Studies in Early Near Eastern
Production, Subsistence, and Environment 20
Neolithic Corporate Identities
Hans Georg K. Gebel
Berlin, ex oriente (2017)
Neolithic Corporate Identities
Hans Georg K. Gebel
Studies in Early Near Eastern
Production, Subsistence, and Environment 20
Berlin, ex oriente (2017)
Studies in Early Near Eastern Production, Subsistence, and Environment (SENEPSE)
Editors-in-Chief: Hans Georg K. Gebel and Reinder Neef
The Studies in Early Near Eastern Production, Subsistence, and Environment are a refereed series.
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dedicated to Klaus Schmidt
who pioneered the change in understanding the Neolithic
The construction of Neolithic corporate identities. Introduction,
by Marion Benz, Hans Georg K. Gebel and Trevor Watkins 1
Neolithic corporate identities in evolutionary context, by Trevor Watkins 13
Human palaeoecology in Southwest Asia during the Early Pre-Pottery Neolithic
(c. 9700-8500 cal BC): the plant story, by Eleni Asouti 21
Society and Economy
Neolithic corporate identities in the Near East, by Hans Georg K. Gebel 57
“Moving around” and the evolution of corporate identities in the late Epipalaeolithic
Natuan of the Levant, by Anna Belfer-Cohen and Nigel Goring-Morris 81
The construction of community in the Early Neolithic of Southern Jordan,
by Bill Finlayson and Cheryl Makarewicz 91
“I am We”: The display of socioeconomic politics of Neolithic commodication,
by Gary O. Rollefson 107
Neolithic “cooperatives”: Assessing supra-household cooperation in crop production at
Çatalhöyük and beyond, by Amy Bogaard 117
Symbols and Media
Changing medialities. Symbols of Neolithic corporate identities, by Marion Benz 135
Cultural memory: Symbols, monuments and rituals sustaining group identity,
by Christa Sütterlin 157
Religion and materialism – Key issues of the construction of Neolithic corporate identities,
by Lisbeth Bredholt Christensen 175
Dress code, hairstyles and body art markers of corporate identities in T-shaped-pillar
sites of Upper Mesopotamia? by Michael G.F. Morsch 187
Hunter into prey. Trying to make sense of the “Media Revolution” at Göbekli Tepe,
by Erhard Schüttpelz 201
M. Benz, H.G.K. Gebel and T. Watkins (eds.), Neolithic Corporate Identities
Studies in Early Near Eastern Production, Subsistence, and Environment 20 (2017): 21-53. Berlin, ex oriente
Human Palaeoecology in Southwest Asia During the Early
Pre-Pottery Neolithic (c. 9700-8500 cal BC): the Plant Story
“According to the materialist conception of history, the ultimately determining element in
history is the production and reproduction of real life … We make our history ourselves, but,
in the rst place, under very denite assumptions and conditions. Among these the economic
ones are ultimately decisive. But the political ones, etc., and indeed even the traditions which
haunt human minds also play a part … In the second place, however, history is made in such
a way that the nal result always arises from conicts between many individual wills, of which
each in turn has been made what it is by a host of particular conditions of life. Thus there
are innumerable intersecting forces, an innite series of parallelograms of forces, which give
rise to one resultant - the historical event. This may again itself be viewed as the product of a
power [that] works as a whole unconsciously and without volition. For what each individual
wills is obstructed by everyone else, and what emerges is something that no one willed.”
Letter of F. Engels to J. Bloch (London, September 21, 1890)
Abstract: This chapter tackles one of the most enduring questions posed by prehistoric archaeology
worldwide attracting the interest of prehistorians, anthropologists, economists, geographers and natural
scientists alike: how and why did late Palaeolithic societies abandon long-lived and highly successful
foraging and hunting economies in order to adopt farming? The chapter provides a critical overview of
how this transformation unfolded in Southwest Asia, the place of origin for some of the economically
most important contemporary plant and animal food staples, at the very end of the Pleistocene and the
beginning of the Holocene some 12,000 years ago. It focuses in particular on the nature of plant manage-
ment practices during this period and how they were intertwined with changes in climate and vegetation,
seasonality patterns, local micro-ecological variability, people’s historical experiences and perceptions
of the landscape, mobility strategies, community interactions, and associated symbolic and ritual be-
haviours. Some of the currently accepted notions about the nature, ecology and economic returns of
predomestication cultivation, the causes and evolution of the morphological domestication syndrome in
crop progenitor species, and the predicted impacts of climate and environmental change on economic
decision-making are critically reviewed and revisited. The chapter concludes by discussing some of the
implications of the Southwest Asian case study for understanding the nature and evolution of prehistoric
human economic behaviours, and the central role that resource ecologies play in determining the direc-
tionality and pace of macroeconomic change.
Keywords: Southwest Asia, Neolithic, domestication, climate change, niche construction theory
1 Department of Archaeology, Classics and Egyptology, University of Liverpool. <E.Asouti@liverpool.ac.uk>
Human Palaeoecology Asouti
Introduction: Setting the Issues
This chapter provides a critical overview and reassessment of the nature and palaeoecology of the Pre-Pot-
tery Neolithic (PPN) subsistence economies of Southwest Asia, one of the most intensively researched
hotspots of the transition from foraging to farming worldwide (Barker 2006). It focuses on the earliest
part of the Neolithic transformation, the rst two millennia of the Holocene. This timespan between the
end of the Younger Dryas at ~11,700 cal BP and 10,500 cal BP, broadly overlaps with the chrono-cultural
horizon identied as the early PPN (Asouti and Fuller 2013; see also Table 1, Fig. 1). In palaeoeconomic
terms, the early PPN witnessed the onset of various pre-agricultural plant and animal management prac-
tices collectively categorized as “low-level food production” (cf. Smith 2001; Asouti and Fuller 2013;
Zeder 2015). “Pre-domestication cultivation” (PDC) conventionally dened as the planting and harvest-
ing of morphologically wild crop progenitor species (Willcox 2013; Willcox et al. 2008) is considered
a typical manifestation of this mode of production in Southwest Asia, and is believed to have developed
independently across the dierent areas of the Fertile Crescent alongside a diverse array of foraging and
hunting strategies. In sociocultural terms, the early PPN is marked by the orescence of symbolic and
ritual behaviours associated with corporate social groups (e.g. lineages, sodalities, age groups, networks)
which are believed to be reected in the communal structures unearthed at several sites in both the south-
ern and the northern Fertile Crescent including Göbekli Tepe, Gusir Höyük, Mureybet, Jerf el Ahmar, Tell
‘Abr, Tell Qaramel, WF16, Tell es Sultan (Jericho) and elsewhere (cf. Kenyon 1981; Stordeur et al. 2000;
Mazurowski and Yartah 2002; Yartah 2005; Schmidt 2007, 2010, 2012; Stordeur and Ibáñez 2008; Kuijt
and Finlayson 2009; Finlayson et al. 2011; Karul 2011). Concentrations of material culture of a distinc-
tive symbolic nature (e.g. animal, human and humanoid reliefs and sculptures, pillars, bucrania and other
animal bone installations, skulls, engraved stones and other artefact caches, etc.) and burials are often
found associated with such structures. For this reason, they have been interpreted by several scholars as
foci of symbolically and/or ritually charged activities such as communal food storage and consumption,
mortuary and feasting rites, and as aggregation sites (see chapters in this volume; also Watkins 2010;
Finlayson et al. 2011; Dietrich et al. 2012; Asouti and Fuller 2013).
The ubiquity of communal buildings across the dierent regions of the Fertile Crescent along-
side the material record of regionally distinctive symbolic and ritual behaviours, are treated by some
scholars as indicators of increasing social complexity in the course of the early PPN (cf. Byrd 2005;
Watkins 2010). Inasmuch as the ecological context of these socioeconomic developments and symbolic
Fig. 1 Map showing the location of key excavated early PPN sites in Southwest Asia.
Asouti Human Palaeoecology
orescence is taken into consideration it has also been hypothesized that they were facilitated by the
rapid climatic improvement that marked the start of the Holocene (Byrd 2005). Early Holocene climatic
amelioration is believed to have prompted the emergence of highly productive and stable resource envi-
ronments. This facilitated the intensication of resource use by groups that lived permanently in favour-
able ecotones and controlled stable, well-dened territories (Zeder 2015). Overall, the current consensus
in the literature is that this unprecedented mix of ecological, socioeconomic and cultural developments
paved the way for the appearance at the end of the early PPN of plant and animal domestication, which
was followed by the establishment and spread of agro-pastoral economies and “village” life during the
late PPN (i.e., from the second half of the 9th millennium cal BC onwards) (see overviews by Harris
2002; Byrd 2005; Zeder 2008, 2015; Asouti and Fuller 2012, 2013).
Much of the ecological argument invoked in support of early PPN resource intensication and
its relation to increasing sedentism, symbolic behaviours and social complexity and the emergence
of regionally distinctive corporate institutions and identities, has been based on the conceptualization
of the rst two millennia of the Holocene as a period of remarkable climatic and ecological stability,
which radically transformed the nature and scale of the regional hunter-gatherer landscape practices.
This is contradistinguished with the pattern of punctuated climatic instability that prevailed during the
late Pleistocene, which is believed to have inhibited the macroevolutionary development of the ecolog-
ical and socioeconomic processes associated with the transition to food production (Richerson et al.
2001). The notion of the importance of resource-rich environments in the formation of food producing
economies is of course not new in the prehistoric archaeology of Southwest Asia. Similar ideas were
tested for the rst time in the 1940s and 1950s by Robert and Linda Braidwood of the Chicago Oriental
Institute, in the context of their eldwork expeditions in northern Iraq. The “Nuclear Zone Hypothesis”
(NZH) (Braidwood and Howe 1960) postulated that post-Pleistocene hunter-gatherers gradually settled
in resource-rich areas where they developed close, symbiotic relationships with preferred plant and an-
imal resources alongside processing and storage technologies that eventually led to their domestication
and the emergence of agriculture. The NZH represents in many ways the archaeological oshoot of the
much earlier theory of the “centres of origin” formulated by the Russian plant geneticist Nikolai Vavilov,
who proposed that present-day hotspots of genetic crop diversity overlap geographically with the an-
cient centres of origin of the domesticated crop species (Vavilov 1992). In Southwest Asia the “hilly
anks” (i.e., the piedmont zone) of the Taurus-Zagros arc were subsequently identied by Harlan and
Zohary (1966) as the primary habitats and centres of origin of the Near Eastern crop progenitor species,
which overlapped with Braidwood and Howe’s (1960) “nuclear zone” of initial domestication.
The NZH represents a classic example of a pull theory, based on the notion that resource abun-
dance (the “pull” factor) is a key precondition of economic, technological, and sociocultural innovation
and progress. As such, it stands in direct opposition to “push” theories positing that resource stress (in-
stead of abundance) played a central role in prehistoric socioeconomic change. Resource-stress theories
rst became popular by Gordon Vere Childe. His “Oasis Hypothesis” (inspired by a similar theory pro-
posed in 1908 by the American geologist Raphael Pumpelly) posited that climate desiccation at the end
of the Pleistocene forced the crowding of humans, plants and animals around shrinking water bodies
Late Pleistocene-Early Holocene
chrono-cultural horizons in Southwest Asia
(calibrated years BC)
Late Epipalaeolithic ~12,000-10,000
Pre-Pottery Neolithic A (PPNA) ~ 10,000/9700-8700
Early PPNB (EPPNB) ~8700-8200
Middle PPNB (MPPNB) ~8200-7500
Late PPNB (LPPNB) ~7500-7000
Pottery Neolithic ~7000-6000
Table 1 Summary of prehistoric chrono-cultural horizons and associated radiocarbon chronologies in
Human Palaeoecology Asouti
(“oases”) eventually leading to the development of symbiotic, domesticatory relationships (Childe 1928).
Later examples of push theories focused primarily on demographics (in particular population pressure) as
the cause of resource depletion and stress rather than negative climate change, including Lewis Binford’s
(1968) “Marginal Zone Hypothesis” and, in relation to it, Kent Flannery’s (1969) “Broad Spectrum Rev-
olution”. Both pull and push models of the probable causes of the Neolithic transformation represent the
intellectual backbone of several inuential theories of global agricultural origins including David Rindos’
Coevolution Theory (1984), hypotheses about the role of the Younger Dryas cold and arid spell in the
onset of early cereal cultivation and domestication (Moore and Hillman 1992; Sherratt 1997; Hillman et
al. 2001; Bar-Yosef and Belfer-Cohen 2002) and, more recently, Optimal Foraging Theory (OFT) (Ken-
nett and Winterhalder 2006; Gremillion et al. 2014) and Niche Construction Theory (NCT) (Smith 2011,
2015; Zeder 2015). OFT and NCT are also based on the same fundamental opposition between conditions
of resource stress (OFT) and resource abundance (NCT) as the prime enablers of prehistoric economic
behaviours and decision-making.
The main theoretical position developed in this chapter is that both pull and push models provide
binary and normative denitions and predictions of resource availability, properties and ecologies that
are unrealistically removed from human experience. This is because they draw on abstract concepts of
environmental stability and instability that are respectively equated with macro-climatic improvement
and deterioration. They thus provide limited tools for understanding periods of human history during
which dynamic, multi-scalar, complex processes of change were manifest in all domains of life, eco-
logical and sociocultural, and in the environment. The early PPN of Southwest Asia represents a prime
example of such a period, for which there are furthermore no viable ecological and socioeconomic an-
alogues in the historical and the ethnographic present (Asouti 2013). As it will be demonstrated later in
this chapter, despite the dramatic climatic improvement that marked the onset of the Holocene, its rst
two millennia were an era of profound short- to medium-term ecological instability that was particularly
pronounced in the semi-arid, continental interiors of Southwest Asia, and impacted signicantly the
distribution and predictability of landscape resources. Therefore, a more realistic reconstruction of early
PPN human palaeoecology requires a more dynamic standpoint, one that takes into account the regional
bioclimatic and ecological diversity, and the complexity and dynamics of the interactions between cli-
mate, resources and economic behaviours. Moreover, in the diverse and rapidly changing environments
of early PPN Southwest Asia economic behaviours did not depend solely or even primarily on expedien-
cy, but also on historical landscape experiences and memories of past events, which informed socialized
forms of environmental knowledge transmission.
Based on these premises, this chapter proposes an explicitly historical-ecological approach focus-
ing on understanding the ecological impacts of short- to medium scale climate oscillations (centennial,
decadal), the seasonality of critical variables (precipitation, temperature), how they aected the balance
of woodland and grassland vegetation, re frequency and herbivory, and their cumulative impacts on
resource distributions, ecology, physiology and phenotypes. Such a survey of the regional ecologies
(rather than the traditional format of palaeoecological investigations in Southwest Asia concentrating
on millennial-scale environmental change) reveals a picture of rich, yet fragmentary and seasonally
unstable resource environments and highly uctuating resource ceilings. It is argued that these eco-
logical constraints severely limited the capacity of PDC to generate sucient and predictable enough
subsistence yields, and therefore the ability of early PPN hunter-gatherer societies to depend on it as
the staple subsistence provider. Meso- to micro-ecological instability, experienced in the context of the
resource-rich environments of the rst two millennia of the Holocene, also provided early PPN societies
with the impetus for developing locally distinctive resilience strategies. These included exible eco-
nomic behaviours alongside social practices that fostered inter- and intra-group cooperation through the
acquisition, storage, and transmission of environmental and landscape knowledge, and the circulation of
material culture and foodstus through community interaction networks. Furthermore, the material cul-
ture record provides evidence for the prominent role of historical experiences of environmental change
in early PPN symbolic behaviours and cosmologies.
A cornerstone of the arguments presented in this chapter relates to the reconstructed impacts of
short- to medium-term climatic instability on the ecology, productivity and predictability of crop pro-
genitor species and associated economic behaviours. With notable exceptions (e.g. Hillman 1996) the
landscape ecology of plant gathering and early cultivation is an issue that has been mostly overlooked
by archaeobotanical studies of PDC and the regional evolution of the domestication syndrome. More
often than not plant domestication in Southwest Asia is approached from a narrowly dened biological
viewpoint, in isolation from its wider ecological and sociocultural contexts. However, these contexts
provide an appropriately inclusive framework for understanding the nature and limitations of early PPN
Asouti Human Palaeoecology
plant management practices and, by extension, the underlying causes of the slow pace of morpholog-
ical domestication observed during this period (see discussion in Asouti and Fuller 2013). This argu-
ment is developed further in this chapter, by drawing on previously overlooked aspects of the regional
palaeoecological and archaeobotanical records. It is proposed that early Holocene climate change and
associated vegetation ecologies (rather than the intensication of PDC by early PPN hunter-gatherer
communities) played a signicant role in the development of larger seed size in cereal crop progenitor
species. The archaeobotanical and ecological indicators traditionally associated with the identication
of PDC are also reviewed in detail, in order to provide a more realistic reconstruction of its nature and
predicted ecological impacts and economic returns. Furthermore, drawing on recent genetic studies, it is
argued that community interactions (rather than the polycentric development of PDC by insular seden-
tary communities) played a pivotal role in the regional spread of socially valued cultivars that gave rise
to domesticated crop species during the late PPN.
The agronomic limitations of PDC were not overcome until the late PPN (i.e., after the mid-9th
millennium cal BC) in a process that overlapped temporally and spatially with the spread of domesti-
cated caprine herding. It is proposed that the use of animal manure as fertiliser was the tipping point,
by enabling the development of xed-plot intensive horticulture and boosting the productivity of cereal
and pulse cultivation while also buering it against seasonal environmental risks. The herding of do-
mesticated caprines altered irreversibly the ecological balance of Southwest Asia through the conversion
of steppe grasslands into pastures. It is argued that it was these direction-changing developments in the
nature, technology and organization of food production, rather than the putative impacts of PDC on the
biology of managed plant species, that enabled the spread of domesticated crop mixtures and anthropo-
genic agroecologies and the establishment of long-lived, sedentary “village” communities dependent on
mixed agropastoral production as the staple subsistence provider. The chapter concludes by outlining a
historical-evolutionary model for the comparative study of global agricultural origins and, more gener-
ally, human economic behaviours during periods of major ecological and socioeconomic change. This
emphasises the need to construct hypotheses that address the interconnectedness of geographically and
historically contingent resource ecologies with the multi-layered ecological, economic and sociocultural
factors that constitute human lifeways.
Climate Change and the Relationship of Resource Ecologies to Resource Choice in Late
Pleistocene and Early Holocene Southwest Asia
Global palaeoclimatic archives derived from marine sediments and polar ice cores have established the
existence of large-scale shifts in global climate (e.g. glacial-interglacial climate cycles) caused by the in-
teraction of the Milankovitch cycles: periodic variations in the Earth’s eccentricity, axial tilt and preces-
sion (lasting ~100 kya, 41 kya and 19 kya respectively) that impact the amount, seasonality and location
of solar insolation around the planet at supra-millennial time scales (Hays et al. 1976; Wunsch 2004).
The last Ice Age was paced by shorter-scale climate oscillations, known as the Dansgaard-Oeschger
(DO) events, occurring in cycles of ~1500 years: in the northern hemisphere these took the form of
decadal-scale warming episodes, followed by gradual cooling over a longer period of time that was then
terminated by another decadal-scale cold and arid episode (Dansgaard et al. 1993; Bond et al. 1997;
Alley 2000). Some of the DO cycles were preceded by rapid cold episodes known as the Heinrich (H)
events that lasted ~1000 years and resulted from the release of ice raft debris in the North Atlantic (Bond
et al. 1992; Bond and Loti 1995) (see also Fig. 2). Some researchers identify the Younger Dryas (dated
in the ice core records at ~12,900-11,500 cal BP) as the last Heinrich event (H0) of the Pleistocene,
while others attribute its rapid onset to the release of large amounts of glacial meltwater from North
America (cf. Bond and Loti 1995; Broecker 1998). Typically, the end of the Younger Dryas is identied
through abrupt increases in temperature and precipitation that marked the onset of the Holocene (Alley
et al. 1993; Severinghaus et al. 1998). This improving trend was reversed at ~11,400 cal BP by a cold
episode known as the Pre-Boreal Oscillation (PBO) that lasted ~200 years and was terminated by anoth-
er decadal-scale warming at ~11,270 cal BP (Bjork et al. 1997; Kobashi et al. 2008). Reconstructions of
surface temperatures have indicated that temperatures after the PBO were warmer than before it, which
has led some authors to hypothesize that this phenomenon may be consistent with observations of a two-
step warming at the onset of the Holocene (Seppä et al. 2002; Kobashi et al. 2008). After ~11,200 cal BP
Holocene climate was characterized by remarkable, by comparison to earlier periods, stability; climatic
conditions reached an optimum by ~9000 cal BP. During this period, the most notable change in north-
ern hemisphere climate was in temperature seasonality. Annual mean changes in surface air temperature
Human Palaeoecology Asouti
were of lower amplitude than seasonal
changes: between ~12,000-8000 cal BP
winters were cooler and summers warm-
er than today, with the maximum win-
ter-summer dierence observed around
9000 cal BP (Brayshaw et al. 2011).
In Southwest Asia, the Younger
Dryas has been identied as a period of
intensely cold and arid conditions. Jones
et al. (2007) have calculated from lake ox-
ygen isotope records that precipitation and
temperature were lower by comparison to
the late Holocene (see also Bar-Matthews
et al. 1997; Table 2). This general precip-
itation pattern appears to have been con-
sistent between Anatolia and the Levant
although, as expected, regional N-S and
E-W gradients are also evidenced in the
available records: the Soreq cave speleo-
thems in the southern Levant record high-
er values than the Eski Acıgöl crater lake
in central Anatolia, and the latter higher
values than lake Van in eastern Anatolia.
Terrestrial pollen records match closely
these regional precipitation gradients: in
the Hula basin catchment (situated in the
Mediterranean Woodland Zone of the Le-
vantine littoral) oak pollen values dropped
from ~70% during the warm and wet
Bølling-Allerød interstadial to ~30% dur-
ing the Younger Dryas stadial, while at the
same time grass pollen values increased to
30% (Baruch and Bottema 1999; Bottema
2002; Wright and Thorpe 2003) (Fig. 3).
This pattern of vegetation response sug-
gests that in the wetter (by comparison to
inland Southwest Asia) Levantine littoral,
grasses were able to compete more eec-
tively with trees for nite ground moisture
resources under the cold and arid condi-
tions of the Younger Dryas. Further north,
in the Syrian Ghab valley catchment its
re-dated pollen sequence points to a more
pronounced decline of oaks during this
period, and the coeval expansion of Arte-
misia-Chenopodiaceae steppe instead of
grasses (Niklewski and Van Zeist 1970;
Wright and Thorpe 2003). Similarly, in the
more continental inland regions of central
and eastern Anatolia the Younger Dryas
was marked by the retreat of grasslands
and the expansion of Artemisia-Chenopo-
diaceae steppe (Roberts et al. 2001; Wol-
dring and Bottema 2002/2003; Wick et al. 2003). The combined pollen and anthracological data also
indicate that, in contrast to the western Levantine littoral, trees never formed a signicant component
of the vegetation of inner Anatolia during the warm and wet conditions of the Bølling-Allerød. Instead,
grassland vegetation (including Cerealia) prevailed associated with a sparse, low-density tree cover
(dominated by members of the Rosaceae family, Pistacia, with low representation of Juniperus and
Fig. 2 INTIMATE Project stratigraphy of Greenland Stadial/
Interstadial (GS/GI) cycles for the last 48 kyr BP (before 2000
AD) based on the Greenland ice core records and GICC05 ice
core chronology (modied after Blockley et al. 2012: Fig. 1).
Asouti Human Palaeoecology
deciduous Quercus) (Roberts et al. 2001; Woldring and Bottema 2001/2002; Litt et al. 2009; Asouti and
Kabukcu 2014; Kabukcu in press) (see also Fig. 4). Further east, on the Zagros mountain range, sparsely
wooded Pistacia grasslands were dominant during the Bølling-Allerød, which were replaced by Arte-
misia-grass steppe during the Younger Dryas (Van Zeist 2008). These important dierences between the
vegetation histories of the Mediterranean and the Irano-Anatolian bioclimatic regions are veried by the
exceptionally long pollen sequences obtained from lakes Van, Urmia and Zeribar, which conrm that,
unlike grasses, trees did not form a prominent element of the vegetation of continental inland Southwest
Asia before mid- to late Holocene times2 (Bottema 1986; Wick et al. 2003; Djamali et al. 2008; Van
Zeist 2008; Litt et al. 2009).
Previous models of the impacts of climate change on the availability of plant resources to late
Pleistocene hunter-gatherers assumed the existence of a positive correlation between stable “climax”
arboreal-grassland habitats (dominated by deciduous oaks and/or Pistacia) and periods of climate im-
provement, and of steppe habitats (dominated by shrubs and herbs) with periods of climate deterioration
(e.g. Zohary 1989; Hillman 1996). However, as discussed above, the available data on the regional Late
Glacial palaeoclimatic and vegetation histories reveal a much more complex picture of terrestrial biome
responses to the Younger Dryas across the dierent bioclimatic regions of Southwest Asia. Recently ob-
tained palaeoclimatic records from the Soreq cave in Israel have provided additional insights into aspects
of climate that are critical for subsistence economies, particularly seasonality: while climate conditions in
the southern Levant were overall arid and cold during the Younger Dryas, seasonal variations in precip-
itation appear to have been of lower amplitude by comparison to the Holocene and the Bølling-Allerød
(Orland et al. 2012). In addition, due to lower temperatures and decreased evaporation rates, major wa-
ter-bodies such as the Dead Sea appear to have been characterized by a more positive water balance, in-
ferred by high lake levels (Litt et al. 2012). In turn, these inferences of decreased precipitation seasonality
point to the possibility that in the southern Levant resource predictability (hence the ability of prehistoric
groups to anticipate and cope with seasonal resource shortfalls) might have been somewhat higher during
2 Djamali et al. (2008) have reported a pre-Holocene peak in oak pollen observed in the long sequence from lake Urmia
during the last interglacial period. However, the presence in the same pollen zone of the mesic, thermophilous, Euxino–Hyrca-
nian element Zelkova caprinifolia (which is notably absent from Holocene pollen spectra) points to climate conditions (milder
winters and more spring or summer rainfall) during the last interglacial that were much more favourable for the expansion and
establishment of woodland vegetation by comparison to the Holocene.
Dates cal BP Key climatostratigraphic
INTIMATE Project no-
menclature for Green-
land Stadial – Intersta-
dial cycles (GS/GI)
General climate trends in South-
21,200-14,700 Last Glacial GS-2 Cold, arid (low precipitation and
14,700-12,650 Bølling-Allerød interstadial GI-1
Warm and wet (increasing tempera-
tures [14.5-18.0oC] and precipitation
[~550-750 mm p.a.])
12,650-11,500 Younger Dryas stadial GS-1 Cold, arid (low precipitation and tem-
11,500-8,200 Early Holocene
Warm and wet (increasing tempera-
tures ~14.5-19.0oC and precipitation
~675-950 mm p.a.); increased sea-
sonality of climate with wet winters
and dry summers.
8,200-present Mid Holocene to present
Establishment of modern climatic
regime (temperatures ~18.0-22.0oC;
precipitation ~450-580 mm p.a.);
increasing aridication impacts com-
pounded by anthropogenic impacts
on the environment
Table 2 Major climatic regimes in the Eastern Mediterranean from the Last Glacial Maximum to the Holocene
(based on Bar-Matthews et al. 1997, 1999; Robinson et al. 2006; Orland et al. 2012; Dean 2014; see also Fig. 2).
Human Palaeoecology Asouti
the Younger Dryas independently
of net resource ceilings. Palae-
oecological records thus add an
important new dimension to on-
going debates about the nature
of the transition from the early
to the late Natuan, and the evo-
lution of the southern Levantine
late Epipalaeolithic subsistence
strategies (see discussion in Henry
2013). This inference of decreased
climate seasonality during the
Younger Dryas seems unlikely to
have applied to other regions of
Southwest Asia. High lake levels
have not been deduced from pal-
aeolimnological records in central
Anatolia (Dean 2014) while, as
discussed earlier, trees and grass-
lands appear to have been equally
negatively impacted by the cold
and arid conditions of the Young-
er Dryas. Evidence for habitation
sites dating from this period is,
tellingly, lacking from the cen-
tral Anatolian plateau (Woldring
and Bottema 2001/2002). In
other parts of inland Southwest
Asia, prehistoric groups adjust-
ed their mobility and subsistence
strategies to the marked shifts in
the availability of woodlands and
grassland biomes, by managing
ecotonal catchments characterized
by more diverse ecologies includ-
ing alluvial-steppe and persistent
arboreal habitats (cf. Savard et al.
2006; Asouti and Fuller 2012;
Riehl et al. 2012; Willcox 2012a;
Henry 2013 and references there-
in). Thus it appears that only in
the harshest environments (e.g.,
in central Anatolia) the Younger
Dryas might have forced prehis-
toric communities to adopt highly
mobile lifeways leaving few if any
The hypothesis that cultivation and domestication rst emerged in the Levant during the Young-
er Dryas as a response to the reduction of wild cereal stands due to adverse climatic conditions, saw its
initial verication in Gordon Hillman’s analysis of the archaeobotanical remains from Abu Hureyra in
northern Syria (Hillman 2000). Hillman based his conclusions on the nds of larger “plump” rye seeds in
late Epipalaeolithic layers and the coeval increase in small-seeded grasses and legumes, which he viewed
respectively as evidence for selection for large seed size under cultivation and the local development of
an arable “weed” ora (Hillman et al. 2001). While the nds of domesticated-type rye grain from the late
Natuan levels of Abu Hureyra were soon afterwards dismissed as intrusive (Nesbitt 2002) the impression
of the site as a Natuan example of PDC has persisted in the literature (cf. Willcox 2012a). However,
in their recent comprehensive re-assessment of the Abu Hureyra archaeobotanical record Colledge and
Conolly (2010) have cast serious doubt on the hypothesis of late Natuan PDC. They argue that the higher
Fig. 3 Select curves from the Lake Hula pollen diagram (shaded area
indicates early Holocene) (dating follows Wright and Thorpe 2003).
Asouti Human Palaeoecology
frequencies of small-seeded grasses and legumes in the late Natuan archaeobotanical samples can be
more parsimoniously explained as plant foods gathered from the wild. Colledge and Conolly propose that
the gradual reduction in the frequencies of large-seeded cereals and legumes, and the coeval increase of
small-seeded grasses and legumes, are more likely to reect the substitution of high-ranked, large-seeded
plants as the preferred plant food subsistence source by a broad spectrum of low-ranked, small-seeded taxa.
They thus interpret the changes observed in relative taxon frequencies as evidence of an increase in di-
et-breadth through time, which was caused by negative climate impacts on the availability of high-ranked,
large-seeded taxa in the environs of Abu Hureyra during the Younger Dryas (Colledge and Conolly 2010:
137 [thus largely following a similar line of argument to that previously proposed by Hillman et al. 1989]).
The Abu Hureyra case study represents a classic example of the conceptual divide between
“environmental determinist” and “optimal foraging” models of prehistoric economic behaviours (cf.
Bettinger 1991; Winterhalder 2001). While the former view prehistoric societies as passively responding to
external impacts on the resource base (exemplied in the case of Abu Hureyra by Hillman’s interpretation
of the pivotal role played by the Younger Dryas in the adoption of cultivation), the latter view economic
decision-making as underpinned by environmentally constrained resource selection. Resource selection
refers to the ranking of resources according to their abundance and preference. In turn, preference is dened
primarily by microeconomic criteria including caloric yields and acquisition (collection and processing)
costs. The interpretation of the Abu Hureyra archaeobotanical sequence proposed by Colledge and
Conolly represents an example of an optimal foraging model, with its emphasis on diet-breadth increase
as a response to the reduced availability of high-ranked resources. However, modelling resource choice
based on microeconomic benchmarks can be misleading on both ecological and economic grounds. For
example, collecting and processing of low-ranked, small-seeded plants is typically predicted to be low-
return in terms of harvested calories and more labour-intensive compared to high-ranked, large-seeded
species (e.g. crop progenitor species). Yet, it has been observed that small-seeded grasses harvested from
the wild tend to have lower seed-cha ratios and a higher number of seeds per stalk compared to large-
Fig. 4 Select curves from the Lake Van (Eastern Anatolia) and Eski Acıgöl (Central Anatolia) pollen diagrams
(original data published by Wick et al. 2003; Litt et al. 2009, and Roberts et al. 2001; Woldring and Bottema
Human Palaeoecology Asouti
seeded glume progenitor species
such as wild-type einkorn,
emmer and barley (Blumler
2002). More recent studies have
found no signicant dierences
in yields and seed-cha ratios
between progenitor and non-
progenitor taxa (Preece et al.
2015). A dierent reading of the
Abu Hureyra archaeobotanical
data might thus emphasize
the greater availability and
abundance of small-seeded
grasses and legumes in the
site environs as the main
determinant of resource
selection by its inhabitants
during the Younger Dryas,
and the opportunities they
aorded for routine subsistence
scheduling. Due to their wider
ecological preferences these taxa are also likely to have returned more predictable yields by comparison
to large-seeded species irrespectively of (assumed) processing costs. Large-seeded species would actually
have been more costly to pursue, because they would have had to be collected from increasingly scarce
and diminishing natural stands or (following Hillman’s interpretation) their seeds planted and tended for
several months under a suboptimal climatic regime before they could actually return a yield.
The assumption of substantial reliance on the management of seed plants, including large-seeded
cereal progenitor taxa, during the Natuan period in the Levant is widespread in the literature (Bar-Yosef
1998; Valla 2000; Byrd 2005). Direct archaeobotanical evidence is available from few sites, of which only
two have provided indications for the signicant presence of wild-type cereals: early Natuan Dederiyeh
cave (Tanno et al. 2013) and late Natuan Abu Hureyra 1 (Hillman 2000) both predating the Younger Dryas.
Other northern Levantine facies contemporary with the Younger Dryas include the later phases (2-3) of
Abu Hureyra discussed above, Mureybet 1 (Van Zeist and Bakker-Heeres 1986) and the Baaz rock-shelter
(Conard et al. 2013). These sites have provided very little or no evidence for reliance on crop progenitor
species. In the southern Levant, phytolith analyses from early Natuan sites located in the Mediterranean
Woodland Zone have indicated that plant-based subsistence derived mainly from tree nuts and fruits rather
than large-seeded grasses; by contrast, phytolith nds from late Natuan sites point to an increased reli-
ance on non-cereal grass taxa (Rosen 2010, 2013; see also overview by Asouti and Fuller 2012). However,
at the same time (and in agreement with the o-site palynological archives discussed earlier) pollen data
have revealed the co-existence of both cereal and tree pollen at several Natuan sites (see Henry 2013 and
references therein). Overall, the combined macrobotanical, phytolith and pollen records point to signicant
variations (alongside some points of convergence) in Natuan plant-based subsistence strategies between
the southern and the northern Levant, which cannot be easily reconciled with the expectations of classic
“push” ‘(i.e., resource-stress) models of prehistoric economic behaviours. Instead, they appear much more
likely to reect regional environmental gradients and diversity in the availability and local ecologies of
plant resources across dierent landscape units (e.g., upland and steppe areas, steppe-woodland ecotones
and the Mediterranean Woodland Zone).
Across Southwest Asia the termination of the Younger Dryas ushered in a period of warm and
wet conditions corresponding to the early Holocene (~11,700-6000 cal BP) (Robinson et al. 2006; Dean
2014). The Younger Dryas-early Holocene transition was markedly abrupt. Recent analyses of annually
laminated lake sediments from Nar Gölü in Cappadocia (central Anatolia) have indicated that the tempo of
the climatic transition was very punctuated, with over half of it occurring within a decade (Dean 2014). A
similarly rapid event of ~12 years is registered in the Soreq cave speleothems (Orland et al. 2012) (Fig. 5).
The Nar Gölü sequence indicates that this extremely rapid, decadal-scale event was followed by a brief
very wet episode lasting for 26 varve years, which was terminated at ~11,400 cal BP by a bimodal cooling
and arid event (corresponding to the PBO) that lasted for 126 varve years (Dean 2014) (Fig. 6). While the
Soreq cave sequence is poorly resolved with regard to the start of the Holocene, the available data never-
theless suggest the prevalence during this period of high-amplitude climate shifts, which did not stabilise in
Fig. 5 The Younger Dryas termination as recorded in the Nar Gölü
stable oxygen isotope sequence (modied after Dean 2014: Fig. 10.6).
Asouti Human Palaeoecology
a Mediterranean-type climatic re-
gime before ~10,500 cal BP (Or-
land et al. 2012). Other regional
palaeoclimatic records alongside
climate modelling have also in-
dicated that the early Holocene
was characterized by heightened
seasonality in surface air temper-
ature, with markedly dry sum-
mer conditions and high levels
of winter precipitation (COHM-
AP 1988; Robinson et al. 2006;
Brayshaw et al. 2011) (Fig. 7).
Thus, although early Holocene
climate was on the whole wetter
and interannual variability was
also lower (Allcock 2013; Dean
2014) at the same time seasonal-
ity was particularly pronounced
with colder and wetter winters
and hotter and more arid summers
by comparison to later periods.
The most noticeable ter-
restrial response to the rapid cli-
matic improvement at the start of
the Holocene was the dramatic
expansion of grasslands, includ-
ing cereal progenitor taxa, which
reached their greatest extent dur-
ing this period particularly in inland Southwest Asia. Grassland expansion in the rst two millennia of the
Holocene also coincided with a peak in wildre signals, deduced from micro-charcoal records and charred
plant macrofossil frequencies in lake sediments (Wick et al. 2003; Wasylikowa 2005; Langer and Wasy-
likowa 2008; Turner et al. 2008, 2010). These studies found very little evidence to suggest that grassland
res were primarily of anthropogenic origin. Their higher frequency ts very well with the reconstructed
seasonality patterns for this period, and most likely resulted from the increased availability of high grass
fuel loads that were extremely susceptible to re disturbance during the hot and dry summer seasons (Turn-
er et al. 2010). Once more it is possible to trace divergent trajectories of regional re histories between the
west Levantine littoral and the semi-arid, continental interiors of Southwest Asia: micro-charcoal records
from the Hula basin in Israel have indicated that woody plants made a greater contribution to micro-char-
coal inux; by contrast, sites in continental inland regions such as Eski Acıgöl (central Anatolia), lake Van
(eastern Anatolia) and lake Zeribar (Zagros) present the reverse pattern with the predominance of grass
charcoals (Wasylikowa 2005; Turner et al. 2010: Fig. 7). Overall, this pattern matches very closely the
available pollen and anthracological and seed archaeobotanical records, which point to signicant regional
variations in the composition, density and structure of early Holocene woodland vegetation: oak-grass
vegetation was dominant in the Levantine littoral while sparsely wooded Pistacia-Rosaceae grasslands
prevailed in semi-arid inland areas (cf. Roberts et al. 2001; Woldring and Bottema 2001/2002; Wick et al.
2003; Wright and Thorpe 2003; Van Zeist 2008; Litt et al. 2009; Asouti and Kabukcu 2014, Asouti et al.
2015; Riehl et al. 2015; Kabukcu in press). Some authors (Roberts 2002; Turner et al. 2010) have hy-
pothesized that these regional dierences in vegetation composition and ecology might reect early PPN
human impacts on woodlands, which delayed the spread of trees, especially deciduous oaks, across inland
Southwest Asia. However, considering also the evidence (already discussed) for the sparseness and open
structure of Late Glacial woodlands in the continental interiors of Southwest Asia, a more parsimonious
explanation is that this time-lag reects the more rapid response of grasslands to the abrupt increases in
moisture and temperature at the start of the Holocene. Cyclic vegetation disturbances, including high-am-
plitude seasonal dierences in precipitation and temperature resulting in higher natural re frequencies,
would have provided grasses with an additional competitive advantage over trees. The prevalence of grass-
lands over woodlands would have been especially pronounced in inland regions that were characterized by
lower annual rainfall averages compared to those of the Mediterranean littoral (Asouti and Kabukcu 2014).
Fig. 6 Comparison of Nar Gölü and NGRIP Holocene stable
oxygen isotope records (modied after Dean 2014: Fig. 10.11).
Human Palaeoecology Asouti
Early Holocene Resource-rich Environments and the
Nature of Early PPN Low-level Food Production
In recent years, resource abundance models of past economic behaviours have provided useful insights
into the structure and ecologies of early Holocene low-level food production. Most prominent has been
Cultural Niche Construction Theory (CNC) (for detailed overviews see Smith 2011, 2015; Zeder 2015).
The basic premise of the CNC is that, instead of passively adapting to environmental conditions, hunt-
er-gatherer groups living on the eve of food production intentionally manipulated and modied their
habitats in order to enhance resource productivity, stability and predictability. Smith (2011, 2015) has
dened the main predictions of the CNC framework for the characterization of early Holocene low-level
food production economies as follows:
(1) They occupied relatively small and few in number settlements located in resource-rich environments
and controlled spatially limited, well-dened resource territories.
(2) Archaeobiological assemblages contain evidence for the harvesting of a broad and diverse spectrum
of species from biotic communities with no evidence for resource depression.
(3) They established various forms of ownership of “wild” (i.e., biologically non-domesticated) resourc-
es and resource-rich territories.
(4) They maintained and consistently updated a comprehensive knowledge of local ecosystems, land-
scape activities and environmental experience, which was encoded in stories, belief systems and cos-
(5) They engineered ecosystems over multiple generations via sustained and repetitive resource man-
agement practices and traditional ecological knowledge transfer, resulting with time in major ecological
and genetic transformations of the biotic components of ecosystems.
(6) They increased the relative abundance, predictability, and availability of targeted wild species within
resource-catchment areas by enhancing their net primary productivity through niche construction, in-
cluding modifying local environments through activities such as burning and clearance.
Several elements of the CNC framework provide an overall good t with the early PPN archae-
ological and palaeoecological records of Southwest Asia. As discussed in the previous section, climatic
Fig. 7 Modelled climate seasonality for the northern hemisphere continents and the Eastern Mediterranean
regions (top) and comparison of insolation changes with aridity trends reconstructed from stable oxygen isotope
records at Nar Gölü (+, -, 0 denote increase, decrease and no change relative to the pre-industrial present
respectively) (modied after Brayshaw et al. 2011: Table 3.2 and Dean 2014: Fig. 10.10).
Asouti Human Palaeoecology
improvement was very rapid at the start of the Holocene resulting in the equally rapid, albeit regionally
variable, expansion and abundance of grasslands, woodlands and water resources. Regional settlement
patterns indicate that habitation sites were relatively small (0.5-1 hectares on average) and widely dis-
persed in the landscape. A degree of residential and/or logistical mobility is suggested by the occurrence
of sites that functioned as transient hunting/foraging camps and activity areas, the relatively limited
lifespan of several habitation sites (often not exceeding a few centuries), and the presence of stratigraph-
ic and radiocarbon discontinuities even at sites with long habitation sequences, which indicate episodes
of site abandonment and re-occupation (see also Asouti and Fuller 2013 and references therein). The
archaeobiological record points to the increasing exploitation of diverse and regionally distinctive reper-
toires of plant and animal species. Signicantly, there is very little evidence for even localised resource
depletion during this period (cf. Starkovich and Stiner 2009; Zeder 2012). It is also interesting to note
that (as least with regard to the available pollen, anthracological and archaeobotanical records) evidence
for signicant early PPN human impacts on the landscape has remained hitherto elusive (cf. Asouti and
Kabukcu 2014; Asouti et al. 2015; Kabukcu in press). It seems doubtful therefore whether relatively
long-lived residential bases managing tightly controlled and temporally stable resource territories were
as common in the early PPN settlement patterns of Southwest Asia as is sometimes implied in the re-
gional archaeological literature. The earliest detectable human landscape impacts in the regional pollen
and anthracological records date from the late PPN (i.e., from the late 9th millennium cal BC onwards).
The available evidence points to the suppression of grasslands and the coeval expansion of semi-arid
woodland pastures, due to the combined eects of domesticated caprine grazing and increasing wood-
land management practices that promoted the spread of preferred rewood species across the semi-arid
regions of the southern Levant, inner Anatolia, and the Taurus-Zagros foothills and mid-altitude slopes
(Asouti and Kabukcu 2014; Asouti et al. 2015; Kabukcu in press). Furthermore, the establishment and
spread of regionally distinctive agroecologies also dates from the late PPN and the Pottery Neolithic
periods (Colledge 2001; Colledge et al. 2004).
One of the most important contributions of CNC in agricultural origins research is its conceptual-
ization of low-level food production as “multigenerational ecosystem engineering” targeted at generating
sizeable and predictable resource yields (Smith 2011; Zeder 2015). In Southwest Asia, PDC forms one
of the most plausible candidates of such a process. In the regional archaeobotanical literature PDC is
widely viewed as representative of the intensication of crop progenitor cultivation by year-round settled
communities, and as a direct precursor to crop domestication and the development of agriculture. Its ar-
chaeobotanical indicators have been attested at several early PPN sites across the Fertile Crescent (cf. Van
Zeist and Bakker-Heeres 1986; Van Zeist and de Roller 1991/1992; Kislev 1997; Colledge 1998, 2001;
Edwards et al. 2004; Tanno and Willcox 2006; Weiss et al. 2006; Feldman and Kislev 2007; Willcox et al.
2008, 2009; White and Makarewicz 2012; Riehl et al. 2012, 2013, 2015; Willcox 2012a). Typically, the
presence of PDC is assessed through a combination of archaeobotanical and ecological criteria including:
(a) an increase in grain size, (b) the decline in the presence and relative frequencies of non-progenitor
seed taxa coevally with the increasing frequencies of crop progenitor species, (c) the identication of
“weed” oras in higher proportions than their expected presence and abundance in natural grassland
vegetation, and (d) the transference of crop progenitor species outside their predicted natural habitats and
geographical distributions (cf. Colledge 1998, 2001; Willcox et al. 2008; Willcox 2012b).
These criteria provide a useful yardstick with which to assess empirically the ecology of PDC,
and the agronomic stability and predictability of its economic returns in early Holocene Southwest Asia.
Beginning with seed size, the predominant view in the regional archaeobotanical literature is that it in-
creased primarily as a response to the favourable conditions generated by cultivation: soil disturbance
via clearance and tillage, and deep seed burial through planting (Fuller 2007; Willcox et al. 2008). How-
ever, it is also the case that large-seeded progenitor species growing naturally on heavy and deep terra
rossa and alluvial soils will display the same plastic response (Blumler 2002). Seed size is furthermore
strongly aected by density stand, inter-specic competition and rapid climate change, all of which
impose selection on plant populations (Neytcheva and Aarssen 2008; Nicotra et al. 2010; Cunni et
al. 2014). The heightened seasonality of the rst two millennia of the Holocene (characterized by wet
winters and pronounced summer aridity) would also have favoured the development of large-seeded
varieties (Blumler 1992). Systematic archaeobotanical evaluations of the regional rates and pace of seed
size increase have indicated that it was a step-like process, with most of it occurring during the early
Neolithic (Willcox 2004). However, at the same time, seed size increase does not appear to have been
associated with other archaeobotanically more reliable phenotypic indicators of domestication, such as
the occurrence in signicant proportions of non-shattering rachises, which are not reported from any of
the sampled early PPN sites. According to Willcox (2004) the absence of evidence for other phenotypic
Human Palaeoecology Asouti
changes reduces the likelihood that seed size increase in cereal cultivars resulted from selection pres-
sures imposed by long-lived cultivation practices. Systematic comparisons of cereal seed sizes recorded
from the later phases of early PPN sites such as Jerf el Ahmar and Dja’dé with those from the Chal-
colithic site of Kosak Shamali (all in northern Syria), have also indicated that there was no signicant
grain size increase after the initial leap manifested during the early PPN (Willcox 2004; see also Fig. 8).
In the absence of other indicators of phenotypic change Willcox (2004) concluded that early PPN seed
size increase might reect the introduction of exotic “plump-grain” varieties into northern Syria from
moister habitats further north in Anatolia, where wild cereals probably grew under more favourable con-
ditions. An alternative explanation for this step-like increase in grain size, which furthermore accounts
for its coeval occurrence in areas outside the northern Euphrates basin, relates to early Holocene uctu-
ations in atmospheric CO2 concentrations and their potential impacts on plant productivity. CO2 values
derived from leaf stomatal index data in northwest Europe have indicated a rapid increase from 210-215
ppmv at the beginning of the Younger Dryas to 270-290 ppmv at the start of the Holocene; after a drop
to 240-250 ppmv during the PBO (~11,400-11,270 cal yr BP) CO2 levels rose again to 270-290 ppmv
until ~10,800 cal BP (Rundgren and Björck 2003; Fig. 9). In turn, the chronology of these uctuations
correlates very well with the beginning of the later early PPN phase at Jerf el Ahmar (11,200 cal BP)
while it also overlaps with the greater part of the habitation at Dja’dé (~11,000-10,300 cal BP). Regard-
less of the ultimate causes of early PPN seed size increase, the inescapable conclusion seems to be that
the potential role of early Holocene climate change in this process has been seriously underestimated;
it appears unlikely that repeatedly practiced, multigenerational, stable cultivation activities and their
assumed ecological and phenotypic impacts were the major contributing factors.
With regard to seed assemblage composition, the available archaeobotanical datasets reveal con-
siderable diversity and variation between sites and across the dierent regions of Southwest Asia. This is
expected if one considers the diversity of the regional climate gradients and associated vegetation ecolo-
gies (see previous section; also overviews of the composition of early PPN archaeobotanical assemblages
in Asouti and Fuller 2012, 2013; Riehl et al. 2013, 2015). Again, the potential role of climate change and
regional ecological variation resulting from natural vegetation disturbance is not suciently emphasized
in the literature. Inter-regional variation in the presence and relative abundance of cereal crop progenitor
species likely relates to climate factors, especially the length of the rainy season (Blumler 1996, 2002).
Grasses (including cereal progenitor taxa) are present in a majority of early PPN sites. The predominant
presence of barley in Levantine sites as opposed to einkorn in the northern Fertile Crescent may reect
the alignment of plant-derived subsistence averages with the general regional gradients in temperature
and the length of the rainy season. By contrast, the dierences observed in assemblage composition be-
tween sites located in the same bioclimatic region may reect local micro-ecological diversity alongside
cultural preferences. Several sites in eastern Anatolia and northwest Zagros (e.g. Hallan Çemi, Demirköy,
Qermez Dere, M’lefaat) and in the southern Levant (e.g., ‘Iraq ed-Dubb, Netiv Hagdud) contained large
quantities of non-cereal taxa and nuts suggesting their preferential management as subsistence mainstays
(cf. Colledge 2001; Savard et al. 2006; Willcox and Savard 2011; Willcox 2012a). At other eastern Anato-
lian sites such as Körtik Tepe large-seeded grasses were abundant, although on the whole progenitor taxa
formed a very small component of the archaeobotanical assemblage (Riehl et al. 2012). Annual legumes
had a signicant presence at several sites in Anatolia and the Zagros (e.g., early PPN phases at Çayönü
and Nevalı Çori, Hallan Çemi, Qermez Dere, M’lefaat; references above, also Van Zeist 1988; Pasternak
1998). Fire disturbance favours the spread of annual legumes by breaking their seed dormancy and the
removal of competing herbaceous growth (Merou and Papanastasis 2009). In addition, legumes are eec-
tive colonisers of heavily disturbed, nitrogen-decient locales due to their capacity to x nitrogen in the
soil (Lajeunesse et al. 2006). Assuming (as proposed already) that the higher incidence of early Holocene
natural vegetation res impacted the dense grasslands of inner Anatolia and the Zagros, recurrent episodes
of re disturbance might explain the increased presence of legumes at sites in these areas. In the eco-
logical literature, it is often assumed that re disturbances always favour the regeneration and spread of
cereal progenitor species, due to the relative protection aorded to grains by their spikelets (that facilitate
post-re soil crack penetration) and the removal of competing perennial grass growth (e.g. Naveh 1974;
Noy-Meir 2001). However, burning of grasslands early in the summer season might have also led to crop
progenitor stand collapse due to high rates of seed mortality especially for barley but also for rye, emmer
and einkorn. A partial exception would have been stands growing near rocky outcrops or on deep alluvial
and terra rossa cracked soils that could have allowed rapid seed burial (Blumler 1992; Hillman 1996:
191). It is thus plausible that crop progenitor populations were seasonally impacted, on occasion even
decimated, by natural res. Being closely linked to the pronounced seasonal variations in temperature
and rainfall, cyclic pulses of re-induced stand collapse likely exerted signicant (if localised) impacts on
Asouti Human Palaeoecology
Fig. 8 Scatterplots comparing seed size measurements of (A) Barley grain from Jerf el Ahmar (JEA), Dja’dé
and Chalcolithic Kosak Shamali (KS); (B) Emmer wheat grain from Çayönü, Aswad II and KS.
Plot A.A shows a clear separation between smaller and larger barley grain sizes from JEA (early) and KS; Plot
A.B shows an increase in grain size between JEA (early) and JEA (late); Plot A.C shows larger on average yet
more dispersed values from Dja’dé, which indicate a greater overlap with JEA (late) and KS. Plot A.D shows the
step-like increase (arrow) in average barley grain size between JEA (early) and JEA (late)/Dja’dé/KS (the further
increase indicated by the KS mean barley grain size measurements is not signicant as it most likely reects
dierences in crop processing methods between early PPN PDC sites and late prehistoric farming sites: in the
latter larger seed sizes appear dominant because crop processing is more systematic (including large-scale
threshing) while coarse sieving (which removes most of the smaller seeds of cereal ears) also takes place o-site.
Plots B.A-B show a similar step-like increase (arrow) from early to late sites. In plot B.A, the greater dispersal of
emmer grain measurements (compared to barley measurements) can be again explained by the preference of
later farming sites for the storage of glume wheats as whole ears (thus resulting in a greater range of preserved
seed sizes). The additional increase in ventral breadth at later sites indicated in plot B.B is likely to reect the
impact of other parameters unrelated to cultivation practices per se (e.g., cultivation of dierent wheat varieties)
(modied after Willcox 2004: Figs. 3-4).
Human Palaeoecology Asouti
resource availability and predictability, by altering the composition and density of grassland patches and
the distribution of preferred grass species near habitation sites, as well as aecting herbivore behaviour.
It would appear therefore that for large parts of inland Southwest Asia the assumption of the existence of
ecologically stable, resource-rich terrestrial environments that could have supported year-round exploita-
tion of spatially delimited territories is unlikely to hold.
The identication of predomestication cultivation “weed” oras presents its own range of ana-
lytical and interpretative challenges. Willcox (2012b) has argued in favour of a taxonomic approach,
identifying as suitable candidates for inclusion in “weed” assemblages taxa which: (a) have no histori-
cally or ethnographically known uses for their seeds, (b) co-occur in archaeobotanical assemblages with
phenotypically wild progenitor species, and (c) belong to the same genus as veried obligatory weeds
of cultivation known from later agricultural assemblages. Based on these criteria, he has proposed a list
of 19 taxa as the most likely candidates for arable “weed” status (Willcox 2012b: Table 2). An important
limitation of the taxonomic approach is that, more often than not, accurate species-level identications
of the carbonized seed remains of wild/“weed” taxa are not feasible. Furthermore, several species be-
longing to these genera are known to occur naturally in grass steppe and woodland vegetation habitats
across Southwest Asia, although Willcox et al. (2008: 322) have argued that outside arable habitats
such taxa normally occur in low frequencies. However, this argument has been contradicted by Gordon
Hillman who noted that, in the absence of very heavy grazing by sheep and goats, several species can
also be found in uncultivated steppe “at densities comparable to those of weed-infested arable elds”
especially after particularly wet winter seasons (Hillman et al. 1989: 253-254). It is therefore possi-
ble that (under the higher winter precipitation regime that characterized the rst two millennia of the
Holocene) wild/“weed” taxa had far wider distributions and ecologies, and occurred in much higher
densities in natural grassland vegetation compared to later periods or present-day conditions. One very
important implication of this observation is that the proportions of non-progenitor taxa in archaeo-
botanical assemblages derived from short-lived habitation phases and/or sites are unlikely to represent
reliable vegetation ngerprints of PDC. As Willcox (2012a) has observed, assigning “weed” status to
non-progenitor taxa should be dependent on the specic characteristics of each archaeobotanical assem-
blage in its entirety (including close monitoring of shifts in their relative proportions through time). For
Fig. 9 CO2 values
reconstructed from leaf stomatal
index data (SW Sweden)
showing rapid increase from
210-215 ppmv at the beginning
of the Younger Dryas to 270-
290 ppmv at the start of the
Holocene, and the sharp drop
to 240-250 ppmv during the
Pre-Boreal Oscillation (PBO;
~11,400-11,270 cal BP) (plot A)
and how they compare to the
CO2 values obtained from
Antarctica ice-core records
(plot B) (modied after Rundgren
and Björck 2003: Fig. 5).
Asouti Human Palaeoecology
this reason, it is probably best applied only to those sites that preserve long habitation sequences that
have been systematically and comprehensively sampled for archaeobotanical remains.
An explicitly ecological approach has been proposed by Colledge (1998, 2001) classifying
wild/“weed” taxa according to modern ecological groupings and monitoring their presence in archaeobo-
tanical assemblages via multivariate statistical techniques. The key assumption is that in archaeobotanical
assemblages which are dominated by crop progenitor species the primary pathway for the inclusion of
wild/”weed” taxa would have been as “contaminants” of cereal harvests. Their ecological groupings are
thus likely to reect the vegetation composition of the habitats in which cereals were growing. Depending
on the nature of these vegetation ngerprints, it might be possible to reconstruct the specic activities that
were associated with PDC (e.g., the regular occurrence of soil disturbance indicators would point to tillage,
etc.) This approach has allowed Colledge (2001) to infer the practice of PDC on naturally fertile alluvial
soils. This proposition has found additional empirical support in recent studies of wild cereal progenitor
functional ecology, which have indicated that wild cereals may eectively exploit sites characterized by
high levels of fertility and disturbance (Cunni et al. 2014). An obvious limitation of this approach is that
its applicability is limited to archaeobotanical assemblages that are dominated by crop progenitor species:
as discussed already this is a condition that is not universally applicable in early PPN Southwest Asia.
The last criterion, the displacement of crop progenitor species from their natural habitats and distri-
butions, is probably the most dicult to evaluate with any degree of certainty based on modern and histor-
ical observations and ecological analogues. Modern climate-vegetation associations are of little utility for
reconstructing prehistoric plant habitats other than assessing the general relationship between present-day
temperature and precipitation gradients and species distributions. This is due to the enormous dierences
observed between present-day and early Holocene climate conditions in all bioclimatic regions of South-
west Asia. Based on modern observations of oristic associations Zohary (1969) had previously suggested
that deciduous oak parklands represent a key primary habitat for cereal and legume crop progenitor spe-
cies. In recent years, however, integrated archaeobotanical and anthracological studies have demonstrated
that the early PPN ranges of progenitor and non-progenitor taxa extended well beyond those reconstructed
for deciduous oak woodlands into the sparsely wooded Pistacia-Rosaceae semi-arid steppe grasslands that
occupied the inland plains and low- to mid-altitude slopes of the Levant, Anatolia and the Zagros foothills
(Asouti and Kabukcu 2014; Asouti et al. 2015; Riehl et al. 2015). Asouti and Kabukcu (2014) have argued
that (outside the Mediterranean Woodland Zone and Euro-Siberian montane refugia) relic associations of
deciduous oak woodlands with grasses on high-altitude slopes and rocky outcrops represent a mid- to late
Holocene phenomenon caused by the retreat of annual grasses from lowland plains, moist steppe habitats
and mid- to low-altitude slopes due to millennia of overgrazing and settlement expansion.
Modern and historical associations of progenitor taxa with specic soil types and edaphic condi-
tions are no less problematic. Wild cereals are reported to thrive on terra rossa, hard limestone and ba-
saltic soils, as well as rocky outcrops (Harlan and Zohary 1966; Willcox 2005). However, other studies
report much wider edaphic tolerances (including more alkaline soils) for several progenitor taxa both in
Anatolia (Karagöz et al. 2009) and in the Levant (Nevo et al. 1992). At present, such associations persist
in areas that are less accessible to livestock; they are thus likely to represent the aggregated result of the
retreat of wild cereals from other habitat types due to persistent overgrazing (Zohary and Brick 1961;
Noy-Meir et al. 1989; Noy-Meir 1990; Valkoun et al. 1998; Waines 1998; Karagöz et al. 2009). His-
torically heavily impacted habitats otherwise capable of sustaining dense concentrations of cereal pro-
genitor species include alluvial plains and moist steppe biomes (Harlan and Zohary 1966; Kimber and
Feldman 1987). Both habitat types have been used as arable and pasture for thousands of years across
all regions of Southwest Asia. The most favourable habitats for wild cereal growth are characterized
by the co-occurrence of several ecological factors besides soil types or slope aspect, including highest
available soil moisture both at the beginning and at the end of the growth season, least competition for
light, highest available soil nitrogen content, and lowest grazing pressures (Noy-Meir et al. 1991a,b).
How “Intensive” or “Reliable” was PDC as a Staple Subsistence Provider?
As noted in the previous section, an empirical assessment of PDC based on the rst three criteria (i.e.,
excluding modern ecological preferences and distributions) is feasible only for sites that preserve long
habitation sequences and have been thoroughly sampled for archaeobotanical remains including mul-
tiple lines of evidence. One such site is Jerf el Ahmar where larger seed size emerged in its later phases
alongside a reduction in the frequencies of small-seeded taxa (Willcox et al. 2008). However, it remains
unclear whether this represents the intensication of crop progenitor cultivation, since a coeval reduc-
Human Palaeoecology Asouti
tion in the frequencies of some progenitor species (einkorn and rye) is also evidenced in the published
dataset. As discussed already, the 1-step increase in seed size observed between the early and the late
phases of Jerf el Ahmar correlates well with the PBO. Its co-occurrence with the shifts in botanical as-
semblage composition may thus not necessarily reect the gradual intensication of crop production by a
year-round settled community of cultivators-foragers. Alternatively, it might signify the reorganization of
plant food procurement and production involving several complementary strategies such as the broadcast
re-seeding of locally available wild barley stands, or the transplantation of both barley and locally scarce
einkorn and rye into shifting alluvial plots of cleared riparian woodland vegetation during the PBO.
Another site that has preserved a much longer habitation sequence is Chogha Golan, on the Ira-
nian southern Zagros foothills, dating from the end of the Younger Dryas to the 8th millennium cal BC
(Riehl et al. 2012, 2013, 2015). To date, the published evidence does not indicate clear directional tem-
poral trends in botanical assemblage composition, grain size or the development of wild/“weed” oras.
All three key variables (proportions of dominant taxa, barley grain size and the presence and relative
frequencies of wild/“weed” taxa) uctuate widely through time, while there appears to be no consistent
pattern of co-variation between them (cf. Riehl et al. 2015: Figs. 5-7). Despite the evidence for the per-
sistent management of wild-type barley since the earliest phases of the site, the rst denitive evidence
for the presence of domesticated-type barley rachises dates from the MPPNB, thereafter to disappear,
while domesticated-type emmer emerges suddenly in the LPPNB (Riehl et al. 2015; see also Fig. 10). In
the view of the present author, this diversity of archaeobotanical signatures at Chogha Golan is sugges-
tive of diverse and of highly variable intensity plant management strategies through time. Such exibil-
ity might have developed in response to pronounced micro-ecological variation (e.g., acute spatial and
temporal uctuations in the availability and distribution of alluvial habitats alongside recurrent episodes
of seasonal vegetation res and disturbance) that likely aected the availability and productivity of crop
Despite their long habitation sequences neither Jerf el Ahmar nor Chogha Golan (or for that matter
any other early PPN site or regional sites cluster in Southwest Asia) has produced convincing evidence
for the local independent development of phenotypically domesticated crop taxa. This archaeologically
veried fact places PDC at odds with CNC’s prediction that “multigenerational ecosystem engineering”
practised by permanently settled groups managing small, resource-rich and ecologically stable territo-
ries would have provided a sucient condition for the emergence of initial plant domestication. To date,
attempts by archaeobotanists to explain the apparent longevity of PDC have focused mostly on a nar-
Fig. 10 Stratigraphic prole from Chogha Golan (left) alongside AMS dates (cal BP) (loci of dated samples in
the prole indicated by blue circles) and archaeological horizons (AH) in Roman numbers. % frequencies of
relevant taxa and groups of taxa (Aegilops sp., Triticum-type taxa, Hordeum spontaneum and “arable weeds”)
were calculated from the total number of identications from each AH (loci of samples in the prole indicated
by small red dots). n=no. of seed and cha records from each horizon. Large yellow dot to the left marks the
sole occurrence of domesticated-type barley cha (which disappears in later AHs). Large red dot marks the rst
appearance of domesticated emmer wheat. To the right are shown select curves from the pollen sequence and
the oxygen stable isotope record of Lake Zeribar (modied after Riehl et al. 2015: Fig. 10).
Asouti Human Palaeoecology
row range of biological and cultural selection pressures (e.g., introgression from wild populations and
harvesting at the dough stage or by beating cereal ears into baskets) (cf. Fuller and Allaby 2009; Fuller
et al. 2011 and references therein). Few have questioned the dominant perception of PDC, at least in the
regional archaeobotanical literature, as a mode of production that was practiced by permanently settled
“village”-like communities in a manner and at a scale that were conceptually similar (if not functionally
identical) to those of later full-time farming societies (see discussion in Asouti and Fuller 2013). The
main criticism of the currently dominant PDC concept is this: if crop progenitor cultivars (cereals in
particular) were intensively managed through annual planting in plots distributed near permanent habi-
tation sites in order to secure and maximise the year-round provision of staple plant foods, then the pace
of the development of the domestication syndrome would have been much faster. That this was not the
case suggests at the very least the existence of diverse, low-intensity plant management practices that
alleviated, and on occasion even reversed, any latent domesticatory pressures (Asouti and Fuller 2013).
Here this argument is developed further by proposing that PDC practices were attuned to the short- to
medium-term ecological instability that characterized much of the terrestrial environments of Southwest
Asia during the early Holocene. Ecological instability arose from the marked seasonality of the early
Holocene climate, its decadal-centennial scale oscillations and the resulting fragility of the regional
grassland biomes. These phenomena were particularly pronounced in the semi-arid continental regions
of inland Southwest Asia. Steppe grasslands, although extensive and also characterized by high species
diversity due to the rapid climatic improvement that marked the start of the Holocene, were susceptible
to climate-paced cyclic re disturbances which led to periodic stand collapse and short-term depletion
pulses, caused by heightened climate seasonality.
Early PPN communities responded to short- to medium term ecological instability by engaging in
exible economic strategies that precluded substantial reliance on delayed-return practices such as seed
crop cultivation. Their landscape practices likely included the residential and/or logistical mobility of
dierent community segments, the management of spatially extensive and ecologically diverse territo-
ries, and sustained social and material investment in the maintenance of long-range community interac-
tion networks (Asouti 2013; Asouti and Fuller 2013; see also next section). Far from being black-boxed
by archaeobotanists as an evolutionary precursor of xed-plot intensive horticulture, PDC can be per-
haps more accurately conceptualized as a constellation of diverse plant management practices including
the harvesting at varying seasonal intensities of wild plant stands that were dispersed across wide terri-
tories, the opportunistic cultivation of plots that were scattered between the most fertile localities (e.g. in
riparian habitats), transient habitat modications (e.g., shifting plots alongside opportunistic small-scale
clearance and tillage), communal grain storage, and translocational seeding (i.e., the exchange and/
or transference of seed corn over long distances). The common denominator of such practices is that
they are all likely to have generated low-intensity ecological and biological (phenotypic) footprints.
More generally, it appears reasonable to infer that in early Holocene Southwest Asia PDC represented a
somewhat dierent mode and scale of low-level food production and hunter-gatherer niche construction
from those predicted by CNC models that were originally developed in the context of the Eastern North
America and Neotropical ecoregions of the New World (cf. Smith 2006, 2012, 2015).
Resilience, Environmental Knowledge Transmission and Community Interactions
As noted in the introduction, the early PPN witnessed a orescence of symbolic/ritual behaviours across
Southwest Asia. While a detailed discussion of the contextual attributes and potential meanings of early
PPN symbolism goes well beyond the scope of this chapter, it is noteworthy that much of its material
manifestations encountered in some of the most celebrated case studies (e.g. at Göbekli Tepe) have dis-
tinctive, if less commented upon, landscape connotations. One characteristic example is “Enclosure” D
at Göbekli Tepe, one of the earliest excavated structures at the site, currently dated between the mid-10th
and the early 9th millennia cal BC (Dietrich et al. 2013). The T-shaped pillars of “Enclosure” D bear
the highest proportion of sculpted snake depictions and the highest diversity of animal representations
from any other excavated structure at Göbekli (Peters and Schmidt 2004: Table 2). Venomous creatures
(scorpions and snakes, the latter strongly reminiscent in shape of the native to the region Vipera lebetina)
are depicted as moving away from wetland birds (pillar 33), while attacking mammals (pillars 20, 33)
or in association with death-related themes (pillar 43) (Fig. 11). Representations on pillar 43 comprise a
narrative of potential cosmological signicance organized in three distinct horizontally arranged panels:
the top panel contains what appear to be habitation structures in a wetland setting, suggested by the pres-
ence of reed-like patterns, a wetland bird and a boar. In the panel below two birds of prey are depicted
Human Palaeoecology Asouti
alongside a sun-like disk and other symbols with less obvious connotations. The panel at the bottom end
of the pillar is dominated by a massive scorpion, which is depicted in ne morphological detail alongside
a snake, possibly a scavenging animal, and a headless male human body that appears to be led away by
another bird of prey.
According to recently published radiocarbon determinations (Dietrich et al. 2013) the begin-
ning of “Enclosure” D is dated to ~9700 cal BC, at the very end of the Younger Dryas and the start
of the Holocene as indicated by the more precisely dated Nar Gölü palaeoclimatic sequence (Dean
2014). The palaeoecological evidence for the magnitude and rapidity of the transition (completed in
the space of a single decade) suggests that its environmental impacts were experienced within indi-
vidual human lifetimes. Memories of the hyper-arid environments of the Younger Dryas (possibly
encoded in death-related themes and stories about swarms of venomous snakes attacking humans
and mammals at a time of increased aridity) that had hitherto shaped people’s landscape experienc-
es likely formed the core of cosmologies and inter-generational knowledge transmission about past
times of adverse conditions and resource stress. Comparative studies of ethnographic accounts of
hunter-gatherer societies living in marginal (arctic, subarctic, arid and semi-arid) environments have
indicated that oral traditions of high-impact, low-frequency episodes of environmental change have
an average lifespan of ~90 years (Smith 1988). After a century has elapsed, such stories become en-
coded in cosmological narratives that are passed down the generations through their incorporation
into highly stressful, even traumatic, once-in-a-lifetime ritual events (e.g. male initiation rites) expe-
rienced collectively by their participants in large aggregation sites. According to Smith (1988) such
events function as vehicles for the storage and inter-generational transmission of collective passive
memory (as opposed to active memory deployed in expedient, daily life tasks) and for establishing
and re-arming lifelong alliances and bonds between participant individuals and social groups.
Snakes, centipedes and birds of prey are depicted on portable objects found in several early PPN
sites of the northern Fertile Crescent (Figs. 12 a, b). At Jerf el Ahmar they occur as engravings on shaft
straighteners (a category of ground stone artefacts traditionally associated with hunting activities) or oth-
erwise unmodied ground stone objects (Stordeur and Abbès 2002). At other sites, such as Körtik Tepe,
representations of snakes and centipedes also appear on stone vessels that might have been used in com-
munal food consumption events (Özkaya and San 2007). Snakes, giant centipedes, scorpions, lizards and
spiders are common in the faunas of semi-arid steppe grasslands and were abundant in the region before
the decimation of their natural habitats by overgrazing, settlement expansion and modern agriculture (cf.
Joger 1984; Crucitti and Cicuzza 2001; Kaltsas et al. 2008; Simaiakis and Mylonas 2008 and references
Fig. 11 T-shaped pillars from “Enclosure” D at Göbekli Tepe (images courtesy of the late Klaus Schmidt).
Asouti Human Palaeoecology
therein).3 Their ubiquity on early PPN portable material culture (especially objects associated with hunt-
er-gatherer mobility and social interactions) might signify the mapping of paths of movement across the
liminal space of the steppe and/or stories of the various dangerous encounters associated with such trips.
The engravings on some ground stone objects are also suggestive of their potential function as mapping/
orientation devices: they often combine snakes, birds of prey and hunted mammals (possible allegories
for the steppe, its dangers and its resources) with xed landmarks such as the round shapes and features
3 A particularly poignant description of the experience of travelling across the steppe grasslands of northern Syria by foot can
be found in the novel “Ariagni” by the Egyptian-Greek author Stratis Tsirkas, where he describes the forced march from Aleppo
to Ar-Raqqah of two battalions of the 2nd Brigade of the Greek Armed Forces in the Middle East in June 1943, following the
mutiny of their republican ocers in April 1943: “Raqqa; the steppe was like a grey yellow sea; full of wild grasses as tall as
reeds, two meters; full of lizards like little crocodiles, poisonous snakes, giant centipedes, scorpions as big as little mice, hairy
spiders; the pain is intolerable and their wounds stink.ˮ (Tsirkas 1983 : 336-337)
Fig. 12a Engraved stone objects: Tell Qaramel shaft-straighteners (1, 4; Mazurowski and Yartah 2002: Fig. 10);
Jerf el Ahmar shaft-straighteners (2, 3, 6, 7; Stordeur and Abbès 2002: Fig. 16); Tell ‘Abr 3 stone plaquette (5;
Yartah 2005: Fig. 7); Körtik Tepe stone vessel (8; Özkaya and San 2007: Fig. 18).
Human Palaeoecology Asouti
resembling standing posts reminiscent of the communal buildings at Jerf el Ahmar, potential representa-
tions of wild plant stands and more abstract motifs (e.g. arrow-like shapes) that might signify orientation
markers (see Fig. 12a: 3, 5-7, Fig. 12b).
By placing these examples of early PPN symbolism and ritual behaviours in their broader
ecological and socioeconomic context, it is possible to begin putting together a larger picture. Hunt-
er-gatherer mobility, material culture symbolism deployed in active and passive landscape memory
storage and knowledge transmission, and community interaction networks sustained by ritually in-
vested communal food consumption events, stood at the core of regional resilience strategies aimed
at mitigating early Holocene ecological instability and associated resource risks. Recently published
plant genetic research has brought out even more poignantly the critical contribution of hunter-gather-
er mobility and community interactions to the macro-evolutionary development of the domestication
syndrome in Southwest Asian early cereal cultivars. Genetic studies suggest that domesticated emmer
wheat has a reticulate rather than a phylogenetic evolutionary relationship with its wild progenitors
(Civáň et al. 2013). Civáň et al. (2013) have proposed that this resulted from hybridization between
dierent lineages eected via the utilisation and cross-pollination of wild grain derived from diverse
sources over long periods of time. This process was ultimately responsible for the development of
predomesticated cultivars that shared phylogenetic signals with emmer populations derived from all
parts of the wild emmer geographical range. Thus, according to the reticulated origins scenario, wild-
type emmer cultivars spread during the early PPN from the southern Levant into northern Syria,
southeast Anatolia and northern Iraq, where their reproductive isolation from parent wild emmer
populations resulted in their morphological domestication. This scenario would explain the phyloge-
netic proximity to the domesticated gene pool evidenced by the wild emmer populations presently
found in Karacadağ (southeast Anatolia) and Sulaymaniyah (northern Iraq), which appear to repre-
sent “the remnants of the cultivated populations from which the rst domesticates evolved” (Civáň
et al. 2013: 9). If veried through further research, these observations may signal the replacement
of both monophyletic and polycentric theories of agricultural origins with a new dynamic reticulate
model: Epipalaeolithic and early PPN hunter-gatherer mobility was the primary determinant of the
geographic distribution and genetic makeup of the wild cereal progenitor species before their initial
domestication. A model of reticulate (vs. polycentric and monophyletic) origins of initial crop do-
mestication also points to the existence of much more dynamic and multifaceted modes of early PPN
niche construction across Southwest Asia (predicated on regional interaction networks and paths of
movement) compared to CNC theory predicting the existence of spatially limited, closely controlled
and temporally stable resource territories.
Fig. 12b Engraved stone objects: Tell Qaramel (1; Mazurowski and Yartah 2002: Fig. 11); Tell ‘Abr 3 (2; Yartah
2013: Fig. 151).
Asouti Human Palaeoecology
Pathways to Agriculture: the Switch from PDC to Agro-pastoral Food Production and its
Socioeconomic and Ecological Impacts
For a delayed-return subsistence strategy such as seed crop cultivation to become established in the
acutely seasonal climatic regimes and unstable vegetation ecologies of early Holocene Southwest
Asia, and supersede long-lived resource management strategies rmly imbedded in social memory,
identities and community interactions, a major innovation in its technology was necessary; one that
would render it viable as a staple subsistence provider. Ethnoarchaeological research on traditional
farming practices in the Eastern Mediterranean has demonstrated that a key condition for the sus-
tainability of crop production is the capacity of cultivation systems to absorb and buer recurrent
seasonal and interannual environmental risks of crop failure. This is achieved primarily through the
intensication of production geared at generating a normal surplus to use in times of need as well as
underwriting social obligations (see discussion in Halstead 1989, 2014). In the context of Southwest
Asia, Abbo et al. (2010) have proposed that a key strategy for mitigating environmental risks was the
cultivation of crop packages (mixtures of cereals and pulses exhibiting variable levels of tolerance to
local micro-ecologies) which would have conferred agronomic stability to early cultivation systems.
The regional archaeobotanical record indicates that the formation of regionally distinctive crop pack-
ages comprising domesticated cultivars was a protracted process that was not complete until the late
PPN (Asouti and Fuller 2012). Moreover, the geographical and chronological pattern of the adoption
and spread of domesticated crop packages follows closely that of the spread of domesticated caprine
herding (Colledge et al. 2004; Peters et al. 2005; Zeder 2008; Asouti and Fuller 2012; Arbuckle and
Atici 2013; Martin and Edwards 2013). This correlation indicates that it was the integration of plant
cultivation with the herding of domesticated animals that likely played a pivotal role in the establish-
ment and spread of farming economies across Southwest Asia (Harris 2002).
A key pathway for this development was the use of animal dung as manure that might have
initially occurred as the unintentional consequence of caprine herds grazing on steppe and alluvial
grasslands following plant harvests. Manuring increased the ecological resilience of crop cultivation
by mitigating some of the risks imposed by early Holocene climate seasonality. The use of dung as
fertiliser provided a major boost to the productivity and reliability of cultivated harvests thus enabling
the intensive annual cropping of xed-boundaries plots in direct proximity to habitation sites (cf.
Harris 2002; Bogaard 2005; Araus et al. 2014). The widespread adoption of small-scale horticulture
integrated with domesticated caprine herding also had lasting eects on the vegetation environments
of Southwest Asia through the development of regionally distinct agroecologies and other types of an-
thropogenic niches (Asouti and Kabukcu 2014; Asouti et al. 2015). Examples of the latter include the
gradual reduction of grasslands and the coeval expansion of semi-arid managed woodlands and wood-
land pastures. Such landscape-scale vegetation changes have been detected in both central Anatolia
and the arid zone of the southern Levant, and resulted from the combined impacts of domesticated
caprine grazing and woodland management activities alongside increasing climatic aridity, especially
from the mid-late 8th millennium cal BC (Asouti and Kabukcu 2014; Asouti et al. 2015).
The societal impacts of the adoption and spread of integrated agropastoral production during
the late PPN were equally far reaching. Communities became perceptibly more settled, occupying
larger habitation sites on a permanent basis and over successive generations (Kuijt 2000; Asouti
2006a). Residential architecture was transformed with the standardization and increasing compart-
mentalisation of building layouts, in order to accommodate a range of functions including storage and
cooking, and heating and food preparation installations (Kuijt 2012; Goring-Morris and Belfer-Cohen
2013). Overall, it is possible to observe a shift away from group-focused social structures and corpo-
rate identities towards the household, which emerges as the principal unit of socioeconomic organ-
ization in the course of the later PPN (Kroot 2014). Evidence for the prevalence of an “egalitarian”
social ethos that might have functioned as a levelling mechanism for emergent social inequalities is
ubiquitous in the late PPN, being principally manifested in the lack of dierentiation in residential
architecture and in burial customs (cf. Kuijt and Goring-Morris 2002; Asouti 2006b; Kuijt 2008; Kuijt
et al. 2011). At the same time, however, the use of communal structures for storage and other social
purposes waned, whereas after the late 9th millennium cal BC most indicators of collective ritual be-
haviours and communal food consumption largely disappear from the regional archaeological record.
Across Southwest Asia late PPN societies were characterized by settlement patterns, mobility strate-
gies, economic practices, and ritual and symbolic behaviours that overall had little in common with
their early PPN antecedents.
Human Palaeoecology Asouti
Climate change at the end of the Pleistocene and the beginning of the Holocene played a pivotal role
in the radical reconguration of the “ecological theatre” in which the “evolutionary play”4 of global
agricultural origins unfolded in the course of the last 12,000 years. The combined archaeological and
palaeoecological records of late Pleistocene and early Holocene Southwest Asia appear to corroborate
the hypothesis that direction-changing, transformative economic shifts and associated technological and
societal innovations are not engendered by conditions of stress (resource, climatic, demographic, or
otherwise). Instead, they seem to uphold the theoretical principle that human societies are primarily
risk-averse, especially when faced with stochastic and/or regime-switching environments (Zhang et al.
2014). When their economic base is challenged by external or internal pressures humans will opt for
behaviours that are focused on security rather than optimization. At the other end of the spectrum, how-
ever, resource abundance per se also does not seem to provide a sucient condition for direction-chang-
ing socioeconomic change. The reason for this is that change is predicated on complex historical cir-
cumstances and contingencies arising from the interplay of varied multi-scalar ecological, social and
historical contexts. In early Holocene Southwest Asia the macroevolution of the regional agricultural
economies ultimately depended on radical innovations in the ecology and technology of early food
production, such as the integration of cultivation with domesticated caprine herding, that emerged quite
separately (more like historical accidents than intended outcomes) from any supposed incremental shifts
in the management, genetic makeup and phenotypes of early PPN cultivars.
The symbolic and ritual orescence that has come to dene the early PPN of Southwest Asia
has been widely portrayed in the literature as intimately linked to the cognitive and sociocultural shifts
deduced for this period (e.g., Cauvin 2000; Verhoeven 2004; Watkins 2006; Hodder and Meskell 2011;
Atakuman 2014). By contrast, the potential contributions of climate change and the environment at large
are often downplayed, or sometimes altogether omitted, from a debate that remains squarely focused
on architecture and material culture. Environmental change is viewed as providing little more than the
ecological background to the main story of sociocultural change and innovation. Contrasting with such
approaches, this chapter argues that the resource-rich, yet ecologically unstable, landscapes of the rst
two millennia of the Holocene likely contributed to the emergence of complex symbolic and ritual be-
haviours in Southwest Asia, which were imbedded in regionally distinctive social identities and corpo-
rate institutions. Such behaviours were deployed in mapping the availability and locations of preferred
resources, and for the storage, exchange and inter-generational transmission of landscape knowledge and
historical experiences of environmental change. Raised resource ceilings in the context of early Holocene
ecological instability also expedited the development of lasting mechanisms of social cooperation, which
fostered the intensication of community interaction networks and the region-wide circulation of socially
prized foodstus, including early cultivars. Increasing degrees of social cooperation are reected in the
ubiquity of ritually invested communal institutions and in behaviours such as communal food consump-
tion that enabled alliance building, the development of shared ideologies and cosmologies, the diusion
of symbolic vocabularies, and the emergence of regionally distinctive group identities.
Cooperation is a major driver of human evolution (Nowak 2006). The critical role of environmen-
tal instability is also widely recognized in evolutionary biology. Building on the dichotomy between var-
iability-selection versus habitat-specic hypotheses (Potts 1998, 2002) the general theoretical principle
is proposed that specialized economic behaviours (such as “agriculture” or “foraging”) are facilitated
by environmental change in a constant direction (whether negative or positive) that renders such behav-
iours adaptive and sustainable in the long-term (e.g., during the Younger Dryas or the Holocene Climatic
Optimum). Conversely, in environments characterized by high degrees of medium- to short-term eco-
logical instability (e.g., the Younger Dryas termination and the onset of the Holocene) which impacts
directly human perceptions of the environment and its resources, exible economic practices and co-
operative social behaviours provide distinct advantages. What dierentiates this proposition from other
ecologically oriented theories of prehistoric economic behaviours (cf. Binford 2001) is its decoupling of
resource abundance and resource stress from idealized notions of environmental stability and instability
respectively. Instead, the emphasis is placed on resource ecologies. For the comparative cross-cultural
study of agricultural origins such a theoretical framework provides an appropriate heuristic environ-
ment for formulating and testing region- and context-specic, historical explanations of the enormous
4 Slightly paraphrasing the title of Evelyn Hutchinson’s famous collection of essays “The Ecological Theatre and the Evolu-
tionary Playˮ rst published in 1965 by Yale University Press.
Asouti Human Palaeoecology
dierences observed in the nature, duration, pace and directionality of socioeconomic change among
early Holocene low-level food producers worldwide (cf. Smith 2001; Barker 2006). This is achieved by
focusing on the distinctive resource ecologies characterizing each world region, in the context of global
climate change, rather than the broad structural similarities of prehistoric economic behaviours.
I wish to express my gratitude to Melinda Zeder, Ceren Kabukcu, Sue Colledge and an anonymous re-
viewer for providing constructive comments and suggestions on earlier drafts of this chapter.
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