining the shape of
horn cores (Horwitz et al., 1990), in which there is a clear distinction
between males and females.
We identified 25 male and 4 female horn cores (NISP) in the
EPPNB assemblage. The male horn cores were mostly broken, but
the female horn cores were complete. This observation suggests
that when the female horns are broken, their remains are too small
and fragile to be identified (see also Bar-Oz et al., 1999, 2004; von
den Driesch and Wodtke, 1997; Horwitz, 2003a). Moreover, a pair of
male horn cores (counted as NISP ¼ 2) was found in relation to
a human burial, and this find suggests that male horn cores were
deliberately saved or collected while female horns might have been
discarded. Therefore, we do not consider horn cores as a good way
of inferring sex ratios in our study.
Table 3
Age classes (after Payne (1973)) of gazelle from EPPNB Motza according to the
eruption and wear stages of the deciduous lower fourth premolar (dP4) and lower
third molar (M3), based on modern specimens of gazelle skulls with recorded age at
death housed in the mammalian collection at the Tel Aviv University Zoologiacl
Museum.
Mandible stage
Age
EPPNB Motza
A
B
C
D
E
F
G
H
I
0–2 mo
2–6 mo
6–12 mo
1–2 yr
2–3 yr
3–4 yr
4–6 yr
6–8 yr
8–10 yr
0
3
3
7
4
2
7
0
2
Fig. 4. Scatterplot of distal humerus measurments (BT versus HTC) of recently sexed
gazelle and fossil gazelle from Earlly PPNB Motza.
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L. Sapir-Hen et al. / Journal of Archaeological Science 36 (2009) 1538–1546
7. Morphometric analysis of Motza gazelles
Possible morphometric changes in gazelle body size or proportions through time can bias the results of the sexing method we
used (Ducos and Horwitz, 1998). Body size changes can result from
selective forces such as coevolution with competitors (Brown and
Wilson, 1956), which can lead to non-random size distributions of
the different species (Dayan et al., 1989, 1990, 1992; Dayan and
Simberloff, 2005), or from climate change in accordance with
Bergman’s (1847) rule (Davis, 1981; Dayan et al., 1991). Another
cause for change in body size or proportions is human-induced
selection, as appears in the process of domestication (CluttonBrock, 1981). The comparison of various skeletal elements from
EPPNB Motza with modern gazelles yielded no significant differences (Table 4, Fig. 5); there was no change in gazelle body size or
skeletal proportions, from the EPPNB in Motza to the present.
A comparison of gazelle body size from EPPNB Motza to the
nearby site of MPNNB Abu Ghosh (measurements were taken by
the first author) reveals no significant difference between the
humerus trochlea (BT) measurements (t ¼ 1.74, p ¼ 0.08) (Table 5).
Moreover, gazelle body size in EPPNB Motza is not significantly
different from the Natufian of el-Wad Terrace, as measured by BarOz et al. (2004). Comparison of the breadth of the humerus trochlea
(BT) reveals no size difference (t ¼ 1.6, p ¼ 0.11) (Table 5). Gazelle
body size in the el-Wad Terrace is significantly bigger than in sites
from the Kebaran and Geometric Kebaran (Bar-Oz, 2004), hence the
gazelle population of Motza maintains the relatively large body size
distribution achieved during the Natufian (Davis et al., 1994), and
does not support the proposed idea of ‘‘proto-domestication’’,
reflected by pronounced dwarfing (Cope, 1991).
8. Discussion
We studied the subsistence economy of the inhabitants of early
Neolithic Motza in order to gain insight into practices of gazelle
hunting, prior to livestock husbandry in the Southern Levant. We
found little evidence for taphonomic biases in the faunal assemblage. It seems that the assemblage formation was not affected by
post-depositional attrition processes. Among possible pre-depositional agents, the main cause for bone fragmentation is marrow
extraction. However, we found no evidence for selective transport
of body parts in relation to the caloric value or marrow yields.
The subsistence economy in PPNB Motza seems to continue in
the Natufian tradition with gazelle as the dominant prey, as found
in sites from the Epipalaeolithic (see review in Bar-Oz, 2004). The
Natufian focus on gazelle hunting has provoked intensive research
concerning exploitation patterns, and their possible implications,
regarding the advent of early domestication. Analysis of the large
assemblage of gazelle remains from the Motza EPPNB layer reveals
that the demographic composition of the archaeological herd, as
Table 4
Student’s t-test analysis results for comparison of measurements from EPPNB Motza
gazelles with recent gazelles.
Element
Measurement
Student’s t-test
Scapula GF
GLP
BG
t ¼ 0.17, P ¼ 0.86
t ¼ 0.42, P ¼ 0.66
Radius
Dp
Bp
t ¼ 0.36, P ¼ 0.71
t ¼ 1.92, P ¼ 0.06
Tibia
Bd
Dd
t ¼ 1.78, P ¼ 0.08
t ¼ 1.21, P ¼ 0.23
Pelvic acetabulum
LA
t ¼ 0.28, P ¼ 0.77
3rd Molar
Width
t ¼ 0.09, P ¼ 0.92
1543
reflected in sex ratios and age distribution, is not significantly
different from that of a modern gazelle population (see Baharav,
1974). This comparison to a modern gazelle population is based on
the assumption that the herd age and sex profile was not different
in earlier periods than today. Humans have a long history in this
region and have influenced both prey and predator populations. In
the past century leopards went extinct in the north of Israel and
wolves have been confined to the Golan Heights for some decades,
although they are currently expanding back into the Galilee. These
species could prey upon adult gazelles. Jackals, major predators of
gazelle young, are currently very common (in fact subject to culling
by conservation authorities) but their populations have fluctuated
in the second half of the 20th century in response to large scale
poisoning. It could be argued that these fluctuations may have
affected the proportion of gazelle young. The data to test this
hypothesis are not in existence, but we note that the 1974 data
ante-dates jackal poisoning by a decade, yet pre-dates their serious
population eruptions by a similar time frame, so use of these
published data make sense as an approximation of ‘natural’
conditions. Additionally, morphometric analysis of different gazelle
body parts of Motza EPPNB shows no change in gazelle body part
proportions, and does not support any hypothesis of proportions
change resulting from cultural control.
A mortality profile composition of an archaeological herd,
resembling that of a living herd in nature, may stem from several
causes and is not trivial to explain as hunting methods differ
depending on the sex and age of the prey animal. Speth and Clark
(2006) suggested that a juvenile frequency that resembles the
living herd would imply selective hunting, as hunters will avoid
taking young animals if they have a choice. Campana and Crabtree
(1990, 1991) suggested that this could be the result of communal
hunting. However, mountain gazelle herds are composed of several
social units of different age and sex, and not of one big group
containing all herd members (Martin, 2000; Mendelssohn et al.,
1995), so hunters were not likely to catch an entire herd together
(see further discussion in Edwards, 1991). Moreover, the assemblage of EPPNB Motza reflects many hunting events along 400
years. Averaging those events could lead to a demographic
composition not different from a living herd that reflects merely
long-term random hunting. This hunting pattern (or, in fact, lack of
pattern) was also found in sites from the previous Epipalaeolithic
period (see discussion in Bar-Oz, 2004). The information concerning early Neolithic gazelle exploitation patterns is very poor (e.g.
Clutton-Brock, 1979; Noy et al., 1980; Tchernov, 1994), but along
with the new information from Motza presented here, no pattern
for selective culling in this period is revealed.
Although there is much interest in the relationship between
gazelles and humans in the Epipalaeolithic, some researchers claim
that the social structure of gazelles make them unsuitable for
a close relationship with humans (Clutton-Brock, 1999; Simmons
and Ilany, 1975–1977; Edwards, 1991). Moreover, Manor and Saltz
(2003) showed that increased human presence has an adverse
impact on gazelle activity, social structure and population performance. These observations cast doubt upon the possibility of
keeping a gazelle herd in captivity for a long time, a difficulty that
later-stage hunters, with years of experience and knowledge of
gazelle behavior in nature, must have been aware of.
The analysis of the EPPNB gazelles from Motza does not reflect
increased hunting pressure from the Natufian to the early Neolithic,
causing changes in the demographic composition or allometric
changes. Therefore, it does not support the proposed idea of ‘‘protodomestication’’, which favored hunting of male gazelles (Cope,
1991; Mithen, 2003). It also does not provide support for the idea of
overexploitation of gazelle populations, in the sense of increased
representation of juveniles.
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L. Sapir-Hen et al. / Journal of Archaeological Science 36 (2009) 1538–1546
Fig. 5. Measurements of recent gazelle and EPPNB Motza gazelles: (a) GLP vs. BG scapula; (b) Bd vs. Dd tibia; (c) Dp vs. Bp radius; (d) LA pelvic acetabulum; (e) width lower third molar.
At the transition to MPPNB Motza, we see the beginning of
a change in the subsistence economy. The frequency of gazelle
remains decreases in the MPPNB, while frequencies of the goat,
boar and aurochs increase. Nevertheless, gazelles are still the
dominant prey species in the MPPNB assemblage; since the goat
remains occur in a low frequency; they are most probably still wild.
Alternatively, they may represent a primitive domestic breed,
possibly imported into the southern Levant (Bar-Yosef, 2000; Peters
et al., 1999). The increase in goat frequencies is perceived as the first
step of herd management (e.g. Clutton-Brock, 1999; Davis, 1987;
Horwitz, 1993; Tchernov, 1993a,b), which will eventually lead to
Table 5
Range, mean, S.D. and results of student’s t-test for measured Humerus BT from
EPPNB Motza vs. Natufian el-Wad terrace and MPPNB Abu Ghosh.
Motza EPPNB
el-Was terrace Natufian
Abu Ghosh MPPNB
Range
Mean
S.D.
N
Student’s t test
23.45–28.43
22–26.4
24.2–28.13
25.51
24.97
26.02
1.15
1.13
1.16
50
15
23
t ¼ 1.6, p ¼ 0.11
t ¼ 1.74, p ¼ 0.08
their full domestication. It could be argued that presence of
domestic livestock in EPPNB Motza may cause a relaxation of
pressure on wild resources, as reflected in the low juveniles
frequency of the gazelles. However, as the goats in EPPNB Motza
comprise only 4% of the ungulates, the likelihood of their affecting
the gazelle exploitation patterns is very low.
In the past years, much research focusing on the Natufian was
aimed at finding the precursor of animal domestication in the
southern Levant. If we assume that the process of goat and sheep
domestication took place in the Taurus-Zagros, where these animals
proliferated, and not in the southern Levant (Hesse, 1984; Legge,
1996; and see review in Bar-Yosef, 2000 and Zeder, 2006), then the
drivers for the process of ungulate domestication should be sought
elsewhere. The central question that should be asked in the
southern Levant is actually concerned with new economic patterns
and what enabled people to adopt them: whether the adoption of
domestication by the inhabitants of the southern Levant was
a reaction to food stress, or was it a cultural issue? It is perhaps not
surprising that we find so little evidence for overexploitation or
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L. Sapir-Hen et al. / Journal of Archaeological Science 36 (2009) 1538–1546
specialized hunting patterns of the chief prey ungulate in the
Natufian. Based on current data on the appearance of domesticated
sheep and goats in archeological sites, it appears that the processes
leading to early domestication (or the adoption of domesticates)
should be studied in periods later then the Natufian. Studies published so far concerning faunal remains of the Pre-Pottery Neolithic
are at present too scarce to allow a comparative study.
Our study provides data concerning gazelle exploitation in the
EPPNB. It adds to information derived from mostly earlier sites,
demonstrating no pattern in gazelle hunting practices from the
Epipalaeolithic throughout the early Neolithic [although earlier
changes do occur (Munro, 2004)]. If caprines were domesticated in
the Taurus-Zagros region, perhaps evidence for food stress and
overexploitation of the environment, mainly gazelles, should be
sought in hunting patterns in sites from that region.
Acknowledgments
We thank R. Rabinovich and T. Shariv for their help with the
reference collections under their care; A. Haber, I. Hershkovitz, L.K.
Horwitz, and N. Munro for helpful conversations; L.K. Horwitz for
the opportunity to measure gazelle bones from Abu Ghosh; A.
Landsman for his assistance. The MS benefited from comments
made by two anonymous reviewers. We thank the Israeli ministry
of Science, Culture & Sport for supporting the National collections
of natural history at Tel Aviv University as a biodiversity, environment and agriculture knowledge center. The research was funded
by the Israel Antiquities Authority, and in part by the Israel Science
Foundation (Grant 147/04).
Appendix 1. NISP and MNE of gazelle bone elements in Early
and Middle PPNB Motza.
EPPNB
MPPNB
NISP
MNE
NISP
MNE
Head
Horn
Mandible
Maxila
43
87
51
24
24
15
0
14
4
0
5
1
Body
Atlas
Axis
Cervical
Thoracic
Lumbar
Caudal
Sacrum
Sternum
Rib
22
48
174
290
324
44
15
5
344
16
19
88
120
149
44
8
2
80
1
4
9
23
32
5
0
0
20
1
3
6
9
20
5
0
0
6
Forelimb
Scapula-GF
Scapula-blade
Humerus-prox
Humerus-dist
Humerus-shaft
Radius-prox
Radius-dist
Radius-shaft
Ulna-prox
Metacarpus-prox
Metacarpus-shaft
127
32
50
134
22
56
29
29
63
49
9
112
10
34
112
/
49
29
/
50
33
/
11
3
4
14
5
5
4
8
8
7
4
10
1
3
10
/
4
4
/
7
5
/
Hindlimb
Pelvis-ilium
Pelvis-ischium
Pelvis-pubis
58
86
83
52
74
79
6
5
5
5
5
5
1545
Appendix 1 (continued)
EPPNB
Pelvis-acetabulum
Femur-prox
Femur-dist
Femur-shaft
Tibia-prox
Tibia-dist
Tibia-shaft
Patella
Astragalus
Calcaneum
Navicular cuboid
Metatarsus-prox
Metatarsus-shaft
Toes
Phalanx 1
Phalanx 2
Phalanx 3
Metapod-dist
Metapod-shaft
MPPNB
NISP
MNE
NISP
MNE
52
111
127
25
60
53
28
23
97
92
29
51
7
39
51
87
/
49
42
/
23
90
74
28
26
/
6
6
3
5
4
9
4
1
8
11
4
10
0
5
3
3
/
4
8
/
1
7
8
4
9
/
248
214
165
184
25
334
232
180
122
/
49
29
23
8
0
41
28
22
6
/
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