International Journal of Osteoarchaeology

Int. J. Osteoarchaeol. (2010)

Published online in Wiley Online Library
(wileyonlinelibrary.com) DOI: 10.1002/oa.1230

Late Epipaleolithic Hunters of the Central

Taurus: Faunal Remains From Direkli
Cave, Kahramanmaraş, Turkey
B. S. ARBUCKLE* AND C. M. EREK
Department of Anthropology, Forensic Science and Archaeology, Baylor University, Waco, TX USA

ABSTRACT Newly initiated research at Direkli cave is helping to define an initial understanding of Epipaleolithic hunter-
gatherer traditions in the central Taurus region of southern Turkey. Detailed analysis of the Direkli faunal
assemblage suggests that in the late Epipaleolithic the cave functioned as a short-term logistical camp used to
intercept wild goats (Capra aegagrus) in the high peaks around the cave, primarily in the late summer and fall.
In addition, hunters opportunistically exploited deer and a variety of other small taxa, including tortoise, in the
forested vicinity of the site. Evidence for low intensity and seasonal occupation of the cave indicates that
Epipaleolithic foragers in the region were highly mobile, utilised a wide range of resources, but primarily
scheduled use of the cave in order to exploit high ranked wild goat resources. This represents the first window
into the nature of foraging systems just prior to the emergence of agricultural economies in this important
region of Turkey. Copyright ß 2010 John Wiley & Sons, Ltd.

Key words: Animal exploitation; Capra aegagrus; Epipaleolithic; Hunter-gatherer; Taurus mountains; Turkey;
Wild goat

Introduction Olszewski & Dibble, 1993; Otte, 1998). Because of its

Newly initiated research at Direkli cave is helping to
define an initial understanding of Epipaleolithic
hunter-gatherer traditions in the central Taurus region
of southern Turkey. Because the Epipaleolithic (c. 20–
11,000 BP calibrated) represents a key period of
cultural and technological change leading up to the
development of Neolithic life-ways, documenting the
regionally specific cultural traditions involved in these
changes has been a major focus of archaeological
investigation across the Near East (e.g. Kuhn et al.,
2009; Bar-Yosef & Valla, 1991; Olszewski & Dibble,
1993; Otte, 1998).
Although the lithic industries and hunting beha-
viours characteristic of Epipaleolithic traditions of the
southern Levant and Zagros regions are fairly well
described, the Epipaleolithic of Turkey remains largely
unknown with vast areas, including most of the central
and eastern Taurus range, unexplored by modern
scientific excavations (Kuhn, 2002; Munro, 2004;
Stiner, 2005; Kartal, 2009; Sagona & Zimansky, 2009;
* Correspondence to: Department of Anthropology, Forensic Science and
Archaeology, Baylor University, Waco, TX USA.
e-mail: benjamin_arbuckle@baylor.edu
strategic and central geographic location the lack of
evidence for the central Taurus Epipaleolithic has
posed a particularly serious problem for attempts to
understand the nature of inter-regional interaction and
innovation at the end of the Pleistocene in the Near
East.
Moreover, excavations at cave sites including
Öküzini, Karain B and Üçağızlı, located along the
Turkish Mediterranean littoral have shown that
hunter–gatherer adaptations associated with terminal
Pleistocene climate change did not necessarily match
those strategies, including sedentism and subsistence
intensification, documented in better known regions
such as the southern Levant (see Munro, 2004; Atıcı,
2007, 2009). This suggests that the Epipaleolithic of
Turkey may represent a rich new source of information
concerning late Pleistocene forager adaptations and for
identifying alternate pathways for the development of
Neolithic life-ways in the Near East.
The rich faunal assemblage recovered from the new
excavations at Direkli Cave provides one avenue for
exploring the nature of late Epipaleolithic adaptations
in the central Taurus region just prior to the emergence
of agricultural economies. Here we report on the initial
results of the analysis of the faunal remains addressing a

Copyright # 2010 John Wiley & Sons, Ltd. Received 17 September 2010
Revised 21 October 2010
Accepted 5 November 2010

range of issues including the spectrum of taxa
represented at the site, assemblage formation pro-
cesses, prey selection and seasonality of the cave’s
occupation, and finally evidence for occupational
intensity at the site. These data represent the first
window into the nature of foraging systems at the
Pleistocene/Holocene boundary in this important
region of Turkey.
The site and its surroundings
Direkli cave is located in the central Taurus mountains
of southern Turkey, approximately 70 km northwest of
the city of Kahramanmaraş near the village of Döngel
(Figure 1). This small cave (12
13 m) is located at an
elevation of c. 1100 m above sea level in a tributary
valley of the Tekir river, which in turn is part of the
larger Ceyhan river drainage (for regional geology see
Karig & Kozlu, 1990; Gül, 2004).
First explored by Kökten (1960) in the mid twentieth
century, excavations at Direkli cave recommenced in
2007 under the direction of C. M. Erek. Although
Kökten reported the presence of a ‘micro-lithic
Aurignacian’ component in the cave, results from the
new excavations have so-far defined eight archae-
ological layers, with layers 3–8 representing late
Epipaleolithic, rather than Upper Paleolithic, occu-
pations. Radiocarbon dates obtained from charcoal
Figure 1. Map showing the location of sites mentioned in the
text.
Table 1. Radiocarbon dates from Direkli Cave
Sample ID Sample source
OS-79556 DM12 [D3/5] charcoal
OS-79558 DM16 [F5/7] charcoal
Beta-276742 DM08 [D3/7] charcoal
Copyright # 2010 John Wiley & Sons, Ltd.
B. S. Arbuckle and C. M. Erek
date the upper part of this occupation (layers 3–5) to
the mid eleventh to mid tenth millennia uncal bp
(c. 10,5009000 cal BC) (Table 1). The dominance of
microlithic elements, especially lunates and backed
bladelets in the lithic assemblage is similar to that seen
in the Late Natufian tradition of the southern Levant,
further supporting the dating of the upper Epipaleo-
lithic layers. Interestingly, the rarity of trapezes and
triangular microliths clearly distinguishes the lithic
assemblage at Direkli from that of the late Epipaleo-
lithic of the Turkish Mediterranean coast as seen at
Öküzini phase 5 and also the Zarzian tradition of the
Zagros region (Kuhn, 2002; Kartal, 2009; Olszewski &
Dibble, 1993). Located above the Epipaleolithic
deposits, layers 0–2 represent a mixture of medieval to
modern deposits. The presence of a thick layer of goat
dung on the cave floor indicates that the cave was used
by pastoralists throughout the second millennium AD.
Methods
The bone sample selected for analysis includes
materials from Geological Horizon II and Archae-
ological levels 4–8 and includes samples selected from
a variety of horizontal units including both the
northern and southern portions of the cave.
Recovery of faunal remains included the use of wet
sieving of all excavated sediments using nested screens
of 5, 3 and 1 mm; resulting in the capture of virtually all
osseous materials. Due to time constraints, a sampling
strategy was devised in which a portion of the
Epipaleolithic assemblage (c. 8000 specimens) was
examined in toto, in order to gather detailed zooarch-
aeological and taphonomic data with which to address
questions concerning assemblage formation processes.
Examination of a smaller portion of the assemblage
(c. 1200 specimens) focused on the recovery of
demographic, skeletal portion and metrical data and
did not include the recording of unidentified shaft frag-
ments or small (<5 mm) unidentified bone fragments.
Faunal specimens were identified to skeletal element
and genus or species whenever possible. Those
specimens that could not be placed in a detailed
taxonomic category (e.g. family, genus) were identified
Uncal bp Cal BC 1 Sigma cal BC
9660  40 9079  130 8900–9160
9500  40 8915  149 8720–9010
10,480  60 10,460  179 10,230–10,590
Int. J. Osteoarchaeol. (2010)
Late Epipaleolithic Hunters of the Central Taurus
with respect to general body-size categories (e.g. large,
medium, small, very small). Long bone shaft fragments
as well as epiphyseal ends were recorded based on the
presence of diagnostic portions including foramina and
muscle attachments (generally following Stiner, 2002).
Maximum length measurement was taken for all
recorded specimens, as was per cent completeness (i.e.
the percentage of the entire skeletal element). The
angle and outline of fracture were also recorded for all
remains of long bones in order to identify fresh versus
post-depositional fragmentation following Villa &
Mahieu (1991). Per cent completeness of shaft
circumference was recorded in order to assess the
importance of human versus carnivore access to the
assemblage as well as assemblage recovery (Bunn,
1983; Atıcı, 2006; Bar-Oz et al., 2008). We also
calculated a modified version of Marean’s (1991)
completeness index which examines the degree of
fragmentation of selected carpals, tarsals, sesamoids
and the petrous bone in order to measure the impact of
post-depositional attritional factors on assemblage
composition.
Modifications including the presence of cut and
percussion marks, rodent and carnivore gnawing, and
burning as well as the state of weathering (following
Behrensmeyer, 1978) were also systematically recorded
although in some loci secondary calcification of faunal
remains hampered the examination of surface modi-
fications.
Although number of identified specimens (NISP) is
used as the basic unit for quantifying taxonomic
abundance in the faunal assemblage, diagnostic zones
are also used in order to correct for differences among
taxa in the number and identifiability of their skeletal
elements. The diagnostic zone method defines a
standard set of skeletal elements common to most
taxonomic groups and its use is described in detail
elsewhere (Watson, 1979; Russell & Martin, 2005;
Arbuckle, 2006). In addition, specimen weight is also
used in order to estimate the contribution of various
mammalian taxa to the diet since bone weight is
roughly correlated with body mass (White, 1953;
Chaplin, 1971; Davis, 1987; Boessneck, 1992; Zeder,
1998).
Skeletal element survivorship is examined by
calculating MNE, MAU and %MAU values (Lyman,
1994). Calculation of MNE follows Binford (1984) and
is based on the greatest number of specimens from a
distinctive portion of a skeletal element (e.g. proximal
end, proximal shaft, midshaft, distal shaft, distal end)
without regard to side. MAU is a measure of the
frequency with which a skeletal portion or element is
present in an assemblage, regardless of side, and
Copyright # 2010 John Wiley & Sons, Ltd.
standardised by the number of times that part or
element is present in one complete skeleton (Binford,
1984). Standardizing MAU (%MAU) further facilitates
identification of differential survivorship or transport of
skeletal elements by comparing observed element
frequencies to their expected values (e.g. Niven et al.,
2004).
Demographic profiles for goats were generated
based on both the state of epiphyseal fusion of long
bones as well as tooth eruption and wear (Payne, 1973;
Zeder, 2006; Helmer et al., 2007). For tooth eruption
and wear we used mandibles that included at least three
cheek teeth as well as both upper and lower loose
deciduous fourth premolars and third molars to
generate survivorship curves (Arbuckle, 2006; Helmer
et al., 2007; Atıcı, 2009).
Due to high fragmentation and the subsequent small
number of measureable specimens, biometrical charac-
teristics of the goat population at Direkli were
examined using the log size index (LSI) method
(Meadow, 1999). In this method, log transformed
measurements taken from archaeological specimens
representing multiple skeletal elements are compared
with those from a standard animal; in this case the
average measurements of male and female wild goats
(C. aegagrus) from the Taurus mountains (Uerpmann &
Uerpmann, 1994).
Finally, in order to address the intensity of the
occupation of the cave we utilise the small game index
following Stiner and Munro (Stiner et al., 2000; Munro,
2003, 2004). This index focuses on the proportion of
slow and easy to capture small game (primarily
tortoise) to fast and more difficult to capture small
game (e.g. hare, birds) in an assemblage as a measure of
the duration and intensity of occupation. Based on
optimal foraging theory, this method predicts that as
occupational intensity increases (i.e. as the number of
human hours at the site per unit time increases) foragers
will be forced to exploit a wider range of lower ranked
taxa with lower energetic return rates (Stiner et al.,
2000).
The faunal assemblage
Faunal spectrum
A summary of the taxonomic and body size class
composition of the Direkli faunal assemblage is
presented in Table 2. Although nineteen mammalian
taxa were identified in the assemblage the majority of
the remains are those of medium artiodactyls, of which
wild goat (C. aegagrus) is the most abundant. Wild goats
Int. J. Osteoarchaeol. (2010)
 B. S. Arbuckle and C. M. Erek

Table 2. Taxa and mammalian size categories identified in the small taxa, which are unlikely to have been consumed
Direkli cave faunal assemblage by humans (including rodents and some small
Taxa/Size categories Common name NISP
carnivores including Martes foina), are removed, the
representation of wild goats increases to 68% (based on
Very small mammal 89 DZ) and 82% (based on weight) of the faunal
Small mammal 121 assemblage.
Medium mammal 7393
Med-large mammal 36 The abundance of this taxon is no surprise given the
Large mammal 36 location of the cave at the base of the steep rocky
Medium artiodactyl 37 slopes of the Delihöbek and Hacıveliler mountains at
Ovis/Capra 301
Ovis sp. Sheep 11 an elevation of c. 1100 m. Although wild goats are no
Capra aegagrus Wild goat 135 longer present in the region, even today, local villagers
Bovid/cervid 316 herd large numbers of goats, which are better suited to
Capreolus capreolus Roe deer 10
Dama dama Fallow deer 29 the broken, rocky environment than other taxa.
Cervus elaphus Red deer 8 Although eleven specimens from the faunal assem-
Sus scrofa Boar/pig 6 blage were identified to the genus Ovis, it is unlikely
sm carnivore 38
Med carnivore 3 that they represent wild sheep (Ovis orientalis). Based on
Large carnivore 5 limited metrical data from these specimens, it seems
Martes foina Stone Marten 20 likely that the sheep remains represent the downward
Mustela putorius Polecat 1
Meles meles Badger 1 movement of domesticates from the latest depositional
Felis sp. Wild cat 1 layers in the cave (layers 0–2) dating to the second
Vulpes vulpes Fox 4 millennium AD. Similarly, the few specimens repre-
Canis sp. Wolf/dog 2
Ursus arctos Bear 12 senting pig (Sus scrofa) are much smaller than modern
Castor fiber Beaver 3 examples of Anatolian boar and also likely represent
Rodents 47 the intrusive remains of later domesticates.
Spalax sp. Mole rat 62
Meriones sp. 1 Following goats, the next most abundant ungulate
Arvicola(?) sp. 1 taxa are deer, including roe, fallow and red deer.
Talpa sp.
Lepus capensis Hare
1
17
Although relatively few in number (NISP ¼ 49) deer
Testudo graeca Tortoise 398 represent 5% of the economically important taxa based
Snake 6 on DZ and 12% based on bone weight, making them
Fish 8 second only to goats in terms of contribution to the
Birds 78
Crustacean 1 diet. Their presence indicates the regular exploitation
9238 of lower altitude forests and valley bottoms located
directly below the cave itself.
Also notable in the Direkli faunal assemblage is the
comprise almost half of the total assemblage identified relatively high representation of tortoise, which makes
to the genus level based on both NISP and diagnostic up almost a quarter of the NISP of specimens identified
zones (DZ). The remains of goats represent 70 and to genus. When DZ are used to quantify the abundance
77.4% of the total bone weight of the assemblage of tortoise, this value drops to 8% of the fauna
indicating that this taxon was of primary economic consumed by humans and only 2% based on bone
importance (Table 3). In addition, when the remains of weight. Tortoises are commonly encountered in the
immediate vicinity of the cave today and it is possible
Table 3. Frequencies of taxonomic groups identified to the that some of the remains represent animals that died
genus level based on NISP, DZ and bone weight. ‘Other’ naturally in the cave. However, tortoises are common
includes pig, hare, beaver, rodents, fish and snakes in Epipaleolithic assemblages in the Near East (e.g.
Stiner et al., 2000; Munro, 2004; Grosman et al., 2008)
Taxa NISP NISP % DZ DZ %
(%) (weight) (%) (weight) and the presence of burning and percussion marks on a
few of the tortoise remains and the generally high
Caprines 47.4 70.0 46.6 77.4 degree of fragmentation of carapace and plastron
Deer 2.9 10.5 3.5 11.6
Carnivores 5.2 3.5 8.8 3.9 specimens suggests that they were regularly exploited
Birds 4.8 0.6 8.1 0.7 by foragers.
Tortoise 24.7 12.8 5.4 2.2 Birds are also an important component of the faunal
Other 14.9 2.7 27.6 4.2
Total N ¼ 1610 4537.4 N ¼ 740 2652.5 assemblage. Due to the lack of appropriate comparative
collections at this early stage in analysis, bird remains

Copyright # 2010 John Wiley & Sons, Ltd. Int. J. Osteoarchaeol. (2010)
Late Epipaleolithic Hunters of the Central Taurus
have been identified only to size category and include
primarily small and medium sized birds with only one
specimen falling within the size range of large birds
(e.g. great bustard/raptors). Although some of the small
birds may have been brought into the cave via
taphonomic processes, the medium and large-sized
birds were likely brought to the cave by humans,
although evidence for burning and cutmarks on bird
bone is rare (see below). Although their contribution to
the diet is underrepresented based on bone weight
measurements (less than 1%), bird remains represent
almost 12% of the economically important taxa based
on DZ.
Carnivores, primarily representing small mustelids
including predominantly the stone marten (M. foina),
make up 5% of the assemblage based on specimen
counts. Although it is possible that some of the small
carnivore remains represent the natural fauna of the
cave, one marten mandible exhibits cutmarks indicat-
ing skinning. In addition, bear remains, which are
present in small numbers, are primarily limited to distal
extremities and teeth suggesting that they were
brought to the site by humans. There is no evidence
to suggest that the cave was used as a denning site for
large carnivores.
The faunal spectrum at Direkli fits with its location at
a moderate elevation on a steep mountain slope in a
forested region. The focus on wild goat and
secondarily on deer indicates that high ranked, high
return mammalian prey located both above and below
the cave were the primary targets of Epipaleolithic
hunters. This focus on high return ungulates including
goats and deer is comparable to the situation
documented for Epipaleolithic assemblages at Uçağızlı
cave, Hatay, where wild goat and roe deer are
dominant, at Ksar Akil, Lebanon, where goat and
fallow deer were hunted, and at Karain B and Öküzini
caves, Antalya, where a combination of wild sheep,
goat and fallow deer were exploited (Kersten, 1987;
Atıcı, 2009; Kuhn et al., 2009).
Assemblage formation processes
One of the important questions to address at Direkli is
who or what was responsible for the formation of the
Table 4. Fragment completeness for small, medium and large mammals. Numbers in parentheses indicate sample size
Completeness 0–25% 25–50%
Small 60.7% (71) 8.5% (10)
Medium 97.3% (7188) 0.4% (30)
Large 88.9% (48) 3.7% (2)
Copyright # 2010 John Wiley & Sons, Ltd.
abundant faunal remains recovered in the cave. This
question can be addressed through examination of the
taxonomic composition of the assemblage as well as
multiple detailed measures of assemblage fragmenta-
tion.
In many Pleistocene cave sites large carnivores
including hyena, brown bear and wolf may have acted
as significant bone accumulators (e.g. Stiner et al.,
1996). Although hyena remains have not been
identified at Direkli, bear and wolf are present in
small numbers (NISPs of 12 and 2) as are the remains of
unidentified large carnivores (NISP ¼ 5). It has been
suggested that c. 20% carnivore is a lower limit for
identifying carnivore denning sites with many sites
exhibiting much higher frequencies (Cruz-Uribe,
1991; Niven, 2007:368). Large carnivores represent
less than 1% of the Direkli assemblage, effectively
ruling them out as significant bone accumulators.
Additional features of carnivore denning sites such as a
high frequency of carnivore gnawing and the presence
of bone ‘tubes’ are also generally lacking (see below).
The Direkli fauna assemblage is highly fragmented
and is comprised primarily of long bone shaft
fragments of medium mammals (primarily wild goat)
that have been reduced to between 20–40 mm in
length. Preservation of spongy bone and skeletal
portions with high cancellous content, such as the
proximal humerus, is very poor. Measurements of
fragment completeness indicate that 97% of medium
mammal and 88.9% of large mammal remains are less
than 25% complete (Table 4). In contrast, small
mammals exhibit a much lower degree of fragmenta-
tion suggesting that the skeletons of medium and large
mammals, the focus of human exploitation, were
intentionally and intensively processed and fragmented
in order to access within bone nutrients including
marrow. In addition, the tarsal and carpal completeness
index is relatively high (3.9 out of 5.0; n ¼ 113)
indicating a moderate to low rate of post-depositional
attrition and suggesting that much of the fragmentation
of the assemblage occurred as the result of intentional
human processing.
In order to understand the role of human agency as
well as post-depositional factors in the survivorship of
skeletal elements within the assemblage we compared
bone survivorship against several other variables
50–75% 75–100% 100%
6.0% (7) 6.0% (7) 18.8% (22)
0.2% (15) 0.6% (42) 1.1% (82)
1.9% (1) 0.0% (0) 5.6% (3)
Int. J. Osteoarchaeol. (2010)
 B. S. Arbuckle and C. M. Erek

Table 5. Measurements of skeletal element abundance for (Table 5). One major factor in bone survivorship is
goats bone mineral density (Binford & Bertram, 1977;
Skeletal elements NISP MNE MAU %MAU
Marean, 1991, 1995). In the Direkli assemblage there
is a positive and significant relationship between bone
Head–bone 36 17 8.5 0.55 density and skeletal portion survivorship (Figure 2a)
Head–teeth 175 31 15.5 1.00 indicating that density mediated attrition had a major
Mandible–bone 40 24 12.0 0.77
Mandible–teeth 231 25 12.5 0.81 impact on the composition of the assemblage. This is
Horn 25 2 1.0 0.06 evident in Figure 3, which graphically illustrates the
Atlas 0 0 0.0 0.00 differential survivorship of skeletal elements. Elements
Axis 0 0 0.0 0.00
Cervical 3-7 0 0 0.0 0.00 of the head, particularly teeth, exhibit the highest
Thoracic 9 5 0.4 0.02 survivorship followed by the metapodials, radius,
Lumbar 3 2 0.3 0.02 astragalus, humerus, tibia and first phalanx. Spongy
Sacrum 1 1 1.0 0.06
Indet vert 40 12 0.5 0.03 elements such as the vertebral column, ribs, proximal
Rib 93 12 0.5 0.03 portions of the humerus and tibia, as well as scapulae
Scapula 11 6 3.0 0.19 and femora are under-represented or completely
Humerus 56 13 6.5 0.42
Radius 56 20 10.0 0.65 absent. Moreover, goat horncores are strongly
Ulna 11 10 5.0 0.32 under-represented and are present at only 6% of the
Radial carpal 2 2 1.0 0.06 expected frequency based on the number of skull
Intermed carpal 3 3 1.5 0.10
Ulnar carpal 4 4 2.0 0.13 remains.
Accessory carpal 3 3 1.5 0.10 In addition, measures of skeletal portion abundance
Carpal 2 þ 3 9 9 4.5 0.29 were compared against Binford’s (1984) Modified
Carpal 4 6 6 3.0 0.19
Metacarpal III þ IV 31 26 13.0 0.84 General Utility index (MGUI) in order to test whether
Innominate 7 5 2.5 0.16 the patterns of element survivorship are the result of
Femur 13 7 3.5 0.23 differential transport of the most nutritious skeletal
Patella 1 1 0.5 0.03
Tibia 28 11 5.5 0.35 elements by hunters (Lyman, 1985) (Fig. 2b). No
Os malleolus 6 6 3.0 0.19 relationship was found to exist between skeletal
Astragalus 15 15 7.5 0.48 portion abundance and the MGUI suggesting that
Calcaneum 11 8 4.0 0.26
Tarsal c þ 4 9 9 4.5 0.29 entire goat carcasses were probably transported to the
Tarsal 2 þ 3 11 11 5.5 0.35 cave for processing and consumption.
Metatarsal III þ IV 31 16 8.0 0.52 Finally, we found that the marrow index, a measure
Metapodial 60 21 5.3 0.34
pxSesamoid 31 31 1.9 0.13 of the volume of the marrow cavity in long bone shafts,
dsSesamoid 5 5 0.4 0.03 is positively and significantly correlated to NISP:MNE,
Phalanx 1 62 48 6.0 0.39 an index of element fragmentation (Fig. 2c) (Bar-Oz
Phalanx 2 49 35 4.4 0.28
Phalanx 3 27 27 3.4 0.22 et al., 2008). This tells us that bones with high marrow
content (e.g. metapodials, humerus, femur) were
subject to greater fragmentation than those with low

Figure 2. Graphs comparing the linear relationships between (A) bone density and bone survivorship (Spearman’s r ¼ 0.562,
p < 0.001); (B) MGUI and bone survivorship (Spearman’s r ¼ 0.274, p > 0.05); and (C) marrow index and NISP/MNE (Spearman’s
r ¼ 0.593, p < 0.01).

Copyright # 2010 John Wiley & Sons, Ltd. Int. J. Osteoarchaeol. (2010)
Late Epipaleolithic Hunters of the Central Taurus

Figure 3. Skeletal element survivorship based on MAU(MNE).

marrow content (e.g. calcaneum), supporting the
interpretation that intensive processing for within
bone nutrients was responsible for the high degree of
long bone fragmentation.
Analysis of fracture outline and edge angle as well
as shaft circumference indicates that the majority of
breakage of medium and large mammal long bones
occurred on fresh bone (Table 6). Abundant green
bone fractures combined with poor preservation of
cancellous-rich epiphyseal ends suggests intensive
bone processing focused on marrow extraction and
possibly grease rendering as well (Bar-Oz et al., 2008;
Munro & Bar-Oz, 2005). However, the presence of a
relatively high proportion of dry breaks indicates that
post-depositional breakage, including trampling, was
also an important factor in assemblage formation.
Low rates of shaft circumference completeness also
suggest human agency as the primary cause of
fragmentation. Long bone specimens with relatively
complete circumferences, often described as ‘tubes’ and
commonly associated with carnivore generated faunal
assemblages (Binford, 1981; Atıcı, 2006) are almost
completely absent from the medium and large mammal
assemblages at Direkli. Shaft completeness is much
higher for small mammals indicating that the long
bones of small mammals were not processed and that
many of these taxa were probably not consumed by
humans.
A relatively small proportion of the assemblage
exhibited clear evidence of surface modifications, the
most common of which included cut and percussion
marks, evidence for burning and carnivore gnawing
(Table 7). Cutmarks were noted on 2.5% of small
mammal, 4.6% of medium mammal and 15.8% of large
mammal remains. Cutmarks were rarely identified on
bird and tortoise remains.
Overall, a host of evidence including the strong
relationship between marrow content and fragmenta-
tion, the low correlation between skeletal element
abundance and food utility, the abundance of green-
bone fractures and the presence of cut and percussion
marks on long bone shafts strongly suggest that
human agency was the major factor in the formation
of the faunal assemblage. Furthermore, it suggests
that wild goat skeletal remains were intensively
processed and virtually all long bones were opened
in order to consume marrow contents. This also
suggests that wild goats were harvested seasonally
when marrow quality was high (Bar-Oz & Munro,
2007) (see below).

Table 6. Frequencies of different types of fracture angle, fracture outline and shaft circumference. Numbers in parentheses indicate
sample size. Small mammals ¼ hare to rodent size; Medium mammals ¼ goat to pig size; Large mammals ¼ cattle and red deer

Fracture angle Fracture outline Shaft circumference

Obtuse or Right angle Indet. Transverse Oblique Indet. <50% >50%

acute (fresh) (dry) (dry) (fresh) complete complete

Small mammals 52.2% (24) 28.3% (13) 19.6% (9) 45.7% (21) 50.0% (23) 4.3% (2) 67.4% (31) 32.6% (15)
Medium mammals 64.0% (812) 29.4% (373) 6.6% (84) 30.7% (388) 64.8% (819) 4.4% (56) 96.3% (1187) 3.7% (46)
Large mammals 82.8% (48) 6.9% (4) 10.3% (6) 19.7% (12) 75.4% (46) 4.9% (3) 89.3% (51) 10.5% (6)

Copyright # 2010 John Wiley & Sons, Ltd. Int. J. Osteoarchaeol. (2010)
 B. S. Arbuckle and C. M. Erek

Table 7. Frequencies of surface modifications on faunal speci- Table 8. State of epiphyseal fusion for the remains of goats
mens. Numbers in parentheses indicate number of occurrences (fusion ages represent estimates for the start of the fusion
of each modification. process following Zeder (2006))

Cutmarks Burning Carnivore Fusion age Unfused Fused %Fused

gnawing (months)

Small mammals 2.5% (4) 4.8% (10) 1.0% (2) Prox radius 2 1 10 90.9
Medium mammals 4.6% (66) 4.9% (407) 0.4% (31) Scapula 6 1 1 50.0
Large mammals 15.8% (12) 4.1% (4) 1.1% (1) Distal humerus 6 1 3 75.0
Birds 1.3% (1) 1.3% (1) 0.0% (0) Prox phalanx 1 12 5 12 70.6
Tortoise 0.8% (3) 2.0% (8) 0.0% (0) Prox phalanx 2 12 7 24 77.4
Metacarpal III þ IV 18 3 1 25.0
Metatarsal III þ IV 18 11 9 45.0
Distal tibia 18 0 8 100.0
Prox ulna 30 2 0 0.0
Prey selection and seasonality Calcaneus 30 4 1 20.0
Distal radius 30 4 2 33.3
In order to address strategies of prey selection and Total 39 71 64.5
seasonality of the Epipaleolithic occupation of Direkli
cave, the age structure for goats is reconstructed based
on both epiphyseal fusion of the long bones (see
Table 8) as well as the eruption and wear of mandibular Survivorship curves for goats were generated from
and maxillary teeth (Figure 4). Because of the small size tooth eruption and wear using both mandibular
of the sample of long bones with epiphyseal ends (following Payne, 1973) and maxillary teeth (following
identified to the genus level and given that the vast Helmer et al., 2007). Both mandibular and maxillary
majority of medium mammal remains represent wild datasets produced almost identical estimates of age of
goats, we combined medium mammal, bovid/cervid, death and were combined to create a more robust
sheep/goat and goat categories in constructing age dataset. These data indicate a peak in culling animals
profiles based on epiphyseal fusion. Although sample between the ages of 6–12 months (25%) and 12–24
sizes are still quite small, they do provide enough data months (18%). Another peak in the goat cull includes
to outline the composition of the kill-off in terms of animals between the ages of 4–6 years (19%)
broad age categories. In addition, the remains of very (Figure 4). Survivorship based on teeth indicates that
young individuals allow us to provide a first estimate of c. 74% of the culled population was older than one year
the seasonality of hunters’ visits to the cave. and 55% were older than two years (figures that match

Figure 4. (A) Goat survivorship and (B) mortality based on the state of wear and eruption of mandibular and maxillary teeth (following
Helmer et al., 2007; Payne, 1973), compared with age data from the Epipaleolithic levels of Öküzini cave 3–5 and Karain B Cave (Atıcı,
2009).

Copyright # 2010 John Wiley & Sons, Ltd. Int. J. Osteoarchaeol. (2010)
Late Epipaleolithic Hunters of the Central Taurus
remarkably well with the estimates from long bone
fusion in Table 8).
Overall, dental data indicate that c. 45% of the goats
culled were juveniles, 44% were prime adults and 11%
were old adults (age estimates following Stiner, 2005).
This pattern of survivorship is similar to that seen in the
Epipaleolithic levels of Karain B and Öküzini caves
(Atıcı, 2009). However, the remains of neonatal
individuals (aged 0–2 months), which are consistently
present in the Karain B and Öküzini assemblages, are
very poorly represented at Direkli. Given identical
recovery techniques and evidence that taphonomic
processes affecting the faunal assemblages from these
three sites were generally comparable (Atıcı, 2009),
this may suggest significant differences in prey
selection and/or seasonality of hunting between Direkli
and the western Taurus sites with a greater focus on
spring and early summer hunting at the latter. In
addition, the high proportion of juvenile goats at
Direkli suggests a hunting strategy targeting demo-
graphically mixed female-kid herds rather than one
focused on selecting only the largest, prime adult
individuals (Stiner, 1990, 2006) (see below).
Examination of teeth and long bones from young
individuals provides a useful initial means to estimate
the seasonality of the occupation of the cave. These
specimens are summarised in Table 9. Using Atıcı and
Stutz’s (2002) age estimates for mandibular deciduous
fourth premolars and assuming May as the modal
birthing month (following Schaller, 1977; Kaya &
Aksoylar, 1992), eight of the nine specimens for which
wear stages could be assigned appear to have been
culled in the late summer or early fall and include
infants as well as yearlings from the previous year’s
cohort. This pattern of a fall cull is also supported by
the more course-grained tooth wear data presented in
Table 9. Estimates of season of death for wild goat based on the state of wear of mandibular deciduous fourth premolars (dp/4) and
early fusing skeletal elements. Dp/4 tooth wear and age estimates taken from Atıcı and Stutz (2002); age estimates for fusion from Zeder
(2006). Estimated season of death assumes birth in May based on modern wild caprines (Kaya & Aksoylar, 1992)
Element Wear/fusion stage Age estimate (months)
dp/4 5 2.5
dp/4 6 4.5
dp/4 6 4.5
dp/4 6 4.5
dp/4 7 4.5
dp/4 7 4.5
dp/4 9 16
dp/4 9 16
dp/4 12 16
Prox radius unfused <2
Prox radius fusing c. 2
Distal humerus unfused <6
Scapula unfused <6
Copyright # 2010 John Wiley & Sons, Ltd.
Figure 4, which indicates peaks in the goat cull at
around 6 and 18 months of age. The single outlier in
Table 9, a deciduous, lower, fourth premolar not yet in
full wear, and suggesting a mid summer cull, may be the
result of variation in the wear of this tooth or a late
season birth (Payne, 1973).
Evidence from early fusing long bones also provides
a means to identify seasonality. Two unfused and fusing
distal humeri provide age of death estimates from
summer through fall which fits well with the
seasonality estimates from tooth wear. One unfused
and one fusing proximal radius, however, derive from
infants suggesting that these two individuals were
culled in the summer.
Overall, the strongest seasonal signal in the Direkli
assemblage suggests that goat hunting was primarily
scheduled in the late summer and early fall months,
although the cave may have been used at other times as
well. It is therefore likely that Epipaleolithic hunters
scheduled their occupation of Direkli cave to maximise
encounters with wild goats, perhaps as they moved
from higher elevation summer home ranges to lower
elevation ranges for the winter months. Hunting
animals in this season would also result in harvesting
animals in good physical condition, which is supported
by the evidence for the intensive processing of long
bone shafts for marrow (Munro & Bar-Oz, 2005). The
limited evidence for a minor summer component to the
occupation of the cave may suggest that the site was
also used opportunistically whenever small herds of
goats or even single males were spotted in the region.
Due to modest sample sizes, examination of the
biometric characteristics of the Direkli goats focused
on LSI values, which represent standardised measure-
ments of archaeological specimens compared to those
of a standard animal (Meadow, 1999) (Figure 5).
Estimated season of death
August (summer)
September–October (late summer/early fall)
September–October (late summer/early fall)
September–October (late summer/early fall)
September–October (late summer/early fall)
September–October (late summer/early fall)
September (late summer)
September (late summer)
September (late summer)
June–July (summer)
July (summer)
June–November (summer-fall)
June–November (summer-fall)
Int. J. Osteoarchaeol. (2010)

Figure 5. LSI values for fused (black) and unfused (grey) wild
goat specimens from early Holocene Ganj Dareh (Hesse, 1978),
Direkli and the Epipaleolithic (Kebaran) levels of Ksar Akil (from
Kersten, 1987). Ganj Dareh measurement based on distal
breadth of the metacarpal.
Measurements clearly indicate that the Direkli goats
are large, comparable in size to modern wild goats from
the region. The Direkli goats are also similar in terms of
overall size range to early Holocene morphologically
wild goat populations from Cafer, eastern Turkey (LSI
mean ¼ 0.04) (Peters et al., 1999: Fig. 9) and Ganj
Dareh, western Iran (Hesse, 1978) (Figure 5). The
Direkli goats are noticeably smaller, however, than the
massive goats from the Kebaran levels of Ksar Akil,
Lebanon (Kersten, 1987). This perhaps reflects a
decline in body size in response to the environmental
changes associated with the transition from the
Bolling–Allerod climatic optimum to the Younger
Dryas (see also Açıkkol, 2006). The presence of several
small specimens (<0.06 on the LSI scale) may
represent the intrusive remains of a few domestic goats
that moved down into the Epipaleolithic strata.
The LSI data from Direkli are characterised by a
bimodal distribution with a major peak at 0.07 and a
minor one at 0.00 on the LSI scale. Based on the degree
of sexual dimorphism seen in modern wild goats
(Zeder, 2001) as well as early Holocene assemblages of
morphologically wild Capra from the region including
Cafer, Ganj Dareh and Aşıklı Höyük, it is likely that
these peaks represent males and females. The
dominance of larger specimens suggests that large
males were targeted for slaughter and the presence of
Copyright # 2010 John Wiley & Sons, Ltd.
B. S. Arbuckle and C. M. Erek
both fused and unfused large specimens indicates that
both immature and mature males were hunted.
Together, demographic and biometric data suggest
that Direkli hunters targeted juvenile and prime aged
adult goats in the late summer and fall months, likely in
and around the rocky slopes that rise 1000þ meters
immediately above the cave to the north and west.
Although the frequency of juveniles in the assemblage
is high, it is comparable to frequencies of juvenile
caprines from the later Epipaleolithic levels of Öküzini
cave (Atıcı, 2009), although the Direkli assemblage
includes the remains of more 6–12 month old kids and
far fewer kids less than 6 months. This high frequency
of juveniles in both western and central Taurus sites
suggests that local populations of wild goats were
under relatively heavy hunting pressure in the late
Epipaleolithic and may indicate hunting strategies
targeting juvenile rich herds (see below).
The age composition of the Direkli goats generally
falls within the demographic range of the living
structure of wild ungulate populations indicating that it
could result either from the accumulation of multiple
nonselective hunting forays, either through stalking or
ambush hunting, or from catastrophic events in which
entire herds were culled at once (Murie, 1944; Klein,
1982; Levine, 1983; Stiner, 1990). However, the
abundance of both mature and immature males and
well as the presence of females suggests that multiple
demographic groups were regularly targeted by Direkli
hunters. Since wild goats segregate themselves
spatially based on age and sex into distinctive herds
with different demographic compositions, it is likely
that ethology plays an important role in explaining
prey selection at Direkli.
For most of the year adult male and female wild goats
inhabit spatially distinct, although overlapping ranges,
with females living in larger ‘female herds’ along with
immature males. Mature males live most of the year in
smaller more mobile ‘bachelor herds’, or are solitary,
while mixed herds including both males and females of
all ages form in the winter during the rut (Schaller,
1977; Kaya & Aksoylar, 1992). Given the presence of
both adult and immature males in the Direkli
assemblage, and given seasonality evidence suggesting
the scheduling of hunting before the formation of
mixed herds in the winter, this suggests that Direkli
hunters preferentially targeted lone males or small
bachelor herds, as well as larger female herds. Given
the prevalence of juveniles in the assemblage it is
suggested that targeting these larger female herds,
which may have been easier to locate and provided a
greater return than lone males, represented a central
component of prey selection strategies.
Int. J. Osteoarchaeol. (2010)
Late Epipaleolithic Hunters of the Central Taurus
Although demographic profiles rich in juveniles and
prime adults are usually interpreted as representing
cursorial or ambush hunting of individuals (Klein,
1982; Levine, 1983), it is also possible that Direkli
hunters culled multiple individuals at once using small-
scale cooperative drives. It is entirely possible that once
a female herd was identified in the region, cooperative
drives were used to move animals either towards
waiting hunters or to constricted areas including steep-
walled box canyons or sheer rocky slopes where they
could easily be culled (Anell, 1969; Kay, 1994; Frison,
2004). Steep, rocky terrain suited for these types of
drives is abundant around the site.
Iconographic evidence from the tenth millennium
site of Göbekli Tepe (southeastern Turkey) suggests
that nets were used to immobilise wild sheep, and large
nets were also used by Paleoindian hunters in North
America for culling mountain sheep (Ovis canadensis)
(Frison et al., 1986). With evidence for cordage
extending back well into the Upper Paleolithic (Soffer
et al., 2000; Kvavadze et al., 2009) it is possible that such
‘soft technologies’, were also used by Epipaleolithic
hunters at Direkli in order to subdue and cull moderate
numbers of wild goats in single kill episodes.
Occupational intensity
The small game index was used to interpret the
intensity of the Epipaleolithic occupation of Direkli
cave. The small game index predicts that as occu-
pational intensity increases the highest ranked and
easiest to harvest resources will be depleted and
hunter–foragers will be forced to exploit harder to
catch, faster reproducing and lower ranked resources
(Stiner et al., 2000; Munro, 2004). The small game
index for Direkli indicates a high reliance on slow
moving, easy to capture, high ranked resources
(tortoise) and relatively low use of fast, harder to
capture, lower return small game such as birds and hare
(Figure 6). This, along with the small size of the cave
and the lack of well-defined architectural features,
suggests that the cave was not used as a long term,
multi-seasonal base camp.
Assemblages that have been interpreted as multi-
seasonal base-camps, such as Karain B and Öküzini 1–4
exhibit much higher frequencies of small fast game,
supporting the notion of higher occupational intensity
during periods of postglacial climate amelioration (the
Bolling–Allerod) (Atıcı, 2009) (Figure 6). During the
Younger Dryas, however, Late Natufian sites such as
Hayonim Cave, Hayonim Terrace and Hilazon Tachtit
show a marked reduction in occupational intensity as
Copyright # 2010 John Wiley & Sons, Ltd.
Figure 6. Small game index showing the frequencies of slow,
small game (tortoise) and fast small game (hare and birds) in
fifteen Near Eastern faunal assemblages. Assemblages include
Üçağızlı B (Upper Paleolithic); Karain B 1 and 2; Öküzini 1–5; El
Wad and Hayonim C (Early Natufian); Üçağızlı Epipaleolithic,
Hayonim C, Hayonim Terrace and Hilazon Tachtit (Late Natufian)
(Atıcı, 2009; Kuhn et al., 2009; Munro, 2004). Assemblages are
roughly ordered from oldest (left) to youngest (right).
foragers restructured mobility patterns in response to
changing environmental conditions (Munro, 2003,
2004).
At Direkli, the small game index is similar to that for
Late Natufian sites, suggestive of a pattern in which
small numbers of hunters scheduled use of the cave for
short periods of time when goats were seasonally
available. Their diet was supplemented with locally and
easily available resources such as tortoise and to a lesser
extent birds and fish. Thus the data from Direkli fit with
a region-wide pattern during the Younger Dryas in
which hunter–gatherers adapted to local environmen-
tal conditions through a combination of high mobility
and targeting high ranked prey, especially wild goats.
Conclusion
Detailed analysis of the Direkli faunal assemblage
suggests that in the late Epipaleolithic period the cave
functioned as short-term logistical camp designed to
intercept wild goats (C. aegagrus) in the high peaks
around the cave, primarily in the late summer and fall.
In addition, hunters opportunistically exploited deer
and a variety of other small taxa, including tortoise, in
the forested vicinity of the site. Evidence for low
intensity and seasonal occupation of the cave indicates
that Epipaleolithic foragers in the region were highly
mobile, utilised a wide range of resources, but primarily
Int. J. Osteoarchaeol. (2010)

scheduled use of the cave in order to exploit high
ranked wild goat resources.
Body part representation suggests that hunters
transported entire carcasses back to the cave, which
in turn suggests that hunting took place on the rocky
slopes to the north and east of the site. The high
frequency of the remains of juveniles and adult females
suggests that female herds may have been a primary
target of hunting parties which may have used stalking,
ambush hunting, or small-scale cooperative drives to
cull multiple individuals at one time. Furthermore, as is
common in many Epipaleolithic assemblages in the
Near East, ungulate skeletons were heavily processed
for within bone nutrients and marrow was removed and
consumed from every available long bone.
Although we are just beginning to understand the
nature of the Epipaleolithic occupation of the central
Taurus mountains, these new results from Direkli
provide an important reference point, which can be
used to further develop our understanding of terminal
Pleistocene hunter–gatherer adaptations in this pre-
viously unknown region.
Acknowledgements
Support for research at Direkli Cave has generously
been provided by the Turkish Ministry of Culture and
Tourism, the General Directorate of Monuments and
Museums; the Kahramanmaraş Archaeology Museum;
Gazi University Faculty of Arts and Sciences; and the
College of Arts and Sciences and the Department of
Anthropology at Baylor University. The authors wish
to thank these institutions for their support. The manu-
script also benefitted from close readings by two
anonymous reviewers, whose comments improved
the quality of the paper.
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