Asikli Hoyuk The Generative Evolution of

Journal of Archaeological Research
https://doi.org/10.1007/s10814-021-09167-z
Aşıklı Höyük: The Generative Evolution of a Central

Anatolian PPN Settlement in Regional Context
Mary C. Stiner1 · Mihriban Özbaşaran2 · Güneş Duru3
Accepted: 8 March 2021

© The Author(s), under exclusive licence to Springer Science+Business Media, LLC, part of Springer Nature
2021
Abstract
The first Neolithic settlements in Southwest Asia began with a dual commitment to
plant cultivation and a sedentary lifestyle. The benefits that foragers-turned-farm-
ers gained from this commitment came with some inescapable constraints, setting
new evolutionary pathways for human social and economic activities. We explore
the developmental process at the early Pre-Pottery Neolithic site of Aşıklı Höyük
in central Anatolia (Turkey), specifically the relationship between internal dynam-
ics and external influences in early village formation. Feedback mechanisms inher-
ent to the community were responsible for many of the unique developments there,
including domestication of a variant of free-threshing wheat and the early evolu-
tion of caprine management, which gave rise to domesticated stock. Gradual change
was the rule at Aşıklı, yet the cumulative transformations in architecture, settlement
layout, and caprine management were great. The many strands of evidence reveal
a largely local (endemic) evolution of an early Pre-Pottery Neolithic community.
However, burgeoning inequalities stemming from production surplus such as live-
stock likely stimulated greater regional interaction toward the end of the sequence.
Keywords Domestication · Niche construction · Social networks · Site formation

processes · Caprine management
* Mary C. Stiner
mstiner@arizona.edu
Mihriban Özbaşaran
ozbasaranmihriban@gmail.com
Güneş Duru
gunes.duru@msgsu.edu.tr
1
School of Anthropology, University of Arizona, P.O. Box 210030, Tucson, AZ 85721-0030,
USA
2
Prehistory Department, Faculty of Letters, Istanbul University, Fatih, 34134 Istanbul, Turkey
3
Faculty of Conservation and Restoration of Culture Property, Mimar Sinan Fine Arts University,
Cumhuriyet Mh. Silahşör Cd. No: 89 Bomonti, Istanbul, Turkey
13
Vol.:(0123456789)
Introduction
The forager-farmer transition, also called the Paleolithic-Neolithic or forager-pro-

ducer transition, is a complex story. Southwest Asia saw a particularly early emer-
gence of village societies, between roughly 11,000 and 9500 years ago (Fig. 1).
There were many geographic variants, yet these cases shared a tendency toward sed-
entism, plant carbohydrate dependency, and proliferation of resource intensification
strategies that included food storage and animal and plant management (e.g., Belfer-
Cohen and Bar-Yosef 2000; Willcox 2013). Rather than moving camp to solve soci-
oeconomic problems as their Pleistocene forebears typically would have done, set-
tled foragers turned to other solutions. One of the most fascinating features of the
forager-farmer transition is the juxtaposition of rapid innovation and the conserva-
tive influences of local history. Innovation requires openness to ideas while local
precedent (such as prior investments, infrastructure, social institutions) can be both
a powerful force for continuity and a highly discriminating filter. Innovation and his-
torical precedent did not function entirely in opposition to one another in emerging
Pre-Pottery Neolithic (PPN) societies, but these communities tended to look inward
for the many of the resources they needed most.
Disagreements about the initial circumstances and causal chains that gave rise
to the first villages and the shift from foraging to food production have occupied
Fig. 1 Map of selected Neolithic and Epipaleolithic sites in Southwest Asia and Cyprus. Inset at lower
right shows the original (black arch) and later (dark gray) geographic conceptions of the “Fertile Cres-
cent” area.
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archaeologists for decades (e.g., Bar-Yosef and Belfer-Cohen 2002; Clem and Serge
2017; Cohen 1985; Gremillion et al. 2014; Gronenborn 2009; Hodder 2018; Kennett
and Winterhalder 2006; Redding 2005; Robb 2014; Smith 2015; Vigne et al. 2011;
Zeder 2009). There is a parallel dispute about what ancient people might have antic-
ipated as the pros and cons of such profound changes (e.g., Abbo and Gopher 2020;
Cohen 2009; O’Brien and Bentley 2015; Zeder 2012a). And, there is the larger ques-
tion of how many times early farming communities emerged quasi-independently
within the narrow time frame of the Pre-Pottery Neolithic (Asouti 2006; Fuller et al.
2012; Rosen 2007; Russell 2011; Vigne 2011; Vigne et al. 2012). Most archaeolo-
gists agree that sedentism had little if any evolutionary precedent for the human spe-
cies and that sedentism facilitated the evolution of highly distinct microhabitats on
the landscape from the end of the Pleistocene through the Holocene (e.g., Goring-
Morris and Belfer-Cohen 2008).
It is clear that early PPN communities arose within regional patchworks that
included mobile and settled forager groups. This situation made for a more heteroge-
neous social landscape than would have existed during the Pleistocene. Information
must have continued to flow between these diverse groups, yet some communities
changed or intensified faster than others. Those that changed most during the early
PPN maintained surprisingly conservative material culture traditions. The plethora
of distinctly local developmental trajectories during this formative period in South-
west Asia suggests that archaeologists’ traditional search for single points of ori-
gin and “the first” animal or plant domestication events is a vain exercise (Arbuckle
2014; Arranz-Otegui et al. 2016; Colledge and Conolly 2007; Fuller et al. 2011).
The local mechanics of PPN village emergence are just as controversial as the
general problem of when, where, and how many emerged at once. Many archaeolo-
gists rightly note “endemic” developmental trajectories as they delve into the details
of one site versus another. How can any overarching theory of change account for
such diversity? Or should we seek common causes at all? Part of the answer to this
question may be had by examining the mechanics of change, as well as some of the
starting conditions in PPN societies. Important practical challenges for archaeolo-
gists are identifying how and when the relative influences of internal dynamics and
external phenomena shifted in these fast-evolving systems. Also critical to under-
standing PPN emergence in Southwest Asia is recognizing which aspects of basic
PPN know-how trace back to the local Epipaleolithic cultures before them (e.g.,
Benz et al. 2015), carried over from mobile to increasingly sedentary patterns of
land use.
We explore these points by taking a close look at the developmental evolution
of Aşıklı Höyük in its larger regional context. Founded on a knoll in the Melendiz
River floodplain roughly 10,400 years ago, this early PPN site grew into a 4 ha
mound that stood 16 m high by the time of its abandonment 1000 years later (Esin
and Harmankaya 1999; Özbaşaran 2011a, b; Özbaşaran and Duru 2015; Özbaşaran
et al. 2012, 2018). Aşıklı is fascinating because, for one thing, it lies well outside the
original perimeter of the “Fertile Crescent” as defined by Breasted (1914), which
did not yet include Anatolia (Fig. 1 inset). Aşıklı nonetheless dates relatively early
among the known PPN sites in Southwest Asia, and it was the first major aggregated
village settlement in central Anatolia of which we know. The continuous cultural
13
sequence represented by Level 5 through Level 2 has yielded a rich and detailed
record of architectural transformations, plant cultivation, domestication of free-
threshing wheat, and rapid evolution of sheep and goat management practices. All
the while, humans made and remade their “domestic” environment within the site.
The settlement also presented novel microenvironments for a variety of small organ-
isms, some of which became major agricultural pests. The benefits that foragers-
turned-farmers gained from plant cultivation and a more sedentary lifestyle came
with some inescapable constraints. Sedentism, for example, largely eclipses the tra-
ditional solution of moving one’s residence to alleviate food shortages and social
stresses. The advantages and challenges that arose in one aspect of settled human
existence must have had consequences for other domains of their lives, forcing or
favoring reorganization of social space and technology. And these changes, them-
selves, had future consequences for other domains of human existence.
Archaeologists increasingly are embracing generative evolutionary perspectives
as conceptual structures for understanding human evolution in all periods. These
perspectives vary considerably in focus but nearly all appeal to co-evolution or
feedback-based interactions as formative forces (reviewed in Stiner 2021). Niche
construction theory (NCT) is particularly discussed because humans radically alter
the organic and inorganic world they inhabit, and this altered world is inherited by
later generations (Laland et al. 2000). Most archaeological applications of NCT
are more paradigmatic than analytical—a general working concept rather than an
explicit basis for hypothesis testing (Stiner 2021; but see, e.g., Laland and O’Brien
2010; Murray et al. 2021; O’Brien and Laland 2012). We hope to do a bit better here
by exploring the developmental feedbacks between human socioeconomics and the
built environment at Aşıklı, emphasizing how certain factors appear to have inter-
acted dynamically while others were more constant, and by gauging the sequence
and magnitude of their respective influences.
The continuity and high quality of preservation of the early PPN record at Aşıklı
allows us to trace 1000 years of change in behavior, economics, and social organiza-
tion at a relatively fine scale. Multidisciplinary studies of mound formation dynam-
ics and construction, for example, reveal nonlinear increases in sedimentary and
biochemical inputs over time, including dung and urine salts from humans and live-
stock, along with recycling of various materials. Other strands of the story, such as
human-animal and human-plant interactions, human diet, child survivorship, mor-
tuary practices, and technology, show different rhythms of change relative to the
transformations of the physical environment of the settlement. Two threads of this
story—architecture and caprine management—were particularly volatile, and inter-
actions between them had profound impacts on the settlement, sometimes in oppo-
sition to social demands. The larger regional context of the Pre-Pottery Neolithic
is also critical to understanding these and other dynamics, especially the degree of
connectedness between communities in the region. Finally, we hope that this essay
can also illustrate some conceptual and technical tools for integrating disparate lines
of evidence.
After providing background on the site and its regional context, we consider
the dynamism of the mound itself. The history of construction, debris accumula-
tion, and disturbance reflects the nature and scale of human activities and other
13
biotic processes within the evolving settlement. We then examine how a number of
dynamic processes might be connected in the development of Aşıklı Höyük and the
strength of those connections. This effort is divided according to four major themes:
human-plant and human-animal co-evolution; the phenomenon of village emer-
gence and the power of human-environment feedback mechanics; social dimensions
of food production and storage; and networks versus sole-source models of Neo-
lithic emergence. Crosscutting these themes are considerations of human health and
diet, space use, mortuary practices, and technologies. We close with a comparison
of the developmental history of the central Anatolian Neolithic settlements to those
elsewhere Southwest Asia, particularly within the Euphrates River system and the
Levant.
Background
Aşıklı Höyük (AH) is the earliest known PPN settlement in central Anatolia (Duru
2002, 2013; Esin and Harmankaya 1999; Özbaşaran 1998; Özbaşaran et al. 2018).
This large mound site is situated on the Melendiz River, roughly midway between
the western margin of the Tuz Gölü Basin and the majestic Hasandağ Volcano in
Cappadocia (Fig. 2). The site was founded around 10,400 years ago and was occu-
pied continuously for about one millennium (Quade et al. 2018). The Cappadocian
landscape is overwhelmingly volcanic, which accounts for its distinctive topogra-
phy of isolated mountains, basins, and “fairy chimney” formations. Ancient volcanic
flows and massive ash fall deposits have been carved over millions of years by tec-
tonic disturbance and erosion (Doğan et al. 2019). Natural areas in the region sup-
port diverse plant communities that generally divide into open woodlands, grassy
steppes, and riparian corridors that cut green ribbons through the Anatolian Plateau.
Rivers and springs fed by snow melt are the main water sources for humans and
Fig. 2 Location of Aşıklı Höyük on the upper Melendiz River (white line), which drains the Melendiz
massif and flows northeastward into lower relief area where Aşıklı Höyük is situated. Obsidian sources
are indicated by smaller dots. Summit of Hasan Dağı volcano is 3268 m asl; Aşıklı Höyük lies at 1118 m
asl (base map adapted from GeoMapApp)
13
animals during the hot dry summers. It was, however, a greener landscape in the
early Holocene than it is today (e.g., Allcock 2013; Kuzucuoğlu et al. 2011).
The Melendiz River valley underwent significant reworking during the Late Pleis-
tocene, changes that had important ramifications for the future inhabitants of Aşıklı.
The river developed a rich, linear marshland toward the end of the Late Pleistocene,
which triggered clay sedimentation that eventually formed a wide, flat valley floor in
the area of the site (Kuzucuoğlu et al. 2018). A drying trend beginning just prior to
the Holocene allowed fertile soils to develop in the valley bottom and on the lower
colluvial terraces. Aşıklı was founded on a low rise in the floodplain, surrounded
by these large tracts of rich moist soil (Kuzucuoğlu et al. 2018). A permanent set-
tlement on the river was unprecedented in the area. Mobile hunter-gatherers of the
region set their camps near permanent water in earlier periods but well above the
youngest river terraces (Baird 2012a, b; Baird et al. 2018; Balkan-Atlı et al. 1999;
Duru 2018a; Duru and Kayacan 2018). It was the early Aşıklans’ interest in plant
cultivation that led them to settle directly on the low river terraces.
Thirty years of archaeological research at Aşıklı has combined a wide range of
scientific and humanistic approaches to what “neolithization” meant for the human
inhabitants and their environment, beginning with the work of Prof. Esin (Esin and
Harmankaya 1999). New scientific methods have been integrated into the research
program since 2010 (Özbaşaran 2011a; Özbaşaran and Duru 2018), including
micro-archaeology (phytolith and residue analyses, sediment micromorphology),
ancient DNA analysis, isotope and sediment geochemistry, and vertebrate taphon-
omy. Systematic fine screening and flotation are practiced throughout the new exca-
vation, which has focused mainly on the earlier occupations (Levels 3, 4, and 5)
but includes further sampling in Level 2. These new applications have been “game
changers” for what can be learned from the site. Meanwhile, decades of systematic
study have produced large bodies of evidence about architecture, macrobotanical
remains, faunal remains, and various technologies. Although the project is ongo-
ing, and not all lines of evidence have reached maturity, the available results war-
rant a synthetic consideration of stasis and change in the history of this early PPN
settlement.
Seventy-seven 14C dates indicate a temporal span of 8350 to 7350 cal BC for the
PPN occupations at Aşıklı (Quade et al. 2018). Most critical are the dates obtained
from charred short-lived plant materials, such as burned twigs and annual seeds, that
were collected from intact hearths and other discrete combustion features (SI Fig. 1).
The cultural chronology can be divided into three general architectural horizons
(Özbaşaran et al. 2018): Level 2 spans 7750 to 7350 cal BC (9.7–9.3 kyr BP); Level
3 spans 8050–7750 cal BC (10–9.7 kyr BP); and Levels 4–5 span >8350–8050 cal
BC (>10.35–10 kyr BP). Dating work on Level 5 continues, but it is clear that this
first occupation is only slightly older than that of Level 4 and may not have been
fully sedentary. Uncertainties of ~200 years surround the transitions between Levels
2–3 and Levels 3–4 due to plateaus in the Holocene radiocarbon calibration curve.
Fortunately, these plateaus do not greatly affect the dates for Levels 4 and 5, when
atmospheric 14C production rates were more uniform. The dates for Level 5 place
the first Aşıklı settlement fairly early in the PPN chronology for Southwest Asia
more generally.
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Mound formation at Aşıklı was closely tied to trends in architecture and spatial
layouts (Duru 2013), which provide a suitable backbone for exploring connections
between space use, economic organization, and social relations over the ensuing
1000 years. The architectural sequence exhibits a transition from rounded to rec-
tangular building contours (SI Fig. 2) and from semisubterranean to exclusively
surface-built residential structures. In this regard, the Aşıklı sequence adheres to
general PPN trends described by Moore (1985) and others (Banning 2003; Özdoğan
1997; Özdoğan and Özdoğan 1989; Stordeur et al. 1997). At Aşıklı, the shift from
a quasi- to a fully permanent settlement was early and quick (from Level 5 to 4),
whereas the shift from rounded semisubterranean to aboveground rectangular build-
ings was gradual and spanned several centuries.
Özbaşaran et al. (2018) identify three provisional organizational stages from
the dominant construction techniques, floor plans, and density of buildings. The
basal cultural layer, Level 5, is provisionally called the Wattle-and-Daub Stage
and lasted two or three centuries at most. This occupation rested directly on the
river alluvium and seems to represent an incipient stage of sedentism. The semi-
subterranean round buildings in Level 5 were sunk deep into the ground, with
roofs standing <0.5 m above the surface. The buildings were made of wooden
posts interwoven with reeds and branches and plastered with kerpiç—a rec-
ipe of mud, tempers, and binding agents (Fig. 3). The floors of these buildings
received several fine coats of mud plaster, but renewals were fewer in number and
the structures less robust than in Level 4. Aboveground structures, constructed
mainly of wood and reeds, were also present in Level 5. Some of these contained
Fig. 3 Plan of Levels 5 and 4 in Trench 4GH at the northern edge of the mound: Level 4 contains Build-
ings 2, 3, and 6, along with activity Spaces 7–9 among others; partly exposed Level 5 contains Buildings
34 and 35 (underneath B.6), burnt circular Structure 33 (under B.2) and Structure 38, and the wattle-and-
daub residential Building 27 that is partly visible under Building 3. Also present in Level 4 were several
pits (removed, but see outlines), the largest of which was filled with cracked cobbles and a discrete char-
coal concentration at one side. Large outdoor activity spaces separate the structures in these early levels
(Aşıklı Höyük Project Photo Archive).
13
remarkable concentrations of seed remains, indicating frequent processing activi-

ties and storage before burning down (Ergun et al. 2018). Other structures, or
layers within them, contain modest concentrations of herbivore dung in primary
context (defecation in situ), based on micromorphological (Mentzer 2018), multi-
element geochemical (Kalkan and Özbal 2018), and phytolith evidence (Tsartsi-
dou 2018). Every one of the aboveground structures was eventually burned to the
ground (SI Figs. 1a and 3), and most were replaced with outdoor activity areas.
The spatial distributions of artifacts and other materials indicate frequent rotation
of outdoor activities that included tanning, butchering, obsidian knapping, cook-
ing, and seed processing (Astruc and Grennet2018; Stiner et al. 2018).
The Oval Buildings Stage in Levels 4–3 clearly represents a permanent set-
tlement. The deposits are thick and minor trends are expressed within them. The
dwellings were semisubterranean as before, but the aboveground walls were
somewhat taller and made of durable sun-dried kerpiç blocks and mortar, rather
than wattle and daub. The use-life of the residential buildings increased in this
period, and their walls and floors were replastered many times over. The inter-
nal features of the residential buildings typically included a central hearth, one
or two large grindstones (some embedded in raised kerpiç benches), small work
areas around the hearth, and human burials under the plastered floors (SI Fig. 4).
Some dwellings contained multiple burials, each placed separately under one or
a succession of floors. Storage pits inside the buildings were small, and shallower
depressions probably supported large baskets. The interiors were spacious, with
an average diameter of about 4 m. The roofs were supported by a radial arrange-
ment of beams that extended from a central post to the outer walls, and these
were covered with a thick layer of reeds and kerpiç. As in Level 5, the residential
buildings of Level 4 were spaced well apart from one another, with ample out-
door activity areas between them (Fig. 3).
Level 3 represents a continuation of many of the architectural characteristics of
Level 4 but with significant upticks in community growth, architectural density,
and debris accumulation. Thick, weathered herbivore dung deposits cover a large
fraction of the top of Level 3 (Mentzer 2018). A shift from overwhelmingly cir-
cular/oval buildings to a mosaic of roundish and quadrangular ones occurred in
the later phases of Level 3 (SI Fig. 2). One structure stands out from the rest due
to its much larger diameter and internal features—a semisubterranean oval struc-
ture (Building 38) that was designed for communal use (SI Fig. 5). Though con-
structed in a manner grossly similar to the residences, the unique internal features
of Building 38 argue against the possibility of it simply being a grander residence
than the other domiciles. Building 38 has benches or platforms along portions of
the interior walls, the surfaces of which are embedded with small pits. No sub-
floor burials have been found. Botanical evidence indicates that plant process-
ing was one of the activities conducted inside the building (Ergun et al. 2018).
The building may have been a gathering area for daily collaborative work and
for social or ritual events. Building 38 was reconstructed more than seven times
on the same footprint, and the floors of each phase were plastered many times
over, indicating that upkeep of the building was demanding and probably done
cooperatively.
13
A longstanding question at Aşıklı concerns the position of entryways to the semi-

subterranean buildings. The excavators never find door-sized openings in the outer
walls. Heating semisubterranean buildings with fireplaces nonetheless requires that
oxygen be brought in at floor level and allowed to exit through the roof. Such ven-
tilation systems are widespread in Levels 5–3 and provide the most compelling evi-
dence for rooftop entryways. Not unlike the ventilation systems of kiva structures
in Ancestral Puebloan sites of the American Southwest, oxygen needed for the fires
was pulled into each Aşıklan building through a semi-vertical shaft along the outer
wall (SI Fig. 6). This ventilation method continued until semisubterranean structures
fell out of use in Level 2.
Subfloor human burials are present throughout Levels 5 through 3, and they lack
grave goods as a rule (Yelözer 2018). Each human body was placed in a pit that was
cut through an existing house floor, usually as a tightly flexed, complete skeleton.
Some of the bodies, especially children, were wrapped in mats made from reeds and
other plant fibers. The floor was replastered once the grave pit was filled, and life
in the building resumed as normal. The practice suggests a powerful symbolic con-
nection between deceased individuals and particular residential buildings from very
early in the history of the settlement. Levels 5 through 3 also contain various above-
ground structures that were not used as residences. Some of them were makeshift
animal pens, others sheltered work areas, and still others may have been dedicated to
food storage. An unusual quadrangular structure (Building 2) in Level 4 was made
by pouring wet kerpiç into an improvised mold, resulting in dense impervious walls
(SI Fig. 7); similar structures has been found recently on the west side of the mound
and none has residential features. Some of the floor layers within Building 2 are rich
in plant food residues; they also contain shallow pits that may have been for seating
storage baskets (SI Fig. 7).
The third and most recent Rectangular Buildings Stage spans Levels 2 through
1 and signals a major reorganization of the settlement. There is little left of Level
1 in the areas that have been excavated, although it is preserved northwest of the
excavation area. Much of Aşıklı’s fame rested until recently on the vast archaeologi-
cal exposures of Level 2 (Fig. 5, SI Fig. 8) accomplished by Esin and her colleagues
(see Esin 1998; Esin et al. 1991). About 100 structures have been exposed in Level
2 (Duru 2013), and this thick layer preserves more than 10 phases. The density of
residential buildings is much greater than before and, at least by the later phases, a
separate group of large special purpose structures was erected. The dwellings were
quadrangular as a rule and constructed on the ground surface of kerpiç block and
mortar, without stone foundations. Standing about 2 m above ground level, the flat
roofs were supported by long wood crossbeams covered with reeds and mud.
The houses of Level 2 had long use-lives, and they were rebuilt completely every
30–50 years on average, usually in the same footprint. The fidelity of buildings to
their original footprint, and the sequences of subfloor burials therein (Fig. 4), indi-
cate a continuation of the symbolic connection between residential buildings and
at least some of their occupants. The number of buildings on the mound increased
rapidly during Level 2, eventually forming dense clusters separated by a few open
areas and passages (see Fig. 5, SI Fig. 8). Individual buildings contained one or two
rooms, rarely three, which were connected internally by doorways at floor level.
13
Fig. 4 Example of many superimposed footprints of a Level 2 residential structure, exposed in the south
wall of Trench 4GH (adapted from Özbaşaran et al. 2018)
Fig. 5 Aerial view of densely packed rectangular buildings in Level 2, exposed on the western side of the
mound. Earlier oval buildings of Level 3 are visible downhill at the extreme left. Burnt remains of Build-
ing 39 in Level 5 are also visible at the base of the slope, in contact with the underlying alluvium (Aşıklı
Höyük Project Photo Archive).
However, there are no doorways through the outer walls. Rectangular hearths lined
with flat cobbles were placed in a corner of the main room (SI Fig. 9), with a small
opening in the lower wall for air intake. As in earlier periods, human burials were
placed in subfloor pits, some covered or wrapped in plant fiber mats. Only in the
later phases of Level 2 (especially 2C-2A) do burials include beads of many types,
sometimes in large numbers but without strong biases by gender or age (Yelözer
2018).
A remarkable group of large, special purpose buildings are situated on the south
end of the mound (SI Fig. 8). We cannot be sure just when this special area was
13
established within the Level 2 sequence, because the earlier phases remain buried.
These ceremonial or public buildings are distinguished by their much larger sizes
and unique construction methods and designs, including dry-lain stone and plastered
interiors painted in vivid shades of red and yellow ochre (Özbaşaran and Duru 2018,
pp. 4–8). The highly specialized complex of Musular, a shallow special purpose site
just across the Melendiz River from Aşıklı (SI Fig.10), also appeared around this
time (Özbaşaran 2000; Özbaşaran et al. 2012).
In summary, Levels 5 through 2 represent a continuous evolution of the settle-
ment, without significant hiatuses in human occupation. The changes were gradual,
but the pace of community growth accelerated with time. Rather than spilling freely
onto the adjacent colluvium above the floodplain, the community tended to pack
itself more densely on the ancestral settlement footprint. The appearance of the set-
tlement thus changed radically from its beginning to its end, 1000 years later.
The Living Mound
By the time of its abandonment, Aşıklı Höyük had grown into a very large mound
composed almost entirely of anthropogenic sediments. Some layers within the
deposits are so dense that they have sealed earlier layers from percolating water.
This situation likely explains the high quality of collagen preservation in the human
and animal bones in the site. Much of the mound is intact, aside from agricultural
tilling of its surface, but a meander of the Melendiz River eventually cut into the
western side (SI Fig. 10).
The Aşıklı mound grew from the relentless accumulation of human and animal
waste of all sorts—debris from daily work and food preparation, dung from captive
live animals, and construction debris dominated by kerpiç blocks, mortar, and mud
plasters. Carbonate minerals are naturally rare in the surrounding volcanic landscape
but are a major constituent of the mound deposits, originating as wood ash from
fireplaces, burned structures, and burned surfaces. The larger the Aşıklı community
grew, the more waste it generated, gradually raising the site high above the valley
floor. Shifts in construction methods may have increased the ratio of construction
waste disproportionately in the later periods, although the continuous borrowing of
old kerpiç to make new blocks and mortar would have damped this effect. Humans
not only created the mound from their many activities, they modified it continu-
ously. Large-scale disturbances within the mound reflect episodes of building, refur-
bishment, clearing, and leveling. Fine-scale disturbances in the form of pits, activity
surface maintenance, and interments of the dead are also common. It is for these
reasons that mound sites present both rich opportunities and daunting challenges to
archaeological interpretation. The great mass of the sediments makes it difficult to
expose deeper cultural layers, and the stratigraphy is complex on several scales. At
Aşıklı and elsewhere, the edges of the mound were unstable, their surfaces changing
continuously as the result of gravitational, sedimentary, and anthropogenic forces.
These many disturbances can be read by archaeologists—sometimes with great
interest and in other cases with dismay—from detailed considerations of the strati-
graphic, radiocarbon, and micromorphologic records of the mound sediments.
13
The geoarchaeological evidence shows us just how “alive” the Aşıklı mound
environment was during the human occupations, both physically and biologically.
Humans are avid bioturbators in general, and Neolithic people, particularly those
who used wattle-and-daub or mudbrick construction, were champions at this. Kerpiç
was the dominant construction material at Aşıklı. Mud was mined from exposures
on the river near the settlement and mixed with a variety of additives. This mix-
ture was used for wattle-and-daub and later mudbrick-and-mortar constructions, and
for plastering walls and horizontal surfaces. The additives varied considerably with
time and application; some mixtures contain cereal chaff and others dung, wood ash
and/or lime (Mentzer 2018). All of these materials contributed greatly to the sedi-
mentary bulk of the mound.
Ungulate dung is another major component of the mound sediments (SI Fig. 11),
appearing in both primary and secondary accumulations. As caprine management
practices evolved (discussed below), the scale of human investment increased expo-
nentially. This is apparent from, among other things, major increases in the accu-
mulation of urine salts on the mound from humans and their livestock (Abell et al.
2019). Three key components of urine are sodium [Na+], chlorine [Cl-] and nitrite
[NO3-]. These elements can also occur naturally in soils, but their concentrations
and ratios in the sediments of Aşıklı are diagnostic of urine. The attribution to urine
is further confirmed by nitrogen isotope signatures in the nitrate (δ15N values per
mil), which closely match values for modern livestock feedlots and are even higher
than typical human septic systems (see Abell et al. 2019, fig. 5). Variation in salt
element concentrations at Aşıklı is summarized by level in SI Figure 12a. The data
are then modeled to reflect increasing rates of sediment accumulation with time
in SI Figure 12b, based on estimated increases in population, intensity of building
activity, and greater volumes of mudbrick per building. Most remarkably, Abell
et al. (2019) document a 10–100x increase in [Na+], [Cl-], and [NO3-] from Level
5 to Level 4, and a 100x–1000x increase from Level 5 to Level 3. The urine inputs
greatly outstrip architectural evidence of human population density in each layer and
thus much of it had to have come from livestock. Urine inputs declined somewhat
from Level 3 to 2, even though the percentage of caprines in the meat diet continued
to increase (Fig. 4c). This and other evidence tells us that livestock corrals were
moved partly or wholly off the mound top around this time, making way for more
residential buildings.
Recycling of materials, such as mud, wood ash, dung, and possibly wood (though
termites limit the use-life of wood in the area), became routine at Aşıklı. Dung
was an important binding agent in mud construction, fuel for fireplaces (Mentzer
2018), and probably also fertilizer for gardens. What waste could not be recycled
was moved into middens. Debris other than fine fragments was seldom allowed to
remain where it first fell to the ground. The fact that the middens became larger and
more communal with time reflects not only an increasing number of consumers but
also the need for greater cooperation in managing waste within the settlement, pre-
sumably so that the residents would not drown in it.
The dynamic nature of the Aşıklı mound—in fact any Neolithic mound—poses
complex challenges to radiocarbon dating its components (Quade et al. 2018). While
the overall stratigraphy of Aşıklı Höyük follows a coherent temporal order based on
13
dates of discrete combustion features, localized 14C anomalies are common. Some of
these anomalies can be blamed on humans reworking the deposits and slumping or
sedimentary inversion near the mound edges. In other instances, however, the prob-
lems arise from contamination of charcoal samples with old carbon that originated
from the very muds that people used to build their structures. This latter problem
is most evident in Levels 5–4, which yielded a few dates of >11,000 years before
present. It is tempting to take the oldest dates at face value, but further analysis
revealed a strong correlation between age and exceptionally low carbon content (so-
called “dirty” samples; Quade et al. n.d.). The implication is that what little carbon
is present in the samples is disproportionately influenced by older carbon inclusions.
Rather than arising from the stereotypical “old wood” problem, the 11,000-plus 14C
dates reflect contamination of samples with microscopic “old carbon” bonded to
clays within the radiocarbon samples. Importantly, the contamination sources only
slightly predate the formation of the archaeological site, namely the terminal Pleis-
tocene clayey sediments that underlie the silty riverbanks and which the Aşıklans
sometimes mined for construction materials. This type of contamination in 14C sam-
ples is widely discussed in soil science (Orlova and Panychev 1993) but seldom in
the dating of Neolithic sites. In the absence of careful control on clay-bonded con-
taminants, the chances of overestimating the antiquity of an early settlement is quite
high. One may wonder how often this happens in early PPN sites in the Middle East,
because of the very kinds of soils on which early cultivators were inclined to settle
and people’s proclivity for using clayey muds as a building material. The problem
is especially relevant for dating the earliest sedentary communities, simply because
PPN people sought out organic-rich Late Glacial soils for cultivating plants.
Human–Plant and Human–Animal Coevolution
A variety of co-evolutionary processes are evidenced at Aşıklı Höyük in the context

of human–plant, human–nonhuman animal, and also human–environment (niche-
constructing) interactions. Many of the biological interactions were essentially pred-
atory, but some were commensal, and a few developed into mutualistic relationships
that included domestication. These interactions are distinctive in that they led to
reciprocal adaptive changes as the result of strong interdependence between humans
and certain other organisms. Predator–prey interactions by definition involve nega-
tive or subtractive relationships, whereas commensals can be benign or somewhat
parasitic. Mutualisms, on the other hand, involve strong positive benefits to each of
the interacting species. Domestication is but one kind of mutualism, a rather extreme
type wherein the interacting species evolve heritable phenotypes that allow them to
thrive in settings that are substantially altered and constrained by one of the partners.
It is important to appreciate that “cultivation” of crop plants and “management”
of prey animals by humans can be pathways to domestication, but things don’t often
get that far (Clutton-Brock 2008; Larson and Fuller 2014; Larson et al. 2014; Zeder
2006). Whereas genetic studies focus mainly on the aftermath of multigenerational
selection and the evolution of clear-cut domestication phenotypes, archaeologists
have the ability to examine some of the initial conditions and developmental contexts
13
of domestication, before many of the obvious genetic and physiological alterations

have emerged. Here we begin with the human–plant interactions at Aşıklı, and then
turn to human–animal interactions. The human–environment interactions, also of
great interest in this paper and strongly related to the biotic interactions, are taken up
in the next section.
Human–Plant Entanglements
Sedentism and the cultivation of annual plants have tended to develop hand-in-hand
in forager societies (see cross-cultural surveys by Binford 2001; Keeley 1995; Kelly
1995). A wide variety of species, including large-seeded annuals, occur naturally
in the plant communities around Aşıklı, and the early Neolithic inhabitants took
advantage of this botanical diversity for food and other resources. Ethnoarchaeo-
logical research conducted in the modern village of Kızılkaya (Ertuğ-Yaraş 1997,
2000), only 1 km from the archaeological site, illustrates the potential richness of
this botanical larder for wild foods, raw materials, fodder, medicinal plants, and fuel.
Macrofossils and phytoliths of many of the species that villagers use today occur
throughout the cultural levels of Aşıklı (Ergun 2016; Ergun et al. 2018). In addi-
tion to cultivating cereal grasses (wheat, Triticum; barley, Hordeum) in moderate
quantities, the Aşıklans also cultivated pulses (lentils, Lens; chickpea, Cicer; pea,
Pisum; bitter vetch, Vicia ervilia) and other annuals. They gathered wild fruits, nuts,
and resins such as bitter and common almonds (Amygdalus), hackberries (Celtis cf.
tournefortii), and mastic from Pistacia atlantica trees (Ergun et al. 2018; Van Zeist
and de Roller 1995, 2003). They also collected wood and leaves for fuel, wild grass
fodder for livestock, reeds (Phragmites australis) and sedges (Capericeae) for bas-
ketry and building construction, and herbs and small shrubs for special uses.
Much of the evidence of plant use at Aşıklı relates to daily agropastoral activi-
ties, but plants served many purposes. The widespread presence of Verbascum (mul-
lein), Taeniatherum (medusahead grass), and Brassicaceae (mustards) at Aşıklı sug-
gests deliberate collection by the residents. Other wild plants in the assemblages are
Bellevalia (hyacinth relatives), Galium (bedstraw), Heliotropium (borage), Malva
(mallow), Thymelea (thyme), Papaver (poppy), and Valerianella (cornsalad). Some
of these taxa may simply have been weeds that grew among the crops in cultivated
fields, but all are valued in many recent cultures as food and/or medicinal plants.
Poaceae grasses are especially common among the wild plant remains, and they
could have been used mainly as animal fodder and perhaps also as bedding material.
Native cereal grasses in Anatolia include several species of wild wheat and barley
(e.g., Özkan et al. 2011; Willcox 2005).
According to Willcox et al. (2009), the natural distributions of wild cereals were
not greatly affected by climate change from the Younger Dryas to early Holocene.
However, the greater climate stability of the early Holocene increased the reliability
of intensive cultivation of the cereal grasses. This is the time when the Aşıklı set-
tlement came into being. The inhabitants’ commitment to cultivating plants at this
location became the anchor for myriad social and economic decisions over the next
1000 years of occupation.
13
The plant diet at Aşıklı was most diverse during the earliest occupations, at
least in terms of evenness in the representation of species, but there was only
a mild decline in plant diet diversity with time (SI Figure 13b). This decline is
explained by somewhat greater use of cereals and pulses in particular (SI Fig-
ure 13a). A combination of botanical (Ergun et al. 2018; Tsartsidou 2018) and
technological evidence (e.g., sickle microblades and milling stones, Astruc and
Grennet 2018) tells us that on-site cereal processing—from threshing and win-
nowing to grinding the kernals—were common activities in the settlement from
its beginning. The cereal assemblages include emmer wheat (T. turgidum dicoc-
cum), einkorn (T. monococcum monococcum), free-threshing wheat (T. aestivum/
durum), “new glume” wheat (defined by Jones et al. 2000; see also Czajkowska
et al. 2020), and wild and domestic forms of barley (Hordeum distichum/vul-
gare). The wheat assemblages are a mixture of wild and “domestic” variants in
all levels, but the proportion of the latter increased mildly with time. The ris-
ing frequency of “domestic” variants is taken to reflect selection within the con-
text of human exploitation, whether deliberate or an artifact of changes in the
ease of collecting and processing the grains. Specifically, the domesticated wheat
variants have a tougher rachis that suppresses the spontaneous shattering of the
ripened seed heads (Tanno and Willcox 2012; Willcox 2013). This trait gives
humans more flexibility in the methods and timing of harvesting. As such, the
cereal assemblages from Aşıklı point to an early stage in the wheat domestication
process. There are also mild increases in barley species in the later levels (e.g.,
SI Fig. 13c, based on chaff remains), including domestic barley (see Ergun et al.
2018).
Van Zeist and de Roller (1995, 2003) were the first to propose cereal domestica-
tion at Aşıklı based on material from Level 2. New data obtained from Levels 5–3
over the last decade demonstrate that wheat domestication was underway from the
earliest phases of occupation, hundreds of years earlier than expected (Ergun et al.
2018). The “new” glume wheat and certain other wheat variants at Aşıklı indicate
the presence of tetraploid hybrids from an early date (Ergun et al. 2018). For the
moment at least, Aşıklı represents the earliest occurrence of the new glume wheat
type in Anatolia, and it ranks among the early examples in Southwest Asia more
generally. Domesticated new glume wheats eventually came to dominate later Neo-
lithic cultivation of wheats in central Anatolia, such as at Çatalhöyük on the Konya
Plain (Bogaard et al. 2013). Wheat domestication in central Anatolia proceeded
slowly and gradually, probably because seed stocks for sowing were replenished
partly from the wild (Tanno and Willcox 2006a; Willcox et al. 2009). The persis-
tence of wild morphotypes in the assemblages could also stem from continued col-
lection of wild cereals, such as einkorn.
In addition to cereals, Level 5 contains remains of at least four species of pulse—
chickpea, pea, lentil, and bitter vetch. The latter two probably were cultivated con-
sistently by the Aşıklans, rather than just gathered from the wild, although the evi-
dence is less clear-cut than for cereals. On the other hand, the relative frequency of
pulses in the assemblages clearly increases in tandem with cereals from Level 5–4 to
Level 2 (SI Fig. 13a), and so the two groups of seed-producing plants seem to have
been nutritional complements.
13
There is also the important question of local (endemic) versus imported (intro-
duced) cultigens at Aşıklı. Bitter vetch and einkorn wheat occur naturally in central
Anatolia today, and pea species have been reported in some areas, though not near
Aşıklı. Early Holocene distributions of these and other species are not known and
could have been somewhat different than today. However, the most common crop
species at Aşıklı was emmer wheat, which is not native to the region but probably
did occur naturally in southeastern Turkey. Lentils are also non-native (Ergun et al.
2018). Emmer wheat and lentils thus appear to have been introduced to Aşıklı from
outside the region, and this happened quite early in the history of the settlement
(Levels 5–4). Rare chickpea remains have also been noted in the earliest levels and
may have originated from a distant source, perhaps as part of a triad of emmer, len-
til, and chickpea (Ergun, personal communication, 2018; Tanno and Willcox 2006b).
We conclude from the early use of cereals and pulses that the founders of Aşıklı
Höyük were dedicated cultivators, who were attracted to the well-developed soils
of the Melendiz floodplain for this purpose. The establishment of this site on the
river represents a significant departure from simply gathering seeds from wild cereal
stands, as foragers had done previously in central Anatolia. Most human–plant inter-
actions at Aşıklı remained stable for the remainder of the stratigraphic sequence.
When it comes to technological uses of plants at Aşıklı, the macrobotanical and
phytolith evidence is just the proverbial tip of the iceberg. Archaeologists catch
small glimpses of what were pervasive and massively important parts of the built
environment, technology, and raw materials in daily PPN life. The great abundance
of, and wear patterns on, osseous tools at Aşıklı testify to widespread fiber work-
ing, and to production of basketry and matting in particular (R. Christidou, personal
communication 2017). Artifacts made from plant fibers (mainly reeds, sedges, and
certain wild grasses) covered many living surfaces within the settlement, enclosed
human bodies, held stored food and other objects (SI Fig. 14), and were components
of human clothing. The phytolith evidence makes plain the great volumes of plant
material inputs into the sediments on the mound as the result of habitual use as fuel,
construction, fodder, storage, and so on. The bulk of this organic contribution has
since disappeared, but inorganic traces in the form of phytoliths are found every-
where in the sediments.
Human–Animal Entanglements
Predation is the most widely studied of human–animal interactions in the Late Pleis-
tocene and early Holocene, and indeed much of the early faunal record of Aşıklı
reflects a continuation of this kind of relationship. But a more intricate web of
human–animal entanglements was set in motion with the founding of the settlement
on the riverbank. New relationships developed with small commensal animals, cap-
tive sheep and goats, hares that predated upon humans’ gardens, and probably cer-
tain invertebrates as well. Dogs were present, based on rare remains found in the
middens, but they may not have been new to the area at the time Aşıklı was founded.
Each kind of human–animal interaction was intimately connected to develop-
ments in the physical nature of the settlement. Spatial analyses of the faunal remains
13
in Aşıklı reveal, for example, that human-built shelters were magnets for toads early
on, when the settlement rested directly on the river terrace, but toads gave way to
small rodents by Level 4 (SI Fig. 15) (Bailey 2018; Stiner et al. 2018). The toads
may have gained food, shelter, and possibly a reproductive advantage in the earli-
est settlement, without harming the human residents and perhaps even benefitting
them slightly by feeding on invertebrates in the refuse piles. More significant for the
inhabitants of Aşıklı were wood mice (Apodemus spp.) and dwarf hamsters (Crice-
tulus, especially C. migratorius), the remains of which dominate the microfaunal
assemblage from Level 4. All were local species; no house mice (Mus musculus)
remains, a non-native commensal originating on the Indian subcontinent that even-
tually swept across Neolithic Asia and Europe (Cucchi et al. 2012), have been found
so far in any of the levels at Aşıklı. The skeletons of wood mice and dwarf hamsters
are common and more or less complete, as would be expected if they lived and died
within the site (Bailey 2018). Both genera are inclined to invade human-built struc-
tures, even though dwarf hamsters can dig their own burrows. Importantly, these
rodents bring no benefits to humans and can cause significant harm. Just a few indi-
viduals can spoil large amounts of stored seed with their excrement. Because the
rodent remains occur mainly inside buildings at Aşıklı, along with a few outdoor
spaces where plant seed remains were highly concentrated (Stiner et al. 2018), there
is good reason to argue that wood mice and dwarf hamsters were a growing problem.
The availability of easy shelter and concentrated food within a settlement sparked
an “arms race” between the rodents and the Aşıklans (in the sense of Tchernov
1984; Weissbrod et al. 2013). The frequency of wood mouse and dwarf hamster
bones (and some preserved coprolites) is greatest relative to other faunal remains in
Level 4, when the structures were more porous and separated by substantial outdoor
spaces. Their numbers decline thereafter (Bailey 2018). Sealed pits and short-lived
structures, such as the pour-molded Building 2 in Level 4, were probably devised
specifically to keep the rodents out of seed and nut stores. The decline in rodent
abundance in the later levels suggests that humans had developed better architec-
tural and other solutions for keeping rodents out of dry-stored foods and building
interiors. Because synanthropic rodents often play host to zoonotic diseases today
(McFarlane et al. 2012), their colonization of early Neolithic sites may also have set
the stage for new trajectories of disease evolution.
Hares (Lepus capensis) are pest of a different sort. They are a highly sustainable
meat source for humans (Stiner et al. 2000) but also a voracious predator of garden
plants. While the overall consumption of small game declined over a millennium of
occupation at Aşıklı (Fig. 6a–c), hares are something of an exception (Fig. 6a; Stiner
et al. 2018, pp. 229–230). A close connection likely developed between humans’
cultivation efforts and hare hunting, also known as “garden hunting” worldwide
(Linares 1976; Smith 2005; Szuter 1991). To protect cultivated plants, hares must
be harvested aggressively near the gardens. Protecting gardens can be a significant
time-sink, and so the persistence of hares in the diet suggests a somewhat altered
relationship with humans.
As for major meat sources, the inhabitants of Levels 5–4 hunted a wide range of
wild ungulate and small animal species. Large prey animals were wild sheep (Ovis
gmelini), wild goat (Capra aegagrus), aurochs (Bos primigenius), red deer (Cervus
13
Fig. 6 Faunal trends from Levels 5 through 2 based on the number of identified specimens (NISP): a
relative frequency of hare, fish (river carp), tortoise and turtle, hedgehog, and birds within the small
game fraction; b relative frequency of aurochs, wild boar, various deer species (red, roe, and rare fallow
deer), and equines (wild horse and onager) among the non-caprine ungulate remains; c changes through
time in overall prey diversity excluding caprines (*to avoid autocorrelation effects), based on the Inverse
of Simpson’s Index (1/D, with potential range 1–9) and the percentage of caprines of total NISP in the
Aşıklı faunal assemblages.
elaphus), wild boar (Sus scrofa), horse (Equus ferus), onager (Equus hemionus), roe
deer (Capreolus capreolus), and rarely fallow deer (Dama dama mesopotamica).
Hares and river fish (especially barbel carp, Barbus plebejus/capito) were consumed
in moderate quantities. Other small animals, such as tortoise (Testudo ibera) and
pond turtle (Emys/Mauremys), hedgehog (Erinaceus concolor), bustard (Otis tarda),
partridge (Alectoris chukar), and various wetland birds, were also used in low
13
quantities. Large birds such as common crane, owls, and raptors were exploited on
occasion, possibly more for raw materials than for food. A trio of large carnivores—
leopard (Panthera pardus), brown bear (Ursus arctos), and wolf (Canis lupus)—
were hunted along with small carnivores, especially foxes (Vulpes vulpes) and less
commonly small felines and mustelids. Exploitation of carnivores was greatest in
the early periods, while fox was hunted somewhat more continuously through time,
possibly because they flourished in the absence of large predators near the settle-
ment and their furs were valued.
The faunal record reveals a remarkable transformation in meat sources from
Level 5 though Level 2. Caprines were only 26% of all prey in Level 5, but their
importance ascended steadily to 92% by the final phases of Level 2 (Buitenhuis et al.
2018; Stiner et al. 2014). The gradual increase in caprines in the faunal assemblages
was offset by a decline in small game resources in particular (Fig. 6a–c). Because
caprines were always an important prey species at Aşıklı Höyük, prey diversity
(1/D) was calculated without caprines in Fig. 6c to avoid the problem of autocorrela-
tion in comparisons of prey diversity to caprine importance (% of all prey). Doing
so reveals that caprine abundance has a weak positive relation to prey diversity (n =
5, r = 0.914, p = 0.030). As the contribution of non-caprines in the diet declined,
the proportions of other prey animals became more evenly distributed in the faunas.
The mild increase in evenness is likely due to the most desirable of the wild species
becoming harder to find, at least where hunters commonly traveled (with or without
their livestock), causing hunters to further diversify their foraging. That said, it is
clear that more and more of people’s efforts were devoted to livestock production as
the millennium wore on.
Management strategies by definition enhance the growth of a subpopulation of
a wild species and allow humans to monopolize its exploitation, or at least make
access to it more predictable. At Aşıklı, sheep and, to a lesser extent, goats were the
focus of management. Higher elevations in the area were prime habitat for native
wild caprines, especially wild sheep. In addition to hunting them, humans began
keeping small numbers of caprines in enclosures between their houses by at least
8300 cal BC if not before. Archaeological evidence for this behavior in Levels 5–4
includes remnants of small corrals and phytolith and micromorphological traces of
in situ dung concentrations that were trampled in place by the captive animals (see
SI Fig. 11) (Mentzer 2018; Tsartsidou 2018). The contextual data, along with zoo-
archaeological evidence for the preferential killing of young males before the age of
6–7 months, tell us that lambs and kids were grown on site for some months before
being slaughtered (Stiner et al. 2014). What is intriguing about the earliest strategy
of caprine management at Aşıklı was the low material investment and the emphasis
on quick nutritional returns.
While stabling may ultimately result in genetic isolation of a caprine subpopu-
lation in the long run, breeding animals in captivity may not have been people’s
plan from the start. Rather, the early stages of caprine management seem to have
provided humans with a more secure meat source during the lean season (late
fall through winter) of any given annual cycle and/or modest opportunities for social
display in the context of feasting events. Lambs and kids could simply have been
taken from the wild when hunters killed their mothers (including late-born infants)
13
and then fed in captivity until they had grown large enough to slaughter. This kind of
flexible, small-scale enterprise is known ethnographically (e.g., for pigs, Dwyer and
Minnegal 2005) and can be very worthwhile in some circumstances. At Aşıklı, this
expedient approach to live meat storage during the times of Level 5–4 appears to
have been one of three means of conserving animal products for delayed consump-
tion, along with drying lean meat and rendering fat and grease from bones. The lat-
ter activity is evidenced by stone-boiling pit features (SI Fig. 16).
Only later did humans achieve substantial reproductive outputs from their captive
caprines. Penning arrangements became more collective during the Level 3 occupa-
tions, when the greater mass and areal extent of dung deposits indicate fewer but
much larger corrals. Toward the end of Level 3, a thick dung accumulation covered a
large area of the mound top (Mentzer 2018). Culling practices continued to empha-
size young individuals, especially males, throughout Level 3. It wasn’t until Level 2
that more animals of both sexes were allowed to live well into adulthood, many to
4 years of age and a few to 6–8 years of age. This late shift in caprine age and sex
structures probably marks the emergence of a full-blown herding system (Buitenhuis
et al. 2018). A shift from capturing and raising young caprines on a small scale to
managing viable herds requires that the animals reproduce easily in captivity and
their offspring survive to adulthood. This was a distinct developmental step in the
human–animal relationship and not necessarily an easy or obvious one for humans.
The shift in age-specific culling in Level 2 implies more attention to building up
the reproductive core of the herd and substantial delays in anticipated returns from
livestock tending. Low-level milk production may also have its roots in this period
(Buitenhuis et al. 2018), although heavy production of milk (and eventually wool)
arose considerably later in the Anatolian Neolithic (Buitenhuis 1997; Greenfield
1988; Halstead and Isaakidou 2013; Helmer et al. 2007).
A comparative study of pathologies on sheep astragali (tarsal bones) by Zimmer-
mann et al. (2018) illustrates some of the early challenges that people faced in man-
aging caprines at Aşıklı. Circumscribed mesoscopic lesions on the articulation sur-
faces of these bones can occur in any ungulate population, wild or captive, but their
severity increases with excessive confinement. At Aşıklı, where a long succession
of faunal assemblages could be studied, the skeletal health of the young sheep was
not very good, and the severity of lesions increased up through Level 3. Significant
improvements in joint health are noted in Level 2, which suggests that caprines were
less confined by then. A δ13C stable isotope study of sheep bone collagen mean-
while suggests a dietary change in the sheep of Level 2, possibly because they were
grazing more in remote pastures rather than just being fed local fodder (Peters et al.
2018).
We have clear evidence of an evolving caprine management system at Aşıklı,
as first suggested by Buitenhuis (1997). Is there also evidence of incipient caprine
domestication at this site? Body size reduction has been suggested to accompany
domestication in many mammals (e.g., Uerpmann 1979), though this may not be
a universal signature and can develop rather late in the process. Wild sheep from
the Epipaleolithic site of Hallan Çemi (10th millennium cal BC) and from Göbekli
Tepe (9th millennium cal BC) were quite large, as one might expect for wild pop-
ulations. Captive sheep from the later Anatolian Neolithic sites of Çatalhöyük,
13
Gürcütepe, Mezraa Teleilat, and Gritille were indeed notably smaller (Buitenhuis
et al. 2018), based on the application of Meadow’s (1999) logarithmic size index.
The Aşıklı sheep and goats from Levels 5–2 combined occupy an intermediate posi-
tion between the comparison size groups. This pattern is consistent with the idea
that Aşıklı represents an early stage on a more drawn-out domestication process.
There is, however, little if any significant body size reduction in the sheep and goats
within the Aşıklı sequence, despite 1000 years of management (Buitenhuis et al.
2018). The lack of a clear diminution trend within the Aşıklı series seems to be the
result of continued genetic introgression between captive and wild stock. Introgres-
sion may slow or arrest the expression of obvious skeletal changes in the captive
population, even while many other traits (e.g., calmness and a variety of selectively
neutral traits) accumulate. Recent studies have shown that introgression is actually
normal to a wide range of domesticating contexts, and it probably is essential to
a successful domestication process over the long term (Clutton-Brock 2008; Lar-
son and Burger 2013; Larson et al. 2014; Marshall et al. 2014). Without continued
genetic input from wild populations, in-breeding undermines the natural diversity of
any small captive gene pool and thus the long-term viability of the subpopulation.
The Aşıklans probably avoided this problem by continuing to capture wild lambs
and kids on occasion. Two important lessons are illustrated by the Aşıklı caprines.
First, a mean reduction is body size emerged later in PPN domestication processes,
whereas essential endocrine and behavioral changes likely preceded them. Second,
continuous introgression was almost certainly necessary to the long-term success of
caprine management at Aşıklı, which developed over a millennium.
A very different line of evidence from ancient DNA confirms that the Aşıklans
participated in the ultimate domestication of sheep and goats in Anatolia and
beyond. Comparisons of mitochondrial and nuclear polymorphisms from sheep sam-
ples dating to the late Pleistocene through early Holocene in central Anatolia suggest
a significant reduction in genetic diversity around or after 7500 BC, a domestica-
tion bottleneck that postdates Aşıklı (Yurtman et al. 2020). Maternal sheep lineages
identified at Aşıklı nonetheless are well represented in modern domestic sheep in
Anatolia and Europe (Peters et al. 2018) and thus survived that population bottle-
neck. Goats were less common than sheep at Aşıklı, but phytolith evidence for tree
leaves and other browse plants in some dung deposits indicate that they were kept
alongside sheep and were subjected to similar selective forces.
Were any other animal species being managed at Aşıklı Höyük? We do not
find clear evidence that cattle or pigs were subjected to management, except pos-
sibly for cattle very late in the cultural sequence. Cattle were never as abundant as
sheep (based on faunal NISP), but cattle continued to be exploited in small numbers
even while most other ungulate species dropped out of the meat supply (Fig. 6b).
Buitenhuis et al. (2018) note more complete body part representation for cattle in
uppermost Level 2 (2C-2A), suggesting that many or most of these large animals
were killed and entirely butchered near the settlement. At Aşıklı, most of the cat-
tle bones in this period are from females. At Musular, the special function site that
appeared just across the river at this time (see SI Fig. 10), 45% of the faunal remains
are cattle, and primary stage butchering activities are apparent. Here, the major-
ity of the Musular cattle are males, and they appear to have been sacrificed. These
13
observations suggest a changing perception of the value of cattle and an increasingly

symbolic association (Buitenhuis et al. 2018, pp. 317–318; see also Russell 2011;
Russell et al. 2005).
Village Emergence and the Power of Human–Environment Feedbacks
As intimated above, human–environment coevolution during the Neolithic shaped

human behavior and biology in some unprecedented ways. This dynamic is also
responsible in large part for the selective environment experienced by certain crop
plants and the two ungulate species that were managed by the Aşıklans. What
sets domesticating interactions apart from other mutualisms is the great extent to
which the nonhuman partner must adapt to the human-altered environment in order
to thrive and reproduce. Getting them to flourish reliably and in greater numbers
requires that humans further adjust the environment to the needs of those plants or
animals—a feed-back process that demands continuous correction. The outcomes of
this process are much more complex than the conscious goals of the human manag-
ers could possibly have predicted.
The theoretical concept of niche construction (Odling-Smee et al. 1996, 2003)
focuses on co-evolutionary feedbacks between organisms and their heritable habitat
(Laland et al. 2000). The evolutionary power of these feedback mechanisms comes
from the asymmetries inherent to problem solving and problem making in the con-
tinuously modified environment. Humans often address problems by resorting to a
domain of culture other than that where the problem first surfaces, particularly if the
cost of switching is lower in the alternative domain and a critical advantage exists
in the former. Examples include social solutions to physical challenges, or changes
in physical efficiencies to solve social dilemmas. These conditions produce “offset,”
or imbalanced, feedback loops that with time may lever human societies into altered
states of existence (e.g., Tainter 2000).
It is clear that some facets of the Aşıklı record changed rapidly, while others
remained relatively stable or shifted only a little. The traditions of rooftop entry-
ways and subfloor burials persisted despite the volatility of other architectural fea-
tures. Chipped stone technology and plant use show surprising continuity through
the cultural sequence. The balance of diverse wild and cultivated plants in the diet
was about the same from Level 5–4 through Level 2; only minor increases in cereal
remains are observed with time, even though selection favoring domesticated vari-
ants of wheat and barley continued. The more conservative facets of archaeological
record could indicate cultural domains that were already sufficiently flexible to serve
the larger system well, or that humans deliberately sheltered these domains from
alteration by being more flexible in others.
Some of the more volatile components of the cultural system changed more or
less in concert with one another. Feedback effects are apparent between shifting soci-
oeconomic arrangements, architectural trends, animal management, waste recycling,
and food storage strategies. It seems that the advantages reaped from one solution
caused perturbations down the line, some of which also had to be accommodated
sooner or later. As such, the human-constructed niche varied rather unpredictably
13
with time and introduced additional selective and neutral conditions for its inhabit-
ants. The prevalence of seed-eating rodents and rodent droppings in Level 4 and the
rise of joint pathologies in captive caprines prior to Level 2 are just two indications
that the food-production learning curve was steep. Though somewhat less visible to
archaeologists, the social benefits of food production and community growth must
also have been accompanied by new sources of tension, perhaps even forcing com-
promises in human health and reproduction.
The pros and cons of livestock keeping for human health and space use at Aşıklı
are a case in point. Architecture and human–animal interactions changed more with
time than any other aspect of the archaeological record examined to date. There is
also reason to think that they were co-evolving in a fairly tightly bound fashion. The
increase in aboveground wall heights from Level 5 though Level 2 was at least partly
stimulated by the need to keep livestock out of buildings and away from delicate
mud-and-reed roofs. A sort of competition between stock-keeping space and human-
dedicated space also developed within the settlement. People initially protected cap-
tive animals from predators or raiders by tucking small numbers of them between
the domestic buildings. Later they kept greater numbers of caprines inside commu-
nal corrals. Architectural infilling took place all the while, leading to an increasingly
dense settlement on the mound top. By the times of Level 2, corrals were shifted
off the mound, thereby accommodating more human residents and dedicated com-
munal spaces. Stock-keeping efforts intensified through the remainder of the cul-
tural sequence, but most animals were no longer held within the dense residential
area and had to be protected by other means.
Human health and meat production suggest yet other interactive feedbacks.
Close proximity to livestock and their waste is known to present health liabilities
for humans. There are hints that the Aşıklans were already experiencing some of
these issues. Eighty-two human skeletons from subfloor burials supply information
on health and survivorship, although most of the individuals examined so far come
from Level 2 (Erdal 2018; Hassett 2018; Pearson et al. 2010). Infants and children
make up about half of this burial sample, a pattern consistent with other Neolithic
communities (Erdal 2018). However, Aşıklı exhibits the lowest infant mortality rate
in all Neolithic Anatolia with the exception of Çayönü (Özbek 2004). Lower infant
mortality may be explained by the early dates of Aşıklı. Infant mortality is known
to have increased along with birth rates (Bocquet-Appel 2011) from the early to the
late Neolithic, in concert with greater agricultural commitments and earlier wean-
ing of children (Erdal 2018; Larsen et al. 2019). Shorter birth intervals require that
mothers replace antibody-rich human milk with a mixed weaning diet, which usually
reduces infants’ access to complete protein and other critical nutrients. Early wean-
ing often compromises the infant’s immune responses to pathogens for this reason.
By later Neolithic standards, the Aşıklı population of Level 2 would seem to have
been doing rather well. Even so, the stresses experienced by children were complex
and protracted in some cases. Most infant deaths occurred within the first two years
of life, especially before the end of the first year (Erdal 2018). Stable nitrogen val-
ues in human bone collagen (Pearson et al. 2010) indicate that weaning commenced
only after 12 months of age, so congenital and parturition problems were likely
factors in the youngest infant deaths. Enamel growth defects in the teeth of some
13
children, on the other hand, point to multiple episodes of stress through the age of
4–5 years (Hassett 2018). These tooth enamel defects cluster bimodally around the
second and the fourth years of life. The first cluster could well relate to weaning-
induced malnutrition due to the birth of a sibling. The later cluster instead points
to a combination of seasonal malnutrition and infectious disease or parasites, two
stresses that are linked to a weakened immune response. Lesions attributed to infec-
tions occur on the cranial and postcranial bones of individuals of all ages, but they
are especially prominent in the Aşıklan infants and children (Erdal 2018).
The settlement’s environment may have shielded the inhabitants from harsh
weather, but it also concentrated smoke, dust, pests, and pathogens around them.
The Aşıklans’ dependence on livestock increased relentlessly, indicating strong pos-
itive selection for continued herd growth in response to a perceived benefit. In fact,
nitrogen isotope data show that the average proportion of animal protein to plant
foods in the Aşıklans’ diets did not decline from Level 4 through the end of Level 2
(Itahashi et al., in press). This is an impressive accomplishment given that the size
of the human population is estimated to have increased roughly 10-fold by the final
phases of Level 2, well in excess of what wild game populations of the area could
support. But benefits experienced at the level of the population do not necessar-
ily translate to benefits for all individuals. Health setbacks due to uneven nutrition,
infections, and vitamin deficiency were fairly common at Aşıklı, yet most people
lived through them (see Wood et al. 1992 on the “osteological paradox”). However,
young children passed through a gauntlet of elevated risk. The human–caprine–envi-
ronment coevolution dynamic at Aşıklı probably was beginning to take a toll on
children, but not to the extent seen in the later Neolithic. For the later community of
Çatalhöyük on the Konya Plain (Fig. 1), a greater reliance on plant carbohydrates,
increased fertility, and greater population density are linked to more severe skeletal
health conditions in the burial population. These maladies are attributed to a combi-
nation of increasing infection rates, poorer diet, and heavy labor (Larsen et al. 2019).
At Aşıklı, waste management strategies show strong responses to the nonlinear
increases in waste generation of all sorts. Whether these were stimulated directly by
health issues is uncertain. We see a marked shift from small opportunistic dumps
scattered widely through the settlement to cooperative use of large communal mid-
dens and greater recycling of livestock dung as fuel, fertilizer, plasters, and con-
struction additives. Waste accumulation and livestock accommodations increasingly
competed with other uses of precious space on the mound top, yet the Aşıklans were
determined to continue residing there at nearly any cost. They responded via archi-
tectural infilling, midden consolidation, and moving corrals to the periphery, rather
than allowing their residences to spill onto the surrounding land. Relocating live-
stock corrals meant significant changes in human time and effort, since these ani-
mals had to be protected. Even so, residential crowding on the mound top was the
preferred solution. There could have been other advantages to architectural infilling
as well, such as community defense (two adults have wounds from bone projectiles)
and certain work-sharing conveniences. The inhabitants’ steadfast interest in resid-
ing on, rather than around, the mound top nonetheless illustrates the mound’s primal
importance as a physical symbol of collective history and household legacies, spe-
cifically the physical connection to house footprints and the burials within them.
13
Another intriguing characteristic that sets Aşıklı apart from other Neolithic settle-
ments is the lack of shared walls as residential structures mutated from roundish to
quadrangular contours and infilling progressed. Level 2 contains 10 distinct building
phases, plenty of time for further innovations to develop, yet even the most tightly
packed residential clusters maintained separate building walls. The practice did not
continue at the later PPN site of Çatalhöyük in central Anatolia, so it is reasonable to
wonder why it persisted through the later phases at Aşıklı. One can propose a variety
of explanations that range from familial divisions to a lack of awareness of potential
structural efficiencies as quadrangular architecture took over the mound top. But the
Aşıklans innovated on so many other architectural fronts that the latter explanation
is hardly satisfying.
A confluence of physics and history may explain the persistence of this unique
style of construction. We know that the narrow spaces between residential buildings
gradually filled with debris that was swept from the rooftops. While most spaces in
the settlement were subjected to regular refuse removal, these narrow between-wall
spaces were not. Seasonal precipitation in Anatolia can be heavy, and a mound site
of Aşıklı’s proportions must have had chronic drainage issues. The roofs were sup-
ported by wood beams of limited length, with a heavy cover of reed and mud plaster.
Experimental evidence indicates that the building roofs were very fragile, subject
to slumping, and needed frequent maintenance (Özbaşaran and Duru 2018). Under
these conditions, the narrow spaces between the buildings would provide a quick
exit for water falling from each roof. The inclination of the drains could be managed
just by controlling the position and shape of the debris cone therein. Once the main
drainage patterns were set within the community as a whole, it would have become
much harder to change them. This situation could explain the continuous practice of
independent walls over multiple centuries at Aşıklı—conservatism rooted in prior
investments, akin to “developmental locks” or generative entrenchment (Arthur
1984; Schank and Wimsatt 1986).
Social Dimensions of Food Production and Storage
We have no reason to believe that the Aşıklans abandoned the egalitarian eth-
ics of their hunter-gatherer forebears. Households were the core of production
and consumption at Aşıklı throughout the millennium of its existence. Extensive
seed-processing areas (see also Asouti and Fuller 2013), cooperative waste dis-
posal, and upkeep of public buildings are some of the indications of communal
activities. Cooperation in seasonal harvesting was also necessary because of the
short time windows of cereal and pulse maturation. However, the patterns of
sharing and reciprocity may have narrowed somewhat to households by the later
phases of Level 2 (see Baird et al 2017; Düring and Marciniak 2006; Kuijt et al.
2011; Marciniak 2008; Winterhalder 1990, p. 82; Winterhalder and Goland 1997,
p. 158; Wright 2014). One important signal of burgeoning inequalities between
households in Level 2C-2A is variation in the stable nitrogen values in human
skeletons. While the overall ratio of animal foods in the diet did not decline with
13
time (see above), these data show that the ratio did vary from household to house-
hold in this late period (Itahashi et al., in press).
Mobile hunter-gatherer groups of the historic era tend to favor relatively
open sharing networks, wherein secrecy and hoarding are suppressed through a
range of social mechanisms (e.g., Kelly 1995; Winterhalder 1990). The limited
privacy of domestic areas in hunter-gatherer camps can serve as a rough proxy
for the relative openness of sharing networks in these societies. Sedentism cou-
pled with an increasing emphasis on food production may alter human coopera-
tive relationships over the long run. The ample outdoor spaces that separated the
residential buildings during the Level 5–4 occupations at Aşıklı could only be
loosely claimed by any one household. At least some of the dry food stores in
this period were kept apart from the residences in small freestanding structures.
These repositories likely served several residential units, and their contents would
not have been secret. The intermediate archaeological levels at Aşıklı display a
combination of indoor and outdoor storage facilities, tilting ever more in favor of
private stores with time. With architectural infilling during the Level 2 occupa-
tions, much of the daily workspace was shifted to flat rooftops. These activity
spaces, though visible to virtually anyone in the community, were clearly defined
at their margins by the supporting walls of each freestanding house. By the final
two centuries of Level 2 (2C-2A), relatively large storage facilities were placed
deep inside each residence, entirely out of public view, and some of these had
large bins in addition to small back rooms. Public stores may also have existed in
the later period, perhaps within one of the large communal buildings, in addition
to the residential storerooms. Lateral sharing among households may have contin-
ued, but the intensive work needed to create food stores seems to have introduced
cul-de-sacs in the reciprocity network that could be controlled by specific indi-
viduals or households.
There is the related question of the overall scale of food storage and, by exten-
sion, the extent to which inequities become possible within a society. Scholars have
proposed a variety of motivations for the evolution of food storage, ranging from
attempts to control subsistence risk to individual quests for social power and influ-
ence (compare Asouti and Fuller 2013; Hayden 2009; Kuijt 2008; Testart 1982;
Winterhalder 1990; Winterhalder and Goland 1997). All of these motives may have
been significant in some way during the Neolithic. What matters most for this dis-
cussion are how the relations of production and food hoarding restructured the food
supply, pitching more of peoples’ efforts toward the social and physical demands
of feeding, protecting, and preserving the concentrated resources. Of course, food
storage was not a Neolithic invention. Food storage practices are widely evidenced,
albeit on smaller scales or focusing on different foods, in Epipaleolithic and Mes-
olithic hunter-gatherer sites across Eurasia (e.g., Testart 1982). However, plant
food storage practices underwent further changes during the Pre-Pottery Neolithic
through Ceramic Neolithic periods, increasing in scale and with a stronger focus on
annual seeds (e.g., Kuijt 2008; Willcox and Stordeur 2012). Under these conditions,
rights of ownership of produced foods and the power to decide how to use surpluses
can shift, effectively pitting notions of the collective rights of the community against
those of individuals or specific households.
13
Given the rising importance of caprine management at Aşıklı, livestock would

also have presented opportunities for generating surplus and accumulating wealth
or social influence. We see in the earliest levels of Aşıklı that just “growing” a
few wild infant ungulates for short periods in captivity could be its own reward,
regardless of whether humans and caprines had entered into a true domestication
relationship (e.g., for pigs at Hallan Çemi, see Rosenberg et al. 1998; Starkovich
and Stiner 2009; also Price and Hongo 2020). Management at this stage could sim-
ply have been about optimizing somatic growth of a few wild individuals. By the
time of Level 3, the richest supply of animal foods no longer came from the wild
but rather from livestock and human investments in their propagation. Here we see
a herding strategy in its infancy. By Level 2, the number of livestock had greatly
increased, equaling or surpassing human biomass in the community. Culling prac-
tices now allowed for the accumulation of many adult animals. If sequestered plant
food stores and live storage of caprine meat improved food security for the Aşıklans,
these practices must also have introduced new potential for inequality (see Bowles
and Choi 2019; Ostrom 2003; Svizzero and Tisdell 2019). Sharing rules and sharing
pools would be affected, particularly if individual or household economic decisions
become less constrained or "entangled" by the decisions of others in the commu-
nity (Stephens and Krebs 1986, p. 7). This question is explored further in regard to
exchange networks in the next section.
Human physiological changes in the Neolithic confirm that workloads, personal
responsibilities, and entitlements were reconfigured by food production. These
adjustments shifted the nature of gendered and age-dependent socioeconomic roles.
Differences in locomotor stresses between men and women are apparent from pat-
terns of bone remodeling and joint pathologies in particular (e.g., Eshed et al. 2004b;
Marchi et al. 2006; Ruff 1987; Scherf et al. 2016). One sees greater lateral symme-
try in these phenomena in Neolithic skeletons than in hunter-gatherer populations.
Locomotor stresses from redundant symmetrical motions and heavy loading are
expressed in many of the joints of adult males and females throughout the Neolithic
period. At the early PPN community of Aşıklı, patterns of joint degeneration and
osteoarthritis in adults indeed suggest differentiation in habitual work patterns and
economic roles by gender. All the adult females and most of the adult males experi-
enced some degree of vertebral degeneration, and about two-thirds of the adults of
both sexes display some degeneration of the shoulder and/or hip joints. There are
greater differences between men and women in other joints—elbows for males pos-
sibly in connection with hunting or portage, and the ankle for females probably from
squatting for long periods (Erdal 2018). The ankle joint pathologies suggest that the
redundant, time-consuming work of plant seed processing was largely performed by
women.
The labor requirements of food production changed what people did with their
time, where they spent it, and even who they shared resources with, setting up new
arenas for further social evolution (see O’Brien and Bentley 2015). The time and
labor commitments of food production may also have potentiated new status con-
figurations and opportunities by gender. We have no reason to think that either sex
dominated food production or the distribution of its products during the early Neo-
lithic (see Peterson 2010), but there were some notable differences between men and
13
women in work emphasis and authority over these kinds of work. The adult sex ratio
at Aşıklı also hints at such a development. Adult females generally outnumber adult
males by a factor of two to one (Ö. Erdal 2018; Y. Erdal 2013). This bias in favor
of women in subfloor burials suggests a powerful affinity of at least some women to
specific household legacies.
Networks versus Sole‑source Models of Neolithic Emergence
The forager-producer transition occurred surprisingly quickly over large areas

of Southwest Asia—so fast in geological time that it is difficult to test conflicting
models of Neolithic origins. A common view has been that PPN lifeways sprang
from just one or a few geographic isolates and then spread with little resistance from
those points of origin. Recent arguments instead propose that the Pre-Pottery Neo-
lithic had many distinct regional roots (compare Arranz-Otaegui et al. 2016; Fuller
et al. 2011; Matthews 2002; Özbaşaran et al. 2018; Stiner et al. 2014; Willcox 2005,
2013; Zeder 2008). These early Neolithic communities were bound by loose infor-
mation networks within and across regions, but they erupted quasi-simultaneously
in widely scattered localities.
The great unevenness in carbon-14 dating work makes it very difficult to compare
just when each sedentary community of plant cultivators was founded and when ani-
mal and/or plant management practices arose therein. To assert that PPN sites across
Southwest Asia have not been dated with equivalent accuracy would be an under-
statement (see Aurenche et al. 2001; Bischoff et al. 2006; Thissen 2002a, b. This
unfortunate situation is due to very limited dating programs at many sites in past
decades and/or the fact that the few dated samples came from ambiguous contexts.
Many key sites are now inundated, destroyed, or lie in politically contested areas and
so cannot be resampled. The likelihood of disparate evolutionary “hotspots” none-
theless is supported by recent data surveys and intensive dating work at a handful
of early PPN sites (e.g., Benz et al. 2012; Quade et al. 2018; Vigne et al. 2012).
The geography of the so-called “Fertile Crescent” has expanded as a result, mostly
westward, transforming the original crescent-shaped area for which it was named
into something that looks more like an amoeba on the move. These many preco-
cious PPN hotspots were only weakly connected to one another for much of their
early histories, as well as to the many complex hunter-gatherer communities lying in
between.
To the extent that results on human ancient DNA are becoming available, the
PPN populations in central Anatolia seem to have originated from within the region
(Chyleński et al. 2019; Feldman et al. 2019). If the people who founded Aşıklı
Höyük already had some familiarity with plant cultivation and growing young wild
animals in captivity, what was the historical foundation of their knowledge? In fact,
many Epipaleolithic and Mesolithic cultures of Eurasia routinely manipulated the
natural distributions and productivity of wild food resources, and they developed
diverse technologies to enhance their nutritional yields (e.g., Asouti and Fuller
2013; Fairbairn et al. 2014; Holst 2010; Munro and Bar-Oz 2005; Richter et al.
2011; Savard et al. 2006; Snir et al. 2015; Willcox 2013; Willcox and Stordeur 2012;
13
Wright 1994). Ancestral knowledge of these strategies was likely as geographically

widespread as it was diverse. This local grassroots knowledge likely explains many
of the broad strategic commonalities that archaeologists observe across burgeoning
Neolithic systems.
Some exchange of knowledge or materials did occur between far-flung commu-
nities, such as during the early centuries at Aşıklı (Levels 5–4), when certain non-
native cultigens, Mediterranean shells, and nonlocal flint artifacts were added to the
local mix. Outside influences dwindled noticeably through Level 3 and were practi-
cally nonexistent through Level 2J–2E. Only toward the end of the Aşıklı sequence
(Level 2D–2A) did new forms of material culture appear. The trends in obsidian
chipped stone and ornament technologies illustrate the variable extent to which the
Aşıklı community participated in larger regional networks over the millennium of
its existence. Obsidian was used for virtually all chipped stone tool production at
Aşıklı. The Level 4 settlement used several of the local sources (Fig. 2) that included
Göllüdağ, Nenezi Dağ, and possibly Acıgöl (Astruc and Grennet 2018). This habit
gave way to a focus on the Göllüdağ-Kayırlı source by Level 2. The different obsid-
ian sources were not favored for specific kinds of tool production in any period. The
dominance of the Göllüdağ-Kayırlı obsidian for tool and blank production in Level
2 simply reflects more regular and possibly more organized visits to this particular
source (Kayacan and Altınbilek-Algül 2018). The Nenezi Dağ source was also used
to some degree in all periods, but it became important only with the latest phases of
Level 2 and in Level 1.
The Aşıklans’ lack of participation in regional obsidian exchange networks to the
east and south is striking, given that some of the greatest obsidian sources in all of
Anatolia (listed above) lay just 20–30 km east of the settlement (see Fig. 2). By the
eighth millennium BC, Cappadocian obsidian was being traded by other groups over
a large area, albeit in small volumes. A major obsidian workshop site geared to trade
production was established at Kömürcü-Kaletepe on the Göllü Dağ, and the mate-
rial produced from this quarry site demonstrates clear trade links between Cappa-
docia and the Levant, and possibly to the middle Euphrates area (Balkan-Atlı et al.
1999, 2008; Binder 2002; Ibáñez et al. 2015). The highly standardized naviform and
prismatic cores of the Kömürcü-Kaletepe workshop could only have been made by
skilled specialists. These cores, and the pressure techniques used to produce them,
are characteristic of the Near Eastern PPNB communities outside of Anatolia but
not at Aşıklı (Balkan-Atlı et al. 1999, 2008; Binder 2002).
The Aşıklans must have been aware of the Kömürcü-Kaletepe quarry workshop,
but they showed no interest in emulating those obsidian working techniques. The
Aşıklan knappers adhered to a distinctly local tradition from Level 5 through 2.
This continuity is apparent in the knapping methods for producing tools, the relative
abundance of typological groups (e.g., oblique truncations and microliths), and the
morphology of the plant harvesting tools (Astruc and Grennet 2018; Kayacan and
Altınbilek-Algül 2018). They used two direct percussion systems known as bidirec-
tional knapping and unipolar knapping. While bidirectional production increased at
the expense of unipolar production from the mid-ninth millennium to the end of the
eighth millennium, this shift unfolded very slowly from one level to the next. Impor-
tantly, the complete spectrum of production debris on site tells us that the Aşıklans
13
made obsidian blades and tools for their own use, and there are no signs of specialist
knappers. It is possible that the Aşıklans exchanged small amounts of obsidian to
communities to the west (i.e., Konya Basin), but there was no specialized production
of cores and blanks for this purpose. Examples of specialized production emerge
only in Level 1, when bipolar production of longer, more elaborate blades began
(Kayacan and Altınbilek-Algül 2018; on Musular, see Kayacan and Özbaşaran
2007).
Bead industries tell a related part of the story. Personal ornaments were produced
and used at Aşıklı throughout the history of the community, but the diversity and
abundance of beads and the range of raw materials increased markedly only in the
final phases of Level 2 (2D-2A, Yelözer 2018). This is also when beads began to be
interred with human bodies, often in large numbers. The bead assemblages in Level
2C-2A are much more varied in the colors, types, and sources of stones and miner-
als represented. Some of these materials were not available locally (Yelözer 2018).
The same is true for a wider range of small artifacts made of stone, bone, and other
materials (Sönmez 2018). A few beads in the final phases of Aşıklı were made from
non-smelted copper (source undetermined, Esin 1999) and carnelian of high qual-
ity (Yelözer 2018). Other exotic goods of this phase include an obsidian bracelet of
eastern Anatolian provenance.
Several authors have documented increases in the complexity and diversity of
ornament assemblages during the later Neolithic, both in central Anatolia (Bains
et al. 2013; Bar-Yosef Mayer et al. 2010; Wright et al. 2008) and the Levant (Bar-
Yosef Mayer and Porat 2008; Wright and Garrard 2003). This trend is thought to
indicate expansion and growth of regional exchange networks and more widespread
participation among the different communities therein. There is no evidence of spe-
cialized bead production at Aşıklı (Yelözer 2018). Yet the proliferation of bead types
and the inclusion of beads in burials in the late phases of Level 2 (2C-2A) attests
to a new pattern of burial symbolism (Yelözer 2018; see Kuijt et al. 2011 on Tell
Halula) in parallel with increased trade connectivity. Given that there are no strong
biases by sex or age in who was buried with beads, the beads may have been mark-
ers of individuality. The increase in bead use in life and in death in the late phases of
Level 2 suggests an important shift in how ornaments were deployed in social com-
munication (in the sense of Kuhn and Stiner 2007). Specifically, a new premium was
set on materials that signalled outside connections—a subtle but significant signal of
status, whether from accumulated wealth or social capital (e.g., Price and Bar-Yosef
2010). It may be no coincidence that these grave goods became important around
the same time as subtle nutritional inequalities among households emerged within
the community.
In summary, the extent of interregional exchange or external cultural influences
varied significantly at Aşıklı from Level 5 through Level 2A. The long middle sec-
tion of the sequence shows the least evidence of network connectivity. Perhaps it
was the introverting demands of the rapidly evolving food production system during
this interval, which in the case of livestock went from being a supplement to the
main meat source. Food production demands can encourage socioeconomic insu-
larity in the sense that less effort is put to “normalizing” long-distance relations
and expanding opportunities for reciprocity with other communities. This energetic
13
“boundedness” can create a unique selective environment for humans that allows
for rapid change from within. If this was indeed the situation, then we must also ask
what rekindled the people’s interest in long-range exchanges later in the settlement’s
history. The Aşıklans’ lack of integration into extra-regional obsidian exchange is
understandable given that the “object” of trade was not so much the raw material as
the elegant pre-prepared blanks that required special skills to produce. The situation
for beads is harder to explain in this way, since the Aşıklans were bead users from
the get-go, and small numbers of exotic types such as marine shells are present in
Levels 5–4. Why did exotic ornament types cease their flow into the settlement until
very late in Level 2? Perhaps the Aşıklans did not have much in the way of high-
quality surpluses to trade prior to this time. The balance may have changed with the
blossoming of a viable herding economy within the time frame of Level 2. This was
also when preliminary isotopic evidence suggests that livestock herds were feeding
on a wider range of foods, including from distant pastures, and that their joint health
had improved due to higher mobility (Peters et al. 2018; Zimmermann et al. 2018).
A more wide-ranging herding system would facilitate the development of special
partners in exchange over long distances, especially if surplus livestock were avail-
able for trade.
Conclusion: Internal versus External Influences
The process of “Neolithization” in central Anatolia grossly parallels that of other

mainland regions of Southwest Asia (compare Baird et al. 2018; Conolly et al.
2011; Ervynck et al. 2001; Helmer 2008; Özdoğan 2011; Özdoğan and Özdoğan
1989; Peters et al. 2013; Willcox 2005) and the island of Cyprus (Vigne et al. 2012).
Yet Aşıklı and subsequent PPN settlements in central Anatolia (Table 1) were not
much like those of the northwestern Zagros, southeastern Anatolia, or the Levant
in the specifics of culture, innovation, or the pace of change (Özbaşaran and Duru
2015). The transition from circular to rectangular buildings occurred around the
same time in central Anatolia as in the other regions. Flannery (2002) famously saw
this transition in the Levant as marking a shift in social organization from nuclear
to extended households. The transition from round to rectangular architecture is
Table 1 Time frame of central Anatolian Neolithic sites discussed in text

Site Time frame (millennium BC) Source
Can Hasan I End of 7th through 6th Fairbairn et al. (2020); French (1998)
Erbaba Second half of 7th Bordaz and Alpers Bordaz (1976)
Çatalhöyük End of 8th through 7th Hodder (2011)
Can Hasan III Second half of 8th French et al. (1972)
Aşıklı, Level 2 Early 8th Quade et al. (2018)
Aşıklı, Level 3 Late 9th to early 8th Quade et al. (2018)
Aşıklı, Levels 5–4 Mid 9th Quade et al. (s2018)
13
clearly evidenced at Aşıklı (the earliest PPN site in the central Anatolian sequence,
Table 1), but indications of a major shift in social relations occurred considerably
later, when distinct neighborhoods emerged in the centuries just before the site was
abandoned (Özbaşaran 2011b).
The architectural innovations first seen at Aşıklı persisted in later central Ana-
tolian settlements (Table 1), but only in this region. There is notable continuity in
the construction methods and in the fidelity to original house footprints over many
cycles of reconstruction (at Aşıklı, see Esin and Harmankaya 2007, pp. 258–262;
at Çatalhöyük, see Hodder 2011, pp. 940–942). The tightly clustered building lay-
outs of the later phases of Level 2 at Aşıklı are repeated at Can Hasan III (French
et al. 1972), Can Hasan I (Fairbairn et al. 2020; French 1998), Erbaba (Bordaz and
Alpers Bordaz 1976), and Çatalhöyük (Hodder 2011). The developmental patterns
elsewhere in Southwest Asia tend to be more abrupt, both prior to and with the
onset of the PPNB “Interaction Sphere” (Bikoulis 2013; Düring 2006; Duru 2018b;
Goring-Morris and Belfer-Cohen 2016), a contrast that is well illustrated at Çayönü
(see Erim-Özdoğan 2011). Equally strong distinctions are found in mortuary prac-
tices, which outside central Anatolia typically included skeletal disarticulation and
secondary burials (Özkaya and Coşkun 2011), skull removal and curation (Kuijt
2009; at Çayönü see Erim-Özdoğan 2011), and elaborate plastering and painting of
defleshed crania (e.g., Tell Aswad, Stordeur and Khawam 2006; Ain Ghazal, Rollef-
son et al. 1999; Schmandt-Besserat 2013). Plastered skulls appeared only very late
at the central Anatolian sites of Çatalhöyük and Köşk Höyük.
The central Anatolian communities maintained very distinct approaches to obsid-
ian working, even though the local obsidian quarries were being exploited at the
time by nonlocal specialists (Balkan-Atlı et al. 1999, 2008; Binder 2002, 2007).
The strongly female-biased adult sex ratio in subfloor burial populations at Aşıklı
and into the Pottery Neolithic in central Anatolia also stand in bold contrast to the
Levantine Natufian-Neolithic pattern, where adult males greatly outnumber adult
females (a ratio of 2 to 1 in the Natufian, and only somewhat less for the Neolithic;
Eshed et al. 2004a). The female bias among adult burials in central Anatolia could
stretch back into the Epipaleolithic, but it likely represents a significant departure
from the land tenure concepts of earlier Paleolithic hunter-gatherer societies. Com-
parable human skeletal data for Paleolithic Turkey are lacking, but strong male
biases seem to have been the rule in other Eurasian Upper and Middle Paleolithic
sites (roughly 2 to 1; see Formicola 2007; Harrold 1980).
The PPN communities of central Anatolia were on their own developmental tra-
jectory, not entirely out of contact with far-flung communities but evolving largely
within their own bounds. Gradual change was the rule at Aşıklı, yet the cumulative
transformations in architecture, settlement layout, and caprine management were
great. None of these major trends can be explained by innovations coming from out-
side of the community. Rather, strong motors worked via feedback (especially posi-
tive feedback) processes from the inside out. Relationships between humans and ani-
mals, and between humans and crop and weed plants, grew increasingly symbiotic.
New efficiencies developed for recycling nutrients and raw materials, which in turn
enhanced crop and livestock productivity. The strategies of caprine management
went through a succession of apparently viable systems, from raising a few young
13
lambs or kids to a true herding system that utilized distant and nearby grazing lands.
The symbioses within the Aşıklı system expanded with time, funneling more energy
into producing food, protecting it, and making it last.
The community maintained a collective way of living. Persistence in communal
patterns of life and work are evidenced by the lack of differentiation among house
plans, building materials, and internal features, among other indications. Longstand-
ing social values of communal work and cooperation seem to have been eroded very
little by the demands and benefits of food production and storage. Social and eco-
nomic forces were at times at odds with one another, such as during structural infill-
ing in Level 2 when the people’s need to maintain a physical connection between
domicile and the deceased forced new solutions for corralling and protecting live-
stock elsewhere. Trends in architectural forms, waste accumulation, contact with
livestock, and population density ultimately affected human health, physiology, and
behavior. Few aspects of life in this settlement were free of the internal web of recip-
rocal influences and contingencies.
That said, the Aşıklı settlement did not develop in a complete vacuum. Its
regional context is essential to appreciating its history, instructive both for its local
uniqueness and its weak but at times influential connections outside central Anatolia.
Aşıklı was one of many village communities that emerged within a short time inter-
val in Southwest Asia. The relative insularity of the Aşıklı community developed
within a few centuries of its founding and persisted until quite late. However, the
influx of exotic ornamental materials and a shift in mortuary practices in Level
2C-2A points to new regional and interregional connections. This was a time when
regional and interregional connections were growing stronger throughout Southwest
Asia (e.g., Bar-Yosef and Belfer-Cohen 1989; Borrell and Molist 2014; Goring-
Morris and Belfer-Cohen 2016; Ibáñez et al. 2015). To the extent that the Level 2
Aşıklı community had anything to ply in these exchange networks, it likely included
surplus livestock. Cereal production increased only mildly with time at Aşıklı, so
surpluses are unlikely to have been regular units of exchange. Other possible items
of exchange could have been basketry, local red ochre, salt from the Tüz Gölü Basin,
and rare volcanic stones. Livestock nonetheless stands out as an inherently valuable
and unique item that was also highly movable.
To understand the Aşıklans’ motivations for entering interregional exchange net-
works after centuries of not doing so, we probably need to consider internal shifts
in social rights and resource control. Trading livestock for other things, particularly
for rare nonutilitarian items, implies personal or household-level ownership of the
surplus to be exchanged for these objects. In other words, livestock had begun to
express differential wealth within the community. Surpluses of livestock could be
transformed into signals of individual or household status, whether in the form of
fancy things, social influence, or ritual authority. Individual or household legacies
may have taken on new layers of meaning in these circumstances, making the physi-
cal connection to building footprints and ancestors on the mound more important
than ever.
We have focused in this essay on how some essential properties of early Neo-
lithic life arose as the result of powerful feedback forces operating within the sys-
tem. Emergent properties can surface at any number of scales, but by definition they
13
are expressed at scales different from the dynamics that produce them. At least three
distinct scales of emergent phenomena have been explored in this paper—the inti-
mate scale of human decision making at the level of the individual or household,
the local scale of the settlement, and regional scale of “neolithization.” Even if the
potential power of feedback interactions seems clear, identifying specific dynam-
ics within extinct cultural systems remains a daunting enterprise. Detailed data on
site formation processes in a long and continuous record are very helpful, but there
are few precedents for how to go about this in archaeological research, and ana-
lysts are only beginning to grapple with the challenges (but see Laland and O’Brien
2010; Murray et al. 2021; O’Brien and Laland 2012; Stiner 2021; Stiner and Kuhn
2016). Nonetheless it seems that many of the trends that we observe at Aşıklı Höyük
over a full millennium were propelled by feedback relationships between humans,
the human-altered environment, and the mix of animal and plant species that they
sought to manage. As such, Aşıklı represents a compelling case of co-evolution via
a niche constructing process, driven largely from within and continuously stimulated
by offsets in social, economic, and physical constraints. The impressive outcomes
of this process did not require human prescience at evolutionary time scales. Peo-
ple’s day-to-day and longer-term decisions were intentional, but the collective con-
sequences of their actions were not. It seems that many grand human “designs” were
not really designed at all.
Supplementary Information The online version contains supplementary material available at https://doi.
org/10.1007/s10814-021-09167-z.
Acknowledgments We would like to express our profound thanks to all the scholars, friends, students,
institutions, and foundations that directly or indirectly contributed to this collective work. Special thanks
also go to Max Price, Steve Kuhn, and four anonymous JARE reviewers for their many insightful com-
ments and corrections to an earlier version of this manuscript. The first author is also grateful to John
Odling-Smee, Claes Andersson, and Kevin Laland for enlightening conversations on NCT over the years,
which have contributed greatly to the conceptual background of this paper. In addition to the community
of Kızılkaya Village, the following institutions deserve special mention for their support: Turkish Min-
istry of Culture and Tourism, General Directorate of Cultural Assets and Museums; Istanbul University
Research Fund (Project ID 25754); Istanbul University Faculty of Letters; Governorship of Aksaray; Gov-
ernorship of Gülağaç District; Public Education Center of Gülağaç Governorship; American Embassy in
Turkey/Ankara. Additional funds for the project were provided to the first author by two consecutive
Archaeology Program grants from the National Science Foundation (BCS-0912148 and BCS-1354138).
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SUPPLEMENTAL INFORMATION – Stiner et al. SI Figures 1-16
Aşıklı Höyük: The Generative Evolution of a Central Anatolian PPN Settlement in Regional Context
Mary C. Stinera, Mihriban Özbaşaranb and Güneş Duruc

a
School of Anthropology, P.O. Box 210030, University of Arizona, Tucson, AZ 85721-0030, USA. Telephone: +1 520-903-7797. Email:
mstiner@arizona.edu
b
Istanbul University, Faculty of Letters, Prehistory Department, 34134 Fatih, Istanbul, Turkey. Email: ozbasaranmihriban@gmail.com
c
Mimar Sinan Fine Arts University, Faculty of Conservation and Restoration of Culture Property, Cumhuriyet Mh. Silahşör Cd. No: 89 Bomonti,
İstanbul, Turkey. Email: gunes.duru@msgsu.edu.tr
SI Figure 1. Examples of combustion features at Aşıklı Höyük: (a) collapsed burnt wooden members of Building 39 in Level 5, exposed in the
western section of the mound; (b) sequentially rebuilt hearth feature in Level 3, also in the west section. (Aşıklı Höyük Project Photo Archive)
SI Figure 2. Schematic of major changes in building contours from Levels 5 through 2 at Aşıklı Höyük. Adapted from Özbaşaran et al. 2018.
SI Figure 3. Reed (a) imprints in kerpiç and (b) intact in-situ reed phytoliths in Level 5, part of burnt surface structure complex in Trench 4GH.
SI Figure 4. View of excavated half of Building 3 in Level 4, showing its internal architectural features: (a-b) sub-floor burials, (c) hearth, (d)
milling stone embedded in kerpiç bench, along with numerous pits. Upper walls were built with kerpiç bricks and mortar. (Aşıklı Höyük Project
Photo Archive)
SI Figure 5. Plan view of an early floor of Building 38 in Level 3, on the western side of the mound. Note bench-like structure near top of photo
with small surface depressions and the very large central hearth. Inset at upper left shows the much larger diameter of Building 38 in comparison
to two adjacent, partly preserved residential structures, B.24 and B.18. (Aşıklı Höyük Project Photo Archive)
SI Figure 6. The ventilation shaft for Building 35 and its successor, Building 34: white arrows indicate the external upper opening and the internal
lower opening indicated by arrows; inset at lower right shows detail of the exterior upper opening.
SI Figure 7. Plan view of one of the excavated floors inside partly exposed Building 2, a non-residential structure in Level 4 (Trench 4GH). The
walls of this unusual structure were made of pour-molded kerpiç. A succession of prepared floors is preserved within this building. Note shallow
cup-like depressions on the floor. (Aşıklı Höyük Project Photo Archive)
SI Figure 8. Plan of the Aşıklı Höyük settlement layout in Level 2C-A. Adapted from Özbaşaran et al. (2018).
SI Figure 9. Example of a residential building interior in upper Level 2, including semi-intact hearth (lower left), stone mortar, and other milling
stones, including one embedded in a long kerpiç bench at upper right. (Aşıklı Höyük Project Photo Archive)
SI Figure 10. Aerial view of Aşıklı Höyük and its satellite sites along the Melendiz river drainage: (1) Aşıklı Höyük, with mid-8 th millennium BC
trench exposures visible on surface; (2) Musular, a special function site coeval with Levels 2C-A at Aşıklı; (3) Ven 7; (4) Gedikbaşı; and (5)
Yellibelen. Melendiz Mountains are visible on horizon at right; modern village of Kızılkaya at upper left. (Aşıklı Höyük Project Photo Archive)
SI Figure 11. Intact layers of ungulate dung in stabling deposits at Aşıklı Höyük: (a-b) thick deposit from Area 2JK exhibits a strongly laminated
fabric viewed (a) in PPL and (b) in XPL, with concentrations of calcareous spherulites that appear as lighter zones; (c) detail of spherulites in a
stabling deposit, XPL. Adapted from Mentzer 2018: Fig.7.
SI Figure 12. Summary of urine salt inputs in the sediments of Aşıklı Höyük: (a) Comparison of mean soluble salt concentrations in
archaeological Levels 5 through 2 and in the natural alluvium at the base of the mound. Sodium [Na+], chlorine [Cl−], and nitrate [NO3−] are given
in moles × 1000 kg−1. Mean values for each archaeological level include samples taken from general middens, dung-dominated middens, and
alleyways; (b) Model-predicted densities of organisms (per m2) required to produce urine-related [Na+], [Cl−], and [NO3−] values, averaged by
level using variable sedimentation rates based on 14C–dated level boundaries; (c) changes in caprine NISP as percent of total taxon-specific NISP
by level. Urine salts data from Abell et al. (2019).
SI Figure 13. Variation in macrobotanical remains by level in Aşıklı Höyük: Relative frequencies (percentages) of (a) pulse seeds and cereal
seeds/grains in the total seed assemblage; (b) wild grass seeds, other herbaceous plant seeds, and fruits/nuts in the total seed assemblage; (c)
specific cereal chaff types in the total chaff assemblage. Chaff type 1 = Triticum monococcum/turgidum/new type (glume base count); chaff type
2 = Triticum turgidum ssp. durum/ T. aestivum ssp. Aestivum (rachis internode count); chaff type 3 = Hordeum vulgare subsp. distichum/subsp.
vulgare/ H. spotaneum (rachis internode count). Data from Ergun et al. (2018).
SI Figure 14. In-situ phytolith traces of plant fiber baskets and mats from Levels 2 through 4: (a) detail of woven mat covering infant burial in
Level 3E; (b) detail of woven basket; (c-d) coiled baskets from residential building floors in Level 2. (Aşıklı Höyük Project Photo Archive)
SI Figure 15. Relative occurrences of each major taxon group in order of body size in intra-mural versus extra-mural features in Levels 5-3. The
mean difference (vertical line) for all taxa from feature contexts serves as the comparative standard. Adapted from Stiner et al. (2018)
SI Figure 16. Outdoor pits in Level 4 in Trench 4GH, the larger filled with fire-cracked cobbles and densely concentrated charcoal at right-hand
margin. This cooking feature was used for stone-boiling.

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Journal of Archaeological Research

Aşıklı Höyük: The Generative Evolution of a Central

Mary C. Stiner1 · Mihriban Özbaşaran2 · Güneş Duru3

Accepted: 8 March 2021

Keywords Domestication · Niche construction · Social networks · Site formation

The forager-farmer transition, also called the Paleolithic-Neolithic or forager-pro-

remarkable concentrations of seed remains, indicating frequent processing activi-

A longstanding question at Aşıklı concerns the position of entryways to the semi-

The Living Mound

Human–Plant and Human–Animal Coevolution

A variety of co-evolutionary processes are evidenced at Aşıklı Höyük in the context

of domestication, before many of the obvious genetic and physiological alterations

observations suggest a changing perception of the value of cattle and an increasingly

Village Emergence and the Power of Human–Environment Feedbacks

As intimated above, human–environment coevolution during the Neolithic shaped

Social Dimensions of Food Production and Storage

Given the rising importance of caprine management at Aşıklı, livestock would

Networks versus Sole‑source Models of Neolithic Emergence

The forager-producer transition occurred surprisingly quickly over large areas

Wright 1994). Ancestral knowledge of these strategies was likely as geographically

Conclusion: Internal versus External Influences

The process of “Neolithization” in central Anatolia grossly parallels that of other

Table 1 Time frame of central Anatolian Neolithic sites discussed in text

Bibliography of Recent Literature

Documenting Domestication: New Genetic and Archaeological Paradigms, University of California

Mary C. Stinera, Mihriban Özbaşaranb and Güneş Duruc

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