Adaptions in subsistence strategy to environment changes across the Younger Dryas – Early Holocene boundary at Körtiktepe, Southeastern Turkey

Abstract

The site of Körtiktepe in southeastern Turkey is one of few sites in the Upper Mesopotamia basin that attests continuous, permanent occupation across the boundary from end of the colder, drier Younger Dryas (YD) into the comparatively wetter and warmer Early Holocene (EH). This allows for the study of the degree of environmental change experienced on a local level over this boundary as well as for the study of the adaptations that the occupants of the site undertook in response to these changes. The mammal assemblage of Körtiktepe remains relatively stable across the YD – EH transition with the
main contributors to diet being mouflon (Ovis orientalis) and red deer (Cervus elaphus) in approximately the same quantities, although the contribution of aurochs (Bos primigenius) increases in the EH. The most significant changes can be seen in the shift in avifauna remains, with a sharp increase of waterbirds during the EH. It is proposed that these shifts reflect changes in the local environment with an increase in woodland cover as well as expansion of local waterways, which is generally consistent with previously published archaeobotanical studies. In terms of species exploited, mortality profiles as well as size
distribution of mammals, a great deal of continuity is observed. This suggests that over this particular period the local impact of the beginning of the Early Holocene was not overly dramatic, allowing for cultural continuity of previously established subsistence strategies.

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https://doi.org/10.1177/09596836221074030
The Holocene
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Introduction
Excavations in the last decades have revealed the importance
of the Upper Mesopotamian basin in understanding the Epipa-
laeolithic-Neolithic transformation that took place in south-
west Asia during the first half of the 10th millennium BC. The
transition from the Late Pleistocene to Early Holocene saw the
beginning of numerous lasting changes in socio-cultural
behaviour including an explosion in visible symbolic iconog-
raphy and the movement to a plant cultivation and livestock
husbandry-based subsistence system. Whilst research into
early farming communities in the Near East was for many
decades concentrated in the Southern Levant, the finding of
sites such as Çayönü Tepesi in the 1960s (Braidwood et al.,
1969, 1971), followed by a wealth of recent work as part of
rescue excavations preceding the construction of numerous
hydroelectric dams in southeastern Turkey has revealed the
extent and nature of early settled communities in this region
(Figure 1). The excavation of Pre-Pottery Neolithic (PPN)
sites such as Nevali Çori (Hauptmann, 1988, 1999, 2011) and
Hallan Çemi in the 1990s (Rosenberg and Davis, 1992; Rosen-
berg et al., 1995), as well as more recently: Hasankeyf Höyük
(Hongo et al., 2019; Miyake et al., 2012), Gusir Höyük (Karul,
2011, 2020), as well as Körtiktepe (Benz et al., 2015; Özkaya
and Coşkun, 2011; Rössner et al., 2018), has demonstrated,
contrary to the previous models, that year-round settlements
indeed preceded the cultivation of plants and the management
of animals (Braidwood and Braidwood, 1953; Childe, 1951:
282; Kenyon, 1965: 44). Archaeozoological work has recently
turned its attention to understanding numerous subsistence
strategies employed by hunter-gatherers at the end of the Ter-
minal Pleistocene and beginning of the Early Holocene in
order to better understand the varied circumstances prior to the
transition to an agriculturally based economy as well as to bet-
ter understand local histories and trajectories (Arbuckle, 2014;
Arbuckle and Erek, 2012; Atici, 2009). One of the many fer-
vent debates that remains in the discussion of the transition to
control over animal and plant reproduction is the role of
Adaptions in subsistence strategy to
environment changes across the Younger
Dryas - Early Holocene boundary at
Körtiktepe, Southeastern Turkey
Stephanie Emra,1 Marion Benz,2 Abu B Siddiq3
and Vecihi Özkaya4
Abstract
The site of Körtiktepe in southeastern Turkey is one of few sites in the Upper Mesopotamia basin that attests continuous, permanent occupation across
the boundary from end of the colder, drier Younger Dryas (YD) into the comparatively wetter and warmer Early Holocene (EH). This allows for the study
of the degree of environmental change experienced on a local level over this boundary as well as for the study of the adaptations that the occupants of
the site undertook in response to these changes. The mammal assemblage of Körtiktepe remains relatively stable across the YD – EH transition with the
main contributors to diet being mouflon (Ovis orientalis) and red deer (Cervus elaphus) in approximately the same quantities, although the contribution of
aurochs (Bos primigenius) increases in the EH. The most significant changes can be seen in the shift in avifauna remains, with a sharp increase of waterbirds
during the EH. It is proposed that these shifts reflect changes in the local environment with an increase in woodland cover as well as expansion of local
waterways, which is generally consistent with previously published archaeobotanical studies. In terms of species exploited, mortality profiles as well as size
distribution of mammals, a great deal of continuity is observed. This suggests that over this particular period the local impact of the beginning of the Early
Holocene was not overly dramatic, allowing for cultural continuity of previously established subsistence strategies.
Keywords
climate change, Early holocene, Körtiktepe, Pre-Pottery Neolithic, subsistence strategies, Younger Dryas
Received 8 September 2021; revised manuscript accepted 13 December 2021
1 Institut für Paläoanatomie, Domestikationsforschung und Geschichte
der Tiermedizin, Ludwig-Maximilians-Universität, Germany
2 Institut für Vorderasiatische Archäologie, University of Freiburg,
Germany
3Department of Anthropology, Mardin Artuklu University, Turkey
4Department of Archaeology, Dicle University, Turkey
Corresponding author:
Stephanie Emra, Institut für Paläoanatomie, Domestikationsforschung
und Geschichte der Tiermedizin, Ludwig-Maximilians-Universität,
Kaulbachstrasse 37/III, Munich 80539, Germany.
Email: stephanie.emra@palaeo.vetmed.uni-muenchen.de
1074030HOL0010.1177/09596836221074030The HoloceneEmra et al.
research-article2022
Research Paper

2 The Holocene 00(0)
climatic change, namely the onset of the Early Holocene (EH)
climatic amelioration (from approximately 11,590 BP/9,640
BC) following the Younger Dryas (YD), a period of worldwide
abrupt climate change characterised by cooler and dryer
weather conditions (Friedrich et al., 2004: 1120). The end of
the Younger Dryas brought with it an increase in average tem-
perature and rainfall which resulted in profound changes in
vegetation and waterways that led to local changes in plant and
animal ecology (Roberts et al., 2001: 732; Willcox et al., 2009:
152).
Körtiktepe (alongside Hallan Çemi) is the earliest site in Ana-
tolia that appears to be occupied year-round and is one of few sites
excavated in this region that attests what appears to be continuous
occupation across the Pleistocene-Early Holocene boundary,
allowing for the study of the effects of this period of climate
change (Benz et al., 2012a: 302, 2016: 231). Presently, only Qer-
mez Dere in northern Iraq has published faunal results spanning
this time period, and whilst showing shifts of emphasis across its
phases, remains relatively stable in taxonomic composition across
its occupation (Watkins et al., 1989, Dobney et al., 1999: 49, Fig
6.2, 55). Additionally, there are the newly found sites of Boncuklu
Tarla, which is currently awaiting further excavations and results
from zooarchaeological analyses, (Kodaş, 2019a: XII; Kodaş,
2019b: 7), as well as Çemka Höyük which is currently unexca-
vated and without precise published dating information (Kodaş
et al., 2020), both sites being located in the Upper Tigris basin of
southeastern Turkey. The site of Hallan Çemi in the foothills of the
Taurus mountains, also within the Tigris River basin, has also
yielded radiocarbon dates from the end of the 11th millennium BP
(uncalibrated) that would correspond to the last part of the Younger
Dryas (Rosenberg, 1994; Rosenberg et al., 1998: 27). However,
the earliest radiocarbon dates display high standard deviations,
and the bedrock of the site was never reached in excavation, so the
dates of the earliest occupation layers remain uncertain, although
it is considered that settlement would have already started in the
terminal Pleistocene (Rosenberg et al., 1998: 28) (For discussion
see: exoriente.org/associated_projects/ppnd_site.php?s=26).
Zooarchaeological results from Hallan Çemi have in any case
largely been published altogether, without chronological differen-
tiation (Starkovich and Stiner, 2009: 49; Rosenberg et al., 1998:
32–33). The exception is Zeder and Spitzer (2016) who differenti-
ated between different building levels, with building level (BL) 3
being the earliest, radiocarbon dated to the very end of the 11th
millennium BC/ beginning of the 10th millennium BC (Higham
et al., 2007: S22–S23; Zeder and Spitzer, 2016: 142, tab.1). As
such, Körtiktepe offers a rare opportunity to investigate the trans-
formation in cultural and subsistence practices that took place at
the Pleistocene-Holocene boundary within a single location. Zoo-
archaeological study across this boundary aims to understand the
degree of continuity over the occupation of the site, the manner
this climate amelioration impacted on the local fauna, and the
response of the inhabitants of Körtiktepe to these potential changes
in their local environment and associated animal resources.
Körtiktepe
Körtiktepe (previously published as Körtik Tepe) (37°48′51.90″N,
40°59′02.02″E) is a tell settlement located in southeastern Anatolia
within the province of Diyarbakır, Turkey. The mound rises
approximately 5 m above the plain with an area of around
100 × 150 m (Özkaya and Coşkun, 2011: 90). Radiocarbon dating
suggests that the site has been continuously occupied since at least
10,400/10,200 cal. BC, the Final Pleistocene, into the Early Holo-
cene, dated to 9,600–9,250 cal. BC (Benz et al., 2012a: 293, tab.1,
2015; Coşkun et al., 2012: 28).
The site is situated by the confluence of the Batman Çayı and
the River Tigris, within a plain on the cusp of the foothills of the
Taurus Mountains. The settlement was likely situated much closer
to the Batman Çayı than it is today as well as another palaeochan-
nel previously located west of the site, placing the settlement on a
kind of peninsula (Benz et al., 2015: 14). Körtiktepe was identified
as an aceramic prehistoric site and excavated as part of rescue
excavations carried out ahead of the building of the Ilısu Dam and
hydroelectric power plant (Özkaya and Coşkun, 2011: 90). From
Figure 1. Map of Pre-Pottery Neolithic sites mentioned in text.

Emra et al. 3
2000 onwards, excavations were directed by Vecihi Özkaya of
Dicle University (Özkaya and Coşkun, 2011: 90) and between
2009 and 2012, as well as in 2014 and 2015 were joined by Ger-
man co-operation partners from the Universities of Freiburg,
Tübingen and Mainz (Benz et al., 2015: 11; Rössner et al., 2018:
17). In 2010 to 2012, epipalaeolithic occupation layers were
reached in some deep soundings of the site (Benz et al., 2015: 11).
Detailed reports of the stratigraphy and architecture of the Younger
Dryas levels of Körtiktepe have been previously published (Benz
et al., 2012b, 2015, 2017; Coşkun et al., 2012).
The buildings excavated at Körtiktepe fall broadly into three
categories, round-planned buildings of approximately 2.3–3 m in
diameter, smaller structures of approximately 1.1–2.1 m in diame-
ter and a small number of larger buildings with a diameter of 3.4–
3.8 m (Özkaya and Coşkun, 2011: 91–92). These have been
interpreted as likely representing respectively: dwellings, storage
units, and potentially a form of public or ‘special’ building (Özkaya
and Coşkun, 2011: 91–92). Already in the Younger Dryas levels
occupation is considered to be year-round with a great deal of
building tradition showing continuity into the Early Holocene,
where it becomes more sophisticated, and the site more densely
occupied (Coşkun et al., 2012: 30; Schreiber et al., 2014: 14).
Especially of note amongst the artefactual finds are over 500
examples of carved stone vessels, many of which display incised
geometric patterns and depictions of plants and animals (Özkaya
and Coşkun, 2011: 96; Özkaya and Siddiq, 2020: 46).
One of the most remarkable aspects of the site is the impressive
quantity of interments that has been uncovered, around 2000 sin-
gle and double burials (Özkaya and Siddiq, 2020: 45). Körtiktepe
contains one of the largest sets of burial data available in the Pre-
Pottery Neolithic Near East, allowing for detailed analysis regard-
ing burial practices, bioarchaeological examination, and grave
good analysis (Benz et al., 2016, 2017; Erdal, 2015; Özkaya and
Coşkun, 2011: 93). A range of burial customs have been observed
including skeletons having been plastered or coloured, with a
small percentage showing cutmarks (Erdal, 2015: 7–8).
As Younger Dryas levels were not reached at Körtiktepe until
2011, previous initial reports and short summaries of the faunal
assemblage have related exclusively to the Early Holocene
(Arbuckle and Özkaya, 2006; Özkaya and Coşkun, 2011: 100–
102; Özkaya et al., 2011). Archaeobotanical analysis has revealed
the changing character of the local environment, as well as subsis-
tence strategies, between the Younger Dryas and Early Holocene
occupation levels (Rössner et al., 2018). The charcoal remains
from the Younger Dryas levels largely reflect the local riverine
environment (Rössner et al., 2018: Tab.5, 23) whereas the other
plant materials show the wider landscape as being dominated by
small-seeded grasses, suggesting a relatively open landscape
(Rössner et al., 2018: 22). The Early Holocene assemblage, how-
ever, shows a shift away from small-seeded cereals towards a more
open oak-woodland environment, although the riverine taxa would
have still contributed significantly to the plant spectrum (Rössner
et al., 2018: 25–26). Possible small-scale cultivation of legumes
and cereals is suggested for the Early Holocene due to their slight
increased importance, the presence of increased quantities of
chaff, as well as cultivation-associated weed flora in the case of the
cereals (Rössner et al., 2018: 25–26). As such, the zooarchaeologi-
cal findings of the study, in combination with the archaeobotanical
results from the site, will be of great value in considering the
impact on the local ecology of these Early Neolithic settlements at
the end of the Pleistocene and in considering the responding adap-
tations in subsistence strategy and social organisation.
Methodology
The vast majority of the faunal remains presented in this study
were excavated in the seasons of 2014 and 2015. Only well-strat-
ified contexts were selected for zooarchaeological identification.
Flotation took place on a limited number of contexts that would
have been excavated after 2009 (mesh size: 0.2 mm), and addi-
tionally numerous microfauna including quantities of small fish
remains were recovered from the bone-washing process. Sieving
was conducted on contexts associated with graves with sieve
sizes of 7–8 mm in addition to sieves 3.5 mm in size. This will
naturally bias the results against smaller elements and taxa.
However, the collection methods for both Younger Dryas and
Early Holocene contexts were the same, so any observed species
change is not due to change in methodology.
Identification was carried out with aid of a modern reference
collection housed at the Institute for Paläoanatomie, Domestika-
tionsforschung und Geschichte der Tiermedizin at the Ludwig
Maximilian University of Munich. Identification of avian remains
was conducted using the reference collection of the Staatssam-
mlung für Anthropologie und Paläoanatomie München (SAPM).
Multiple bones judged to be coming from the same individual
were collectively given a number of identified specimens (NISP)
of one. Antler fragments were not included within the statistics as
the specimens may have been collected shed. Material was
recorded using Ossobook, a specialist database for zooarchaeo-
logical findings (Kaltenthaler et al., 2021). Where possible all
information relating to taphonomy, ageing or pathology were
recorded. Measurements followed von Den Driesch (1976). Dif-
ferentiation of mouflon and wild goat was based on published lit-
erature (Boessneck et al., 1964; Halstead et al., 2002; Zeder and
Lapham, 2010; Zeder and Pilaar, 2010).
Tooth eruption and mandibular wear stages (Grant, 1982;
Munro et al., 2009; Payne, 1973) were recorded, but due to the
small sample sizes, are not presented in this study. It was not pos-
sible to construct mortality profiles based on teeth wear and erup-
tion stages for any taxa due to low sample sizes of mandibular
teeth, likely resulting from differential transportation of carcass
portions. As such, mortality profiles have been reconstructed using
post-cranial epiphyseal fusion data, which is only intended to sug-
gest general mortality trends. For caprines, the epiphyseal
sequence and approximate age of fusion follows Zeder (2006)
based on a modern collection. The percentage of epiphyses fused
is expressed for each age category, independent of one other. For
the construction of the Körtiktepe mortality profiles for mouflon,
both elements positively identified as mouflon, as well as those
identified as being mouflon/wild goat (Ovis orientalis/Capra
aegagrus) were used. This approach is aimed to increase the sam-
ple size of elements used, as well as to mitigate the bias towards
elements where distinction between Ovis and Capra are more eas-
ily made (e.g. humerus, coxa, metapodials). Additionally, some
morphological features used for distinguishing between Ovis and
Capra only develop with age, this then allows for the inclusion of
these younger specimens where this has not yet developed. It is
considered that despite the potential occasional inclusion of wild
goat into the mortality profile of mouflon through this method, the
overall trends should still be valid as mouflon appears to outnum-
ber wild goat considerably in the assemblage (sheep-to-goat ratio:
YD: 28: 1, EH 5: 1). This was also done in the case of the mortality
profile constructed for Göbekli Tepe, as no wild goats were found
within the assemblage, so all undetermined caprines are most
likely mouflon, but in the cases of Gusir Höyük and Gürcütepe II
and III, only elements identified as being mouflon were used due
to a larger percentage of the assemblage being wild goat.
The use of epiphyseal fusion for red deer (Cervus elaphus) is,
however, more problematic than in the case of caprines, with few
studies looking into the timing of fusion of the skeleton (Calderón
et al., 2019; Habermehl, 1985; Mariezcurrena and Altuna, 1983).
The problem is compounded by the relatively wide range of ages
when particular epiphyses fuse, which is for instance affected by
sex, with females generally fusing at an earlier age than males
(Calderón et al., 2019: 205). As such, the fusion age categories of
red deer have been collapsed into three phases, being: epiphyses

4 The Holocene 00(0)
fused by 20 months, those at 36 months and those above 48 months.
The elements assigned to each of these categories are summarised
in Appendix 1. Individuals under 36 months can be considered
juvenile (Calderón et al., 2019: 207).
Due to limited sample sizes, biometric data have been pre-
sented using the logarithmic size index (LSI) method, comparing
archaeological data against standard modern individuals
(Meadow, 1999). The standard animal used for comparison with
mouflon is a female published in Uerpmann and Uerpmann
(1994). Red deer measurements were compared to a male stan-
dard from Upper Bavaria kept in the Staatssammlung für
Anthropologie und Paläoanatomie München (SAPM) (SAPM-
MA-01295) (Appendix 2). The standard used to compare aurochs
is based on a Pinzgau cow from Austria from the same collection
(SAPM-MA-01259), published in Manhart (1998): table 103).
Measurements for wild goat (Capra aegagrus) and wild boar
(Sus scrofa) were too few to present LSI information, but the raw
measurements are presented in Appendixes 3–11. In the case of
the red deer remains from the Körtiktepe Early Holocene con-
texts, a number of outlying, smaller values were generated from
measurements from the scapula. It is considered that this is due
to the scapula being an early fusing bone, which then continues
some appositional growth, so despite being fused, may be still
subadult (Habermehl, 1985: 36–37; Mariezcurrena and Altuna,
1983: 184, Tab. 54). Due to this possibility, values generated
from scapula measurements have not been used for either the
Younger Dryas or Early Holocene phases. Measurements from
the scapula were included in the Gusir Höyük data, the LSI val-
ues being 0.14 and 0.02 (still fusing), well within the general
expected range.
In order to assess the impact of the Early Holocene climatic
transformation on the taxonomic diversity or heterogeneity of the
mammalian assemblage the Shannon entropy, or ‘archaeobiodi-
versity’ (aD), was calculated (for aD methodology see Pöllath
et al., 2008: 65). Heterogeneity in this context is meaning the mea-
sure of both richness (the number of taxa present or NTAXA) as
well as the ‘evenness’, how evenly the quantity of different taxa
are distributed (Lyman, 2008: 175,192). High heterogeneity is cor-
related with taxonomic richness and evenness, making the identity
of any one random individual difficult to predict (Lyman, 2008:
176–177). Archaeobiodiversity was calculated considering mam-
malian and avian faunal material separately. Within the mammalia,
microfauna (smaller than hedgehog) was excluded and caprines
were considered together.
Where possible, the results from Körtiktepe were compared
with other zooarchaeological datasets from PPN sites in the region,
those being: Gusir Höyük, Göbekli Tepe, and Gürcütepe II and III.
These sites were selected for comparison due to the availability of
data (including measurements, sexing, and epiphyseal data) and
comparable recording methodologies.
Gusir Höyük is located beside Gusir lake near the
intersection with the Tigris and Botan rivers, in the foothills of
the Taurus mountains and, with the currently excavated remains
dating to the Early Holocene, although as yet the earliest
remains have yet to be excavated (Karul, 2011: 1–2,
2020: 78–79). Preliminary zooarchaeological results from
Gusir Höyük provide an example for comparison of subsistence
strategies from a nearby, near-contemporary community
(Kabukcu et al., 2021; Neuberger et al., 2019). Göbekli Tepe is
located approximately 15km NE of Şanlıurfa on top of a lime-
stone ridge rising from the Harran Plain (Schmidt, 1995: 9,
2000: 45). The site consists of a number of buildings, the oldest
and largest having rings of T-shaped pillars gathered around
two central pillars, many of which are decorated with carvings
of animals (Peters and Schmidt, 2004; Schmidt, 2006, 2011,
2012). The site dates to the PPNA through to the middle of the
PPNB, but as the majority of the faunal remains come from the
infilling of the large buildings, the dating of these remains can-
not be certain (Dietrich, 2011). Gürcütepe consists of a series of
hills in the Harran plain, 4km SE of Şanlıurfa with settlements
II and III dating to the late PPNB (Beile-Bohn et al., 1998: 5,
13–14; Schmidt, 1995: 9). This site provides a comparison with
an assemblage considered to consist of largely domestic ani-
mals (von Den Driesch and Peters, 1999: 32). The first zooar-
chaeological data published from Göbekli Tepe and Gürcütepe
can be found in von Den Driesch and Peters (1999). Subsequent
publications on the archaeozoology of Göbekli Tepe can be
found in Peters and Schmidt (2004), Peters et al. (2005) and
Pöllath et al. (2018).
The faunal assemblage
Taphonomy
A total NISP of 21,783 was recorded from the Younger Dryas
contexts with a weight of 27.8 kg, of which 2950 (27.2 kg)
could be identified to a size-category or better taxonomic
level. From the Early Holocene contexts, a total NISP of
21,936 (49.1 kg) was recorded, with 4018 (47.7 kg) being iden-
tified to a size-category or better level. Overall, the remains
were well preserved, with frequent finds of more fragile bones
such as bird elements. However, despite the excellent bone
surface preservation, the remains were brittle and frequently
showed modern breaks, which inhibited, in particular, the tak-
ing of measurements. This brittle nature is seen similarly in
both the Younger Dryas and Early Holocene contexts with
86.3% from Younger Dryas contexts being unidentified mam-
mal remains and 81.5% in the Early Holocene. Bone surface
preservation was also similarly good in both phases (Table 1).
Percentage of elements with calcareous deposits on the surface
of the bone is, however, higher in the Early Holocene assem-
blage, which is disproportionately higher on remains of
aurochs likely effecting the ability to identify surface modifi-
cations (Table 2). The higher percentage of calcareous deposits
found on EH remains may be due to the level of the water table
at the site. Both rodent and carnivore gnawing is low in both
phases and elements with some degree of burning being a little
higher in the Younger Dryas phases (Table 2). The good condi-
tion of the bones, as well as frequent finds of bones from the
same context that could be re-articulated, suggests that the
material was rapidly buried after deposition resulting in mini-
mal attritional effects. A small proportion of remains, that are
considered to be intrusive, are from domestic species, as deter-
mined by their extremely small size (Appendix 12). These
intrusive specimens were found in various contexts across the
site, at various depths, and were introduced likely as the result
of animal or root action. A potential source of these specimens
is the medieval remains that are also present on site, and some-
times disturbed the Neolithic layers (Özkaya and Coşkun,
2011: 90).
Mammals
The mammal remains from both the Younger Dryas and Early
Holocene contexts are summarised in Table 3, the full species list
can be found in Appendix 12. In Appendix 12, where a species was
recorded as, for example, ‘probably Ovis aries’, this has been
combined with the category ‘Ovis aries’.
Table 1. Percentage of level of bone surface preservation of
mammal bones (without unidentified mammal remains) by phase
(YD: n = 2509, EH: n = 3123).
Good Medium Poor
Younger Dryas 62.3 33.8 3.9
Early Holocene 60.8 28.3 10.9

Emra et al. 5
In both the YD and EH phases the most abundant taxon by
NISP is red deer, although if it is assumed that the vast majority
of the undetermined caprine remains are belonging to mouflon,
then in both phases the number of red deer and mouflon remains
are found in similar proportions (Table 3). The main changes
between the two phases are a marked increase in the importance
of aurochs as well as a minor increase in the wild boar (Sus
scrofa) in the EH. The increased importance of aurochs is par-
ticularly seen when considering the relative weight of the
remains which is of a near-negligible amount in the Younger
Dryas (2.7%), rising to over one third of the identified mammal
remains in the Early Holocene contexts. In both phases mouflon
significantly outnumber the finds of wild goat (Capra aegagrus)
with the YD having a mouflon to wild goat ratio of approxi-
mately 28: 1 and the EH 5 : 1, although the number of wild goat
may be too low to consider the change in ratio between phases
to be particularly meaningful.
Size reconstruction
Red deer. The size of red deer remains relatively stable across
the late Pleistocene/Early Holocene boundary with the YD
having a median LSI value of 0.08 and the EH 0.07 and a mode
of 0.09, 0.08 and 0.08 respectively. However, whilst the upper
end of the LSI range is the same for both the YD and EH data
(0.15), a number of smaller individuals are present in the EH
(Figure 2). The Younger Dryas data appears to show a dimor-
phic distribution with a peak that would correspond to the
males at 0.08 and 0.09 (Figure 2). A second, smaller peak is
also perhaps seen at 0.05 which may represent a lesser number
of females being hunted. In the Younger Dryas there is a clear
pattern of more males being hunted, whereas in the Early
Holocene there is a slight shift towards an increasing propor-
tion of females being represented in the assemblage. In the
Early Holocene data, the same larger peak at 0.09 is also seen,
perhaps representing the males, and a second peak being at
around 0.06, although it is not as clear as in the Younger Dryas
data. Too few remains were able to be sexed in order to be used
to help reconstruct sex-specific patterns, with the coxae from
the YD levels numbering 1 male, 6 female and 2 probable
females. In the Early Holocene 4 male were identified, 6
female and 1 probable female. The LSI data from Körtiktepe
was compared with the nearby late PPNA site Gusir Höyük.
The overall LSI size range exhibited is similar in both sites,
having a median LSI value of 0.07 and the mode for Gusir
Höyük being 0.07, against 0.08 in Early Holocene Körtiktepe.
Gusir Höyük has a more unimodal distribution than Körtik-
tepe, which may be the result of different hunting strategies,
Table 2. Percentage of mammal bones with taphonomic modifications (without unidentified mammal remains) by phase and species (YD:
2509, EH: n = 3123).
Percentage with calcareous
deposits (%)
Percentage carnivore
gnawed (%)
Percentage rodent
gnawed (%)
Percentage
burnt (%)
Younger Dryas
Bos primigenius 9.1
Ovis orientalis 0.8
Cervus elaphus 2.5
Total mammal 1.1 0.0 0.2 9.2
Early Holocene
Bos primigenius 36.4
Ovis orientalis 11.3
Cervus elaphus 12.8
Total mammal 10.0 0.1 0.1 3.3
Table 3. Identified mammal taxa from Körtiktepe.
Younger Dryas Early Holocene
NISP Weight (g) NISP (%) Weight (%) NISP Weight (g) NISP (%) Weight (%)
Bos primigenius 21 684.0 1.8 2.7 203 14,528.5 13.3 33.9
Ovis orientalis 304 3904.6 26.5 15.7 151 2829.2 9.9 6.6
Capra aegagrus 11 197.1 1.0 0.8 29 878.4 1.9 2.0
Ovis orientalis/Capra aegagrus 191 2046.2 16.6 8.2 350 4310.3 22.9 10.1
Gazella subgutturosa 16 151.7 1.4 0.6 35 360.6 2.3 0.8
Cervus elaphus 476 16,242.1 41.4 65.2 534 16315.6 34.9 38.0
Dama dama 2 68.0 0.2 0.3 8 192.5 0.5 0.4
Cervidae 68 775.4 5.9 3.1 99 790.7 6.5 1.8
Sus scrofa 15 473.1 1.3 1.9 63 1751.1 4.1 4.1
Equus sp. 2 85.0 0.2 0.3 13 734.7 0.9 1.7
Canis lupus/Canis sp. 2 26.6 0.2 0.1 2 19.0 0.1 0.0
Vulpes vulpes 9 23.5 0.8 0.1 11 51.1 0.7 0.1
Felis silvestris 0 0.0 0.0 0.0 6 19.1 0.4 0.0
Lepus capensis 14 23.0 1.2 0.1 14 9.9 0.9 0.0
Castor fiber 7 149.1 0.6 0.6 2 27.3 0.1 0.1
Rodent 5 0.1 0.4 0.0 4 0.7 0.3 0.0
Other mammal 6 46.8 0.5 0.2 5 64.7 0.3 0.2
1149 24,896.2 1529 42,883.3

6 The Holocene 00(0)
perhaps with Gusir Höyük having less of an emphasis on large
males. Too few measurements for red deer were available from
Göbekli Tepe and Gürcütepe II and III for comparison.
Mouflon. In both the Younger Dryas and Early Holocene con-
texts mouflon at Körtiktepe have a median LSI value of 0.07
and mode of 0.08. Both data sets show an upper peak at 0.08,
likely representing the males (Figure 3). In the Younger Dryas
a peak perhaps representing the females appears to be around
0.04 and 0.05, and whilst less convincingly present in the Early
Holocene, there also appears to be this second peak at around
0.05. Whilst generally there is a lot of continuity between the
Figure 2. LSI values for red deer (Cervus elaphus) from YD and EH Körtiktepe and Gusir Höyük. The black line and arrow represent the
median value.
Figure 3. LSI values for mouflon (Ovis orientalis) from YD and EH Körtiktepe, Gusir Höyük, Göbekli Tepe and Gürcütepe II and III. The
black line and arrow represent the median value.

Emra et al. 7
two periods, there is perhaps a perceptible shift towards more
females represented in the Early Holocene. Sexed skeletal ele-
ments are only few in number but show an even split between
the sexed with the YD having five males, five females and one
that was probably female. The EH levels likewise show a near
even split with five females and four males. The size ranges
from Körtiktepe fits well with the size ranges observed at the
late PPNA site of Gusir Höyük, the site having a median of 0.06
and mode of 0.08, with 0.08 also likely being the upper peak,
representing rams (Figure 3). A similar range is also expressed
at Göbekli Tepe, although there appears to be more females
than males, a tendency also expressed in the sexed coxae with
10 elements identified as being female to six being male. As
such, the median and mode are much lower both being 0.06.
Whether the emphasis on female mouflon from Göbekli Tepe is
the result of targetted hunting, or from the remains representing
early animal management is a topic for future study. In com-
parison, at LPPNB Gürcütepe II and III, where the majority of
Ovis sp. are from domesticated animals, the median and mode
are again considerably lower at 0.01 and 0.02. The size of mou-
flon at Körtiktepe fits well with the broadly contemporary
Round-Grill building subphases at Çayönü and at PPNA Hasan-
keyf (von Den Driesch and Peters, 1999: 28, tab.4; Hongo
et al., 2005: 117, fig.3; Hongo et al., 2019: 165, Fig.7a). Grad-
ual size diminution at Çayönü in sheep is seen from the Chan-
nelled subphase onwards with a definitive shift in the Large
Room subphase (LPPNB) representing the beginnings of cap-
rine husbandry at the site (Hongo et al., 2005: 116–117, 120).
As such, the sizes of mouflon of Körtiktepe are clearly in the
size ranges of the wild animals of the region, with no indication
yet of the size diminution evidenced at other sites indicative of
animal management taking place.
Aurochs. Due to the small number of aurochs (Bos primige-
nius) remains identified from the Younger Dryas levels only
one measurement was available for LSI which gave the value
(0.00), putting it on the lower end of the expected size range.
From the Early Holocene contexts, one value was considerably
smaller than would be expected (−0.04, from a fused calcaneus)
and so is presumed to be intrusive cattle and removed from
considerations. Other than this specimen, the aurochs at Körtik-
tepe display a wide variation in size from an LSI value of 0.01–
00.22. Overall, Early Holocene Körtiktepe, Gusir Höyük and
Göbekli Tepe show relatively similar sizes with the medians
being 0.11, 0.13 and 0.10 respectively. Körtiktepe, Gusir Höyük
and Göbekli Tepe all have a mode of 0.10 (Figure 4). Although
only a small number of measurements are available from the
previously published PPNA round building phase of Çayönü
(n = 9), the general size also appears to be similar, having a
median of 0.12 (Öksüz, 2000: 162, tab.1). This is compared
against LPPNB Gürcütepe II and III, by which time it is consid-
ered that cattle have come under human control (Peters et al.,
2017: 9). The size range is, however, roughly similar to that of
the wild animals, also with a median of 0.10, which has previ-
ously been attributed to a lack of genetic isolation of the man-
aged animals (von Den Driesch and Peters, 1999: 30). Sexed
elements from Gürcütepe suggest a predominance of females
with two male and nine female coxae being found, with a gen-
eral shift chronologically towards more females being inter-
preted as a potential result of a degree of human control (Peters
et al., 2017: 9). No sexed elements were available from Körtik-
tepe, so whilst the domestication status cannot be indepen-
dently determined from either size or sex ratio, from what is
known in the region, it is considered that these specimens rep-
resent wild individuals.
Mortality profiles
Red deer. Whilst only a rough picture can be gained from the
mortality profiles construction from post-cranial information,
generally in both the Younger Dryas and Early Holocene
phases, relatively few juvenile individuals are represented, with
the majority making it past 4 years, suggesting the targetting of
prime-aged individuals (Figure 5). An attritional or
Figure 4. Box plot of LSI values for Aurochs (Bos primigenius) from the Early Holocene contexts of Körtiktepe and Gusir Höyük, Göbekli
Tepe as well as the LSI values of Bos taurus and Bos taurus/Bos primigenius from Gürcütepe II and III.

8 The Holocene 00(0)
ambush-hunting model would more closely resemble the living
structure, with a greater emphasis on younger individuals than
that has been exhibited here (Stiner, 1990: 323). These general
trends are found in both the Younger Dryas and Early Holocene
levels, indicating that hunting strategies did not greatly change
over this transition. At Gusir Höyük few young individuals are
represented, and few also made it past 4 years of age, suggest-
ing the targetting of young adult individuals. The LSI measure-
ments from Gusir Höyük, as well as the sexed elements
(excluding antler), suggests a slight emphasis on the hunting of
females (male: 6, female: 10). Too few red deer remains from
Göbekli Tepe and Gürcütepe were recovered in order to provide
a comparison.
Mouflon. In both phases of Körtiktepe, relatively few infantile
individuals were identified with instead there being steady attri-
tion of the age classes, until in both cases only just over half are
making it into maturity (Figure 6). Generally, with caprines,
after a high mortality of lambs, survivorship of yearlings and
older age-classes is high (Hesse, 1982: 407). The significant
proportion of sub-adults here suggests that nursery herds may
have been targetted by hunters. LSI data suggests that amongst
adult animals, males were predominantly targetted. As such
these mortality profiles might be the result of the hunting of a
mixture of both nursery herds that would contain females and
juveniles as well as either bachelor herds or lone males. As in
the case of red deer, the overall profile remains relatively similar
Figure 5. Mortality profile for red deer based on post-cranial fusion. Körtiktepe Younger Dryas (n = 192), Körtiktepe Early Holocene (n = 162)
and Gusir Höyük Early Holocene (n = 234) ‘Percentage fused (%)’ refers to the proportion of elements that would fuse at that age group, it is
independent of other age groups. The elements included in each epiphyseal group are given in Appendix 1.
Figure 6. Mortality profiles for combined mouflon (Ovis orientalis) based on post-cranial fusion (Zeder, 2006: 107, fig.15) Körtiktepe Younger
Dryas (n = 118), Körtiktepe Early Holocene (n = 149) Gusir Höyuk Early Holocene (n = 359), Göbekli Tepe (n = 363) and Gürcütepe (n = 347).
‘Percentage fused (%)’ refers to the proportion of elements that would fuse at that age group, it is independent of other age groups.

Emra et al. 9
in both the YD and EH levels. At Gusir Höyük, survivorship of
sub-adults is even higher, but there a significant drop-off by
3–4 years, suggesting the targetting of adult individuals. As sug-
gested by the LSI results, sexed elements (mostly coxae, but
also based on sexed atlas and axis vertebrae and horncores) also
show that males were particularly targetted, with over double
the number of males identified than females (male: 67, female:
28, probably male: 1, probably female: 1). This suggests that the
hunters of Gusir Höyük particularly targetted bachelor herds
and/or lone males. Göbekli Tepe also exhibits a similar pattern
with the main targets being individuals around 2 years and as
well as adults above 3–4 years. Unlike in the other two sites
however, the sexed elements suggest a similar ratio of males and
females (male: 15, female: 19). In contrast, the post-cranial
remains from LPPNB Gürcütepe II and III show a high survi-
vorship into maturity and sexed elements shows a slight prefer-
ence of females (male: 91, female: 112). Previously published
mortality profiles constructed using tooth wear data from Gür-
cütepe II, however, indicates that the majority did not reach
2 years of age (Peters et al., 2017, 8: 16.2). This discrepancy in
results between the tooth wear and epiphyseal fusion data maybe
be due to taphonomic considerations such as the role of dogs on
the site. Dogs are likely to preferentially select the softer ele-
ments of juveniles, particularly the meatier post-cranial ele-
ments, and carry them off-site for consumption. The consumption
of domestic animals by dogs at Gürcütepe has already been
strongly indicated by isotopic data that suggests that dogs con-
sumed remains from animals that were pastured with C4-plants
(Grupe and Peters, 2011: 74). This in opposition to the signature
demonstrated by the wild animals of the site that were feeding
exclusively on C3 plants, such as cereals (Grupe and Peters,
2011: 74). The preferential destruction of juvenile post-cranial
elements by dogs may lead to this result where the mortality
profiles constructed by tooth-wear or post-cranial fusion may
widely differ. With this in mind, post-cranial fusion-based mor-
tality profiles for all sites should be considered cautiously.
Nonetheless, from what can be considered, both phases at Kör-
tiktepe suggest an emphasis on male adults, indicating that the
assemblages are the result of hunting rather than the early man-
agement of mouflon.
Wild goat. In both the Younger Dryas and Early Holocene, the
sample size for wild goat was too small to make anything but very
general observations about the slaughter profile. In both cases very
few juveniles were found (Appendix 13). Little sexing information
is available; one wild goat was sexed as female from the Younger
Dryas, and one female, one probable female and four males were
sexed from the Early Holocene contexts. The small numbers of
goat and slight tendencies towards older and male individuals per-
haps suggests that these findings are the result of hunting of lone
males or small ram bands.
Aurochs. No fusion information was available for the Younger
Dryas due to limited sample size. The remains from the Early
Holocene only gave a small dataset to consider mortality profiles
for aurochs, which is presented in Appendix 14.
Wild boar. In the case of wild boar, there were again too few spec-
imens from the Younger Dryas to consider age of death, and only
a small number from the Early Holocene, these are summarised in
Appendix 15. A number of quite young individuals are presented
including one ulna from a neonate. Little sexing information is
available, with just a find of a mandible and a lower canine, both
from males. Dominance of an assemblage of juveniles does not
necessarily reflect any sort of human management and instead
may reflect a natural kill-off pattern from hunting (see discussion
on the Hallan Çemi boar remains (Lemoine, 2012; Peters et al.,
1999: 40–41; Starkovich and Stiner, 2009: 56).
Avifaunal remains
Bird remains from the Younger Dryas and Early Holocene levels
are summarised in Figure 7, with the full species list found in
Appendix 12. In both phases the Great Bustard (Otis tarda) is the
most commonly found species, comprising around one third of
the assemblages. Between the Younger Dryas and Early Holocene
there is, however, a major shift towards the exploitation of water-
birds, which increases from 20.3% in the YD to 59.6% of the EH
bird assemblage. This is largely driven by the Greylag goose
(Anser anser), which increases from 3.8% to 27.1%. In contrast,
in the Younger Dryas layers there is more of an emphasis on rap-
tors which makes up 26.6% of the YD remains, as opposed to
6.0% in the EH. Despite the smaller assemblage size of bird
remains coming from the YD, the range of bird species repre-
sented is wider with 24 different species identified compared to
19 in the EH.
Figure 7. Overview of avifaunal remains from Körtiktepe.

10 The Holocene 00(0)
Seasonality
Seasonality has been exclusively addressed through the presence of
avian species, due to a lack of suitable data from the mammalian
assemblage. Due to a general lack of cranial elements being found
at Körtiktepe, addressing seasonality through mammalian tooth
wear (as per. Gourichon, 2004) is not possible. There are also very
few neonate specimens found on site (one neonate bone from the
YD level and one from the EH, both being caprine humeri) and so
shows some evidence for winter/spring hunting activities. How-
ever, the lack of neonate remains might be easily explained by a
lack of targetting of very young individuals, or a lack of survival of
young bones due to a variety of taphonomic reasons. Additionally,
at this time isotopic data for the animals of Körtiktepe has not been
published, so seasonality cannot be addressed through oxygen and
strontium isotopic signatures (e.g. Lang et al., 2013).
Avifaunal assemblages can help establish the seasonality of
archaeological sites by considering the migratory habits of the vari-
ous species present. This is a particularly important issue to consider
for this period of time as Körtiktepe is considered to be one of the
earliest sedentary sites in the region (Özkaya, 2009: 3–4; Schreiber
et al., 2014). The geographical limits of ancient populations of birds
as well as migratory patterns are in most cases based on what is
known about the species’ modern habits. In a number of cases how-
ever, modern migratory patterns and population extent will be dif-
ferent from those seen at the end of the Pleistocene and in the Early
Holocene due to changes in climate, as well as through human
action such as over-hunting or the destruction of habitats. This prob-
lem is compounded by gaps in knowledge on population extent and
migration patterns of modern birds in some parts of the world, with
many areas of southeastern Turkey having only been systematically
surveyed relatively recently. As such, modern seasonality data must
be considered with a great deal of caution.
Seasonality at Körtiktepe has been addressed in previous pub-
lications, with year-round occupation suggested on the basis of
substantial architectural investment at the site, presuming that the
remains of a seasonal camp would have been more ephemeral, as
well as evidence from strontium isotopic data (Benz et al., 2016)
(although see Asouti and Fuller, 2013). Using avifaunal data,
Arbuckle and Özkaya (2006) concluded that Körtiktepe shows
evidence of multiple-season occupation, on the basis of the pres-
ence of numerous winter visitors to the site, as well as the presence
of the mallard (Anas platyrhynchos), considered to be a summer
visitor (based on Harrison, 1982). Taking into account more
regionally specific and modern survey data however, we would
consider the mallard to be a year-round resident, as well as a win-
ter visitor (Biricik and Karakas, 2012: 99, tab.2; Karakaş, 2010:
474, tab.2; Karakaş and Kiliç, 2004: 304, tab.2). Additionally,
Özkaya and Coşkun (2011) argue that seasonality is implied by the
presence of the greylag goose and the great bustard, which are
breeding in the region in spring and summer (Özkaya and Coşkun,
2011: 101). Nowadays there are only small groups of the greylag
goose that ‘probably represent no more than relicts of a once much
more widespread breeding population’ in the region (Scott and
Rose, 1996: 10), but generally these populations are sedentary or
‘locally dispersive’ (Scott and Rose, 1996: 73). Whilst it has previ-
ously been stated that the graylag goose vacates their breeding
grounds in eastern Turkey during the winter (Scott and Rose,
1996: 73), this species has more recently been found at the reser-
voir at the Göksu dam, south of Diyarbakır, there apparently being
an important winter-feeding ground for this species (Karakas and
Kilic, 2002: 49). With the population extent of the graylag goose
seemingly having greatly changed in the recent past, the presence
of this species is not considered to be a reliable indicator for
seasonality.
The migratory habits of the great bustard, being the most com-
mon bird species found in both the YD and EH occupations, is
poorly understood (Kirwan and Martins, 1994: 91). It is however
considered to be a resident species, although until recently (1980s)
it is reported that their numbers used to hugely swell in the winter
in this region (Kirwan et al., 2008b: 177; Kirwan and Martins,
2000: 25). As such, its presence cannot be used as a seasonal indi-
cator, but if in the ancient past this species also frequented south-
east Anatolia as a wintering ground, this bird could have been
considered an important winter resource.
The majority of bird species from the Younger Dryas levels,
including the most-commonly found (great bustard and the Eur-
asian eagle-owl) are both known to be locally resident species,
although, as mentioned, the numbers of the great bustard may be
larger in the winter (Kirwan et al., 2008a: 171, 251). There are a
small number of species that are known to be winter visitors
including the hen harrier (Circus cyaneus) and the greater white-
fronted goose (Anser albifrons) (Kirwan et al., 2008a: 132, 60). No
species that are found in southeastern Anatolia exclusively in the
summer were identified in the Younger Dryas levels. A number of
taxa are locally resident, but their numbers will be larger in the
winter when migrating populations will join them, such as the
northern pintail (Anas acuta), the greylag goose, the common
crane (Grus grus) and the common buzzard (Buteo buteo) (Kirwan
et al., 2008a: 71, 61, 166, 139). This profile is much the same
within the Early Holocene period, with a large overlap in terms of
species found, with the addition of the presence of other winter
visitors such as the red-breasted goose (Branta ruficollis) and the
grey heron (Ardea cinerea) (Kirwan et al., 2008a: 63, 110).
The lack of presence of summer visitors is however not that surpris-
ing for southeastern Anatolia. With southeastern Anatolia being at a
relatively low latitude, comparatively few bird species would be exclu-
sively summer visitors (Newton, 2008: 370–371). Southeastern Anato-
lia is however an important stopover point for migrating birds, which is
reflected by the higher numbers of birds and higher range of species
observed in the spring today (Biricik and Karakas, 2012:104, fig.2;
Karakaş, 2010, 479, fig.2, fig.3; Karakaş, 2015, 109, fig.2; Karakaş and
Kiliç, 2004: 307, fig.2). Overall, in both the YD and EH levels, bird
exploitation, and by extension site occupation can be inferred for the
winter months. Summer occupation of the site cannot be inferred from
the observed bird species. However, the lack of strong seasonality in
terms of which birds would be present at different points in the annual
cycle means that it is difficult to use birds as a seasonality indicator in
this region. The lack of species indicating a summer occupation is how-
ever not necessarily evidence for the site being a seasonal encampment,
but is more a reflection of the habits of birds in this region. Presuming
year-round occupation, local bird populations may have provided a
valuable food resource through winter and spring, as well as during the
spring/autumn seasonal migrations, when bird populations in the region
swell, and likely played a less important subsistence role in the summer,
when bird numbers are lower.
Reptilia
The reptile taxa from Körtiktepe are summarised in Table 4. The
most common species found in both phases is the Greek Tortoise
(Testudo graeca). It has been previously published that a number
of tortoise carapaces have also been found within burials (Özkaya
and Coşkun, 2011: 94). Additionally, from the Early Holocene
Table 4. Reptile taxa from Körtiktepe.
Younger Dryas Early Holocene
NISP Weight (g) NISP Weight (g)
Lacertidae 1 0.0
Ophisaurus apodus 1 0.8
Testudo graeca 18 266.4 5 367.7
Total 18 266.4 7 368.5

Emra et al. 11
contexts several skull elements from a European glass lizard
(Ophisaurus apodus) and a fragment of a vertebra from the family
of wall lizards (Lacertidae) were identified, both of which are
potentially intrusive.
Fish
A total of 436 fish specimens from the Younger Dryas were
recorded and 758 from the Early Holocene contexts. The vast
majority of fish specimens could not be identified to a species
level, with 53.9% and 26.1% of fish remains being recorded sim-
ply as Pisces in the Younger Dryas and Early Holocene respec-
tively. Where species could be determined, all were freshwater
taxa, unsurprising due to the location of Körtiktepe, on the bank of
the modern-day Batman Çayı and the Tigris River. In both phases
the vast majority of taxa are from the carp family (Cyprinidae)
with the most commonly identified species being the mangar
(Luciobarbus esocinus) (Table 5). Mangar can reach lengths of
over 2 m and weights of in excess of 150 kg (Coad, 2021:1163-
1164). A small number of particularly large vertebrae from EH
Körtiktepe suggests that mangar of this size were being caught.
The Mangar’s environmental requirements are not well under-
stood but they are generally found in modern times in rivers,
streams, lakes, marshes and reservoirs (Coad, 2021:1164). Apart
from members of the carp family, a few examples of the spiny eel
(Mastacembelus mastacembelus) were also identified. The spiny
eel is a freshwater fish living in both still and running water, usu-
ally found near the bank amongst rocky crevices or plant stumps
(Coad, 2015: 6).
Whilst too few fish specimens were identified to a species level
in order to convincingly speak on differences in taxa between the
Younger Dryas and Early Holocene at Körtiktepe, potentially there
is some shift in the size of fish being caught between the two
phases. Figure 8 shows that the height of pre-caudal vertebrae of
cyprinids, as a rough indicator for the size of fish, is potentially
increasing between the Younger Dryas to the Early Holocene, with
certainly all of the larger specimens being recovered in the later
stage of the settlement.
Downstream from Körtiktepe, Hasankeyf Höyük also reports
an ichthyological assemblage dominated by Cyprinidae, most
commonly Barbus sp. (although these may have been reclassified
into genera such as Arabibarbus sp. or Luciobarbus sp.), as well as
Capoeta sp. (Itahashi et al., 2017: 42; Hongo et al., 2019: 160).
Similarly, Cyprinidae was the most commonly found family at
Gusir Höyük with the mangar also being the most commonly iden-
tified species.
Molluscs
Finds of molluscs were exceptionally rare in both phases of the
site. These were limited to one find of Theodoxus sp. a small fresh-
water snail, from the Younger Dryas, a small number of fragments
belonging to freshwater mussels (Unionidae) in both phases, and
single case of (Unio tigridis) being identified in a context from the
Younger Dryas levels (Table 6). It should be noted, however, that
finds of molluscs as grave goods in form of beads were extremely
common, the full analysis of beads found is forthcoming (Özkaya
and Coşkun, 2011: 101).
Taxonomic diversity
It has been proposed that in the Late Epipalaeolithic/Early Neolithic
communities of the Near East, domestication was preceded by an
increase in diet diversity, termed the Broad Spectrum Revolution
(BSR) (Flannery, 1969). The BSR model has been adopted into stud-
ies of early domestication worldwide but has been intensely studied
and debated in the context of the wider Mesopotamian region, particu-
larly in regards to whether this broadening should be attributed to
resource stress, or resource richness (Dobney et al., 1999; Starkovich
and Stiner, 2009; Stiner, 2001; Stiner et al., 2000; Stutz et al., 2009;
Wallace et al., 2019; Zeder, 2012, 2016). If the BSR were to be seen
taking place at precisely this transition to the Early Holocene, hetero-
geneity is expected to increase.
As has been previously discussed in Peters et al. (2013), south-
eastern Turkey (largely sites from the Euphrates basin) sees a
decrease in mammalian archaeobiodiversity (aD) over time. This
increasing specialisation in terms of mammal species exploited is in
particular related to the development of animal husbandry (Peters
et al., 2013: 89: Fig.5.5). This pattern is, however, not observed
within their ‘eastern Turkey’ assemblages (largely sites from the
Upper Tigris Basin) (Peters et al., 2013: 89: Fig.5.5). The analysis
for eastern Turkey however heavily relies on samples with relatively
small sample sizes (<800 NISP), and a low sample size will mean
that low heterogeneity is likely to be observed. It also heavily relies
on the data from just a small number of sites to see what happens in
the later stages of the PPN (particularly Çayönü and Cafer Höyük).
Çayönü is also notably further west than the other sites compared in
the Tigris valley assemblage, and Cafer Höyük is in the Euphrates
basin, so may not fit into any regional pattern, if there is one to be
Table 5. Fish taxa from Körtiktepe. Percentage NISP calculated only using specimens identified to a species level.
Scientific name English name Younger Dryas Early Holocene
NISP Weight (g) NISP % NISP Weight (g) NISP %
Acanthobrama marmid Mesopotamian Bream 2 1.3 11.3 1 0.0 6.3
Arabibarbus grypus Shabout 0 0.0 0.0 1 2.1 6.3
Leuciscus vorax Tigris asp 3 2.5 19.0 0 0.0
Luciobarbus kersin Kersin barbel 1 0.4 4.8 0 0.0
Luciobarbus subquincunciatus Mesopotamian barbel 2 4.2 21.5 0 0.0
Luciobarbus esocinus Mangar 14 15.7 102.5 10 100.0 62.5
Luciobarbus kottelati Menderes barbel 3 0.5 11.9 1 4.1 6.3
Luciobarbus xanthopterus Yellowfin barbel 1 0.5 5.1 1 0.6 6.3
Capoeta trutta Longspine scraper 1 0.1 3.7 0 0.0
Cyprinidae Carp 172 60.8 544 197.1
Planiliza abu Abu mullet 1 0.0 3.5 0.0
Mastacembelus mastacem-
belus
Euphrates spiny eel 1 0.0 3.5 2 0.2 12.5
Pisces 235 2.8 198 3.9
Total 436 88.8 758 308.0

12 The Holocene 00(0)
observed. It was noted by the authors that Çayönü in particularly
retains a high level of diversity throughout its occupation, which
may be due to a location in a particularly lush environment, with
many species readily available to exploit (Peters et al., 2013: 89).
This would be in congruence with the knowledge that Çayönü is the
only known site in the Tigris Valley basin to be continuously inhab-
ited throughout the PPNB (Clare and Kinzel, 2020; Erim-Özdoğan,
2011: 62). The particular local environments and site-specific strate-
gies tailored to those environments may then be very individualistic,
rather than conforming to a wider regional trend. Körtiktepe, like
Çayönü, offers an opportunity to consider these strategies over a
longer period of time.
Körtiktepe sees only a small increase in NTAXA from the YD to
EH from 14 to 15 mammal species, although the sample size from
the EH is almost 50% higher. Nonetheless, the Shannon Entropy or
aD increases from 3.0 to 4.3, showing that whilst the number of taxa
utilised has remained more-or-less the same, there is a broadening
on the focus of diet, with less dependence on just one or two taxa
(Figure 9). The opposite can, however, be said in the case of the
avifaunal remains with a decrease in NTAXA from 18 to 15 and
corresponding decrease in the aD from 8.4 to 3.9, reflecting the
increasing emphasis in particular on geese in the Early Holocene.
The observed decrease in variance of δ13C values as well as δ15N in
males found buried at the site, suggests a restricting of diet spectrum
from the YD to EH, is then likely not driven by a change in the
mammal spectrum, but instead perhaps from the relatively limited
range of birds exploited in the EH, or by a narrowing of the plant
diet (Benz et al., 2016: 241). Whilst a broadening of the mammal
spectrum can be observed from the YD to EH, with the YD levels
showing similarly lower levels of heterogeneity as later sites practic-
ing animal husbandry, a uniform pattern is not observed, with the
site of Gusir Höyük also displaying lower levels of aD. The restric-
tion in bird species exploited, and indications from isotopic data also
do not show conclusive evidence for a diet broadening at Körtiktepe
in the PPNA. However, the slight widening of the mammalian spe-
cies spectrum, with a restriction of the bird species exploited might
be seen in terms of more opportunities, with more choice over which
species hunted leading to a broadening of diet in some realms, but
specialisation in others. This would then suggest that in our study
area any observed broadening of diet was a result of climate amelio-
ration allowing for experimentation in strategy, rather than necessity
forcing the utilisation of low-ranking foodstuffs. This limited broad-
ening of diet at Körtiktepe, largely being driven by an increase in
aurochs rather than the ‘low-ranked’ species that would be expected
in a BSR scenario, as well as there being little evidence of a broad-
ening of diet in the PPNA more regionally, argues against evidence
for a BSR in the Upper Mesopotamian basin, in line with previous
studies’ results (Arbuckle, 2015: 227; Peters et al., 2013: 89: Fig.5.5;
Starkovich and Stiner, 2009: 57–58). It should however be kept in
mind that sieving of sediment was not widely undertaken at Körtik-
tepe, except for burials, and so smaller species are likely to have
been generally underrepresented in both the YD and EH contexts.
Discussion
The increase in humidity, temperature, rainfall, and forest expan-
sion that characterises the beginning of the Early Holocene in this
region is clearly reflected in the relative proportion of species
exploited at Körtiktepe. A major shift, particularly within the avi-
faunal assemblage to an emphasis in the exploitation of waterfowl
in the Early Holocene, suggests changes in local waterways took
Figure 8. Percentage NISP (%) of the height of pre-caudal vertebrae of Cyprinids from Younger Dryas (YD) and Early Holocene (EH) of
Körtiktepe.
Table 6. Mollusc taxa from Körtiktepe.
Younger Dryas Early Holocene
NISP Weight (g) NISP Weight (g)
Theodoxus sp. 1 0.0
Unio tigridis 1 8.0
Unionidae 3 2.5 2 18
Total 5 10.5 2 18

Emra et al. 13
place that became particularly favourable for these species. This
may have included an increase in marshland with large reed beds
and wet meadows or the formation of lakelets (Del Hoyo et al.,
1992: 582). In the mammalian assemblage, these changes are also
indicated by an increase in aurochs remains, a species that likely
inhabited largely riverine, marshy, and forested environs (van
Vuure, 2005: 236, 239–240, 245). This is accompanied by a mod-
erate increase in wild boar, which whilst is an adaptable species,
particularly thrives in gully side vegetation, oak forest, and mixed
wood, but tends to avoid open spaces, making the local Early
Holocene landscape more suitable than that of the Younger Dryas
(Abaigar et al., 1994: 208; Wilson and Mittermeier, 2011: 290).
The appearance, although in small numbers, of the European
wildcat (Felis silvestris) is also indicative of expanding woodland
in the vicinity of the settlement. Whilst riparian gallery forest may
have increased generally in the EH, regionally the, if anything,
slight decrease in bird species associated with woodlands, sug-
gests that any such new woodland areas in the vicinity of Körtik-
tepe in the EH, still likely remained relatively sparse.
Changes in local waterways were also likely responsible for the
increase in the size of fish found in the Early Holocene. Whilst it
is not possible in this assemblage to know whether this increase is
due to a change in the species available and exploited, or in the size
of the same species, it nevertheless indicates a change in either the
location or character of the local rivers or the number and size of
the bodies of water. This is however caveated that this shift might
be a reflection of changes in human choice or fishing technique.
For instance, in one of the later layers of Körtiktepe a fishing hook
with two holes in it was discovered, and it has been proposed that
if the two holes were connected with string the hook would be bet-
ter protected from breaking when catching large fish (Coşkun
et al., 2010: 62). The importance of the role of human choice, cul-
tural preference or changes in hunting strategy is exemplified by
the avifaunal assemblage at Gusir Höyük, which despite the settle-
ment’s location on the edge of a lake and being not far from two
large rivers, is almost exclusively dominated by partridges (Alec-
toris chukar, Perdix perdix) whose preferred habitats are rocky
hills and grasslands respectively (Kabukcu et al., 2021: 4; Kirwan
et al., 2008a: 50, 53).
Small increases in frequency of a number of thermophile mam-
mals such as the goitered gazelle, equids, and potentially also in
fallow deer hints at the beginnings of a changing faunal spectrum,
the result of warming at the beginning of the Early Holocene (Table
3). The increases in these species are only very slight, representing
only the very beginnings of the climatic amelioration, with the fau-
nal spectrum changing only after a delay as a period of adjustment.
Overall, however, there is a great deal of continuity with species
such as mouflon and great bustard remaining well represented in
the assemblages. The continued presence of these species suggests
continued access and exploitation of steppe-like, grassland habitats
in the vicinity of the site, which is also supported by the archaeobo-
tanical results (Rössner et al., 2018: 25).
Whilst being of relatively lesser importance, wetland environ-
ment was however exploited in the Younger Dryas, with various
species of ducks, waders, and raptors such as harriers (Circus sp.)
often associated with wetland habitats (Harrison, 1982: 101). Gen-
erally lacking from the Early Holocene layers, but making up a
sizeable percentage (12.7%) in the Younger Dryas, however, is the
Eurasian eagle-owl (Bubo bubo), a species that would primarily
roost in rocky environments such as the nearby Raman mountains
approximately 6 km east of the site (Harrison, 1982: 171; Kartal
et al., 2018: 92). Exploitation of mountainous terrain is also per-
haps indicated in both periods by the hunting of wild goat, as well
as the presence of a find of Chamois (Rupicapra rupicapra) in the
YD. These taxa were perhaps exploited especially in the winter
when they descend from the mountains, particularly to avoid
heavy snowfall (Ballo, 2010: 36; Nowak and Paradiso, 1983:
1299). The lack of Eurasian eagle-owl in the EH period may sug-
gest either a movement away from the utilisation of this environ-
ment, or lack of interest in hunting this particular species.
Overall, whilst in both the Younger Dryas and Early Holocene
multiple ecozones (wetland, grassland, mountainous habitats) were
exploited, a feature that has been observed to characterise early
Holocene sites in this region (Zeder and Spitzer, 2016: 144), a shift
towards the exploitation of species associated with wetland envi-
ronments suggests a change in the nature of the waterways in the
surroundings of the site. The higher representation of wetland asso-
ciated species (aurochs, boar and waterbirds) indicates that the
increased rainfall of the Early Holocene has translated into an
increase in marshland and/or fluvial lakes locally. This new eco-
logical habitat, in addition to the presence of the steppes and moun-
tainous regions that the inhabitants of the site had previously
Figure 9. Shannon Entropy/ archaeobiodiveristy (aD) of Körtiktepe mammalian fauna and comparison with PPN sites in the Upper Tigris and
Euphrates basins. Hallan Çemi data from (Starkovich and Stiner, 2009: 50, tab.3), Mezraa Teleilat data from (İlgezdi, 2008: 85, tab.11), Cafer
Höyük and Çayönü after Peters etal. (2013).

14 The Holocene 00(0)
exploited, allowed for the broadening, and change in emphasis of,
the mammalian taxa exploited, leading to higher heterogeneity, as
indicated by the increase in archaeobiodiversity, seen in the Early
Holocene. Conversely, the introduction of more geese in the local
environment led to an emphasis on the hunting on these species,
particularly the greylag goose, meaning a decrease in the archaeo-
biodiversity amongst the bird remains. From a purely optimal for-
aging theory perspective, geese make an attractive prey due to their
relatively large size and their habit of flocking in relatively large
numbers, allowing for the prospective of multiple catches in one
hunt. Their size also means large feathers, which would be another
attractive product to the hunters of Körtiktepe.
The increasing archaeobiodiversity amongst mammals, and
restricting aD amongst the avifaunal at Körtiktepe, as well as a
lack of noticeable trend over time in the Upper Tigris Basin sug-
gests a lack of overarching trends in this region regarding climate
change and heterogeneity. Instead, amongst the PPNA sites, a
wide range of strategies can be seen from the relatively special-
ised, such as at Gusir Höyük (aD = 2.71), compared to the more
diverse, such as at Hallan Çemi (aD = 4.55) or EH Körtiktepe
(aD = 4.26). Instead, the particular circumstances of the site
seems to be the most important factor, with the EH amelioration,
in the case of Körtiktepe, allowing for the introduction of a more
diverse range of mammalian taxa, and in the case of birds allow-
ing for the specialisation on taxa that suits the inhabitants needs,
either in terms of calories, convenience, use of fat or feathers, or
ritual uses.
This change in local environment does not, however, translate
into substantial change in mammal size ranges with both red deer
and mouflon showing remarkable continuity in this respect across
the Late Pleistocene/Early Holocene boundary. The few examples
of smaller red deer individuals that appear in the Early Holocene is
unlikely to be explained by the environment changing to one that
is unsuitable for the species, as red deer are thermophiles that
thrive particularly in the interface between woodland and grass-
land, which more closely characterised the local Early Holocene
conditions rather than that of the Younger Dryas (Wilson and Mit-
termeier, 2011: 422). Over-hunting, particularly of large males
leading to selection of smaller individuals might be considered,
although the largest sizes of red deer in the Early Holocene are still
of the same size as those in the Younger Dryas layers, making this
scenario unlikely. It may also be explained by a change in hunting
strategy or herd composition, leading to more females being
hunted and a more complete representation of their body size
range in the LSI data. Alternatively, this might be the beginning of
some size diminution due to Bergmann’s rule, which observes that
in populations members of the same species will exhibit smaller
body sizes in warmer climates and conversely, larger body sizes in
colder conditions (Bergmann, 1847; Meiri and Dayan, 2003; Post
et al., 1997). It has also been found that Bergmann’s rule leads to
size diminution of female red deer in warmer climates, but size
increase in males, which might explain why the upper size range
has not also reduced (Post et al., 1999). Bergmann’s rule is how-
ever controversial and the mechanisms that might lead to changes
in body sizes are numerous and particular to different species (for
controversy over Bergmann’s rule see: Teplitsky and Millien,
2014; Watt et al., 2010). This size diminution is additionally not
replicated in mouflon, whose size range remains stable across the
transition. The smaller individuals in red deer may then simply be
a result of relatively low sample sizes, and further recording may
show that these smaller individuals are present in the landscape in
both periods. It is not possible to consider size change of aurochs
across the YD-EH boundary due to low sample size. However, the
wide size variation of EH aurochs could also be explained by simi-
lar reasons as the variation seen in red deer. Size diminution due to
rising temperatures may be responsible for the smaller individuals,
but not being far into this process, the larger individuals are also
present. Alternatively, the comparatively wide size range may be
due to the sex ratio. If the sex ratio is more balanced compared to
the other sites, a greater size range may be observed due to the less
targetted hunting strategy. However, with no sexed elements being
available, it is not possible to confirm this possibility.
Continuity is observed in the mortality profiles of both red
deer and mouflon, which in both species remains stable across
time. This may indicate that the movements of particular sex and
age cohorts of the population (bachelor herds, nursery herds as
well as lone males) continued without substantial interruption
across this time span, meaning hunting strategies persevered
uninterrupted. This may be due to the resilience of the taxa, but
also the relatively slow change in vegetation in this region, despite
the rapid change in humidity and temperature at the onset of the
Early Holocene (Wick et al., 2003: 670–671). The radiocarbon
dates available for Körtiktepe’s early Holocene contexts begin
only just after the end of the Younger Dryas and should still be
considered to be within this transition phase. As such the lack of
indication of change in animal movements or prey selection may
then not be considered so surprising, but more dramatic changes
to animal behaviour, as well as species composition, may have
been observed in the assemblage had the occupation at Körtiktepe
not been abandoned.
Conclusion
Körtiktepe is one of few sites within the Upper Mesopotamian
basin that exhibits occupation across the Late Pleistocene/Early
Holocene boundary, and thus gives a rare opportunity to consider
climatic impact on the fauna at the onset of the Early Holocene
amelioration and the human response to it. In both the Younger
Dryas and the Early Holocene occupation levels the people of
Körtiktepe primarily exploited red deer and mouflon, as is typical
of this region (Peters et al., 2014: 140). The size of these species,
as reconstructed through LSI data, remains relatively stable across
the transition, with the exception of a small number of smaller red
deer individuals being present in the Early Holocene levels. The
culling profiles of both red deer and mouflon also remain similar
in both periods, suggesting that the migration patterns and hunt-
ing strategies of these species remained stable, and that the envi-
ronment remained hospitable for these taxa. Whilst early
small-scale cultivation is suggested for Körtiktepe in the early
Holocene, at present there is no indication of early animal man-
agement, although it is not possible to assess the status of wild
boar and aurochs due to small sample sizes.
In the Early Holocene there was however a shift to greater
exploitation of aurochs as well as a major shift towards geese
within the avifauna. This indicates changes in the local environ-
ment, such as increased woodland in the case of aurochs, or
changes in the waterways, perhaps the emergence of more marsh-
land, fluvial lakes or larger or more permanent rivers. This is con-
sistent with both the archaeobotanical evidence from the site as
well as more general environmental proxies that the region
received greater precipitation with the onset of the Early Holo-
cene (Rössner et al., 2018: 24–25; Wick et al., 2003: 667–670).
The results then show both a reflection of the wider environmen-
tal changes happening in the region, as well as a very local story
and response.
Whilst a great deal of continuity is evidenced in both the imme-
diate surroundings of the site, as well as in hunting strategy, the
hunting of aurochs would have required a larger group of hunters in
order to kill and process the carcasses of these colossal beasts (see
Pöllath et al., 2018 for discussion on aurochs hunting strategies).
The hunting of aurochs likely had a deeper socio-ritual meaning,
with the necessity for large group co-operation combined with the
danger that aurochs impose. The ritual significance of these species
is well documented in the neighbouring Euphrates basin during the
later PPNB period (Helmer et al., 2004: 157; Stordeur, 2010: 124),
but is, for instance, also indicated for the Tigris basin by the find of

Emra et al. 15
an aurochs skull at a communal building at nearby PPNA Hallan
Çemi, as well as finds of an aurochs horncores at Körtiktepe along-
side an intramural burial ( Özkaya and Coşkun, 2011: 112, fig.6;
Rosenberg, 2011: 63).
The setting of Körtiktepe allowed for the exploitation of grass-
land, aquatic, riverine, and conceivably mountain-dwelling spe-
cies, which facilitated year-round occupation supported by a
hunter-gatherer-fisher strategy. This strategy was largely main-
tained across a period of relatively rapid climatic change, largely
without dramatic upheaval, but perhaps with the addition of small-
scale cultivation, and a shifting emphasis to species that would
likely have been found more frequently under the new climatic
conditions. Under these conditions, the site flourished, to be then
suddenly abandoned sometime in the Late PPNA for unknown rea-
sons, although the threat of flooding will be explored as a possible
reason in future studies.
Acknowledgements
We owe thanks to the General Directorate of Cultural Heritage
and Museums, Ministry of Culture and Tourism, Republic of Tur-
key, and our colleagues and the students of the Körtiktepe team
for their hard work and valuable support. Many thanks to Jörg
Schibler and Francesca Ginelli (IPNA Basel) for their support in
re-sorting and handling of the assemblage. We are also thankful
to Joris Peters and Nadja Pöllath for supplying unpublished data
for the sites Gusir Höyük, Göbekli Tepe and Gürcütepe, as well
as their valuable feedback. We are also grateful to the valuable
feedback and kind comments of two anonymous reviewers.
Funding
The author(s) disclosed receipt of the following financial support
for the research, authorship, and/or publication of this article: We
also gratefully acknowledge the financial support of the Deutsche
Forschungsgemeinschaft (DFG) awarded to Joris Peters - Project
number 165831460.
ORCID iD
Stephanie Emra https://orcid.org/0000-0003-0175-9169
References
Abaigar T, Del Barrio G and Vericad JR (1994) Habitat preference
of wild boar (Sus scrofa L., 1758) in a Mediterranean environ-
ment. Indirect evaluation by signs. Mammalia 58(2): 201–210.
Arbuckle BS (2014) Pace and process in the emergence of animal
husbandry in Neolithic southwest Asia. Bioarchaeology of the
Near East 8: 53–81.
Arbuckle BS (2015) Large game depression and the process of
animal domestication in the Near East. In: Kerner MS, Dann
R and Bangsgaard P (eds) Climate Change in Ancient Soci-
eties. Copenhagen, Denmark: Museum Tusculanum Press,
pp.215–243.
Arbuckle BS and Erek CM (2012) Late epipaleolithic hunters
of the central Taurus: Faunal remains from Direkli Cave,
Kahramanmaraş, Turkey. International Journal of Osteoar-
chaeology 22(6): 694–707.
Arbuckle BS and Özkaya V (2006) Animal exploitation at Körtik
Tepe : An early Aceramic Neolithic site in southeastern Tur-
key. Paléorient 32(2): 113–136.
Asouti E and Fuller DQ (2013) A contextual approach to the emer-
gence of agriculture in southwest Asia: Reconstructing Early
Neolithic plant-food production. Current Anthropology 54(3):
299–345.
Atici L (2009) Implications of age structures for epipaleolithic
hunting strategies in the western Taurus mountains, southwest
Turkey. Anthropozoologica 44(1): 13–39.
Ballo M (2010) Spatial and vertical distribution of chamois (Rupi-
capra rupicapra tatrica) in the western Tatra mountains. Oeco-
logia Montana 19(1-2): 35–41.
Beile-Bohn M, Gerber C, Morsch M et al. (1998) Neolithische
Forschungen in Obermesopotamien-Gürcütepe und Göbekli
Tepe. Istanbuler Mitteilungen 48: 5–78.
Benz M, Coşkun A, Hajdas I et al. (2012a) Methodological impli-
cations of new radiocarbon dates from the early holocene site
of Körtik Tepe, southeast Anatolia. Radiocarbon 54(3-4): 291–
304.
Benz M, Coşkun A, Rössner C et al. (2012b) First evidence of
an Epipalaeolithic hunter-fisher-gatherer settlement at Körtik
Tepe. Kazı Sonuçları Toplantısı 34(1): 65–78.
Benz M, Deckers K, Rössner C et al. (2015) Prelude to village life.
Environmental data and building traditions of the epipalaeoli-
thic settlement at Körtik Tepe, southeastern Turkey. Paléorient
41(2): 9–30.
Benz M, Fecher M, Scheeres M et al. (2016) Results of stable
isotopes from Körtik Tepe, southeastern Turkey. Arkeometri
Sonuçları Toplantısı 31: 231–252.
Benz M, Willmy A, Doğan F et al. (2017) A burnt pit house, large
scale roasting, and enigmatic epipaleolithic structures at Kör-
tik Tepe, southeastern Turkey. Neo-Lithics 17(2): 3–12.
Bergmann C (1847) Ueber die Verhältnisse der Warmeökonomie
der Thiere zu ihrer Grösse. Göttinger Studien 3: 595–708.
Biricik M and Karakas R (2012) Birds of Hasankeyf (south-east-
ern Anatolia, Turkey) under the threat of a big dam project.
Natural Areas Journal 32(1): 96–105.
Boessneck J, Müller HH and Teichert M (1964) Osteologische
Unterscheidungsmerkmale zwischen Schaf (Ovis aries Linné)
und Ziege (Capra hircus Linné). Kühn-Archiv 78: 1–129.
Braidwood RJ and Braidwood L (1953) The earliest village com-
munities in southwestern Asia. Journal of World History 1(2):
278–310.
Braidwood RJ, Çambel H, Redman CL et al. (1971) Beginnings
of village-farming communities in southeastern Turkey. Pro-
ceedings of the National Academy of Sciences of the United
States of America 68(6): 1236–1240.
Braidwood RJ, Çambel H and Watson PJ (1969) Prehistoric inves-
tigations in southeastern Turkey. Science 164(3885): 1275–
1276.
Calderón T, DeMiguel D, Arnold W et al. (2019) Calibration of
life history traits with epiphyseal closure, dental eruption and
bone histology in captive and wild red deer. Journal of Anat-
omy 235(2): 205–216.
Childe VG (1951) Man Makes Himself. New York, NY: New
American Library.
Clare L and Kinzel M (2020) Response to comments by Ian Hod-
der and Christian Jeunesse with notes on a potential Upper
Mesopotamian ‘Late PPNA Hunter-Crisis’. In: Gebaer AB,
Sørensen L, Teather A, et al (eds) Monumentalising Life in
the Neolithic: Narratives of Continuity and Change. Oxford:
Oxbow Books, pp.61–68.
Coad BW (2015) Review of the spiny eels of Iran (family Masta-
cembelidae). Iranian Journal of Ichthyology 2(1): 1–12.
Coad BW (2021) Carps and Minnows of Iran (Families Cyprinidae
and Leuciscidae. Available at: http://www.briancoad.com/Spe-
cies%20Accounts/Carps%20of%20Iran%2010Sept2opt1.pdf
(accessed 19 January 2022).
Coşkun A, Benz M, Erdal YS et al. (2010) Living by the water–boon
and bane for the people of Körtik Tepe. Neo-Lithics 2(10): 60–71.
Coşkun A, Benz M, Rössner C et al. (2012) New results on the
younger dryas occupation at Körtik Tepe. Neo-Lithics 1(12):
25–32.
Del Hoyo Elliott A and Sargatal J (1992) Handbook of the Birds of
the World, vol. 1. Barcelona, Spain: Lynx edicions.
Dietrich O (2011) Radiocarbon dating the first temples of man-
kind: Comments on 14C-dates from Göbekli Tepe. Zeitschrift
für Orient-Archäologie 4: 12–25.
Dobney K, Beech M and Jaques D (1999) Hunting the broad
spectrum revolution: The characterisation of Early Neolithic

16 The Holocene 00(0)
animal exploitation at Qermez Dere, northern Mesopotamia.
In: Driver CC (ed.) Zooarchaeology of the Pleistocene/Holo-
cene Boundary, vol. 800. London: BAR International Series,
pp.47–57.
Erdal YS (2015) Bone or flesh: Defleshing and post-depositional
treatments at Körtik Tepe (southeastern Anatolia, PPNA
period). European Journal of Archaeology 18(1): 4–32.
Erim-Özdoğan A (2011) Çayönü. In: Özdoğan M, Başgelen N
and Kuniholm I (eds) The Neolithic in Turkey. New Excava-
tions and New Research. The Tigris Basin. Istanbul, Turkey:
Archaeology and Art Publications, pp.89–127.
Flannery KV (1969) Origins and ecological effects of early domes-
tication in Iran and the Near East. In: Ucko PJ and Dimbleby
GW (eds) The Domestication and Exploitation of Plants and
Animals. London: Duckworth, pp.73–100.
Friedrich M, Remmele S, Kromer B et al. (2004) The 12,460-year
Hohenheim oak and pine tree-ring chronology from central
Europe—a unique annual record for radiocarbon calibration
and paleoenvironment reconstructions. Radiocarbon 46(3):
1111–1122.
Gourichon L (2004) Faune et saisonnalité: l’organisation tempo-
relle des activités de subsistance dans l’Épipaléolithique et
le Néolithique précéramique du Levant nord (Syrie). Unpub-
lished PhD thesis. Université Lumière-Lyon II, France.
Grant A (1982) The use of tooth wear as a guide to the age of
domestic animals. In: Wilson B, Grigson C and Payne S (eds)
Ageing and Sexing Animal Bones From Archaeological Sites,
vol. 109. Oxford: BAR British series, pp.91–108.
Grupe G and Peters J (2011) Climatic conditions, hunting activi-
ties and husbandry practices in the course of the Neolithic
transition: The story told by stable isotope analyses of human
and animal skeletal remains. In: Pinhasi R and Stock JT (eds)
Human Bioarchaeology of the Transition to Agriculture. New
York, NY: John Wiley & Sons, pp.63–85.
Habermehl KH (1975) Altersbestimmung bei Haus- und Labortie-
ren. Hamburg, Germany: Paul Parey.
Habermehl KH (1985) Altersbestimmung bei Wild- und Pelztieren:
Möglichkeiten und Methoden. Hamburg, Germany: Paul Parey.
Halstead P, Collins P and Isaakidou V (2002) Sorting the sheep
from the goats: Morphological distinctions between the man-
dibles and mandibular teeth of AdultOvis and Capra. Journal
of Archaeological Science 29(5): 545–553.
Harrison CJO (1982) An Atlas of the Birds of the Western Palaearc-
tic. New York, NY: HarperCollins.
Hauptmann H (1988) Nevali Çori: Architektur. Anatolica 15(1):
99–110.
Hauptmann H (1999) The Urfa region. In: Özdoğan M (ed.) Neo-
lithic in Turkey: The Cradle of Civilization; New Discoveries.
Istanbul, Turkey: Arkeoloji ve Sanat Yayınları, pp.65–86.
Hauptmann H (2011) The Urfa region. In: Özdoğan M, Başgelen N
and Kuniholm I (eds) The Neolithic in Turkey: The Euphrates
Basin. Istanbul, Turkey: Arkeoloji ve Sanat Yayınları, pp.85–138.
Helmer D, Gourichon L and Stordeur D (2004) À l’aube de la
domestication animale. Imaginaire et symbolisme animal dans
les premières sociétés néolithiques du nord du Proche-Orient.
Anthropozoologica 39(1): 143–163.
Hesse B (1982) Slaughter patterns and domestication: The begin-
nings of pastoralism in western Iran. Man 17(3): 403–417.
Higham TFG, Bronk Ramsey C, Brock F et al. (2007) Radiocar-
bon dates from the Oxford AMS system: Archaeometry dat-
elist 32. Archaeometry 49(s1): S1–S60.
Hongo H, Arai S, Takahashi R et al. (2019) Transition to food pro-
duction suspended—A remarkable development in the eastern
Upper Tigris valley, South Anatolia. In: Peters J, McGlynn G
and Goebel V (eds) Animals: Cultural Identifiers in Ancient
Societies. Rahden: VML Verlag Marie Leidorf, pp.155–172.
Hongo H, Meadow RH, Öksüz B et al. (2005) Sheep and goat
remains from Çayonü Tepesi, southeastern Anatolia. In:
Archaeozoology of the near East VI. Proceedings of the sixth
international symposium on the archaeozoology of South-
western Asia and adjacent areas (eds Buitenhuis H, Choyke
AM, Martin L, Bartosiewicz L and Mashkour M), London,
UK, 30 August - 1 September 2002, pp.112–123. London:
University College London.
Itahashi Y, Miyake Y, Maeda O et al. (2017) Preference for fish
in a Neolithic hunter-gatherer community of the upper Tigris,
elucidated by amino acid δ 15 N analysis. Journal of Archaeo-
logical Science 82: 40–49.
İlgezdi G (2008) The domestication process in southeastern Tur-
key. The evidence of Mezraa-Teleilat. Unpublished PhD the-
sis.
Kabukcu C, Asouti E, Pöllath N et al. (2021) Pathways to plant
domestication in southeast Anatolia based on new data from
aceramic Neolithic Gusir Höyük. Scientific Reports 11(1): 2112.
Kaltenthaler D, Lohrer J, Kröger P et al. (2021) OssoBook v20.1.
München, Germany: Basel. Available at: http://xbook.vetmed.
uni-muenchen.de/(accessed 1 January 2021).
Karakaş R (2010) Bird diversity in Bismil Plain IBA’S with new
records for south-eastern Anatolia, Turkey. European Journal
of Wildlife Research 56(4): 471–480.
Karakaș R (2015) Current status and distribution of diurnal raptor
species in the south-eastern Anatolia region, Turkey. Slovak
Raptor Journal 9(1): 105–113.
Karakas R and Kilic A (2002) Birds of Göksu Dam (Diyarbakır) and
new records in south-east Turkey. Sandgrouse 24(1): 38–43.
Karakaş R and Kiliç A (2004) The birds of Dicle dam (Diyarbakır).
Turkish Journal of Zoology 28(4): 301–308.
Kartal M, Kartal G, Coşkun A et al. (2018) Chipped stone assem-
blages of Körtik Tepe (Turkey). Journal of Archaeological
Science Reports 19: 92–99.
Karul N (2011) Gusir Höyük. In: Özdoğan M, Başgelen N and
Kuniholm I (eds) The Neolithic in Turkey: The Euphrates Basin.
Istanbul, Turkey: Arkeoloji ve Sanat Yayınları, pp.85–138.
Karul N (2020) The beginning of the Neolithic in southeast Anato-
lia. Documenta Praehistorica 47: 76–95.
Kenyon K (1965) Archaeology in the Holy Land. London: Benn.
Kirwan GM, Boyla K, Castell P et al. (2008a) The Birds of Turkey.
Texas: Helm.
Kirwan GM, Özen M and Demirci B (2008b) Turkey bird report
2002–06. Sandgrouse 30: 166–189.
Kirwan GM and Martins RP (1994) Turkey bird report 1987-91.
Sandgrouse 16(2): 76–117.
Kirwan GM and Martins RP (2000) Turkey bird report 1992-1996.
Sandgrouse 22(1): 13–35.
Kodaş E (2019a) Yukarı Dicle’de Yeni Bir Çanak Çömlek-
siz Neolitik Yerleşim Yeri: Boncuklu Tarla Kazıları ve İlk
Gözlemler. Arkeoloji Ve Sanat Dergisi1 57: VII–XX.
Kodaş E (2019b) A new Aceramic Neolithic site in the upper Tigris val-
ley: Preliminary results of Boncuklu Tarla. Neo-Lithics 19: 3–15.
Kodaş E, Genç B, Çiftçi Y et al. (2020) Çemka Höyük: A late epi-
palaeolithic and pre-pottery Neolithic site on the upper Tigris,
southeast Anatolia. Neo-Lithics 20: 40–46.
Lang C, Peters J, Pöllath N et al. (2013) Gazelle behaviour and
human presence at early Neolithic Göbekli Tepe, south-east
Anatolia. World Archaeology 45(3): 410–429.
Lemoine X (2012) Pig (Sus scrofa) Exploitation at Hallan Çemi,
Southeastern Anatolia: Proposing an alternative model. Paper
1, Unpublished Honors Theses, Portland State University, USA.
Lyman RL (2008) Quantitative Paleozoology. Cambridge: Cam-
bridge University Press.
Manhart H (1998) Die vorgeschichtliche Tierwelt von Koprovic
und Durankulak und anderen prähistorischen Fundplätzen in
Bulgarien aufgrund von Knochenfunden aus archäologischen
Ausgrabungen. Documenta Naturae 116: 129–149.
Mariezcurrena K and Altuna J (1983) Contribución al cono-
cimiento del desarollo de la dentición y el esqueleto post-
craneal de Cervus elaphus. Munibe. Ciencias Naturales 34:
203–246.

Emra et al. 17
Meadow RH (1999) The use of size index scaling techniques for
research on archaeozoological collections from the Middle East.
In: Meadow RH, Becker C, Manhart H et al. (eds) Historia Ani-
malium ex Ossibus. Beiträge zur Paläoanatomie, Archäologie,
Ägyptologie, Ethnologie und Geschichte der Tiermedizin. Rah-
den, Germany: VML Verlag Marie Leidorf, pp.285–300.
Meiri S and Dayan T (2003) On the validity of Bergmann’s rule.
Journal of Biogeography 30(3): 331–351.
Miyake Y, Maeda O, Tanno K et al. (2012) New excavations at
Hasankeyf Höyük: A 10th millennium cal. BC site on the
upper Tigris, southeast Anatolia. Neo-Lithics 1(12): 3–7.
Munro ND, Bar-Oz G and Stutz AJ (2009) Aging mountain gazelle
(Gazella gazella): Refining methods of tooth eruption and wear and
bone fusion. Journal of Archaeological Science 36(3): 752–763.
Neuberger FM, Balasse M, Pöllath N et al. (2019) Diet of wild
versus culturally controlled caprines in Early Neolithic Anato-
lia based on stable carbon isotope analysis of bone apatite. In:
Peters J, McGlynn G and Goebel V (eds) Animals: Cultural
Identifiers in Ancient Societies. Rahden, Germany: VML Ver-
lag Marie Leidorf, pp.155–172.
Newton I (2008) The Migration Ecology of Birds. Amsterdam, the
Netherlands: Elsevier.
Nowak RM and Paradiso JL (1983) Walker’s Mammals of the World,
vol. II. Baltimore and London: Johns Hopkins University Press.
Öksüz B (2000) Analysis of the cattle bones of the PrePottery
Neolithic settlement of Çayönü. In: Archaeozoology of the
near East IVA proceedings of the fourth international sympo-
sium on the archaeozoology of southwestern Asia and adja-
cent areas (eds Mashkour M, Buitenhuis H, Choyke AL and
Poplin F), Paris, France, 29 June – 3 July 1998, pp.154–161.
Todmorden: ARC Publication.
Özkaya V (2009) Excavations at Körtik Tepe. A new pre-pottery
Neolithic a site in southeastern Anatolia. Neo-Lithics 9(2): 3–8.
Özkaya V and Coşkun A (2011) Körtik Tepe. In: Özdoğan M,
Başgelen N and Kuniholm I (eds) The Neolithic in Turkey.
New Excavations and New Research. The Tigris Basin. Istan-
bul, Turkey: Archaeology and Art Publications, pp.89–127.
Özkaya V, Coşkun A, Benz M et al. (2011) Körtik Tepe 2010
Yılı Kazısı. In: Dönmez H and Ötgün Ö (eds) Kazı Sonuçları
Toplantısı, vol. 33(1). Ankara, Turkey pp.315–338.
Özkaya V and Siddiq AB (2020) A heart-shaped bone artifact from
Körtiktepe. Antropoloji 40: 44–55.
Payne S (1973) Kill-off patterns in sheep and goats: The mandibles
from Aşvan Kale. Anatolian Studies 23: 281–303.
Peters J, Arbuckle BS and Pöllath N (2014) Subsistence and
beyond: Animals in Neolithic Anatolia. In: Özdoğan M,
Başgelen N and Kuniholm P (eds) The Neolithic in Tur-
key Vol.6. 10500-5200 BC: Environment, Settlement, Flora,
Fauna, Dating, Symbols of Belief, With Views From North,
South, East, and West. Istanbul, Turkey: Archaeology and Art
Publications, pp.135–203.
Peters J, Buitenhuis H, Grupe G et al. (2013) The long and winding
road: Ungulate exploitation and domestication in Early Neo-
lithic anatolia (10000–7000 cal BC). In: College S, Connolly
J, Dobney K et al. (eds) The Origins and Spread of Domestic
Animals in Southwest Asia and Europe, vol. 59. Walnut Creek,
CA: Left Coast Press, pp.83–114.
Peters J, Helmer D, von Den Driesch A et al. (1999) Early animal
husbandry in the northern Levant. Paléorient 25(2): 27–48.
Peters J, Pöllath N and Arbuckle B (2017) The emergence of live-
stock husbandry in Early Neolithic Anatolia. In: Albarella U,
Rizzetto M, Russ H et al. (eds) The Oxford Handbook of Zoo-
archaeology. Oxford: University Press Oxford, pp.247–265.
Peters J and Schmidt K (2004) Animals in the symbolic world of
pre-pottery Neolithic Göbekli Tepe, south-eastern Turkey: A
preliminary assessment. Anthropozoologica 39(1): 179–218.
Peters J, von den Dreisch A and Helmer D (2005). The Upper
Euphrates-Tigris Basin: cradle of agro-pastoralism?. In: Vigne
J-D, Peters J and Helmer D (eds) The first steps of animal
domestication. New Archaeological Approaches, pp.96–124.
Pöllath N, Dietrich O, Notroff J et al. (2018) Almost a chest hit:
An aurochs humerus with hunting lesion from Göbekli Tepe,
south-eastern Turkey, and its implications. Quaternary Inter-
national 495: 30–48.
Pöllath N, Peters J and Jousse H (2008) Archaeobiodiversity of
ichthyofaunas from the Holocene Sahel. In: Badenhorst S,
Mitchell P and Driver JC (eds) Animals and People. Archaeo-
zoological Papers in Honour of Ina Plug. Oxford: Archaeo-
press BAR International Series 1849, pp.57–74.
Post E, Langvatn R, Forchhammer MC et al. (1999) Environ-
mental variation shapes sexual dimorphism in red deer.
Proceedings of the National Academy of Sciences 96(8):
4467–4471.
Post E, Stenseth NC, Langvatn R et al. (1997) Global climate
change and phenotypic variation among red deer cohorts. Pro-
ceedings of the Royal Society of London. Series B: Biological
Sciences 264(1386): 1317–1324.
Roberts N, Reed JM, Leng MJ et al. (2001) The tempo of holocene
climatic change in the eastern Mediterranean region: New
high-resolution crater-lake sediment data from central Turkey.
The Holocene 11(6): 721–736.
Rosenberg M (1994) Hallan Çemi Tepesi: Some further observa-
tions concerning stratigraphy and material culture. Anatolica
20: 121–140.
Rosenberg M (2011) Hallan Çemi. In: Özdoğan M, Başgelen N and
Kuniholm I (eds) The Neolithic in Turkey: The Euphrates Basin.
Istanbul, Turkey: Arkeoloji ve Sanat Yayinlari, pp.61–78.
Rosenberg M and Davis M (1992) Hallan Çemi Tepesi, an early
aceramic Neolithic site in eastern Anatolia: Some preliminary
observations concerning material culture. Anatolica 18(1):
1–18.
Rosenberg M, Nesbitt M, Redding RW et al. (1995) Hallan Çemi
Tepesi: Some preliminary observations concerning Early
Neolithic subsistence behaviors in eastern Anatolia. Anatol-
ica 21: 3–12.
Rosenberg M, Nesbitt RM, Redding RW et al. (1998) Hallan Çemi,
pig husbandry, and post-pleistocene adaptations along the Tau-
rus-Zagros Arc (Turkey). Paléorient 24(1): 25–41.
Rössner C, Deckers K, Benz M et al. (2018) Subsistence strate-
gies and vegetation development at Aceramic Neolithic Körtik
Tepe, southeastern Anatolia, Turkey. Vegetation History and
Archaeobotany 27(1): 15–29.
Schmidt K (1995) Investigations in the upper Mesopotamian Early
Neolithic: Göbekli Tepe and Gürcütepe. Neo-Lithics 2(95):
9–10.
Schmidt K (2000) Göbekli Tepe, southeastern Turkey: A prelimi-
nary report on the 1995-1999 excavations. Paléorient 26:
45–54.
Schmidt K (2006). Animals and a headless man at Göbekli Tepe.
Neo-Lithics 2(6):, 38–40
Schmidt K (2011) Göbekli Tepe. In: Özdoğan M, Başgelen N and
Kuniholm I (eds) The Neolithic in Turkey: The Euphrates
Basin. Istanbul, Turkey: Archaeology and Art Publications,
pp.41–83.
Schmidt K (2012) Göbekli Tepe: A Stone Age Sanctuary in South-
Eastern Anatolia. Berlin, Germany: Ex Oriente.
Schreiber F, Coşkun A, Benz M et al. (2014) Multilayer floors in
the early holocene houses at Körtik Tepe: An example from
house Y98. Neo-Lthics 2(14): 13–22.
Scott DA and Rose PM (1996) Atlas of Anatidae Popula-
tions in Africa and Western Eurasia. Oxford: Information
Press.
Starkovich BM and Stiner MC (2009) Hallan Çemi Tepesi: High-
ranked game exploitation alongside intensive seed processing
at the Epipaleolithic-Neolithic transition in southeastern Tur-
key. Anthropozoologica 44(1): 41–61.

18 The Holocene 00(0)
Stiner MC (1990) The use of mortality patterns in archaeological
studies of hominid predatory adaptations. Journal of Anthro-
pological Archaeology 9(4): 305–351.
Stiner MC (2001) Thirty years on the “Broad Spectrum Revolu-
tion” and paleolithic demography. Proceedings of the National
Academy of Sciences of the United States of America 98(13):
6993–6996.
Stiner MC, Munro ND, Surovell TA et al. (2000) The tortoise
and the hare: Small-game use, the broad-spectrum revolu-
tion, and paleolithic demography. Current Anthropology
41(1): 39–79.
Stordeur D (2010) Domestication of plants and animals, domesti-
cation of symbols. In: Bolger D and Maguire LC (eds) Devel-
opment of Pre-state communities in the Near East. Oxford:
Oxbow Books, pp.123–130.
Stutz AJ, Munro ND and Bar-Oz G (2009) Increasing the resolu-
tion of the broad spectrum revolution in the southern Levan-
tine Epipaleolithic (19-12 ka). Journal of Human Evolution
56(3): 294–306.
Teplitsky C and Millien V (2014) Climate warming and Berg-
mann’s rule through time: Is there any evidence? Evolutionary
Applications 7(1): 156–168.
Uerpmann M and Uerpmann HP (1994) Animal bone finds from
excavation 520 at Qala’at al-Bahrain. In: Hojlund WF and
Andersen HH (eds) Qala’at al-Bahrain, Vol. 1: The Northern
City Wall and the Islamic Fortress. Aarhus, Denmark: Aarhus
University Press, pp.417–444.
van Vuure C (2005) Retracing the Aurochs: History. Morphology
and Ecology of an Extinct Wild Ox. Sofia and Moscow: Pen-
soft Publishers.
von Den Driesch A (1976) A guide to the measurement of ani-
mal bones from archaeological sites: as developed by the
Institut für Palaeoanatomie, Domestikationsforschung und
Geschichte der Tiermedizin of the University of Munich, vol.
1. Harvard: Peabody Museum Press.
von Den Driesch A and Peters J (1999) Vorläufiger Bericht über
die archäozoologischen Untersuchungen am Göbekli Tepe und
am Gürcütepe bei Urfa, Türkei. Istanbuler Mitteilungen 49:
23–39.
Wallace M, Jones G, Charles M et al. (2019) Re-analysis of
archaeobotanical remains from pre- and early agricultural
sites provides no evidence for a narrowing of the wild plant
food spectrum during the origins of agriculture in south-
west Asia. Vegetation History and Archaeobotany 28(4):
449–463.
Watkins T, Baird D and Betts A (1989) Qermez Dere and the
early aceramic Neolithic of N. Iraq. Paléorient 15: 19–24.
Watt C, Mitchell S and Salewski V (2010) Bergmann’s rule; a con-
cept cluster? Oikos 119(1): 89–100.
Wick L, Lemcke G and Sturm M (2003) Evidence of lateglacial
and holocene climatic change and human impact in eastern
Anatolia: High-resolution pollen, charcoal, isotopic and geo-
chemical records from the laminated sediments of Lake Van,
Turkey. The Holocene 13(5): 665–675.
Willcox G, Buxo R and Herveux L (2009) Late pleistocene and
early holocene climate and the beginnings of cultivation in
northern Syria. The Holocene 19(1): 151–158.
Wilson DE and Mittermeier RA (2011) Handbook of the Mammals
of the World, Volume 2: Hoofed Mammals. Barcelona, Spain:
Lynx Ediciones.
Zeder MA (2006) Reconciling rates of long bone fusion and tooth
eruption and wear in sheep (Ovis) and goat (Capra). In: Rus-
cillo D (ed.) Recent Advances in Ageing and Sexing Animal
Bones, 9. Oxford: Oxbow Books, pp.87–118.
Zeder MA (2012) The broad spectrum revolution at 40: Resource
diversity, intensification, and an alternative to optimal forag-
ing explanations. Journal of Anthropological Archaeology
31(3): 241–264.
Zeder MA (2016) Domestication as a model system for niche con-
struction theory. Evolutionary Ecology 30(2): 325–348.
Zeder MA and Lapham HA (2010) Assessing the reliability of crite-
ria used to identify postcranial bones in sheep, Ovis, and goats,
Capra. Journal of Archaeological Science 37(11): 2887–2905.
Zeder MA, Lemoine X and Payne S (2015) A new system for com-
puting long-bone fusion age profiles in Sus scrofa. Journal of
Archaeological Science 55: 135–150.
Zeder MA and Pilaar SE (2010) Assessing the reliability of crite-
ria used to identify mandibles and mandibular teeth in sheep,
ovis, and goats, Capra. Journal of Archaeological Science
37(2): 225–242.
Zeder MA and Spitzer MD (2016) New insights into broad spec-
trum communities of the early holocene Near East: The birds
of Hallan çemi. Quaternary Science Reviews 151: 140–159.
Appendix
Appendix 1. Epiphyseal fusion stages used for red deer based on:
(Habermehl, 1985: 36, 37; Mariezcurrena and Altuna, 1983: Tab. 54,
184).
Age of fusion Element
<20 months Proximal and distal humerus
Proximal Radius
Proximal Femur
Acetabulum
Distal Scapula
20–36 months Proximal Ulna
Distal Metapodials
Phalanx I and II
Calcaneus
>36 months Proximal and Distal Tibia
Distal radius
Distal Femur
Appendix 2. Red deer standard, male, (SAPM-MA-01295),
Oberbayern.
Red deer standard
Measurement
code
Measurement
(mm)
Log
value
Scapula LG 42.3 1.626
SLC/KLC 34 1.531
Humerus BT 49.8 1.697
Bd 54.4 1.736
Bp 71.3 1.853
Radius BFp 47.8 1.679
Bp 52.8 1.723
Bd 48 1.681
Metacarpus Bp 40.9 1.612
Bd 41.1 1.614
Tibia Bd 48.1 1.682
Bp 73.5 1.866
Calcaneus GB 31.9 1.504
Metatarsus Bp 33.7 1.528
Bd 41.1 1.614
Talus Bd 31.9 1.504
Os centroquartale GB 40.2 1.604

Emra et al. 19
Appendix 3. Measurements of Cervus elaphus from Younger
Dryas contexts.
Cervus elaphus – Younger Dryas
Scapula ID number S LC LG
1893-2012-1-2012 45.58
332-2012-1-2012 35.33 46.8
339-2012-1-2012 55.33
866-2017-1-2012 52.42
877-2017-1-2012 51.33
Humerus BT Bd Bp
107-2073-1-2012 62.67
1664-2012-1-2012 56.49 60.85
1665-2012-1-2012 58.27
1934-2012-1-2012 79.96
1951-2012-1-2012 57.83
1986-2012-1-2012 60.9
2071-2012-1-2012 59.5
341-2012-1-2012 60.37
47-2073-1-2012 57.67 55
49-2073-1-2012 62.87
528-2017-1-2012 62.62
838-2017-1-2012 59.49
839-2017-1-2012 65.44 66
878-2017-1-2012 55.99 57.5
Radius BFp Bp Bd
1101-2017-1-2012 57.08 60.09
1913-2012-1-2012 27.28
1989-2012-1-2012 61.3
2851-2012-1-2012 59.09 61.25
2880-2012-1-2012 58.52 61.23
753-2017-1-2012 61.08 63.03
931-2017-1-2012 57.52
Metacarpus III + IV Bp Bd
1103-2017-1-2012 49.59
113-2073-1-2012 46.54
184-2073-1-2012 45.93
2015-2012-1-2012 50.31
2072-2012-1-2012 49.91
2853-2012-1-2012 45.86
638-2017-1-2012 52.78
881-2017-1-2012 51.67
948-2017-1-2012 53.44
995-2017-1-2012 44.74
Tibia Bd Bp
1002-2017-1-2012 89.22
1397-2012-1-2012 56.41
1725-2012-1-2012 54.62
173-2073-1-2012 80.41
276-2012-1-2012 53.5
491-2012-1-2012 62.01
543-2017-1-2012 53.34
643-2017-1-2012 54.8
73-2073-1-2012 52.79
899-2017-1-2012 53.22
953-2017-1-2012 55.13
984-2017-1-2012 55.44
Calcaneus GB
1003-2017-1-2012 36
2747-2012-1-2012 34.5
280-2012-1-2012 43.8
954-2017-1-2012 39
970-2017-1-2012 38
Metatarsus III + IV Bd
151-2012-1-2012 45.66
1730-2012-1-2012 55.19
Appendix 4. Measurements of Cervus elaphus from Early
Holocene contexts.
Cervus elaphus – Early Holocene
Scapula ID number SLC LG
1058-2012-1-2012 35.18
1059-2012-1-2012 31.09 35.23
155-2017-1-2012 47.27
2450-2012-1-2012 39.58
3076-2012-1-2012 31.39
367-2012-1-2012 55.16
435-2012-1-2012 54.31
799-2012-1-2012 37.75 49.76
816-2012-1-2012 43.65
Humerus Bd BT
1095-2012-1-2012 63.23 60.87
1621-2012-1-2012 55.37 52.4
1699-2012-1-2012 55.54
1851-2012-1-2012 68.76 61.01
Cervus elaphus – Younger Dryas
747-2017-1-2012 50.03
934-2017-1-2012 49.65
Talus Bd
1005-2017-1-2012 39.97
1006-2017-1-2012 38.98
1060-2017-1-2012 42.35
1111-2012-1-2012 38.32
1159-2012-1-2012 39.53
1160-2012-1-2012 38.81
118-2012-1-2012 43.78
12-2073-1-2012 40.43
1410-2012-1-2012 36.71
1411-2012-1-2012 41.7
154-2073-1-2012 40.69
1661-2012-1-2012 38.37
1886-2012-1-2012 39.67
190-2073-1-2012 41.36
1927-2012-1-2012 37.19
2069-2012-1-2012 35.66
2301-2012-1-2012 40.28
254-2012-1-2012 41.57
262-2073-1-2012 40.72
2749-2012-1-2012 36.4
2857-2012-1-2012 38.82
2886-2012-1-2012 42.05
2887-2012-1-2012 40.49
492-2012-1-2012 42.95
548-2017-1-2012 39.06
549-2017-1-2012 39.86
58-2073-1-2012 39.44
884-2017-1-2012 39.15
956-2017-1-2012 40.11
Centroquartale GB
1300-2012-1-2012 56.36
1662-2012-1-2012 49.23
1663-2012-1-2012 46.55
263-2073-1-2012 44.52
281-2012-1-2012 57.31
331-2012-1-2012 52.88
493-2012-1-2012 48.86
546-2017-1-2012 46.48
971-2017-1-2012 47.14
(Continued)
Appendix 3. (Continued)
(Continued)

20 The Holocene 00(0)
Cervus elaphus – Early Holocene
2007-2012-1-2012 75.4 65.4
2261-2012-1-2012 57.74
2262-2012-1-2012 57.19
2314-2012-1-2012 57.74
3127-2012-1-2012 56.6
379-2012-1-2012 58.65 55.01
444-2012-1-2012 60.5 59.14
Radius BFp Bp Bd
1337-2012-1-2012 60.14 62.49
1457-2012-1-2012 57.7 64.11
1772-2012-1-2012 64.75
1964-2012-1-2012 55.25 58.02
1965-2012-1-2012 60.6 64.86
2050-2012-1-2012 55.46
3128-2012-1-2012 53.96 56.41
445-2012-1-2012 55.85
641-2012-1-2012 63.21
734-2017-1-2012 56.68
737-2017-1-2012 48
829-2012-1-2012 58.68
905-2012-1-2012 49.09
907-2012-1-2012 52.47
Metacar-
pus III + IV
Bp Bd
1339-2012-1-2012 49
1443-2012-1-2012 55.78
210-2017-1-2012 48.81
2628-2012-1-2012 43.1
3133-2012-1-2012 47.83
410-2012-1-2012 50.4
449-2012-1-2012 50.98
54-2012-1-2012 50.74
Tibia Bd
1127-2012-1-2012 56.22
1440-2012-1-2012 52.82
1571-2012-1-2012 57.83
1761-2012-1-2012 61.53
2081-2012-1-2012 53.6
25-2012-1-2012 58.82
3060-2012-1-2012 53.65
468-2012-1-2012 56.97
719-2017-1-2012 55
Calcaneus GB
1618-2012-1-2012 39.96
187-2017-1-2012 31
2356-2012-1-2012 35.9
318-2012-1-2012 35.5
3564-2012-1-2012 38.5
404-2012-1-2012 34.5
862-2012-1-2012 41
Metatar-
sus III + IV
Bd
189-2017-1-2012 47.46
860-2012-1-2012 43.65
Talus Bd
1236-2012-1-2012 36.6
1600-2012-1-2012 34.99
165-2017-1-2012 43.01
1841-2012-1-2012 39.23
1959-2012-1-2012 35.27
2027-2012-1-2012 37.25
209-2012-1-2012 39.7
239-2017-1-2012 39.6
2407-2012-1-2012 41.3
28-2012-1-2012 38.68
Cervus elaphus – Early Holocene
3062-2012-1-2012 39.47
3599-2012-1-2012 41.14
470-2012-1-2012 36.89
508-2012-1-2012 39.09
53-2012-1-2012 44.23
75-2012-1-2012 40.54
793-2012-1-2012 40.64
794-2012-1-2012 39.66
Centro-
quartale
GB
1524-2012-1-2012 56.47
210-2012-1-2012 47.49
Appendix 5. Measurements of Ovis orientalis from Younger
Dryas contexts.
Ovis orientalis – Younger Dryas
Scapula ID number SLC GLP LG BG
1073-2017-1-2012 21.27
130-2012-1-2012 43.5 32.44 28.33
1307-2012-1-2012 25.25
1489-2012-1-2012 19.69 27.59 23.24
46-2073-1-2012 22.27 38.6 30.53 26.34
67-2073-1-2012 24.35 40.13 29.51 27.07
730-2017-1-2012 35.84 28.33 23.92
867-2017-1-2012 26.33 43.41 31.69 28.83
946-2017-1-2012 21.97 35.09 28.03 23.31
Humerus ID number Bp SD Bd BT
110-2073-1-2012 36.05 34.14
16-2073-1-2012 38.19 37.27
1896-2012-1-2012 33.87
1897-2012-1-2012 34.62 34.76
1984-2012-1-2012 43.98
208-2073-1-2012 35.24 33.36
2544-2012-1-2012 38.66 36.92
2848-2012-1-2012 38.89 37.17
736-2017-1-2012 36.93 34.7
84-2073-1-2012 17.33 35.44 35.1
993-2017-1-2012 35.93
Radius ID number Bp BFp Bd BFd
1264-2012-1-2012 36.06 34.54
147-2073-1-2012 34.54 33.94
534-2017-1-2012 32.82 27.2
535-2017-1-2012 32.84 27.88
Metacarpus
III + IV
ID number SD Bd Dd
1112-2012-1-2012 32.96
1267-2012-1-2012 31.65
2816-2012-1-2012 32.28
51-2073-1-2012 31.94 19.98
537-2017-1-2012 20.19 19.95
Phalanx 1
ant.
ID number Bp SD Bd GLpe
1257-2012-1-2012 15.11 11.63 14.21 48.67
1910-2012-1-2012 14.7 11.29 13.91 47.52
1928-2012-1-2012 14.82 10.9 14.52 46.79
20-2017-1-2012 14.25
235-2073-1-2012 15.45 13.15 16.02 51.56
236-2073-1-2012 14.61 12.04 15.75 50.04
470-2017-1-2012 11.45 14.11
Femur ID number Bd
169-2073-1-2012 51.43
Tibia ID number Bp Bd
2238-2012-1-2012 32.01
274-2012-1-2012 30.54
(Continued)
(Continued)
Appendix 4. (Continued) Appendix 4. (Continued)

Emra et al. 21
Ovis orientalis – Younger Dryas
275-2012-1-2012 49.38
Phalanx 1
post.
ID number Bp SD Bd GLpe
1106-2017-1-2012 14.35 11.58 14.1 47.1
1107-2017-1-2012 13.72 13.56 43.34
1258-2012-1-2012 18.54 12.41 14.48 46.5
149-2012-1-2012 14.58 11.51 13.33 51.39
1892-2012-1-2012 14.13 11.55 13.82 46.34
1911-2012-1-2012 14.02 10.87 13.27 47.37
1929-2012-1-2012 13.9 10.89 12.77 46.3
1982-2012-1-2012 16.24 12.02 14.01 53.22
2218-2012-1-2012 15.28
2881-2012-1-2012 12.91 10.71 12.21 46.93
894-2017-1-2012 15.91 13 15.8 53.15
Phalanx 1
ant./post.
ID number Bd
2217-2012-1-2012 14.67
2731-2012-1-2012 13.59
Talus ID number GLl GLm Dl Bd
1800-2012-1-2012 35.67 34.1 20.88 24.64
1949-2012-1-2012 36.96 34.97 21.47 24.43
2012-2012-1-2012 33.54 31.84 19.38 22.22
2070-2012-1-2012 33.37 31.24 19.16 22.4
212-2073-1-2012 35.14 32.6 19.96 22.05
2622-2012-1-2012 36.48 35.6 21.31 23.42
2623-2012-1-2012 35.72 35.5 21.28 23.48
273-2012-1-2012 31.52
2771-2012-1-2012 34.7 32.63 19.45 21.95
473-2017-1-2012 37.1 35.36 21.4 25.39
Calcaneus ID number GL GB
1299-2012-1-2012 23.54
150-2012-1-2012 25.75
1926-2012-1-2012 21.02
330-2012-1-2012 73.63 20.5
901-2017-1-2012 70.5
955-2017-1-2012 80.34 20
Appendix 6. Measurements of Ovis orientalis from Early
Holocene contexts.
Ovis orientalis – Early Holocene
Scapula ID number SLC GLP LG BG
1148-2012-1-2012 25.18 44.41 31.05 28.29
1486-2012-1-2012 20.49
1569-2012-1-2012 22.95 41.28 27.49 26.23
1570-2012-1-2012 24.78 42.98 28.52
1767-2012-1-2012 28.2 42.07 32.53 27.91
217-2012-1-2012 21.66 39.22 31.07 24.32
218-2012-1-2012 18.81 22.98
219-2012-1-2012 40.43 32.82 26.26
2325-2012-1-2012 38.54 31.55 26.31
405-2012-1-2012 24.75 40.88 29 27.08
50-2012-1-2012 41.3 30.29
51-2012-1-2012 21.42 38.27 26.59 26.07
549-2012-1-2012 24.14 41.96 29.77 26.14
637-2012-1-2012 37.55 32.08 23.96
699-2012-1-2012 35.22 28.2 22.84
700-2012-1-2012 23.26 37.71 25.48 24.23
714-2012-1-2012 21.83 37.4 30.56 25.15
818-2012-1-2012 26.99 44.44 33.06 29.3
819-2012-1-2012 28.03 44.13 36.36 28.47
871-2012-1-2012 37.99 28.86
872-2012-1-2012 40.86 30.77 26.34
968-2012-1-2012 41.36 30.21 27.97
Ovis orientalis – Early Holocene
Humerus ID number SD Bd BT
1449-2012-1-2012 37.63 34.91
1450-2012-1-2012 36.85 36.12
1527-2012-1-2012 35.79 33.59
185-2017-1-2012 36.2 35.35
1962-2012-1-2012 37.44 34.39
2122-2012-1-2012 20.27 31.99
2331-2012-1-2012 33.66 32.86
432-2012-1-2012 38 36.64
610-2017-1-2012 36.97 36.12
676-2012-1-2012 36.62 35.27
741-2017-1-2012 42.05 41.86
746-2017-1-2012 35.21 33.5
946-2012-1-2012 34.2 32.19
Radius ID number Bd BFd
15-2012-1-2012 31.72 27.83
196-2012-1-2012 31.95
49-2012-1-2012 29.79
Metacar-
pus III+IV
ID number Bd
1125-2012-1-2012 31.15
1143-2012-1-2012 33.62
2353-2012-1-2012 34.72
47-2012-1-2012 32.16
553-2012-1-2012 31.82
Phalanx 1
ant.
ID number Bp SD Bd GLpe
3072-2012-1-2012 3.55 11.08 13.46 45.99
Femur ID number Bp DC Bd
2279-2012-1-2012 25.56
231-2017-1-2012 43.12
3596-2012-1-2012 25.71
858-2012-1-2012 52.93 23.65
Metatar-
sus III+IV
ID number Bp Bd
1697-2012-1-2012 26.98 32.82
509-2012-1-2012 29.08
510-2012-1-2012 31.9
670-2012-1-2012 29.14
861-2012-1-2012 30.05
Tibia ID number Bp SD Bd
2019-2012-1-2012 51.29 19.88 33.5
234-2017-1-2012 48.67
507-2012-1-2012 30.22
Phalanx 1
post.
ID number Bp SD Bd GLpe
1378-2012-1-2012 11.59 13.69
488-2012-1-2012 15.28
634-2012-1-2012 13.84 12.01
80-2012-1-2012 15.86 12.34 15.16 49.34
Phalanx 1
ant./post.
ID number Bd
1328-2012-1-2012 14.92
143-2017-1-2012 13.36
797-2012-1-2012 14.89
Talus ID number GLl GLm Dl Bd
1327-2012-1-2012 36.53 34.51 24.48
188-2017-1-2012 32.98 23.58
1960-2012-1-2012 36.65 33.86 20.92 23.71
2278-2012-1-2012 33.58 32.07 19.16 22.22
2357-2012-1-2012 36.23 33.58 20.42 23.29
3061-2012-1-2012 36.09 33.76 30.37 24.35
3601-2012-1-2012 34.06 31.1 19.84 22.43
48-2012-1-2012 33.42 31.43 19.11 21.81
576-2012-1-2012 33.64 31.66 21.93
629-2012-1-2012 36.82 34.01 21.58 24.69
745-2017-1-2012 35.56 32.86 24.32
(Continued)
(Continued)
Appendix 5. (Continued) Appendix 6. (Continued)

22 The Holocene 00(0)
Ovis orientalis – Early Holocene
Calcaneus ID number GL GB
163-2017-1-2012 76.74 20
1957-2012-1-2012 73.35 17.8
1958-2012-1-2012 77.31 21.2
2310-2012-1-2012 72.95 18
2448-2012-1-2012 21.23
976-2012-1-2012 75.42 25.08
Appendix 7. Measurements of Bos primigenius from Younger
Dryas contexts.
Bos primigenius - Younger Dryas
Tibia ID number Bp
1430-2012-1-2012 91.54
Appendix 8. Measurements of Bos primigenius from Early
Holocene contexts.
Bos primigenius – Early Holocene
Humerus ID number BT
194-2017-1-2012 114.28
2427-2012-1-2012 101.83
Metacarpus III + IV ID number Bp SD Bd
1224-2012-1-2012 88.07
1412-2012-1-2012 87.82
1427-2012-1-2012 67.15
1834-2012-1-2012 55.57 29.31
615-2017-1-2012 73.26
733-2017-1-2012 82.7
Phalanx 1 ant. ID number GLpe Bp SD Bd
1222-2012-1-2012 74.94 38.38
1445-2012-1-2012 77.71 45.66
1603-2012-1-2012 67.15 37.63 32.17 34.44
970-2012-1-2012 72.69 37.53
Phalanx 2 ant. ID number GLpe Bp SD Bd
2412-2012-1-2012 44.53 34.02 26.71 29.42
Tibia ID number Bp Bd
1422-2012-1-2012 139.71
2447-2012-1-2012 72.12
Metatarsus III + IV ID number Bp Bd
1424-2012-1-2012 55.75
2452-2012-1-2012 62.44
2453-2012-1-2012 72.01
616-2017-1-2012 80.5
804-2012-1-2012 67.15
805-2012-1-2012 72.63
Phalanx 1 post. ID number GLpe Bp SD Bd
1221-2012-1-2012 70.18 33.01 30.62 34.77
2429-2012-1-2012 39.66 34.52 34.85
Phalanx 2 post. ID number Bp SD
2413-2012-1-2012 34.28 28.35
Talus ID number GLl GLm Dl BC/BD
1019-2012-1-2012 76.67 71.53 43.89 48.28
1181-2012-1-2012 67.54 40.35 42.16
1223-2012-1-2012 83.22 48.36 52.2
2011-2012-1-2012 85.53 79.14 52.56 63.81
2319-2012-1-2012 86.32 80.89 43 57.26
2383-2012-1-2012 70.73 49.56
2437-2012-1-2012 82.82 74.99 51.57
357-2017-1-2012 84.44 77.22 43 54.12
358-2017-1-2012 76.4 37
863-2012-1-2012 71.91
Bos primigenius – Early Holocene
Calcaneus ID number GL GB
1018-2012-1-2012 115.8 39.6
1617-2012-1-2012 141 56.85
1700-2012-1-2012 176 61
1751-2012-1-2012 39
Appendix 9. Measurements of Sus scrofa from Early Holocene
contexts.
Sus scrofa – Early Holocene
Scapula ID number GLP LG BG
434-2012-1-2012 44.03 42.48 34.48
Humerus ID number Bd BT
22-2017-1-2012 52.24 38.98
Radius ID number Bd
433-2012-1-2012 40.7
Ulna ID number BPC
249-2012-1-2012 27.31
Tibia ID number Bd
1439-2012-1-2012 41.02
Talus ID number GLl GLm Dl
2332-2012-1-2012 46.97 44.61 24.89
485-2012-1-2012 41.49 22.41
Calcaneus ID number GL_unfused
360-2017-1-2012 67.41
Appendix 10. Measurements of Capra aegagrus from Younger
Dryas contexts.
Capra aegagrus – Younger Dryas
Scapula ID number HS SLC GLP LG BG
1940-2012-1-2012 16.54 33.16 27.15 20.96
529-2017-1-2012 20.09
868-2017-1-2012 19.4 38.68 28.05
928-2017-1-2012 150.9 20.73 36.95 28.97 23.5
Humerus ID number BT Bd
672-2017-1-2012 31.5 31.9
Phalanx 1 post. ID number Bp
997-2017-1-2012 17.58
Phalanx 2 ant./
post.
ID number GL SD Bp
2896-2012-1-2012 30.81 9.98 13.75
Appendix 11. Measurements of Capra aegagrus from Early
Holocene contexts.
Capra aegagrus – Early Holocene
Scapula ID number SLC GLP LG BG
1149-2012-1-2012 21.65 39.35 28.16 25.18
1448-2012-1-2012 20.5 37.67 25.12 24.05
1709-2012-1-2012 19.32 37.71 29.42 22.12
3587-2012-1-2012 39.9 30.92 25.65
675-2017-1-2012 20.64
78-2012-1-2012 22.96 43.94 31.49 27.65
958-2012-1-2012 21.61 39.7 31.64 27.33
959-2012-1-2012 38.24 31.3 27.23
967-2012-1-2012 29.25
Humerus ID number BT Bd
2031-2012-1-2012 37.21 37.19
2032-2012-1-2012 37.21 37.19
2405-2012-1-2012 38.22
Radius ID number SD Bp BFp
2123-2012-1-2012 22.28 35.36 33.38
(Continued) (Continued)
Appendix 6. (Continued) Appendix 8. (Continued)

Emra et al. 23
Capra aegagrus – Early Holocene
Tibia ID number SD Bd
1178-2012-1-2012 36.97
1760-2012-1-2012 18.06 31.27
Talus ID number Bd
1180-2012-1-2012 27.03
Metatarsus III+IV ID number Bd
253-2012-1-2012 23.71
Phalanx 2 ant./post. ID number GL SD Bp Bd
145-2017-1-2012 30.86 10.93 13.93 11.79
Appendix 12. Körtiktepe taxa.
Younger Dryas Early Holocene
NISP Weight NISP Weight
(g)
Mammalia
Bos primigenius 21 684 203 14,528.5
Bos taurus 2 128.1
Bos taurus/Bos primi-
genius
2 48.3
Ovis aries 1 24.8 5 31
Ovis aries/Capra hircus 3 21.3
Ovis aries/Ovis ori-
entalis/Capra hircus/
Capra aegagrus
3 30.8 5 27.1
Ovis orientalis 304 3904.6 151 2829.2
Ovis orientalis/Ovis
aries
17 165.5 5 36.4
Capra aegagrus 11 197.1 29 878.4
Capra hircus 4 117.2
Capra hircus/Capra
aegagrus
2 12.6 1 13
Ovis orientalis/Capra
aegagrus
191 2046.2 350 4310.3
Rupicapra rupicapra 2 45.37
Gazella subgutturosa 16 151.7 35 360.6
Cervus elaphus 476 16242.1 534 16,315.57
Dama dama 3 268.2 8 192.5
Cervidae 68 775.4 99 790.7
Sus scrofa 15 473.1 63 1751.1
Equus hemionus onager
| Equus hydruntinus
2 85 13 734.72
Canis lupus 1 24.5 2 19
Canis sp. 1 2.1
Vulpes vulpes 9 23.5 11 51.1
Felis silvestris 6 19.1
Ursus arctos 1 58.6
Meles meles 2 0.74 2 5.2
Carnivora 1 0.02
Lepus capensis 14 22.95 14 9.85
Erinacaeus europaeus 1 0.9
Erinaceidae 2 0.02
Hemiechinus auritus 1 0.64
Castor fiber 7 149.13 2 27.3
Rodent (large) 2 0.02 1 0.5
Rodent (small) 1 0.01 1 0.1
Rodentia 1 0.01 1 0
Mus musculus 1 0.01 1 0.06
Mammalia 18803 355.37 17870 456.49
Mammalia < hare 32 0.54 16 0.29
Mammalia > cattle/deer 6 27.3 81 223.8
Mammalia size of cattle/
deer
942 1333.4 909 2673.2
Younger Dryas Early Holocene
NISP Weight NISP Weight
(g)
Mammalia size of hare 13 2.71 7 8.2
Mammalia size of sheep 200 321.59 484 687.7
Total 21,170 27,371.01 20,923 47,355.4
Mollusca
Gastropoda 1 0.01
Unio tigridis 1 7.96
Unionidae 3 2.51 2 18
Total 5 10.48 2 18
Others
Brachyura 2 0.02
Potamon potamios 1 0.14
Vertebrata 3 0.12 6 0.01
Total 6 0.28 6 0.01
Pisces
Acanthobrama marmid 2 1.28 1 0.01
Arabibarbus grypus 0 0 1 2.1
Leuciscus vorax 3 2.5 0
Luciobarbus kersin 1 0.4 0
Luciobarbus subquin-
cunciatus
2 4.24 0
Capoeta trutta 1 0.08 0
Cyprinidae 161 60.8 528 197.07
Planiliza abu 1 0.01
Luciobarbus esocinus 14 15.73 10 99.95
Luciobarbus kottelati 3 0.45 1 4.1
Luciobarbus xanthop-
terus
1 0.48 1 0.6
Mastacembelus masta-
cembelus
1 0.01 2 0.19
Pisces 235 2.8 198 3.93
Total 425 88.78 742 307.95
Reptilia
Lacertidae 1 0.01
Ophisaurus apodus 1 0.83
Testudo graeca 18 266.44 5 367.7
Total 18 266.44 7 368.54
Aves
Anas acuta 1 0.9 1 1.1
Anas crecca 1 2.1 0 0
Anas penelope 0 0 1 0.8
Anas platyrhynchos 2 1.6 2 1.3
Anas querquedula 1 0.1 0 0
Anas sp. 2 1.1 4 4
Aythya fuligula 0 0 1 0.7
Anser albifrons 1 1.9 11 32.3
Anser anser 3 15.1 45 159.6
Anser anser | Anser
albifrons
0 0 3 10.2
Anser sp. 1 0.2 24 43.4
Branta ruficollis 0 0 3 5
Cygnus olor 1 15.2 0 0
Ardea cinerea 0 0 1 2.2
Botaurus stellaris 1 0.3 0 0
Ciconia sp. 0 0 1 2.4
Grus grus 2 3.2 1 7
Otis tarda 31 138.66 54 202.2
Numenius arquata 0 0 1 0.2
Gyps fulvus 1 4.7 1 3.7
Aquila heliaca 0 0 1 1.6
Haliaeetus albicilla 1 6.8 0 0
(Continued) (Continued)
Appendix 11. (Continued) Appendix 12. (Continued)

24 The Holocene 00(0)
Younger Dryas Early Holocene
NISP Weight NISP Weight
(g)
Buteo buteo 1 0.7 0 0
Circus aeruginosus 0 0 1 1
Circus aeruginosus |
Circus cyaneus
1 0.1 0 0
Circus cyaneus 1 0.2 0 0
Circus pygargus 1 0.1 0 0
Circus sp. 2 0.45 0 0
Circus sp. | Circus
cyaneus
2 0.3 0 0
Falco sp. 1 0.7 0 0
Falconiformes 0 0 1 0.2
Bubo bubo 10 20.3 5 16.5
Strix aluco | Asio otus 0 0 1 0.2
Perdix perdix 2 2 2 0.5
Columba livia 1 0.2 0 0
Streptopelia turtur 1 0.1 0 0
Sturnus sp. | Sturnus
vulgaris
1 0.1 0 0
Turdus sp. 1 0.01 0 0
Corvus sp. 3 3.22 0 0
Pyrrhocorax pyrrhoco-
rax
0 0 1 0.3
Passeriformes 1 0.01 0 0
Passeriformes | Prunella
sp.
1 0.01 0 0
Aves 38 1.29 35 2.2
Aves large 8 6.4 26 12.8
Aves middle sized 27 7.12 13 7.2
Aves small 9 0.45 1 0.1
Total 161 235.62 241 518.7
Grand total 21,785 27,972.61 21,921 48,568.6
Appendix 13. Number post-cranial bones fused in each age
category for wild goat (Capra aegagrus) based on (Zeder, 2006:107
fig.15).
Younger Dryas Early Holocene
Unfused Fused Unfused Fused
6–12 m 1 14 1 7
12–18 m 0 1 0 2
30–48 m 0 4 0 0
48+0 0 0 1
Appendix 14. Percentage of post-cranial bones fused in each
age category for Aurochs (Bos primigenius) based on (Habermehl,
1975: 104–105).
Unfused Fusing Fused
<20 months (Scapula, Coxa, Proximal
Radius, 2nd Phalanx, distal humerus)
5 0 11
20–36 months (1st Phalanx, distal Tibia,
Metapodials, Calcaneus)
11 2 20
>36 months (Proximal Femur, Humerus
and Tibia. Distal Femur, Radius. Proxi-
mal and Distal Ulna)
4 1 5
Appendix 15. Sus Scrofa post-cranial fusion from Early Holocene
Körtiktepe (n = 15). Age fusion stages and approximation of age of
fusion from Zeder etal. (2015), p. 139, table 3.
Fusion age class Early Holocene
Approximate age of fusion Unfused Fusing Fused
D7–8 m 2 2
E8–18 m 0 1
G24–36 m 0 1
H36–48 m 2 1 1
I48–60 m 4 1
Appendix 12. (Continued)